Tetrapod Zoology - SoundSage

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The enormous two-day event that was DinoCon 2026 – the second of our dinosaur-focused comic con-style meetings – is over, and wow. It went so well, the attendance being around twice that of our first event of last year.

**Caption:** a DinoCon 2026 montage, showing our banner, a packed hall during a talk, and vendor stall (Jed Taylor’s) during set-up on Friday. Images: Darren Naish.

DinoCon 2026 was held at the Hilton Birmingham Metropole over the weekend of July 25th and 26th and mostly occurred in two giant halls and an additional four or so smaller rooms, all located within a minute’s walk of each other. The venue did us well and the fact that many (albeit not all) attendees were staying on site meant that we had an excellent social atmosphere, people staying late in the hotel’s bars and restaurants. Remember that you are helping us run the event by being in, and spending money in, the hotel.

As was the tradition with TetZooCon before it (which ran from 2014 to 2024), DinoCon (which I co-run with Nathan and Annie Barling and a team of helpers) is a fusion of both popular and technical content on dinosaurs and other animals with palaeoart-themed events, a quiz, exhibitions and more. As per last year, we were joined by the Dinomania team and thus a number of life-sized juvenile ankylosaurs, a small Triceratops, and a few large theropods were wandering about the venue. Jurassic Park-themed jeeps and other vehicles were parked out front.

**Caption:** one of the several *Jurassic Park* vehicles we had parked at the front of the venue. Numerous people had fun getting their photos taken alongside. Image: Coco Ellenbogen.

Vending and vendors. What started as two tables of books and plush toys back in 2014 has expanded to a gigantic and slightly terrifying vendor hall, this year housing around 70 vendors, vaiously selling books, fossils, toys, models, sculptures, apparel, art and more. The quality and quantity was astounding and we know that many vendors did extremely well. I was moderately sensible in what I purchased and part of me wishes I was more reckless. Maybe next year.

**Caption:** stalls at DinoCon 2026. At left, *Carnotaurus* skull model at the **EXEtinct** stall. It had sold by the end of the event. At right, books on sale at the **Warwick Books** stall. Photos by Alfred Barwick.

No event is ever perfect and let me say upfront that we learnt a lot abut managing and running meetings of this size. Queuing and room size was an issue for some events and we did have a few problems with our AV setup. Rest assured that we’re very aware of these things and are putting measures in place to improve the experience for the future.

Talks and lectures. We’re now at the stage where so much is happening that I can’t hope to see all of it and there are many talks that I never got to see, nor did I even get to properly catch up with the speaker. Some talks – those we imagine to be more popular – were held in the gigantic Palace room while others – those we imagine to be a bit more on the technical side and thus maybe not as well attended – were held in the Library (it’s not actually a library, it’s a room that has a libraryesque feel). But we got things a bit wrong, since some technical talks were so popular that the Library was over-full, with numerous people standing at the back and sides of the room. We live and learn.

Talks on Saturday started with Matt Dempsey’s outstanding presentation on modelling dinosaurs: on how musculature is added, on the possible masses of select dinosaurs, and on what muscularity means for agility and locomotion. Happening in parallel was Sue Judd on ‘It Takes a Team to Build a T. rex’, a discussion on the design, creation and marketing of the new Tyrannosaurus model made by Everything Dinosaur (Sue is one half of the Everything Dinosaur team). Next up, Dean Lomax and Bob Nicholls gave a joint presentation on ‘The Secret History of Dinosaurs’, this given in conjunction with the recent publication of their excellent book of the same name (Lomax 2025). Both did a book signing later in the day and had to deal with a very substantial queue.

**Caption:** Dean Lomax on the DinoCon 2026 main stage, here discussing his own background. Image: Alfred Barwick.

Chelsey Kynff’s talk – ‘A Bone to Pick’ – discussed important women in the history of palaeontology, among them Mignon Talbot, Annie Alexander (of thalattosaur fame), Mary Leakey and others on a tour across history. David Unwin showcased new data gleaned from pterosaurs via the use of UV light while recently minted PhD Vicky Coules reviewed the cinematic events that brought dinosaurs and prehistory in general to the screen during the early 20th century.

**Caption:** Chelsey Kynff on stage, here discussing Mary Anning. Image: Alfred Barwick.

  **Caption:** Vicky Coules on stage at DinoCon 2026, here discussing one of the several 20th century silent movies she presented (and showed). Several were comedic efforts involving prehistoric people. Image: Alfred Barwick.

A real standout performance last year came from filmmaker and artist David James Armsby who not only spoke about the making of his films (in particular SAURIA and Dinosauria) but also released one of his films as an exclusive premiere. So good was this presentation and the audience reception that we opted to host David as our 2026 Guest of Honour, and once again he put on an incredible show. The premiere film this time – Great Maia – is an outstanding piece of work and our audience did us proud in showing their appreciation.

**Image:** David James Armsby thrills the audience with the backstory to his *Great Maia* film, an exclusive at DinoCon 2026. Extended and rapturous applause occurred… more than once. Image: Alfred Barwick.

  **Image:** David James Armsby on stage at DinoCon 2026, here with one of his sketchbooks. Some of the material included therein is obviously relevant to the films we’ve seen, but some is for projects not yet released or abandoned. Image: Alfred Barwick.

Other events. Saturday also featured a live recording of the Terrible Lizards podcast, run by Dave Hone and Iszi Lawrence, another book signing (Dave Hone and Mark Witton on their Spinosaur Tales), and a meet and greet event with David Armsby. Did I mention that David brought one of his sketchbooks along?

**Caption:** Iszi Lawrence and Dave Hone record an episode of the Terrible Lizards podcast in front of a DinoCon 2026 audience. The recording isn’t out (at the time of writing) but the back catalogue of the podcast is **here**. Image: Alfred Barwick.

  **Caption:** Mark Witton (at back) and Dave Hone sign copies of their *Spinosaur Tales* at DinoCon 2026 while people wait patiently in line. The queue extended out the door and down the corridor outside. Image: Darren Naish.

Towards the end of Saturday we hosted a discussion panel whereby Nathan spoke to Emma Nicholls, Maria McNamara, Mathew Wedel and Mike Taylor about ‘Thriving in Palaeontology’: on how they’ve remained involved with the field and how they have, or have not, made careers out of their interests. Mike is a vocational palaeontologist employed in another field and Nathan runs a gaming and publishing company, so there are many ways to be involved in palaeontology even when not employed in academia.

**Caption:** Maria McNamara holds the floor during the panel event, this year devoted to thriving in palaeontology. It’s a notoriously difficult field to get into, with very very few opportunities. Nevertheless, it’s clearly possible. Image: Alfred Barwick.

Costumes and quiz. And then there was the costume competition. Once again, we welcomed people on stage as they displayed costumes, puppetry and other accoutrements, this culminating in judging and prize-giving.

**Caption:** suit-wearers and puppeteers on stage during the DinoCon 2026 costume competition. It’s interesting that theropods (maniraptorans in particular) are, by far, the most popular objects of focus for people building suits. Everyone here won something. Image: Alfred Barwick.

The quiz this year wasn’t wholly focused on Mesozoic dinosaurs as it was last year but instead included sections on Cenozoic mammals and birds, Mesozoic marine reptiles and Palaeozoic animals. Nathan is, I suppose quite rightly, of the opinion that palaeontology involves geology and therefore made the argument that our quiz should include a section on that topic. And so it was that we also had a section on sedimentology, mineralogy and such.

**Caption:** some quiz images. At left, the Age of Dinosaurs placard that accompanied the Sandbox VR tickets we gave out at prizes (we also showed the trailer, which you can watch **here**). At right, Max Stringer of **ReptiMax** discusses his *Hatzegopteryx* model, visible in its box in the middle of the image. Images: Alfred Barwick.

The prizes for the quiz were very special. In attendance this year was Max Stringer from South Africa, aka ReptiMax. Max very kindly donated a boxed version of his spectacular, big Hatzegopteryx model. That was first prize, as was a set of tickets for the Age of Dinosaurs Sandbox VR experience (facilities for which exist in Birmingham, plus in London and elsewhere in the UK). Second and third prizes were some special and sought-after dinosaur models kindly donated by Everything Dinosaur, plus more Age of Dinosaurs tickets for second place. James Appleby won first place, Tim Versteeg was second, Joschua Knüppe third. Thanks to everyone who took part. Knowing how to pitch a quiz for an audience of this sort is never easy given the level of expertise we have in the crowd. If it were even moderately easy, we’d have tens of very high scorers.

**Caption:** the DinoCon 2026 quiz was won by James Appleby, and here he’s holding the ReptiMax *Hatzegopteryx* model. Videos about this model can be found on the **TetZooTowers\_collection TikTok**, incidentally. Image: Alfred Barwick.

The art exhibition. Running at the same time as the quiz was our art exhibition, once again showcasing work by attending artists interested in putting things on display. Some of the work was on sale. The whole thing was spectacular, featuring the work of around 20 world-class artists including established names (like Mark Witton, John Conway and Anthony Hutchings) as well as newcomers. I exhibited the giant Crocodylomorph Empire poster (copies of which were on sale at the Tet Zoo stall) alongside a compilation of images and captions on its backstory and compilation.

**Caption:** at left, my Crocodylomorph Empire poster; it was on sale at the event (a digital version will be on sale **here** soon). At right, part of the DinoCon 2026 art exhibition showing the poster as well as images and notes relevant to its backstory. Images: Darren Naish.

More talks: theropods, biomolecules and sea reptiles. Sunday was packed too, morning talks in the Palace including those by Emma Nicholls on ‘Bookending 200 Years of Megalosaurus’, Maria McNamara on ‘Could Jurassic Park Really Happen?’ and Dave Hone on ‘Spinosaurus – Ecology and Controversy’. All were phenomenally well attended, and I know that the Q&A for Dave’s talk extended something like an hour beyond the talk’s scheduled finish. He was ok with this and we had to force him to stop answering questions.

**Caption:** Emma Nicholls discusses our evolving knowledge of *Megalosaurus*, starting with its scientific recognition in the 1820s. Image: Coco Ellenbogen.

My own talk – ‘Mesozoic Marine Reptiles, the View From the Mid-2020s’ – happened in the Library and concerned exciting recent discoveries made about ichthyosaurs, plesiosaurs and mosasaurs, it serving as a sort of update to my Ancient Sea Reptiles (which is on sale here at the Natural History Museum) (Naish 2022). A tangent on the claimed giant size of certain pliosaurids was made all the more timely thanks to the new paper by Eddy Bartlett and colleagues (Bartlett et al. 2026). Immediately on finishing I ran over to another part of the building for a book signing, not just of Ancient Sea Reptiles but for whatever books people wanted me to scribble in. It went well and I was kept busy for the next two hours, yikes. There is immense pressure to get through a queue, but I managed it. I’m always impressed by how gracious people are about waiting in line for so long.

Workshops. Occurring throughout the weekend were a number of palaeoart workshops, those for beginners being led by Dane Pavitt and those for people with some degree of experience being led by Joschua Knüppe. The latter had to be booked in advance but that wasn’t so for the former, and we substantially underestimated the interest there would be in attending. We apologise to the people who wanted to get in but couldn’t. We’re going to have to implement a ticketing system or some other way of limiting numbers in future.

Auctions. Our Live Charity Auction also happened on Sunday. This year we were raising funds for the Turtle Foundation. I get that an event like DinoCon is a huge exercise in escapism, but I feel strongly that we mustn’t move away from efforts to help preserve the natural world and we’ll continue to host auctions for conservation charities in future years. Harrison Watler of the Turtle Foundation gave a talk on the foundation’s work before the auction began.

**Caption:** Nathan and Darren on stage during the live auction, the lot here being an original Luis Rey painting, kindly donated by Russell Batten. It went for an amount over £300, as is appropriate for original art. Image: Coco Ellenbogen.

Auction items included bespoke and unique dinosaur models, original artwork (including by John Sibbick, Steve White and Luis Rey) and a signed first edition of Katrina van Grouw’s The Unfeathered Bird. A silent auction, involving books, art prints, dinosaur models and more also occurred. Massive thanks to everyone who donated or made bids.

Talks on palaeomedia. On afternoon talks, that by Paul Barrett – on ‘Zimbabwe’s Dinosaur Heritage’ – unfortunately didn’t happen as Paul was unable to make it through no fault of his own. We look forward to seeing him attend in future. If you pay attention to dinosaur-themed news you might have seen the brand-new publication of the sauropodomorph Musango matusadonaensis from Zimbabwe; Paul is one of the describers (Barrett et al. 2026). I wonder if he was planning to mention it, as an exclusive reveal, at DinoCon… I guess we’ll never know.

**Caption:** Tom Fletcher at DinoCon 2026… Tom discussed the making of palaeo-themed TV shows at Silverback Films. At right, promo image for the Netflix series *The Dinosaurs*, first broadcast in March 2026. I enjoyed it a lot more than I did *Life On Our Planet*. Images: Coco Ellenbogen; © Netflix.

Our final two talks focused on big-budget TV series about prehistoric life. First up was Tom Fletcher of Silverback Films who discussed the ups and downs of making, firstly, Life on Our Planet and then The Dinosaurs. It was a brilliantly fun talk that showed great (and exclusive) clips and segments.

Next up was Tim Haines, who absolutely got the most incredible introductory audience reception, his talk being ‘Surviving Earth – Why?’. This discussed the thinking behind his new series and the background to the extinction events it portrayed. By now, it’s no secret that Tim said – right at the end of his talk – things that were interpreted as an endorsement of generative AI, his primary point being that this is now part of the CGI pipeline. The audience did not respond well but this didn’t stop Tim from receiving appropriate applause at the end; if you’ve heard otherwise, it’s not accurate.

**Caption:** Tim Haines on stage at DinoCon 2026. One moment of *Surviving Earth* shows an adult pliosaur thrown ashore and pinned up in the air by some trees. Tim noted that this was very much inspired by Bob Nicholls’s image of a deceased mosasaur posed the same way, and appeared through agreement with Bob himself. Image: Coco Ellenbogen.

We closed by thanking the many people who worked with us in making it all come together – the AV team, the hotel staff, our volunteers and assistants and everyone who attended – before announcing the amount raised for charity by the audience. It was over £3200, a respectable amount that we’re pleased to pass on to the Turtle Foundation.

**Caption:** a fun and ever-present feature of DinoCon is the Dinomania team and their various charges. The two ankylosaurs Bramble and Conker proved really popular. Image: Coco Ellenbogen.

Goodbye DinoCon 2026! Packing up, tidying, and heading out involved a lot of work, but we did it in good time and I finally got to spend appropriate time in the bar on Sunday night. While there were assorted hiccups and scares of the sort that always occur in running these events, and while there are many things that could have been done better, DinoCon 2026 was a massive success and we owe huge thanks to everyone who attended for making it what it was. My co-organisers worked so, so hard in bringing it all together, as did the hotel staff, our volunteers, all of the many vendors, our photographers and videographers, and our speakers and presenters. Thank you all!

**Caption:** Sam Neil was mentioned in our opening ceremony, and we also had these two books of remembrance present throughout. Massive thanks to Vicky Robinson for working with us in setting this up. Image: Darren Naish.

It remains to be said that DinoCon has undergone such growth since the first one that we do expect things to become even bigger and better in future. Yes, we want YOU if you’re interested in vending, in attending, in speaking, or in helping as a volunteer. So, goodbye DinoCon 2026… watch our website and social media presence for news on what might happen in 2027!

**Caption:** goodbye DinoCon 2026! Image: Coco Ellenbogen.

For previous articles on DinoCon and its ancestor – TetZooCon – see these articles….

The events of TetZooCon 2014, July 2014
TetZooCon 2015 Is On, July 2015
The Events of TetZooCon 2015, November 2015
Coming Soon: TetZooCon 2016, September 2016
The Day After TetZooCon, October 2016
The Fourth TetZooCon, September 2017
The TetZooCon of 2017, October 2017
Reasons to Attend TetZooCon 2018, September 2018
TetZooCon 2018: Best TetZooCon So Far, October 2018
Announcing TetZooCon 2019 – the Biggest Yet, August 2019
Final Call For TetZooCon 2019, October 2019
The Sixth TetZooCon, October 2019
TetZooCon 2020 + Zoom = TETZOOMCON 2020, December 2020
TetZooMCon 2020 an Unbridled Success, December 2020
Cronch Cats, Beasts of Gévaudan, Dinosauroids, Mesozoic Art and Much More: TetZooMCon 2021 in Review, September 2021
The Ninth and Largest of the Tetrapod Zoology Conventions, December 2022
Announcing the 10th Tetrapod Zoology Convention, October 2023
At TetZooCon 2023, November 2023
TetZooCon 2023 the report, December 2023
The Last TetZooCon, October 2024
DinoCon 2025 Has Left the Building, August 2026

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

Refs - -

Barrett, P. M., Botha, J., Sciscio, L., Stuart, B. P., Lovegrove, J., Munyikwa, D., Zondo, M., Broderick, T. J., Edwards, S. F., Mbambo, E., Chapelle, K. E. J., Dollman, K. N., Tolan, S. & Choniere, J. N. 2026. A new sauropodomorph dinosaur from the Pebbly Arkose Formation (Upper Triassic: Norian) of Kariba, Zimbabwe. Journal of Systematic Palaeontology 24, 2678610.

Bartlett, E. O., Martill, D. M. & Smith, R. E. 2026. Re-evaluating body size in the Middle Jurassic pliosaur Liopleurodon ferox Sauvage, 1873. Journal of Vertebrate Paleontology e2691149.

Lomax, D. R. 2025. The Secret Lives of Dinosaurs. Columbia University Press, New York.

Naish, D. 2022. Ancient Sea Reptiles. Natural History Museum, London.

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Regular readers of Tet Zoo will – I hope – be familiar with taxonomic vandalism…

Put simply, this is the phenomenon whereby rebellious individuals, typically working outside of institutional academia, seek to formally name or rename the organisms they’re interested in, most usually by way of self-published works that have a decidedly ‘non-standard’ feel and approach. I’ve published on this subject, not just at Tet Zoo (here) but also in both the semi-technical literature (Naish 2025) and as a contributor to technical cases that seek to minimise or negate the acts of specific vandals (Rhodin *et al*. 2015).

Among those organisms most frequently affected by taxonomic vandalism are reptiles, in particular those of Australasia. And you may know already that one of the most active of the various battlefields is that involving one Mr Raymond Hoser, his work building on that of his predecessors, among them Richard Wells and Ross Wellington.

So interesting and frighteningly esoteric are these various characters and their exploits, adventures and battles that this entire subject looks over-ripe as the perfect fodder for a book, and that’s why we’re here: Snake Men by Zach St. George, published by W. W. Norton & Co, is out now and on the shelves, and I’m in the fortunate position of owning an advance copy.

Snake Men is fun and fast-paced, focusing mostly on Hoser and what it’s like to spend one-to-one time with him as he speeds about the Melbourne suburbs in his Nissan Micra, at times applauding pedestrians “for their fitness” and leaning “out the window to woof at dogs” (St. George 2026, p. 141). St. George notes that he wasn’t able to spend as much time with Hoser as he might have liked due to Hoser being busy with, err, “legal efforts” (St. George 2026, p. 130).

**Caption:** Raymond Hoser, a scrupulously ethical individual who definitely doesn’t run numerous different sock puppet accounts on social media nor engage in a worrying amount of projection. Here he is on Twitter/X in 2021, falsely accusing me of stuff.

Use of the term Snake Men as the title is not, as St. George notes deep within the book, without a little irony given that Hoser claims legal ownership of this widely used term, something he has no right to do.

Before getting to Hoser, we learn about the history of formal zoological taxonomy and the herpetologists, amateur and professional, who shaped the road before Hoser came along. I was fascinated to learn that Richard Wells – a polymath and inveterate collector of books as well as a dedicated herpetologist – claims to have gained his special ability to process and understand complex things following a night-time encounter with a mysterious mobile light (St. George 2026, pp. 91-93).

**Caption:** Richard Wells (in green shirt) with just part of his vast collection of books. This screengrab is from a documentary devoted to Wells, viewable **here on YouTube**.

Working together with Ross Wellington, Wells published articles that have to be regarded as among the most controversial within the history of Australasian herpetology (Wells & Wellington 1983, 1985). How Wells and Wellington came to be outcast from formal zoology is pegged, in their version of events, on Ross Sadlier at the Australian Museum but let’s just say that what Wells describes isn’t exactly consistent with the recollections of others, most notably Hal Cogger, author of the standard tome on Australia’s reptile and amphibian diversity (St. George 2026, pp. 100-101). Anyway, there’s a lot to say about the long-running furore that Wells and Wellington initiated and I thoroughly enjoyed St. George’s coverage of it.

It was Wells and Wellington’s work, predictably enough, that inspired Hoser to do what he does, though Wells, at least, doesn’t speak highly of Hoser within the book. But then, not many do. Hoser’s assorted legal dealings, chaotic entertainment business and debates and disputes with academic herpetologists – in particular the group that Hoser terms the ‘Wüster gang’ (after Bangor University snake expert Wulfgang Wüster) – are the focus of several chapters. The ICZN case is covered, St. George at one point taking 14 pages to list the taxonomic names that Hoser published between 2013 and 2021 alone (St. George 2026, pp. 181-194).

**Caption:** at left, opening text of Kaiser *et al*. (2013), a key paper in the fight against taxonomic vandalism in herpetology. At right, Black-necked spitting cobra *Naja nigricollis*, type species for *Afronaja*, the taxon also named ‘Spracklandus’ by Hoser and pivotal in a case submitted to the ICZN on Hoser names. Image: Warren Klein, CC BY-SA 3.0 (**original here**).

All in all, Snake Men is a highly entertaining, well-written overview of one furiously over-active taxonomic vandal in particular and I really enjoyed it. It’s required reading for those interested in the bizarre life and times of Raymond Hoser, and St. George never has to work hard to show Hoser’s true colours. The book says enough about taxonomic vandalism in general to be of broader interest too, and I strongly recommend it to anyone interested in the more arcane side of taxonomy or in reptiles or biology in general.

My thanks to W. W. Norton & Company for provision of an advance review copy.

St. George, Z. 2026. Snake Men: Rebels, Reptiles & the Race to Name Earth’s Creatures. W. W. Norton & Company, New York. ISBN 9781324134084, hardcover, pp. 272. $29.99. *Here from the publishers.*

Hoser names and taxonomic vandalism have been mentioned or discussed on a few previous occasions at Tet Zoo. See…

Scolecophidians: seriously strange serpents, May 2008
The Freshie: Australian crocodile, seemingly from the north (crocodiles part V), July 2012
Ray Hoser, Number 1 Taxonomic Vandal, May 2024

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for TetZoo and the more productive I can be on those long-overdue book projects. Thanks!

Refs - -

Kaiser, H., Crother, B. I., Kelly, C. M. R., Luiselli, L., O’Shea, M., Ota, H., Passos, P. Schleip, W. & Wüster, W. 2013. Best practices: in the 21st Century, taxonomic decisions in herpetology are acceptable only when supported by a body of evidence and published via peer-review. Herpetological Review 44, 8-23.

Naish, D. 2025. Taxonomic trolling. The Biologist 72 (2), 26-29.

Rhodin, A. G. J., Kaiser, H., van Dijk, P. P., Wüster, W., O’Shea, M., Archer, M., Auliya, M., Boitani, L., Bour, R., Clausnitzer, V., Contreras-MacBeath, T., Crother, B. I., Daza, J. M., Driscoll, C. A., Flores-Villela, O., Frazier, J., Fritz, U., Gardner, A., Gascon, C., Georges, A., Glaw, F., Grazziotin, F. G., Groves, C. P., Haszprunar, G., Havaš, P., Hero, J. M., Hoffmann, M., Hoogmoed, M. S., Horne, B. D., Iverson, J. B., Jäch, M., Jenkins, C. L., Jenkins, R. K. B., Kiester, A. R., Keogh, J. S., Lacher Jr., T. E., Lovich, J. E., Luiselli, L., Mahler, D. L., Mallon, D., Mast, R., Mcdiarmid, R. W., Measey, J., Mittermeier, R. A., Molur, S., Mossbrugger, V., Murphy, R., Naish, D., Niekisch, M., Ota, J., Parham, J. F., Parr, M. J., Pilcher, N. J., Pine, R. H., Rylands, A. B., Sanderson, J. G., Savage, J., Schleip, W., Scrocchi, G. J., Shaffer, H. B., Smith, E. N., Sprackland, R., Stuart, S. N., Vetter, H., Vitt, L. J., Waller, T., Webb, G., Wilson, E. O., Zaher, H. & Thomson, S. 2015. Comment on Spracklandus Hoser, 2009 (Reptilia, Serpentes, ELAPIDAE): request for confirmation of the availability of the generic name and for the nomenclatural validation of the journal in which it was published. (Case 3601; see BZN 70: 234–237; 71: 30–38, 133–135, 181–182, 252–253). Bulletin of Zoological Nomenclature 72 (1): 65-78.

St. George, Z. 2026. Snake Men: Rebels, Reptiles & the Race to Name Earth’s Creatures. W. W. Norton & Company, New York.

Wells, R. W. & Wellington, C. R. 1983. A synopsis of the Class Reptilia in Australia. Australian Journal of Herpetology 1, 73-129.

Wells, R. W. & Wellington, C. R. 1985. A classification of the Amphibia and Reptilia of Australia. Australian Journal of Herpetology, Suppl. Ser. 1, 1-61.

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Among the most handsome and striking of seabirds is the Razorbill.

**Caption:** Razorbill profiles. Note that the eyes are reflective enough to be picked out from the plumage, that the white stripe on the head leads toward the middle part of the eye and is not above it, and that the upper part of the bill is deep. Images: **Marek Szczepanek**, CC BY-SA 4.0 (original **here**); **Charles J. Sharp**, CC BY-SA 4.0 (original **here**).

If you don’t know this bird already, its incredibly sharp, perfect black and white lines can look almost unreal, and it’s for this reason that a number of AI images have recently been widely shared online, sigh. Anyway…

The Razorbill Alca torda is a North Atlantic auk, its largest breeding colonies being those around the shores of Britain, Scandinavia, western Greenland, Iceland and north-east Canada. It dislikes ice, avoids brackish water, and prefers rocky islands with broad ledges as nesting areas. It’s a large, stout-bodied auk, and is longer-tailed than other large auks, like guillemots. Long tails in seabirds typically relate to improved aerial manoeuvrability: I can’t find any comments in the literature on the aerial manoeuvrability of the Razorbill relative to other auks (all of which have much shorter tails), but – as we’ll see – an adaptive reason for the long tail might have been identified.

**Caption:** Razorbill pair photographed in Skomer Island, Wales. Image: **Charles J. Sharp**, CC BY-SA 4.0 (original **here**).

The English common name of this species comes from the superficial resemblance its deep, laterally compressed bill has with an old-fashioned razor (its old name was ‘Razorbilled auk’), and it uses this to grab and hold sand-eels, capelin and other mid-water schooling fishes. As many as 20 fish can be held in the bill at any one time. People often wonder how puffins – which manage the same trick – can hold fish while continuing to catch others. I presume Great auks did the same thing, but unfortunately virtually nothing is known of their diet or feeding habits (for data on possible prey species see Olson et al. 1979). Anyway, the birds apparently use their tongue, the horny papillae on the palate, and the mobile prokinetic hinge zone to retain fish at the back of the bill while continuing to grab new ones at the front. It’s still hard to appreciate how this might work, but it obviously does.

The inside of the mouth is yellow. The Great auk also had a yellow mouth interior according to some reports, though others said that the inside of its mouth was red or orange, so we’re not fully sure. I tell you, it’s shocking how little we know of the Great auk: a species that only went extinct in 1844 (or thereabouts).

**Caption:** a Razorbill in Norway demonstrating the presence of a yellow mouth interior. The rhamphothecal grooves are obvious here too and you can also seen the rows of papillae on the soft choanal folds of the palate. A subtle feature is that the white stripe leading from the bill to the eye is approximately continued posterior to the eye by a shallow groove in the feathers. Image: (c) Diego González Dopico, CC BY 4.0 (original **here**).

What’s with the lateral grooves on the Razorbill’s bill? We don’t know, but a few suggestions have been made. One idea is that they play a role in sexual display, and this seems likely given that the bill is used extensively for signalling during courtship. Rather more interesting, however, is the hypothesis that the grooves “function rather like the sights on a rifle as the bird dives in pursuit of its prey” (Freethy 1987, p. 58). This seems unlikely to me given data showing that few birds can see the sides of their own bills (e.g., Martin et al. 2004). Again, highly similar structures were present in the Great auk (and presumably in the fossil species intermediate between these two). Debate exists as to whether the grooves of the Great auk were white or not.

**Caption:** a museum Razorbill skull that I photographed in 2009, still with the rhamphotheca present on the upper jaw (it has broken away from the lower jaw and is now attached to the upper part). Prominent rhamphothecal grooves are present and the deep, curved form of the beak tissue overall is obvious. Note also the shallow and large concavities over the eyes (only visible here in the anterior view). These are the fossae for the supraorbital salt glands. Images: Darren Naish.

Another interesting feature is the narrow channel that extends backwards from the eye. This is better known in guillemots, where – in so-called bridled individuals of Uria aalge – it’s picked out by white feathers. Gaston & Jones (1998) hypothesised that this channel might help “aid the flow of water over the eye while the birds are swimming rapidly” (p. 71), though this intriguing function remains untested so far as I know. Furthermore, if this were true I would expect the Great auk to have such a channel: there’s no mention of it in the Great auk literature (Fuller 1999). Then again – as just mentioned – our knowledge of the Great auk as a live animal is pitiful and it’s not possible to be confident about such a subtle, easily overlooked feature. The ‘lazy’ hypothesis – that the channel might function in communication – was deemed unlikely as the channel is usually all but invisible (Gaston & Jones 1998). It’s been claimed that prominent supra-orbital ridges in the Razorbill might help resist deformation of the eyeball caused by deep-diving: well, maybe. However, some deep-diving seabirds lack prominent supra-orbital ridges, and I’m not sure that Razorbills have prominent supra-orbital ridges anyway.

**Caption:** Razorbill in flight off Skomer Island. Note how shallow the wing is relative to the body (or how long the body is relative to the wings). Image: **Charles J. Sharp**, CC BY-SA 4.0 (original **here**).

I was surprised to learn that as much as 42% of the Razorbill diet might be made up of crustaceans and annelids, and 10% by molluscs (W. E. Collinge, cited in Freethy 1987). This is surprising because the Razorbill is almost exclusively a pursuit-diver, swimming underwater with feet and half-open wings, and its generally short dive time (mostly less than one minute) and maximum dive depth indicate that it doesn’t spend time foraging at or near the sea-floor. Maybe these benthic prey come from the stomachs of the fish it eats. These data might not be accurate, however, as some sources only discuss fishes (and predominantly mid-water fishes) as forming Razorbill diet (Gaston & Jones 1998).

As with so many other seabirds, it turns out that Razorbills dive much more deeply than used to be thought. Older references say that dives are typically of 2-3 m, and that 10 m might be exceptional. More recent work indicates that average dive depth is 25 m, with the range being 11-38 m (Wanless et al. 1988, Barrett & Furness 1990). However, Piatt & Nettleship (1985) reported a maximum dive depth of 120 m for the species and Jury (1986) then reported 140 m, thereby making a mockery of all previous estimates. These deep dives can’t, I assume, have been done in “less than one minute” as noted above, so the bird can clearly stay down for longer than we’ve generally thought.

**Caption:** there are no Razorbills in this photo (so far as I can tell) but the species was present at the location (South Stack, Anglesey) when this photo was taken (August 2016). If you’re interested in seabirds, seeing these giant cliffside breeding ‘cities’ are an incredible experience that you have to seek out. The birds on the ledges here are all Common guillemot *Uria aalge* but Atlantic puffins *Fratercula arctica* and Razorbill were seen on the cliffs and water surface nearby. Image: Darren Naish.

Equally surprising is the fact that the Razorbill is adept at kleptoparatisism, and it frequently steals from Atlantic puffins. Razorbills attack puffins in flight, but more frequently “swim beneath the puffin and torpedo it from below or actually pursue the puffin under the water” (Freethy 1987, p. 71). One might predict that species which indulge in aerial piracy need to be particularly manoeuvrable: is this why the Razorbill has such a long tail?

**Caption:** a really great photo of a Razorbill vs a European shag *Phalacrocorax aristotelis*, taken in Norway. The shag is a fantastic and striking seabird as well, looking something like a slender-jawed miniature dragon. Image: (c) Diego González Dopico, CC BY 4.0 (original **here**).

This brief text is borrowed from an article originally published at Tet Zoo ver 2 back in January 2009 (original here), written following my observation of a razorbill skull in a museum collection.

For previous Tet Zoo articles on seabirds, see…

Hacks Vs Wildlife: the Eternal Vilification of Gulls, July 2019
Pete Dunne and Kevin T. Karlson’s Gulls Simplified: A Comparative Approach to Identification, October 2021

Refs – -

Barrett, R. T. & Furness, R. W. 1990. The prey and diving depths of seabirds on Hornoy, North Norway after a decrease in the Barents Sea capelin stocks. Ornis Scandinavica 21, 179-186.

Freethy, R. 1987. Auks: An Ornithologist’s Guide. Facts on File, New York.

Fuller, E. 1999. The Great Auk. Harry Abrams, New York.

Gaston, A. J. & Jones, I. L. 1998. The Auks. Oxford University Press, Oxford.

Jury, J. A. 1986. Razorbill swimming at depth of 140 m. British Birds 79, 339.

Martin, G. R. & Coetzee, H. C. 2004. Visual fields in hornbills: precision-grasping and sunshades. Ibis 146, 18-26.

Olson, S. L., Swift, C. C. & Mokhiber, C. 1979. An attempt to determine the prey of the Great auk (Pinguinus impennis). The Auk 96, 790-792.

Piatt, J. F. & Nettleship, D. N. 1985. Diving depths of four alcids. Auk 102, 293-297.

Wanless, S., Morris, J. A. & Harris, M. P. 1988. Diving behaviour of guillemot Uria aalge, puffin Fratercula arctica and razorbill Alca torda as shown by radio-telemetry. Journal of Zoology 216, 73-81.

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Salamanders have been covered at Tet Zoo on numerous occasions, there being articles on newts and other salamandrids, plethodontids (lungless salamanders) of numerous sorts, and also ambystomatids (the group that includes the Axolotl and other mole salamanders). It’s the very weird salamanders that bring in the most interest, however, and today we’re going to look briefly at them again….

**Caption:** amphiumas in life. At right is Cuddles, the Two-toed amphiuma *Amphiuma means* formerly kept at the Centre for Fortean Zoology in Exeter, UK. A very big amphiuma can exceed 1.1 m in length. Images: Darren Naish.

Amphiumas are a remarkable group of eel-shaped, long-bodied, aquatic salamanders with internal gills and reduced limbs. They’re represented by just three extant species in one genus (Amphiuma), all restricted to the south-eastern USA. Fossils show that the group has been present in North America since the Late Cretaceous at least. I have written about them in the past. Today, we’re looking briefly just at their cranial anatomy, because boy is it interesting.

**Caption:** diagram of a Two-toed amphiuma *Amphiuma means* skull that accompanies Iordansky (2001) online… yet doesn’t appear in this unlabelled form in the paper. Things to note: this is a robust, well-ossified, relatively long-snouted skull with prominent teeth, a tall coronoid process on the mandible and a prominent, elongate occipital condyle. Image: Iordansky (2001).

In general, the amphiuma skull is “firmly constructed” (Iordansky 2001, p. 180) with an unusually long, narrow and most un-salamander-like snout. Amphiumas lack eyelids, as is typical of aquatic amphibians, and also lack a tongue. The mouth is made shorter than might be expected from skull anatomy thanks to extensive lateral skin folds that cover the jaw margins: here’s a reminder that the structures termed ‘lips’ in the vernacular have other terms in the anatomy literature, with ‘inferior labial fold’ and ‘superior labial fold’ used for the structures extending from the lower and upper jaws, respectively.

**Caption:** facial soft tissues and musculature in an amphiuma, depicted by Iordansky (2001). The small eye is obvious, as is the superior labial fold (lfs) and inferior labial fold (lfi). Moving to muscles, the Maes (adductor mandibulae externus superficialis) and Mdm (depressor mandibulae) and their aponeuroses (like the ca, the coronar aponeurosis) are prominent. Image: Iordansky (2001).

Despite those soft tissue folds, amphiumas can open the jaws extremely wide, in part because the occipital condyles (which are normally paired convexities on either side of the foramen magnum) extend posteriorly as short stalks. This allows the cranium to be rotated much further dorsally (and ventrally) than is normally the case. When writing about all of this in the past, I had to make do with ineffective photos of skull found online. I’ve since discovered that Nikolai Iordansky published a very useful paper in 2001 that includes good images of amphiuma skull anatomy (Iordansky 2001), and hence…

**Caption:** amphiuma skull in dorsal (left) and ventral views, from Iordansky (2001). Some areas (marked with crosses) are cartilaginous, others tendinous (marked with dotted lines). The stalked occipital condyles are obvious; note also the vomerine tooth rows, large shallow areas on the skull roof for jaw muscles, and the big, flattened palatal area, mostly formed by the parasphenoid (prs). Images: Iordansky (2001).

The skull is also kinetic, specifically pleurokinetic thanks to mobile zones between the maxillae and the nasals and premaxilla, and the prefrontal and frontals. These allow the maxillae (which form much of the sides of the snout) to rotate outwards and inwards by around 15° (Iordansky 2001). The premaxillae are fused into a single element, perhaps to reinforce the snout and allow for more powerful bites, though a role in burrow-digging has also been suggested. The sides of the snout bones have a distinctive sculptured texture.

Amphiumas will simply grab prey with a rapid bite, but high-speed photography has shown that they can also use extremely rapid jaw-opening and throat expansion to generate suction (Erdman & Cundall 1984). The adductor and depressor muscles are substantial; the adductors are attached to multi-part, tendinous sheets (aponeuroses) that cover much of the sides of the rear part of the skull and even well posterior to it.

**Caption:** I like this photo of a captive amphiuma as it emphasizes the fact that salamanders can ‘elevate’ (or extend) the head and neck relative to the long axis of the vertebral column. Image: (c) Andri Pogo, archived version **here**.

The teeth have distinct pedicels (flexible zones separating the crowns from the roots) and there are rows of teeth on the vomerine bones on the palate. Amphiumas are reported to bite readily and nastily and are sometimes referred to as to the only amphibians within their range that can pose any sort of physical (as opposed to chemical) danger to humans.

For previous Tet Zoo articles on salamanders (there’s a lot, but much of it is now only findable on the internet archive and a whole list of articles I just checked are all ruined and lacking images) see…

Nobody Expects the Korean Crevice Salamander UPDATE.. now titled The Korean Crevice Salamander, August 2015
A Need for News on Nototriton, August 2015
The Pacific Giant Salamanders, October 2016

Erdman, S. & Cundall, D. 1984. The feeding apparatus of the salamander Amphiuma tridactylum: morphology and behaviour. Journal of Morphology 181, 175-204.

Iordansky, N. N. 2001. Jaw apparatus of the permanent-aquatic Urodela: paedomorphosis, neoteny, and feeding adaptations. Russian Journal of Herpetology 8, 179-194.

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Long-time readers of Tet Zoo will perhaps be aware of my efforts, beginning in 2012 or something like that, to depict in a montage the diversity of crocodylomorphs extinct and extant. This whole thing actually started way back in 2001 when I published a terse but adequate review article of crocodylomorph diversity in Geology Today (Naish 2001).

**Caption:** low-res section of the Crocodylomorph Empire poster. **Now on sale, right now only for collection at DinoCon 2026.**

My aim since then has been to publish a big review of crocodylomorph diversity and fossil history. Maybe that’ll happen one day but you can understand that the topic is considered sufficiently niche that publishers generally don’t want to commit to an entire book on the group. Anyway, various versions of the montage have appeared since I compiled it, including in museum displays, on a T-shirt and in online articles.

**Caption:** old versions of my big croc montage, this being the ‘vertical’ version designed for t-shirts… hence the hilarious text at right. Yes, there actually are people who own t-shirts with this very design. I do.

  **Caption:** the more horizontal version of the montage, this one showing several variants of the animals that I later became unhappy with and replaced, the metriorhynchids in particular.

In 2014, Jeff Martz – best known for his work on aetosaurs, an armour-plated group of Triassic croc-line archosaurs – liked the montage sufficiently that he opted to colour it, meaning that we have a new and improved version, much more attractive than my black-and-white original. It looked so good that Jeff and I discussed plans to make it the main focal point of a big, croc-themed poster. And, thus, more than ten years later… here we are. Yes, it’s taken me years to put this together, but I’ve finally done it. The montage is at the core and the main feature, but there are discussions of anatomy, skull diversity, croc nomenclature and phylogeny overall, and individual pieces of text on the featured taxa too. It should be obvious that this is a visual treat to those interested in these animals, or in archosaurs or reptiles more broadly.

A downside of the fact that Jeff worked with my 2014 version is that a few of the animals look technically incorrect for 2026, mostly because my drawings were based on information that’s now been superseded by new finds and publications. This is all explained within the text of the poster: it’s a necessary consequence of effort to depict fossil animals in life that views become out of date.

**Caption:** three enormous, diverse crocodylomorph groups are wholly extinct, though notosuchians persisted to relatively recently (the Late Miocene or maybe even the Pliocene). The life reconstructions and skull drawings here all feature on the poster.

Crocodylomorph, crocodyliform, crocodylian; and what is a ‘crocodile’? A topic I’ve aimed to emphasize here at Tet Zoo – thus far with limited success due, in part, to the loss of so much material published at ver 2 and 3 – is the massive diversity of fossil crocodylomorphs. The archaic ‘sphenosuchians’ of the Triassic and Jurassic and the ‘protosuchians’ of the Triassic, Jurassic and Cretaceous (neither group appears to be a clade) are extremely different from living members of this group, and the thalattosuchian, notosuchian and tethysuchian radiations – all wholly extinct – have proved enormous and complex and all contain species, again, very different from their extant cousins.

**Caption:** a simplified cladogram from the poster, used to show the difference between the terms Crocodylomorpha, Crocodyliformes, and Crocodylia.

The terminology of these animals – I’ve addressed this several times at Tet Zoo but still need to keep doing it – is confusing but the majority of workers follow the Benton & Clark (1988) system. ‘Crocodilia’ has been sufficiently vague across its history that Clark (in Benton & Clark 1988) opted to restrict Crocodylia (which is the more correct spelling in view of the generic name it’s based on) to the crown-group, Crocodyliformes being used for all ‘crocodylian-like’ lineages and Crocodylomorpha for the whole lot. This is why I refer to all of these animals as ‘crocodylomorphs’: I opt not to use ‘crocodilian’ at all anymore, and ‘crocodylian’ should only be applied to the crown.

Adding further complexity is that some palaeontologists – including the Benton of Benton & Clark (1988)! – have pushed back on this system and have argued that we should use Crocodylia in place of Crocodylomorpha. I agree with Brochu et al. (2009) that this is a stupendously bad idea at this point in the game. And I wish that palaeontologists would wean themselves off the frustrating habit of just calling everything a ‘crocodile’. An alligator is unambiguously not a crocodile, ergo extinct crocodylomorphs sure aren’t either. Crocodiles are a specific group within Crocodylia, and not an especially old one in geological terms.

**Caption:** how many crocodiles are in this montage? Answer: ZERO. It is categorically wrong to call alligatoroids of any sort ‘crocodiles’. Ergo, the term ‘crocodile’ has a strict and specific meaning and we should push back against the tendency in palaeontology to just slap it on all members of Crocodylia, Crocodyliformes or Crocodylomorpha. Images: Darren Naish.

The poster is now on sale. Anyway… if you want to learn more about crocodylomorph diversity, anatomy and evolutionary history, I hope you’re interested in getting the poster yourself. Yes, years in the making, representing a huge amount of work! It’s big, at 1.4 m long and 76 cm deep.

The good news is that it’s available for sale right now, initially *only for collection in person at DinoCon 2026*. Yes, DinoCon, happening this year on July 25th-26th at the Hilton Birmingham Metropole (tickets and accommodation available here). Once DinoCon is out of the way we’ll be selling the poster more widely, but please be patient.

Thanks for your interest, and more on crocodylomorphs here in due time.

Your usual lament. Crocodylomorphs of several sorts have been covered here on a reasonable number of occasions but most of this material is now lost due to the death of ScienceBlogs and Scientific American blogs. Consequently, only a small number of articles are easily findable today. I aim to recover and republish my older articles here, in time. Sigh. Anyway…

Crocodiles Attack Elephants Then, Now, and Still, August 2025
Koumpi the Wealden Croc: Tons of New Data and Phylogeny Resolved, April 2026
‘Shanklin Croc’ and the Dawn of the Tethysuchian Radiation, April 2026

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

Refs - -

Benton, M. J. & Clark, J. M. 1988. Archosaur phylogeny and the relationships of the Crocodylia. In Benton, M. J. (eds) The Phylogeny and Classification of the Tetrapods, Volume 1: Amphibians, Reptiles, Birds. Clarendon Press (Oxford), pp. 295-338.

Brochu, A. C., Wagner, J. R., Jouve, S., Sumrall, C. D. & Densmore, L. D. 2009. A correction corrected: consensus over the meaning of Crocodylia and why it matters. Systematic Biology 58, 537-543.

Naish, D. 2001. Fossils explained 34: Crocodilians. Geology Today 17 (2), 71-77.

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I feel the need to publish something new on rodents, and with no time to produce anything lengthy or all that complex, here are brief thoughts on one of my favourite rodent groups: voles!

**Caption:** voles alive! At left, a Short-tailed field vole *M. agrestis* I encountered in rocky scrub at Lepe Beach, southern England. At right, Lusitanian pine vole *M. lusitanicus*, an Iberian endemic. Images: Darren Naish; José Ramón Pato Vicente, CC BY-SA. 2.5 (**original here**).

Voles (Arvicolinae or Arvicolidae), as a generalisation, are mostly small (on average, 10-11 cm long, with a tail of 3-4 cm), blunt-headed, short-tailed, herbivorous muroids. They’re predominantly animals of Northern Hemisphere forest floors, grasslands, steppes and rocky places, many constructing near-surface tunnels and burrows. Bog and marsh dwellers, and amphibious species, are part of the group too: the unusually large and long-tailed muskrats (Ondatra) are voles, as are the cold-adapted, tundra-dwelling lemmings (the subgroup Lemmini). Collared or varying lemmings (Dicrostonyx), which are a distinct group from true lemmings, are cold-adapted voles of North America and Eurasia that undergo colour change across the seasons and – uniquely – grow massive, double-pronged digging claws during the colder parts of the year.

This is a highly speciose group, with around 155 extant species classified in about 30 genera. They’re related to cricetids (hamsters, New World mice and kin) and included with them in the same ‘family’ by some authors. ‘Classic’ voles, at least here in Europe, include water voles (Arvicola) and field voles and their many relatives (Microtus). Those last two are familiar and widespread in Eurasian faunas and the fact that their massive populations have long been of significance to agriculture, and undergo great cycles of boom and bust, have made them the subject of substantial scientific study.

**Caption:** artistic depictions of (at left) a *Phenacomys* heather vole; at right, winter and summer looks for a *Dicrostonyx* varying lemming. Images: Darren Naish, colouring by Gareth Monger.

North America has a list of voles that are very unusual by Eurasian standards. Some are arboreal (the Arborimus tree voles) and some are inhabitants of upland heaths (the Phenacomys heather voles). I wrote about North America’s more distinctive voles in a 2014 article at Tet Zoo ver 3. Several different phylogenetic hypotheses have been published for voles, some of these showing collared lemmings, true lemmings and heather voles outside of a clade that includes muskrats and microtine voles (Chaline & Graf 1988, Chaline et al. 1999, Conroy & Cook 1999, Cook et al. 2004, Galewski et al. 2006).

**Caption:** a simplified vole cladogram, this depicting a topology where heather voles and collared lemmings are outside the clade that includes voles like *Microtus*. Image: Darren Naish.

Voles have an excellent and complicated fossil record from the Miocene onwards. This involves numerous immigration events from Eurasia into North America; events that involve the opposite – the moving of North American groups into Eurasia – were evidently far rarer, maybe occurring just once in the case of collared lemmings. Triangular or T-shaped occlusal surfaces are typical of their teeth as is the presence of a distinctive muscle scar for the medial part of the masseter and a so-called ‘arvicolid groove’ on the mandible (Martin 2007). Changes in the abundance of certain lineages and the evolution of their teeth can be linked to climatic shifts and a trend towards hypsodonty is present with some species possessing rootless molars. Ondatra has an excellent North American fossil record with a series of forms demonstrating increasing body size, hypsodonty and tooth complexity.

**Caption:** at left, upper and lower cheek teeth of a Bank vole *Clethrionomys glareolus* showing the zig-zagging form of the occlusal surfaces. At right, an open-rooted, constantly growing cheek tooth of a Short-tailed field vole *M. agrestis* on the left, and a close-rooted Bank vole tooth. Images: Lawrence & Brown (1967).

And that will do. There is lots on rodents in the Tet Zoo archives… as ever, mostly ruined by the downfall of ver 2 and 3. One day I might renovate it for republication here. Among those articles that are still worth checking out are…

Osgood, Fuertes, and mice that swim and mice that wade, March 2006
Australia, land of placentals (part I), December 2007
Australia, land of placentals (part II), December 2007
New, obscure, and nearly extinct rodents of South America, and… when fossils come alive, November 2008
Of vole plagues and hip glands, January 2014
North America: land of obscure, freaky voles, January 2014
A brief history of muskrats, September 2014
Cricetomyines: the African pouched rats and mice, November 2014
African Climbing Mice and the Congo Link Rat, October 2015
Meet the Shreats, July 2016
The Sigmodontines, March 2017
Cloudrunners and Other Cloud Rats of the Philippines, June 2020

Refs - -

Chaline, J., Brunet-Lecomte, P., Montuire, S., Viriot, L. & Courant, F. 1999. Anatomy of the arvicoline radiation (Rodentia): palaeogeographical, palaeoeocological history and evolutionary data. Annales Zoologici Fennici 36, 239-267.

Chaline, J. & Graf, J. D. 1988. Phylogeny of the Arvicolidae (Rodentia): biochemical and paleontological evidence. Journal of Mammalogy 69, 22-33.

Cook, J. A., Runck, A. M. & Conroy, C. J. 2004. Historical biogeography at the crossroads of the northern continents: molecular phylogenetics of red-backed voles (Rodentia: Arvicolinae). Molecular Phylogenetics and Evolution 30, 767-777.

Conroy, C. J. & Cook, J. A. 1999. MtDNA evidence for repeated pulses of speciation within arvicoline and murid rodents. Journal of Mammalian Evolution 6, 221-245.

Galewski, T., Tilak, M.-K., Sanchez, S., Chevret, P., Paradis, E. & Douzery, E. J. P. 2006. The evolutionary radiation of Arvicolinae rodents (voles and lemmings): relative contribution of nuclear and mitochondrial DNA phylogenies. BMC Evolutionary Biology 2006 6: 80.

Lawrence, M. J. & Brown, R. W. 1967. Mammals of Britain: Their Tracks, Trails and Signs. Blandford Press, Poole, Dorset.

Martin, R. A. 2007. Arvicolidae. In Janis, C. M., Gunnell, G. F. & Uhen, M. D. (eds) Evolution of Tertiary Mammals of North America, Vol. 2. Cambridge University Press, Cambridge, pp. 480-497.

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I am extremely pleased to announce that my Dinosaurs: How They Lived & Evolved – the Natural History Museum, London’s flagship dinosaur book, co-authored with Professor Paul Barrett – is once again in print, once again as a new edition…

Yes, this is the *fourth* edition and its publication ten years after the book’s first appearance in 2016 means that it’s a special, expanded tenth anniversary version (Naish & Barrett 2026). I’ve been in possession of a personal copy for a few months but a recent visit to the NHM (I was there for the opening of the *Jurassic Oceans* exhibition) has allowed me to confirm that it’s out and on sale, and available on the shelves. Buy your own copy here.

**Caption:** proof that the 4th edition is now out and findable on shelves… at least, in the Natural History Museum. I found copies both in the museum’s main shop (left) and in the smaller shop adjacent to the dinosaur exhibition. Images: Darren Naish.

I’ve mentioned in previous discussions on books I’ve authored that keeping a book in print over several editions is a hugely significant thing in personal terms. My thanks to everyone who’s bought Dinosaurs: How They Lived & Evolved (DHTLE) – whatever edition that was – and of course to my coauthor Paul, to all of our colleagues in the publishing team, and to the many artists, scientific colleagues, photographers and designers whose work we used.

Maybe I’m weird, but my favourite part of the book – and the part I most enjoyed compiling – is the preface. Yes, with ten years and three previous editions behind us, the addition of a preface seemed appropriate. The idea that we might include one comes from Gower et al.’s (2023) Snakes: Their Diversity, Ecology and Behaviour, another Natural History Museum book that’s part of the same series as DHTLE. I really liked the fact that the preface featured within that book (Gower et al. 2023, pp. 5-6) describes the NHM’s noble history of producing snake-themed books and of being globally important in snake research. For the fourth edition of DHTLE we should, I mused, do the same: not only has the NHM been crucial to our understanding of dinosaurs ever since Richard Owen, it also has a history of publishing mainstream books on these animals, books that have done a good job of bringing the scientific developments of the time to the public. And if you want to know more on what I’m getting at, buy the book and find out for yourself.

**Caption:** **Naish & Barrett (2026)**, and its previous editions, is the successor to Alan Charig’s *A New Look at the Dinosaurs* (in press between 1979 and the early 1990s) and Tim Gardom and Angela Milner’s *The Natural History Museum Book of Dinosaurs* (in press between 1993 and the 2000s). Not shown here is William Elgin Swinton’s 1934 *The Dinosaurs: A Short History of a Great Group of Extinct Reptiles*. Image: Darren Naish.

Once again, we have outstanding new cover art by Bob Nicholls. Our previous editions have featured theropods (twice) and an ornithischian, so this time it was only right to feature a sauropodomorph. Bob did us proud and has created a convincing new-look version of Argentina’s Amargasaurus, the animal looming toward the viewer, its fantastically textured surface being displayed in detail. The reconstruction takes account of Cerda et al.’s (2022) argument that the long bony neck spines of this sauropod (and presumably related taxa too) weren’t horn-covered spines after all, but more likely united in paired sails… as was proposed initially, back when the late Brian Franczak first created a life reconstruction of Amargasaurus in 1992.

**Caption:** so, we generally don’t talk about the cover of the first edition (at far left), but the covers of the second edition (the heterodontosaurid one at far right) and third edition (featuring the British spinosaurid *Ceratosuchops*, at centre) look fantastic. We have a great working relationship with Bob Nicholls, who created both of these images.

As a tenth anniversary special edition, it only seemed right to use this as an opportunity to expand the book. And expand it we did: there are new sections on dinosaur brains and intelligence (things on this front have, after all, been interesting of late…) and also ecological modelling. Numerous updates are included throughout, including on elasmarians and other ornithopods, facial tissues in sauropods, tyrannosauroids and other dinosaurs, spinosaurids (of course), the end-Cretaceous extinction, neornithine bird evolution, and more. We went to press just a little too late to make allowance for the reappraisal of Nanotyrannus but – even prior to the publication of Zanno & Napoli (2025) – the writing was on the wall and hence our wording isn’t as ‘against’ the hypothesis of Nanotyrannus being valid as it might have been.

We were, however, able to take account of Cau’s (2024) argument that various small theropods conventionally grouped together as compsognathids might be the juveniles of megalosauroids, tyrannosauroids and such. Ergo, the hypothesis is now in the ‘mainstream’, non-technical literature.

You know, I’ve heard a few academics over the years say that they don’t consider a hypothesis or proposal worth paying attention to until it works its way into the popular literature. That might seem odd, and I don’t agree with it myself, but it does make you think. If that is how some people decide which scientific ideas and arguments are and are not worth absorbing into the canon… well then, with great power comes great responsibility, since those of us writing and publishing books of this sort therefore have control in terms of choosing which results and hypotheses from the technical literature ‘make it’ into popular texts. Make of that what you will.

**Caption:** the section on dinosaur life appearance now includes images of the famous Los Angeles *Edmontosaurus* skull (LACM 23502) shown at right, famously preserved with part of its original beak tissue in place (the peculiar absence of the tissue on the animal’s right is the result of its accidental destruction by a preparator). These animals weren’t ‘duck-billed dinosaurs’ at all (a term we owe to Barnum Brown, who first used it in 1900): the rhamphotheca was downturned and massive and the form of the underlying bony rostrum was not present in life. **I wrote about this at Tet Zoo ver 3 in 2018** but it is not in the least bit original to me, since Versluys in 1923 and Morris in 1970 both emphasized the same thing. All those authors and artists pushing ‘duck-billed’ hadrosaurs throughout the latter part of 20th century were simply unaware of what was in the literature the whole time. Images: Darren Naish.

We also used this as an excuse to fully overhaul the illustrations, meaning that new images of fossils, extant animals, mounted skeletons and more are present throughout. We’ve also incorporated a good number of new artistic reconstructions, including by Andrey Atuchin, Brian Choo, Edyta Felcyn-Kowalska, Anthony Hutchings, Bob Nicholls and Mark Witton. I owe a massive debt of thanks to the extremely hard-working Marcin Ambrosik for his production of a number of excellent new colour pieces of art, variously featuring maniraptorans and sauropods. An oversight means that Marcin is not adequately thanked in the acknowledgements, a mistake on my part that I’ll correct in future.

**Caption:** some of the new art that appears in the book. At left, displaying caudipterids from the maniraptoran section, by Bob Nicholls. At right, titanosaur hatchlings from the section on dinosaur reproduction, by Marcin Ambrozik. Images: Bob Nicholls; Marcin Ambrozik.

There’s more that could be said but that’ll do. It’s gratifying to see a book that took so much work sell out, and then get the green light to be released via a new edition four times over. If you own previous editions of Dinosaurs: How They Lived & Evolved, now’s the time to get an updated, expanded one (the book is not expensive!). And if you haven’t obtained the book before, now is the time.

Finally… if you’re aware of DinoCon (happening this year on July 25th-26th at the Hilton Birmingham Metropole, book tickets and accommodation here), this new book might have a presence have. I said might.

For articles on the previous editions of Dinosaurs: How They Lived & Evolved, see…

Naish and Barrett's Dinosaurs: How They Lived and Evolved, November 2016
The Second Edition of Naish and Barrett’s Dinosaurs: How They Lived and Evolved, November 2018
The Third Edition of Naish and Barrett's Dinosaurs: How They Lived and Evolved, February 2024

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

Refs - -

Cau, A. 2024. A unified framework for predatory dinosaur macroevolution. Bollettino della Società Paleontologica Italiana 63, 1-19.

Cerda, I. A., Novas, F. E., Carballido, J. L. & Salgado, L. 2022. Osteohistology of the hyperelongate hemispinous processes of Amargasaurus cazaui (Dinosauria: Sauropoda): implications for soft tissue reconstruction and functional significance. Journal of Anatomy 240, 1005-1019.

Gower, D., Garrett, K. & Maddock, S. 2023. Snakes: Their Diversity, Ecology and Behaviour. The Natural History Museum, London.

Naish, D. & Barrett, P. M. 2026. Dinosaurs: How They Lived & Evolved (Fourth Edition). The Natural History Museum, London.

Zanno, L. E. & Napoli, J. G. 2025. Nanotyrannus and Tyrannosaurus coexisted at the close of the Cretaceous. Nature 648, 357-367.

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It’s time for another article in my Tet Zoo Reviews Zoos series, and this time we look at a zoo in that troubled and terrifying nation known as the USA, specifically in the north-western state of Oregon…

**Caption:** an Oregon Zoo montage, featuring animals seen on my visit of late 2022. Images: Darren Naish.

My Oregon Zoo trip wasn’t recent but happened way back in November 2022; I just haven’t had time until now to finish the article. I should mention that I was able to visit thanks to the hospitality of my host, the very excellent Nico Spadafore, so big thanks to him. Nico himself worked at the zoo back in the distant past and thus gave me some great insider info that I wouldn’t have had otherwise.

**Caption:** the zoo’s main entrance is relatively understated. The iron gates (just visible here) feature nice metal sculptures, including of penguins, otters, monkeys, elephants, rhinos, flying frogs(!) and more. Image: Darren Naish.

Oregon Zoo was founded in 1888, making it the oldest zoo in the entire west of the US, but it changed locations and names a few times prior to its 1959 relocation to Washington Park. It’s involved in a good number of conservation efforts, several involving local species like Oregon spotted frog Rana pretiosa, Pygmy rabbit Brachylagus idahoensis (specifically those of the Columbia Basin population) and Oregon silverspot butterfly Speyeria zerene hippolyta. The zoo has a leading role in California condor Gymnogyps californianus conservation, both in terms of breeding and releasing birds into the wild and in leading educational campaigns to reduce the quantity of lead shot that exists in condor habitat. Some of the condors involved in the zoo’s programme are on exhibit but others are kept at the off-site Jonsson Center in Clackamas County. They’ve then been released in northern California and the hope is that they’ll naturally expand their range to recolonise Oregon.

**Caption:** each California condor is identified by its numbered tag, and here’s one of the several individuals (a young one) that was on show at the zoo during the time of my visit. Fossils show that California condors were far more widespread in the geological past, formerly occurring east to west across North America. The habitat we associate with them today is by no means one the species is specialised for. More on condors below. Image: Darren Naish.

Oregon Zoo houses six zones: Great Northwest (devoted to North American wildlife), Pacific Shores, Primate Forest, Elephant Lands, Africa and Discovery Zone. My coverage here will discuss the animals and enclosures as they were encountered by Nico and I as we walked through the zoo, rather than in phylogenetic order or anything like that. Our visit occurred on an overcast, sometimes rainy, day and this affected both my chances for photography and the tendency of the animals to be out and about.

Great Northwest. Immediately upon entering the zoo, you’re in the North American ‘Great Northwest’ section, the trees and landscaped and planted areas surrounding the walkways being of Oregonian sort. Puma Puma concolor was on show (the zoo uses ‘Cougar’) and nearby were two Bald eagles Haliaeetus leucocephalus. It looked to me as if both birds had damaged eyes and this made me think that they were rescued birds that couldn’t be released into the wild.

**Caption:** two Bald eagles on a tree on a damp day. The Bald eagle would be better termed the White-headed eagle (‘balde’ is an old term for ‘white’), and of course versions of this name are used in some languages. At right, one of two reclining Puma seen during our visit. I assume that they’re of local, North American origin, since the species is distributed north to south throughout the mainland of the Americas. Images: Darren Naish.

Close by was a Rocky Mountain goat Oreamnos americanus, a new species for me and one I was very happy to see up close. As you can see, the individual I photographed was both co-operative and charismatic and provided a good view of its tongue (for some reason) as well as its horns and bodily profile. Oreamnos is a strange animal and isn’t really a ‘goat’ at all: it’s not a close relative of the Old World animals most properly associated with that name (Capra hircus and kin), instead perhaps being closer to takins.

**Caption:** maybe you’ve seen a Rocky Mountain goat before... but have you seen its tongue? The tongue is smooth and greyish, with a slightly notched tip. Image: Darren Naish.

  **Caption:** as a collector of toy and model animals, I’ve always been bothered by the fact that several toys of this species give it light brown horns with transverse rings or ridges. As you can see, they’re dark grey and smooth. Image: Darren Naish.

Nearby was another new species for me but one highly familiar to many North Americans: a Black bear Ursus americanus. I haven’t spent enough time in North America to ever see a Black bear, either wild or in captivity, and this one was not only of the very dark sort typical for the species but also a highly robust – I’m not going to fat shame – individual with substantial adipose deposits about the body. The animal was engaging in an interesting behaviour that I’ve elsewhere seen in pet cats: it was dunking its paws in a trough and then licking off the water. Presumably (and hopefully) this reflects the fact that it wanted to drink, since behaviour of this sort can reflect boredom, a real problem for captive bears as I’ve said before.

**Caption:** the bad lighting of the day has resulted in poor photos, many a bit washed out (though the real excuse is that I’m a poor photographer with no good understanding of how cameras work). Here’s one of the zoo’s several American black bears, seen from an overhead walkway and while it was reclining at a drinking trough. Images: Darren Naish.

  **Caption:** any and all bears can be dangerous, but this comparison at the zoo emphasizes how a black bear (not all populations of which are black) is a very different beast from a grizzly, which is an American version of the brown bear *Ursus arctos*… at least, according to the taxonomy we use today. I really must write some time about the bear taxonomies that were used in the past. Image: Darren Naish.

A covered walkway, landscaped to look like a natural tunnel, was home to a Pacific lamprey Entosphenus tridentatus exhibit and a large, glass-fronted display featured trout and sturgeon. Northwestern garter snake Thamnophis ordinoides and Western toad Anaxyrus boreas had small enclosures nearby. The toad sign used the ‘traditional’ name Bufo boreas for that species; I’m an advocate of the Frost et al. (2006) taxonomy for toads.

**Caption:** at left, large, landscaped aquarium, featuring trout of several species as well as White sturgeon *Acipenser transmontanus*. At right, one of many Pacific lamprey on show. Yes, I have close-up photos of their mouths but opted not to share them here. Images: Darren Naish.

Condors. A highlight was the large California condor exhibit. Several British bird collections house Andean condors Vultur gryphus but I can’t recall seeing the North American species before. Juvenile (albeit adult-sized) and adult condors were both on show, and all were busy feeding on a carcass (I think of a domestic horse foal). A sign warning people to be prepared to witness the consumption of a domestic mammal carcass was located at the entrance to this section, which is fair enough; you have to respect that some people really are bothered by the sight of dead animals. An impressive condor sculpture made of waste plastic was also near the entrance.

**Caption:** adult California condor with bill partly inside a large mammal carcass. It’s well known that condors (and cathartids/vulturids in general) can’t grasp or carry items with their feet: note how long the toes are and how the talons are nowhere near as sharply curved as they are in eagles and falcons. Image: Darren Naish.

  **Caption:** my first view of California condor. I was impressed with lots of things, but something I hadn’t thought about before was the profound difference of pigmentation between the very black youngsters and the carotenoid-heavy, brightly coloured adults. Their extensive facial soft tissues are of major interest too. Images: Darren Naish.

  **Caption:** good zoos include good art, and Oregon had this great display on plastic pollution near its condor exhibit. Image: Darren Naish.

  **Caption:** a good edutainment feature used by several zoos and wildlife parks… compare your arm-spread with the wingspans of several birds, in this case raptors. California condors can have a wingspan of 3 m but there are unverified claims of birds measured at 3.4 m. So far as we can tell, they’re outclassed in wingspan by the Andean condor, record-holders of which have been measured at 3.3 m. Image: Darren Naish.

North American ducks. I’m a fan of wildfowl, so I was pleased that several species were on display and viewable at close range due to windows positioned at water level on the enclosure side and human standing level on the other. None of the birds were substantially exotic to a denizen of northern latitudes, but I know what I like and I like ducks. I’m also relatively inexperienced when it comes to the ducks of North America, at least some of which don’t have ordinary residential status here in Europe.

**Caption:** a North American duck montage part 1, showing (clockwise from top left) Ring-necked duck, Ruddy duck and Bufflehead and Hooded merganser. At lower left is an image showing how part of the exhibit allows you to see the ducks underwater, a really nice feature. I’m not fully sure which species is visible there but I think it’s an aythyin (a diving duck, like the Ringed-necked). Images: Darren Naish.

The species I remember seeing are Ruddy duck Oxyura jamaicensis, Hooded merganser Mergus cucullatus, Wood duck Aix sponsa, Ring-necked duck Aythya collaris, Bufflehead Bucephala albeola, Northern shoveler Spatula clypeata, Green-winged teal Anas crecca and Redhead Aythya americana. A sign said that Cinnamon teal S. cyanoptera was on show too, but I didn’t knowingly see that species. A shame, as it would have been new to me.

**Caption:** a North American duck montage part 2, showing (clockwise from top left) Carolina duck, Green-winged teal, Northern shoveler, Redhead. My apologies for focusing on the more gaudy, boldly marked males. Images: Darren Naish.

North American beaver Castor canadensis, Snowy owl Bubo scandiacus, River otter Lontra canadensis and Northwestern pond turtle Actinemys marmorata were in this part of the zoo too.

**Caption:** I’ve seen American River otter in captivity in the UK, so seeing one in the US wasn’t a first for me. At right, two beavers in the indoor den part of their enclosure, which they share with the selection of American wildfowl discussed above. The zoo has had involvement with various local beaver rescue and release projects. Images: Darren Naish.

Pacific Shores. We followed the walkway south and moved to the Pacific Shores section, a crown jewel of the zoo. A second bear species – Polar bear U. maritimus – was on show here. As typical for Polar bear exhibitions, the enclosure had a blue-walled pool and was fronted by glass. One of the bears spent time right at the front, allowing close-up views. Again, I saw a damaged eye; I don’t know if this is a coincidence or reflects the zoo taking on animals that can’t be elsewhere.

**Caption:** part of the polar bear enclosure, showing a low cliff and other (fake) rocky structures and a deep blue pool. I didn’t get to see the bears swimming while I was there, unfortunately. I’ve seen polar bears in captivity several times but can’t recall ever seeing them in the water. Image: Darren Naish.

  **Caption:** two bears were on show at the time of my visit, one dozing on the rocks and one right up close to the glass. The polar bear is a member of the brown bear clade but exactly where it fits within that group varies among studies. I understand that technical work does mostly still find it nested within a specific clade of grizzly bears. Images: Darren Naish.

More unusual and more noteworthy (no offence, polar bears) was the sea otter Enhydra lutris display. However… the only sea otter I recall seeing (2022 is a long time ago) was a statue. I do remember seeing sea otters on the same trip, but they were at the Oregon Coast Aquarium, a venue I’ll discuss in a different article, if time allows. Anyway, an adjacent Harbour seal Phoca vitulina section featured a nice landscaped ‘cove’ above water, and an impressive underwater section with rocky tunnels, overhangs, submerged logs and caverns. From underground floor-to-ceiling windows, you could watch the seals close up as they swam back and forth, sometimes upside down as is so common for pinnipeds. Also in this section was a coastal ‘general marine life’ exhibit, with crabs, urchins, sea stars, chitons and anemones, rockfishes and surfperches.

**Caption:** a strength of the zoo is the presence of life-sized statues, and here’s one showing a sea otter doing the standard ‘using the chest as a table’ thing. Sea otters are diverse in terms of what they eat but studies show that they mostly stick to the kind of prey they were introduced to by their mothers. Image: Darren Naish.

  **Caption:** the above-water ‘Steller Cove’ area of the harbour seal exhibit, showing rock walls and haul-out sites well suited for pinnipeds. It looks appropriately NW Pacific I think. Image: Darren Naish.

  **Caption:** underwater viewing area of the seal display, two seals visible swimming at right. I really like the landscaping here; it’s a great exhibit. The Harbour (or Common) seal is a widespread species that occurs throughout the coastal North Atlantic and North Pacific and the two have traditionally been recognised as distinct subspecies (*P. v. vitulina* in the Atlantic and the larger *P. v. richardii* in the Pacific). Image: Darren Naish.

A central rotunda – very close to the literal centre of the zoo – was devoted to penguins, though only Humboldt penguins Spheniscus humboldti were there. Inca tern Larosterna inca and Grey gull Larus modestus were present too. I’ve commented before on the relative rarity of gulls in zoo displays, so Oregon Zoo is on the short list concerned. The building – with blue interior walls and a mostly concrete rocky ‘island’ for the penguins in the middle – wasn’t great and didn’t show the birds in an interesting or pleasing setting. I really hope it gets a revamp.

**Caption:** interior view of the rather sorry penguin exhibit. You might just be able to see a few Inca terns at far right. The yellow toy ducky is a nice touch. Image: Darren Naish.

  **Caption:** Oregon Zoo is very strong on signage and art that informs visitors about the animals they’re looking at, and various items in the penguin rotunda are devoted to the penguins of the world. Several zoos showcase penguin diversity in this way. It’s also not infrequent to see some representation of extinct species, focusing of course on the giant species of the Paleogene. Image: Darren Naish.

We moved next to the Primate Forest, where there are chimpanzees Pan troglodytes, Bornean orangutan Pongo pygmaeus (and, formerly, Sumatran P. abelii too) and White-cheeked gibbon Nomascus leucogenys. Tall indoor and outdoor enclosures provided a lot of climbing space and meant that the apes could stay out of view if they wanted to, and I didn’t see any of the chimps. A male orang was suitably active near the indoor windows when I was there.

**Caption:** the zoo’s adult male Bornean orangutan Bob at left, who came to the zoo from Greenville Zoo in South Carolina in 2014. A Sumatran orangutan female kept at Oregon Zoo – Inji – was 61 when she died in 2021, and the oldest orangutan in captivity. At right, a Schmidt’s red-tailed monkey *Cercopithecus ascanius schmidti*, a Central African guenon. Images: Darren Naish.

  **Caption:** many zoos today have a feature where you can ‘insert’ yourself (or a companion) into a montage depicting our closest relatives, and here’s one at Oregon. Bonobos and gorillas aren’t kept at the zoo but gibbons, orangs and chimpanzees are. As for Nico… he’s only there occasionally. Image: Darren Naish.

Elephants. Virtually the whole of the zoo’s east and south-east is occupied by the massive Elephant Lands section, this housing Asian elephants Elephas maximus and including two indoor display areas and a large outdoor arena with a deep pool. The zoo is strongly associated with Asian elephants and has housed individuals since 1953; a remarkable 29 calves have been born at the zoo. One individual – a male called Samudra – is a third-generation captive-born animal, the only such elephant in the US.

**Caption:** elephants outdoors, doing interesting things. I can’t remember what the adult female at left was doing, but the young animal at right is reaching under the fence to pluck grass from the verge. Note how this behaviour involves lying part of the trunk on the ground, rather than plucking the grass with the trunk tip alone. Images: Darren Naish.

The zoo has had its fair share of elephant-related controversy, most notably in 2000 when a keeper was found guilty of seriously abusing the female elephant Rose-Tu. The fallout resulted in the Rose-Tu law of 2001, which basically strengthened the power of the law to prosecute animal abusers in the state. On keeping elephants in general… naturally, no zoo can really give elephants sufficient space relative to what they have when living wild and the zoo has been in the news on several occasions due to protests about its keeping of elephants. There is, of course, a lot that could be said on the ethics of zoos and this isn’t the place. As someone invested in conservation and efforts to help animals persist into the future, I will say, however, that zoos are necessary, in cases essential, whatever our personal feelings on life-time captivity for their occupants.

**Caption:** the zoo’s Elephant Lands is really big on information panels and displays that provide information on elephant anatomy, biology and lifestyle, and there are impressive features on feet, tusks, and on the history of Oregon Zoo’s elephants specifically. Image: Darren Naish.

  **Caption:** view showing part of the outdoor arena of Elephant Lands. Oregon is cool or cold for part of the year and at or close to freezing in the winter. Elephants are very resistant to low temperatures but modern zoos of course provide them with heated buildings. Image: Darren Naish.

Africa Zone. We next moved on to the south-west quadrant, the Africa zone. Outdoor enclosures feature Giraffe Giraffa camelopardalis (both Masai G. c. tippelskirchi and Reticulated G. c. reticulata), Black rhino Diceros bicornis, Cape hunting dog Lycaon pictus (the zoo uses ‘Painted dog’), African lion Panthera leo, Mountain or Eastern bongo Tragelaphus euryceros isaaci and Bontebok Damaliscus pygargus. Bontebok are a big deal in view of the conservation history of this taxon*, (in)famously reduced to a relict population of less than 20 South African individuals.

You will of course be aware of the argument that these should be recognised as species, not subspecies.

The conventional view that the Bontebok is conspecific with the Blesbok D. phillipsi would mean that it was an endangered subspecies, not a species, but views differ on what to do with the taxonomy of these antelope (Castelló 2016).

**Caption:** perhaps because of the weather, the Cape hunting dog group were all huddled together in their rock shelter during my visit. At right, cheetah on the move. As usual for zoos, this is a cheetah of east African origin. The far rarer Asiatic cheetah is scarcely not kept in captivity at all; sources disagree, but there are something like five or six in captivity, I think all of them in Iran. Images: Darren Naish.

  **Caption:** the Cape hunting dog enclosure with rock shelter (and resting dog pile) very obvious. Over-use of the same areas of ground is evidenced by the heavily worn, de-grassed areas. Image: Darren Naish.

  **Caption:** these images give some idea of how close you can get to the lions thanks to the glass partition that allows viewing inside their rest area. I think that the male here is Zawadi, who has only just (April 2026) died. He was 18, making him another unusually old animal kept at the zoo. Many zoos here in the UK now have Asiatic, rather than African, lions, so seeing African animals these days often feels special. Images: Darren Naish.

  **Caption:** I think the photos here show two different Bontebok individuals. The animal on the right has its ears in an odd position because this photo was taken just before it shook its head (due to light rain). Images: Darren Naish.

Both Greater flamingo Phoenicopterus roseus and Lesser flamingo Phoeniconaias minor shared an enclosure with Fulvous whistling duck Dendrocynga bicolor, White-faced whistling duck D. viduata, African spoonbill Platalea alba and Gadwall Mareca strepera. Other animals I recall seeing here include Common dwarf mongoose Helogale parvula and Red-tailed monkey Cercopithecus ascanius schmidti. A number of heated buildings featured Red-billed hornbill Tockus erythrorhynchus and Blue-bellied roller Coracias cyanogaster as well as Baja blue rock lizard Petrosaurus thalassinus and Philippine sailfin lizard Hydrosaurus pustulatus (both very much not at home in an African section), African plated lizard Gerrhosaurus validus, Malagasy Henkel’s leaf-tailed gecko Uroplatus henkeli and more.

**Caption:** at left, a Greater flamingo photobombs my whistling duck photo, both White-faced and Fulvous species visible here. At right, two different Gadwall, I think a male (top) and female. These photos are quite washed out and the browns on these birds should be brighter and richer. Images: Darren Naish.

  **Caption:** flamingo and duck exhibit, with Lesser flamingo at left and two Greaters at right. Whistling ducks, a spoonbill and gadwalls are visible too. Image: Darren Naish.

  **Caption:** a bird montage with bonus Henkel’s leaf-tailed gecko. Clockwise from upper left: Golden-breasted starling *Cosmopsarus regius*, Northern red-billed hornbill *Tockus erythrorhynchus*, Grey gull (from the penguin rotunda discussed above), and gecko seen from underneath while it was clinging to the side wall of its terrarium. Images: Darren Naish.

  **Caption:** two big, impressive squamates housed in the Africa section, even though they’re very much not from that part of the world. Sailfin lizards like that at left are from Indonesia and the Philippines; Crocodile monitors *Varanus salvadorii* are from New Guinea. The Crocodile monitor is one of the world’s rarest monitors, but these days it’s one of the species most often encountered in captivity, due predominantly to a successful captive breeding programme. Images: Darren Naish.

The large Aviary Understory (it should be ‘Understorey’), another rotunda, featured a large area of tropical plants and free-flying Crested coua Coua cristata, Golden-breasted starling, Purple glossy starling Lamprotornis purpureus, Emerald starling L. iris and much else. The birds I most wanted to see – Maccoa duck Oxyura maccoa, Allen’s gallinule Porphyrio alleni and Magpie shrike Corvinella melanoleuca – were a no-show for me, alas.

**Caption:** from left to right, Blue-bellied roller, Crested coua and the Afro-Arabian Spur-winged lapwing or Spur-winged plover *Vanellus spinosus*. The Spur-winged plover is the species apocryphally claimed to clean the teeth of crocodiles, though this role is also associated today with the Egyptian plover *Pluvianus aegyptius*‍ ‍(a bird that isn’t a plover, despite its name). Images: Darren Naish.

  **Caption:** Hadada ibis at left, a large ibis widely distributed across Africa and named for its distinctive call. It’s been introduced to peninsula Malaysia, which is weird. All *Bostrychia* ibises are odd, often being comparatively short-billed and with unusual ornamentation. At right, African spoonbill. Images: Darren Naish.

I really enjoyed seeing Hadada ibis Bostrychia hagedash, a splendid bird with a deep base to its bill and striking eyes where the pupil has a slightly oval form (the horizontal axis being the longest one). If you think of ibises as a fairly samey group of birds, it’s well worth checking out the diversity they include, since there’s a lot going on there. I don’t think I’ve ever written about them at length.

**Caption:** Southern ground hornbill *Buceros leadbeateri* at left, Speke’s gazelle *Gazella spekei* of Somalia at right. The hornbill isn’t nesting (like all hornbills, it’s a cavity-nester). Speke’s is one of the most unusual of gazelle, what with its inflatable nose, but is today among those species most often seen in captivity. Image: Darren Naish.

Another indoor display area was devoted to Naked mole-rat Heterocephalus glaber (though read on). As is standard in displays of this species, they were housed in a human-made system of tunnels and galleries and could be viewed through small windows. I like Naked mole-rats and enjoy looking at them, but it’s somehow ironic that the most specialized and unusual member of this group – there are around 30 mole-rat species – is essentially the only one you ever seen in captivity. I guess that both weirdness and fame account for this. You might argue that their sociality and use of complex burrow systems recalling those of social insects are also plus points for display in captivity, though this would work for at least some other, non-naked mole-rat species. Incidentally, some workers argue that H. glaber of traditional is a species complex and consists of taxa that started diverging during the Late Miocene (Uhrová et al. 2026). It might be a while before we start seeing this proposed taxonomy – if it proves valid – reflected in zoos, I think.

**Caption:** naked mole-rats are fun to watch in their tunnels, but they never actually do that much. Fossils show that these animals have been around since the Miocene, with molecular data indicating divergence from other mole-rats even earlier than this. I wrote about mole-rat evolution **back here in 2016** and I’m pleased to see that the article is intact, its images still in place! Image: Darren Naish.

Tortoises, Spotted-necked otter Hydrictis maculicollis, fruit bats (Rodrigues fruit bat Pteropus rodricensis, Egyptian fruit bat Rousettus aegyptiacus and Straw-coloured fruit bat Eidolon helvum) and lemurs were in this part of the zoo too, and also indoors was a Slender-snouted crocodile Mecistops cataphractops. If you’re a crocodylian fan, this is a major tick. Several collections in the UK house Mecistops, so I have seen them before. Then again, there’s more than one species, so not all Mecistops are alike. Spotted-necked otter is rarity but wasn’t showing, perhaps because of the rain. A big African lungfish Protopterus annectans lived in a landscaped enclosure near the crocodile.

**Caption:** the glass-fronted crocodile enclosure allows you to look at the *Mecistops* while it’s submerged, which is great. It seems odd today to think that *Mecistops* was included within *Crocodylus* just a few decades ago. Molecular data published since around 2006 has shown that the old genus name published for it by J. E. Gray back in 1844 should be reinstated, and that it might be an osteolaemine. Some anatomical data even supports a close relationship with the extinct *Euthecodon*. Image: Darren Naish.

  **Caption:** *Mecistops* in profile. The steepness of the forehead is interesting, as is the relative shortness of the skull deck. The nictitating membrane is sufficiently transparent (albeit not fully) that it clearly allows good underwater vision, though I wonder if submerged crocodylians see the world through a slightly milky lens. Image: Darren Naish.

  **Caption:** African lungfish co-operatively showing at the front of its enclosure. There are four extant species of *Protopterus* and this is specifically *P. annectans*, the so-called West African lungfish. It does live in the west, but in the east as well. It’s large, reaching 1 m in length. Image: Darren Naish.

The lions were busy reclining, albeit awake, but the Black rhinos Diceros bicornis were extremely active. The two individuals we watched had been housed together in the hope that they might mate. Courtship in black rhinos seems to involve the male pursuing, harassing and mounting the female until she eventually gives in. Nature is the way it is and many ‘natural’ behaviours are unfair and involve pain and distress; also, the world needs more baby rhinos. But you have to feel sorry for a female when, as here, there isn’t the opportunity to ever get away.

**Caption:** black rhino pair interacting. There are supposed to be at least five black rhino subspecies, two of which are recently extinct. For some animals, we always get to learn which subspecies we’re looking at; for others – and the black rhino is an example – we very often don’t. I had assumed that most black rhinos in zoos are of the south-central subspecies *D. b. minor* but these individuals are the critically endangered Eastern black rhino *D. b. michaeli* (which was only awarded taxonomic distinction in 1965). Image: Darren Naish.

  **Caption:** this sort of behaviour is all part of courtship, though the male here has a sheepish and embarrassed look and seems to know that he’s doing wrong. The male rhino is called King (he was born at Lincoln Park Zoo, Chicago, in 2013); the female is Jozi (she was born at Pittsburgh Zoo in 2012). Images: Darren Naish.

Overall, that’s a pretty impressive selection of African (and Malagasy) species, involving zoo classics as well as species only rarely seen.

**Caption:** Siberian/Amur tiger enclosure, located in the Discovery Zone and hence in the far west of the zoo. You should be able to see the lone tiger close to the centre of the image. Two were present at the time of my 2022 visit: Eloise and Bernadette, both originally from Milwaukee County Zoo in Wisconsin. The zoo has kept several Siberian tigers in recent years and one of them, Mikhail, was the oldest captive member of his species. On his death in 2018 he was one day short of his 20th birthday. Image: Darren Naish.

Finally… the Discovery Zone includes an Insect Zoo as well as Siberian/Amur tiger Panthera tigris altaica (the zoo goes with ‘Amur’). And that about sums things up. I was really impressed both with the landscaping and general look of the zoo and also at its selection of animals, and also with its really well-designed and informative signage and art. I’ve said before that you can judge a zoo from the amount and quality of information it provides its visitors, and Oregon Zoo is very, very strong on this front. A number of animal statues throughout the zoo improved the surrounds and were appreciated as well.

**Caption:** I really enjoyed the landscaping and sculpted features of the zoo, the rocks here (from the Great Northwest section) being a good example. At right, one of at least two pika statues findable within the same section. Images: Darren Naish.

  **Caption:** more very nice metal statues from the Great Northwest section. A skunk at left, Rocky Mountain goat at right. Images: Darren Naish.

As ever, I’ll end with my wholly idiosyncratic scoring system…

Selection of species: 9 out of 10
Zoo nerd highlights: Rocky mountain goat, California condor, Slender-snouted crocodile, Sea otter, Spotted-necked otter
Quality of signage: 10 out of 10
Value for money: 9 out of 10
Overall worthiness: 9 out of 10

For previous articles in my zoo reviews series, see…

Tet Zoo Reviews Zoos: Edinburgh Zoo, August 2016
Tet Zoo Reviews Zoos: Colchester Zoo, October 2017
Tet Zoo Reviews Zoos: the Isle of Wight Zoo, February 2020
Tet Zoo Reviews Zoos: ZSL Whipsnade Zoo, June 2021
Tet Zoo Reviews Zoos: Bristol Zoo, July 2021
Tet Zoo Reviews Zoos: Tripoli Zoo in Libya, January 2023
Tet Zoo Reviews Zoos: New Forest Wildlife Park, April 2024
Tet Zoo Reviews Zoos: Tokyo’s Ueno Zoo, September 2024
Tet Zoo Reviews Zoos: Tama Zoological Park, Tokyo, February 2025

My technical research and my writing here at the blog continues with your kind support via patreon. Many thanks to those who assist my projects. Please consider assisting if you can.

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Castelló, J. R. 2016. Bovids of the World: Antelopes, Gazelles, Cattle, Goats, Sheep, and Relatives. Princeton University Press, Princeton and Oxford.

Frost, D. R., Grant, T., Faivovich, J., Bain, R. H., Haas, A., Haddad, C. F. B., De Sá, R. O., Channing, A., Wilkinson, M., Donnellan, S. C., Raxworthy, C. J., Campbell, J. A., Blotto, B. L., Moler, P., Drewes, R. C., Nussbaum, R. A., Lynch, J. D., Green, D. M. & Wheeler, W. C. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297, 1-370.

Uhrová, M., Mikula, O., Bryja, J., Frýdlová, P., Zemlemerova, E. D., Elmi, H. S. A., Štolhoferová, I., Tymlová, V., Maštera, V., Frynta, D., Lavrenchenko, L. A. & Šumbera, R. 2026. More than one species of the naked mole-rat, a new biomedical model. Communications Biology 9, 70.

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Inspired by the recent publication here of thoughts on the new Koumpiodontosuchus paper… and by other work, in prep… I felt it appropriate to rescue another article from the archives, specifically from ver 3. Here we go…

**Caption:** a Shanklin croc montage, showing the actual fossil at upper right. At left is a not entirely serious reconstruction by Vladimir Dinets (colourised by Darren Naish).

The article concerned was published in 2014 (it’s here at wayback machine) and revolves around the publication of a technical paper co-authored by myself and colleagues, led by marine crocodylomorph guru Mark Young (Young et al. 2014).

**Caption:** the Shanklin croc specimen (NHMUK PV OR36173) in (at top) right lateral, (at middle) ventral, and (at bottom) dorsal view. Images from Young *et al*. (2014). It's not the prettiest fossil in the world, but it's still interesting.

This paper describes another of those frustrating fossils that combines anatomical novelty and possible phylogenetic significance with a most unsatisfactory degree of completeness. The fossil concerned is an ugly chunk of bone (going by the catchy accession number NHMUK PV OR36173), about 13 cm long, that represents the anterior part of the incomplete right dentary bone of a reasonably large, long-snouted Cretaceous crocodylomorph. It comes from Shanklin on the south-east coast of the Isle of Wight, England. While my innovative co-author Lorna Steel proposed the nickname Shanklin Shocker for the creature, I’ll simply refer to it from hereon as the ‘Shanklin croc’.

**Caption:** Richard Lydekker (1849-1915): biogeographer, anthropologist, zoologist, palaeontologist, here photographed in or around 1900. Image: (c) but allowed under fair use. **Original here.**

History and provenance. The specimen isn’t completely new to the literature. We know that it was purchased by the then British Museum (Natural History) (now the Natural History Museum, or NHM) in 1861 from a Mr Simmons. Palaeontologist, zoologist and famed ‘lightning cataloguer’ Richard Lydekker then wrote about it in 1889 but (mis)identified it as the premaxillary bone of the plesiosaur Polyptychodon. Fast forward about a century, and Leslie Noé – known to marine reptile workers for his (still predominantly unpublished) PhD work on pliosaurids – left a note in the NHM collections saying that it isn’t a plesiosaur at all, but more likely a crocodylomorph of some sort.

It’s this that brought it to the attention of croc-worker Lorna Steel; Lorna has been working for some years with Mark Young on new (and/or previously overlooked or under-appreciated) crocodylomorphs. Mark and Lorna then assembled a team of people who work on relevant crocodylomorph groups. I became involved because we needed to determine whether the specimen was anything to do with the goniopholidids, a group of long-snouted Jurassic and Cretaceous crocodyliform crocodylomorphs well represented in the fossil record of the Isle of Wight. Regular readers will perhaps recall that Steve Salisbury and I reviewed the Wealden crocodyliforms back in 2011 and therein named or renamed several new species (Salisbury & Naish 2011). This isn’t the first time crocodylomorph remains have been identified as those of a plesiosaur, by the way (e.g., Buchy 2008).

**Caption:** locational and stratigraphic data on the Shanklin croc. It comes from the south-east coast of the Isle of Wight (C shows Luccombe Chine, D shows Knock Cliff), though exactly which Cretaceous stratum it came from is uncertain. Knock Cliff has also yielded **a fragmentary theropod that I’ve worked on**, namely the holotype of *Vectaerovenator inopinatus* Barker *et al*., 2020. This figure is from the paper (Young *et al*. 2014).

The precise provenance of the Shanklin croc is somewhat uncertain and there are several different sedimentary strata it could have come from. Based on the fossil’s colour, it seems most likely that it came from the ‘Malm rock’ of the Upper Greensand Formation, a geological unit deposited during the late Albian during the Early Cretaceous. A few other units that crop out in the same area and are Aptian or Albian in age are also possible sources for the specimen, meaning that we’ve given its age as ‘?Aptian-Albian’ (Young et al. 2014). Upper Albian seems most likely.

Incidentally, if you’re confused as to why I’m seemingly inconsistent in my use of ‘lower’/‘upper’ and ‘early’/‘late’, it’s because ‘lower’ and ‘upper’ refer to the position of rock layers while ‘early' and ‘late’ refer to geological time. So, the Shanklin croc lived during the late Albian of the Early Cretaceous, but its fossil was found in the upper Albian of the Lower Cretaceous.

What sort of crocodylomorph might the Shanklin croc be? Could it belong to any of the groups already reported from the Isle of Wight Cretaceous? These are atoposaurids, goniopholidids, bernisartiids, and hylaeochampsids. Short answer: no. Long answer: read the paper [contact any of the authors if you want a pdf]. Intermediate-length answer: as revealed by comparisons made between the Shanklin croc and the dentaries of those other groups, none possess the specific alveolar configuration or mandible shape that the Shanklin croc does and all possess anatomical peculiarities not present in the Shanklin croc (Young et al. 2014). We could also exclude several other Cretaceous crocodylomorph groups, including gavialoids, teleosaurids, metriorhynchids and the several lineages often grouped together in Pholidosauridae (Young et al. 2014). So... what is it?

**Caption:** the Shanklin croc specimen (A) compared with the anterior dentaries of assorted other crocodylomorphs, including thalattosuchians, dyrosaurids, goniopholidids, hylaeochampsids and gavialoids. Diagram from Young *et al*. (2014).

Based on the presence of enlarged anterior alveoli, the shape of the bone’s outer margin, the large and widely spaced foramina on the lateral and ventral surfaces of the bone and other features, it seems most similar to species included within Dyrosauridae, a group of long-snouted crocodyliforms of the Cretaceous and Paleogene. However, it remains unusual enough compared to undoubted dyrosaurids that we have some reservations about referring it to Dyrosauridae outright. Our conservative conclusion is that the Shanklin croc should be identified as Tethysuchia incertae sedis, Tethysuchia being the crocodyliform clade that includes dyrosaurids, pholidosaurids and allied taxa.

Meet the tethysuchians. Tethysuchians haven’t been much (if at all) discussed on Tet Zoo before, so this is a good time to say some things about them. Experts have long recognised several Jurassic, Cretaceous and Paleogene crocodylomorph lineages that all look roughly alike in being long-jawed, strongly aquatic predators that have a substantially ‘archaic’ osteoderm configuration compared to living crocodylomorphs (the crocodylians).

**Caption:** the three crocodylomorph groups originally assembled in the 'longirostrine clade' by Clark (in Benton & Clark 1988). Thalattosuchia includes the two main groups Teleosauroidea and Metriorhynchoidea. Images: Darren Naish.

At the core of the assemblage are the dyrosaurids of the latest Cretaceous and Paleogene and the pholidosaurids of the Early and Late Cretaceous. Benton & Clark (1988) and Clark (1994) grouped these together with the sea-going thalattosuchians into a ‘longirostrine clade’ (they didn’t give it an official name). Goniopholidids have also been allied with the ‘longirostrine clade’ in some studies (Andrade et al. 2012). Brand-new at the time of writing is Jouve et al.’s (2026) proposal that the name Coelognathosuchia be used for the goniopholidid + tethysuchian clade, and Stenorhynchosuchia for the thalattosuchian + tethysuchian clade. These names are mutually exclusive: Coelognathosuchia can only be used when goniopholidids are close to tethysuchians (it cannot include thalattosuchians), and Stenorhynchosuchia can only be used when thalattosuchians are close to tethysuchians (it cannot include goniopholidids) (Jouve et al. 2026).

**Caption:** a highly simplified version of a phylogenetic hypothesis supported in several studies, where thalattosuchians are well away from crown-crocs (Crocodylia) and are not close to tethysuchians. This tree also shows goniopholidids close to Tethysuchians, in which case we could use the name Coelognathosuchia for the relevant clade. That name was first published by **Martin *et al*. (2014)** and recently given a phylogenetic definition by Jouve *et al*. (2026). Image by Darren Naish.

On that note, do all of these longirostrine crocodylomorphs really go together? Well, this is controversial. If thalattosuchians really are close relatives of dyrosaurids and pholidosaurids, this makes thalattosuchians ‘advanced’ crocodyliforms, not all that far in phylogenetic distance from crown-crocs. But other studies don’t support a close relationship between thalattosuchians, goniopholidids and the other longirostrine groups, instead finding thalattosuchians far, far away from the crown (Pol & Gasparini 2009, Sereno & Larsson 2009, Young et al. 2012, Montefeltro et al. 2013), and perhaps not even in Neosuchia at all. This story was summarised a while back at Tet Zoo [UPDATE: in another article now only findable at wayback machine]. Most character evidence puts goniopholidids closer to crown-crocs than are dyrosaurids and pholidosaurids (e.g., Salisbury et al. 2006, Montefeltro et al. 2013), but not all studies find this. Anyway, the longirostrine ‘core’ consists only of dyrosaurids, pholidosaurids and close kin.

Buffetaut (1982) was first to make the case for Tethysuchia and coined that name. Its use is widespread in crocodylomorph studies today. Pholidosaurids have sometimes been found to represent a paraphyletic series of outgroups to Dyrosauridae but more recent work has supported their monophyly.

**Caption:** skeletal reconstruction of *Dyrosaurus* to scale with a person. Note the long supratemporal fenestrae and tall neural spines in the shoulder region. The very long, strongly curved retroarticular process (the curving structure at the posterior end of the lower jaw) is interesting too. Image: LiterallyMiguel, CC BY 4.0 (**original here**).

Thoughts on dyrosaurids. Dyrosaurids haven’t really had their monophyly doubted and are now a large and complex group. They’re predominantly associated with Africa (and likely originated there) but species also inhabited southern Asia and both North and South America. They were generally slender-snouted animals that grabbed fish but some (like giant Phosphatosaurus from the Eocene of Tunisia) have robust jaws and blunt teeth and look capable of handling robust prey items. Others (like Cerrejonisuchus from the Paleocene of Colombia) are especially robust-snouted and yet others (like Anthracosuchus, also from the Paleocene of Colombia) are comically short-snouted.

Dyrosaurid skulls are reasonably easy to recognise. The eye sockets are dorsally placed and the supratemporal openings are about twice as long as they are wide. Dyrosaurids represented by good postcranial remains reveal unusual proportions relative to those of crocodylians. The forelimbs are long, slender and sometimes longer than the hindlimbs, tall neural spines on the dorsal vertebrae must have created a deep chest and somewhat hump-backed appearance and the tall is especially deep and narrow.

Schwarz-Wings et al. (2009) reconstructed dyrosaurid musculature and reported evidence for unusually large and strong limb, body and tail musculature. They suggested that dyrosaurids below a certain body size might have been especially strong walkers, that their ability to generate thrust by the tail exceeded that of crocodylians, and that their powerful body and tail allowed them to better move and forage in strong currents. These features could conceivably have been specialisations for life in tidal, coastal habitats. Not all dyrosaurids are alike though: the relatively small, short-snouted Cerrejonisuchus has features suggestive of a semi-terrestrial way of life (Scavezzoni & Fischer 2021).

**Caption:** as is the case for so many extinct crocodylomorph groups, dyrosaurids are actually not that similar to crocodylians when we really pay attention to their proportions and various traits. This graph, from **Scavezzoni & Fischer (2021)**, shows how dyrosaurids (red) differ notably from crocodylians (yellow) in anatomical proportions, as do thalattosuchians (green). Image: **Scavezzoni & Fischer (2021)**, CC BY 4.0 (**original here**).

What, then, to make of the Shanklin croc? Our identification of the Shanklin croc as some sort of dyrosaurid-like tethysuchian makes it especially interesting, because it’s geologically older than expected based on what we know about tethysuchian history. Dyrosaurids are well known from the Maastrichtian onwards but pre-Maastrichtian specimens – some have been reported from the Cenomanian and Campanian (Buffetaut et al. 1990, Churcher & Russell 1992, Churcher 1995) – are fragmentary. Furthermore, the skull fragments concerned resemble those of some pholidosaurids and thus might not be from dyrosaurids after all; Buffetaut et al. (1990) did report some vertebrae that look very dyrosaurid-like, however. We conclude for now that there are no definite records of dyrosaurids in the ‘middle’ Cretaceous, the implication from the fragmentary remains reported so far – the Shanklin croc included – being that several ‘proto-dyrosaurid’-type taxa might have been in existence from the Aptian/Albian onwards, but that better remains are needed before we can say anything more.

As for which specific tethysuchian taxon the Shanklin croc represents, it’s something new: a new species and genus that needs a name. You might argue that we should have named it in the paper (part of me thinks that we should have). However, people are generally discouraged from attaching names to fragmentary specimens such as this. As ever, we hope that newer and substantially better material will turn up in time and that better remains from this part of the Cretaceous will help us pin down the origins and biogeographical history of one of the most important of marine crocodylomorph radiations.

**Caption:** stratigraphic ranges of relevant Lower Cretaceous crocodylomorph lineages, plotted against time and compared with the possible age range of the Shanklin croc. Dotted lines represent parts of lineages for which fossils are unreported; arrows show that the lineage concerned persisted beyond the end of the Turonian. Diagram from Young *et al*. (2014).

Crocodiles Attack Elephants Then, Now, and Still, August 2025
Koumpi the Wealden Croc: Tons of New Data and Phylogeny Resolved, April 2026

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

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Andrade, M. B., Edmonds, R., Benton, M. J. & Schouten, R. 2012. A new Berriasian species of Goniopholis (Mesoeucrocodylia, Neosuchia) from England, and a review of the genus. Zoological Journal of the Linnean Society 163, S66–S108.

Buchy, M.-C. 2008. Reevaluation of the holotype of Plesiosaurus (Polyptychodon) mexicanus Wieland, 1910 from the ?Upper Jurassic of Mexico: a thalattosuchian, not a sauropterygian. Revista Mexicana de Ciencias Geológicas 25, 517-522.

Buffetaut, E. 1982. Radiation évolutive, paléoécologie et biogéographie des crocodiliens mésosuchiens. Mémoires de la Société Géologique de France 60, 1-85.

Buffetaut, E., Bussert, R. & Brinkmann, W. 1990. A new nonmarine vertebrate fauna in the Upper Cretaceous of northern Sudan. Berliner Geowissenschaftliche Abhandlungen A 120, 183-202.

Churcher, C. S. 1995. Giant Cretaceous lungfish Neoceratodus tuberculatus from a deltaic environment in the Quseir (=Baris) Formation of Kharga Oasis, Western Desert of Egypt. Journal of Vertebrate Paleontology 15, 845-849.

Churcher, C. S. & Russell, D. A. 1992. Terrestrial vertebrates from Campanian strata in Wadi el-Gedid (Kharga and Dakhleh Oases), Western Desert of Egypt. Journal of Vertebrate Paleontology 12 (Supplement), 23A.

Clark, J. M. 1994. Patterns of evolution in Mesozoic Crocodyliformes. In Fraser, N. C. & Sues, H.-D. (eds) In the Shadow of the Dinosaurs – Early Mesozoic Tetrapods. Cambridge University Press (Cambridge, NY, Melbourne), pp. 84-97.

Jouve, S., Young, M. T., Hastings, A. K. & Salih, K. 2026. The systematics and nomenclature of Tethysuchia (Archosauria: Crocodylomorpha) under the International Code of Phylogenetic Nomenclature. Zoological Journal of the Linnean Society 206, zlag045.

Martin, J. E., Lauprasert, K., Buffetaut, E., Liard, R. & Suteethorn, V. 2013. A large pholidosaurid in the Phu Kradung Formation of north-eastern Thailand. Palaeontology 57, 757-769.

Montefeltro, F. C., Larsson, H. C. E., de França, M. A. G. & Langer, M. C. 2013. A new neosuchian with Asian affinities from the Jurassic of northeastern Brazil. Naturwissenschaften 100, 835–841.

Pol, D. & Gasparini, Z. 2009. Skull anatomy of Dakosaurus andinensis (Thalattosuchia: Crocodylomorpha) and the phylogenetic position of Thalattosuchia. Journal of Systematic Palaeontology 7, 163-197.

Sereno, P. C. & Larsson, H. C. E. 2009. Cretaceous Crocodyliforms from the Sahara. ZooKeys 28, 1-143. doi:10.3897/zookeys.28.325

Salisbury, S. W., Molnar, R. E., Frey, E. & Willis, P. M. A. 2006. The origin of modern crocodyliforms: new evidence from the Cretaceous of Australia. Proceedings of the Royal Society of London B 273, 2439-2448.

Salisbury, S. W. & Naish, D. 2011. Crocodilians. In Batten, D. J. (eds). English Wealden Fossils. The Palaeontological Association (London). pp. 305-369.

Scavezzoni, I. & Fischer, V. 2021. The postcranial skeleton of Cerrejonisuchus improcerus (Crocodyliformes: Dyrosauridae) and the unusual anatomy of dyrosaurids. PeerJ 9, e11222.

Schwarz-Wings, D., Eberhard Frey, E. & Martin, T. 2009. Reconstruction of the bracing system of the trunk and tail in hyposaurine dyrosaurids (Crocodylomorpha; Mesoeucrocodylia). Journal of Vertebrate Paleontology 29, 453-472.

Young, M. T., Brusatte, S. L., de Andrade, M. B., Desojo, J. B., Beatty, B. L., Steel, L., Fernández, M. S., Sakamoto, M., Ruiz-Omeñaca, J. I. & Schoch, R. R. 2012. The cranial osteology and feeding ecology of the metriorhynchid crocodylomorph genera Dakosaurus and Plesiosuchus from the Late Jurassic of Europe. PLoS ONE 7(9): e44985. doi:10.1371/journal.pone.0044985

Young, M. T., Steel, L., Foffa, D., Price, T., Naish, D. & Tennant, J. P. 2014. Marine tethysuchian crocodyliform from the ?Aptian-Albian (Lower Cretaceous) of the Isle of Wight, UK. Biological Journal of the Linnean Society 113, 854-871.

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Yes, the time is right to discuss the possibility of an expanded, ten-year-anniversary edition of my 2016/2017 Arcturus book Hunting Monsters…

**Prologue.** *Hunting Monsters*, for those who don’t know it, mostly focuses on the issue of *how* modern views on specific cryptids were compiled *and* how the foundational evidence fares today (or, at least, when I wrote the text in the 2010s). It’s sceptical and advocates critical appraisal, both of the aforementioned evidence and its mostly sensationalist proponents. The strongest criticism of the book is that it doesn’t provide affirmation for those who want something more satisfying in emotional terms. You can decide whether that’s a valid thing to say about a work of non-fiction.

**Caption:** 2016 promotion from Arcturus Publishing for the first edition of the book.

One more bit of preamble: there’s a popular view (both among proponents of monster existence and society at large) that people with a sceptical approach to subjects like cryptozoology are hateful, joyless bourgeoisie devoid of ‘real world’ life experience and existing only to stamp on the dreams of the hopeful. This entire narrative is bullshit. The majority of sceptics love the subjects they comment on and are deeply invested in them; they’ve come to reject the base-tier, literal position of ‘believers’ for good reason. But, like the keenest of YouTubers and Nessie bloggers, we sceptics would, actually, really like it if Bigfoot and Nessie were real. Many of us were once ‘believers’ too.

**Caption:** at left, the 2016 ebook cover (which I think is rad as hell). At right… there was a time when I had a huge store of hardcopies, but alas. These were mostly sold at TetZooCon 2017. A screengrab of my favourite amazon review is featured. Images: Darren Naish.

Incidentally, there are two publication dates for Hunting Monsters because it appeared as an ebook in 2016 (Naish 2016) and in hardcopy the following year (Naish 2017). These two versions are slightly different and hence count as different editions.

What to change? On social media (mostly BlueSky), I've been compiling a list of the specific changes I'd want to include should a new, third edition get the green light. It hasn’t, despite efforts, so this is all hypothetical for now. I have five motivations to see such a version published...

There are assorted typos and small errors of logic I’d like to see corrected.
If my maths works out, Hunting Monsters is ten years old. Over its lifetime it’s received a fair amount of comment, both from those who agree with its arguments and those who don’t. It has been cited a lot, has been discussed in books and articles on the subject, has been critiqued by detractors and supporters, and is seen as a notable contribution to the field. I’m especially pleased with the discussions devoted to it in Binns (2017) on the Loch Ness Monster, Hoyland (2018) on the yeti and Bille (2022) in his overview of cryptozoology-themed literature. In 2019, Fortean Times devoted an entire article to the book, which is quite the accolade I can tell you (The Hierophant’s Apprentice 2019). It’s also been good to see it fairly cited in Mullis (2019), Greenfield (2023), Witton & Hing (2024), Shine (2024), Paxton (2025) and Lewis & Bartlett (2026).

Among articles that provide a less supportive view of its value is Henry Bauer and Roland Watson’s ‘Failings of Nessie debunkers and of debunkers in general’ (Bauer & Watson 2024); this work functions as part of the cryptozoological apologetics already typical of these authors and doesn’t fare well if you reject (as I do) all the cornerstones of their ‘Case for Nessies’. Watson also published very negative reviews of the book (actually, its Nessie chapter specifically) at amazon and at his blog, and I find his view so naïve and biased that I published this article in response. The late Jeff Meldrum was also not a fan – this time for reasons relating to Bigfoot rather than Nessie of course – but I can’t see that he ever published a review. If a 'ten year anniversary' version were to appear, I would want it to include a section that discusses this reception: an introspective.

**Caption:** the published version of *Hunting Monsters* includes 20 images, among which are those shown here. But the original plan was to use about three times as many. Images: Darren Naish.

Several topics in Hunting Monsters are covered in very cursory fashion (due to word count) and I would really like the opportunity to expand on them. In the section on sea monsters, for example, I glossed over the debate on what relevance the 1930s discovery of the supposedly extinct coelacanths has for the claimed persistence of animals like plesiosaurs... it would be appropriate to add new material that alleviates this and other shortcomings. Ideally, I would want a new edition to be expanded relative to the first.
On that note, any opportunity to include more pictures would be great. I would supply these; there isn't anything that would need to be bought from picture agencies.

Caption: my Hunting Monsters thoughts on the 1962 Pensacola incident – always one of my favourite sea monster stories – have proved mistaken, if, that is, you buy the interpretation provided by David Goudsward in his 2020 Sun, Sand, and Sea Serpents and his 2020 article in Fortean Times. Thus update is required.
Finally, several specific monster cases I discuss have been re-evaluated since I wrote about them, meaning that we have new interpretations that I’d like to incorporate and discuss. I’d prefer not to share all those details here because… spoilers but I have already outed the bulk of them on social media. They include the seminal 1848 Daedalus, 1893 Umfuli and 1905 Valhalla sea serpent accounts and the 1962 Pensacola sea serpent attack story. I also want to push back against the ‘King Kong’ hypothesis regarding the Spicer’s Nessie sighting; in addition, the ‘swan hypothesis’ for Hugh Gray’s Nessie photo needs update. I should add that much new evaluation of alleged cryptid photos is already going into another in-prep book.

Caption: given my tetrapod (and vertebrate) bias, it’s ironic that among my favourite sections of Hunting Monsters are devoted to invertebrates. The section on giant pancake monsters now needs slight expansion….

Caption: we made a mistake in Hunting Monsters when discussing the Hugh Gray Nessie photo and showed the wrong sort of swan. It can’t have been a Mute swan Cygnus olor, as shown here at left, but a Whooper C. cygnus. As you can see from the montage at right, I still think that this is a good interpretation.

Anyway, in an ideal world, I'd like to make enough changes to create a substantially augmented, modified version. Right now, I have to report that nothing is happening and there are no indications that a ten-year-anniversary edition might appear. The printed, 2017 version is now a collector’s item that sells at relatively high price (this is nothing to do with me; I have procured a few copies and do have them for sale), but the ebook version is still on sale and available here (here in the UK).

That’s where I’ll end. Next: more crocodylomorphs!

The text here originally appeared at the Tet Zoo patreon. Support me there and you get to see in-prep and unreleased work, much of it exclusive. You’re also supporting the persistence of this blog and my work and research in general.

For previous relevant articles on cryptozoology, see…

Books on the Loch Ness Monster 1: Ronald Binns’s The Loch Ness Mystery Reloaded, March 2019
Books on the Loch Ness Monster 2: Gareth Williams’s A Monstrous Commotion, March 2019
Usborne’s All About Monsters, April 2019
Sea Monster Sightings and the ‘Plesiosaur Effect’, April 2019
A Review of Robert L. France’s Disentangled: Ethnozoology and Environmental Explanation of the Gloucester Sea Serpent, November 2019
The 1972 Loch Ness Monster Flipper Photos, August 2020
Morgawr and the Mary F Photos, February 2021
A Cryptozoologist’s Bibliography: Matt Bille’s Of Books and Beasts, February 2024
Nessie Point and Counterpoint; Who Owns the ‘Facts’ on the Loch Ness Monster?, November 2024
Yowies and the Marsupial Hominoid Hypothesis – Neil Frost’s Fatfoot: Encounters With A Dooligahl, February 2025
Adrian Shine's A Natural History of Sea Serpents, March 2025

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Bille, M. 2022. Of Books and Beasts: A Cryptozoologist’s Library. Hangar 1 Publishing.

Binns, R. 2017. The Loch Ness Mystery Reloaded. Zoilus Press.

Greenfield, T. 2023. Of megalodons and men: reassessing the ‘modern survival’ of Otodus megalodon. Journal of Scientific Exploration 37, 330-347.

Hoyland, G. 2018. Yeti: An Abominable History. William Collins, London.

Lewis, J. & Bartlett, A. 2026. Bigfooters and Scientific Inquiry. Routledge, Abingdon, UK.

Mullis, J. 2019. Cryptofiction! Science Fiction and the Rise of Cryptozoology. In Caterine, D. & Morehead, J. W. (eds) The Paranormal and Popular Culture: A Postmodern Religious Landscape. Routledge, Abingdon, UK, pp. 240-252.

Naish, D. 2016. Hunting Monsters (ebook). Arcturus, London.

Naish, D. 2017. Hunting Monsters (hardcopy). Arcturus, London.

Paxton, G. M. 2025. Weird things some scientific skeptics say about science. Skeptical Inquirer 49, 54-57.

Shine, A. 2024. A Natural History of Sea Serpents. Whittles Publishing, Caithness.

The Hierophant’s Apprentice. 2019. Building a Fortean library No 47. There ain’t so such animal. Fortean Times 382, 56-57.

Witton, M. P. & Hing, R. A. 2024. Did the horned dinosaur Protoceratops inspire the griffin? Interdisciplinary Science Reviews 49, 363-388.

Hunting Monsters: Cryptozoology and the Reality Behind the Myths

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A very long article on a very small croc…

**Caption:** a *Koumpi* montage, showing images by Mark Witton and from **Barker *et al*. (2026)**.

If things go to plan – and that’s a big if – 2026 will see various announcements here relating to crocodylomorphs, that fantastic group of archosaurian reptiles that includes the crocodylians and their numerous extinct relatives. Today sees the first of those, and it relates to a publication, brand-new as of yesterday, of a super-detailed, extensive study focused on the small Wealden crocodyliform Koumpiodontosuchus aprosdokiti. Koumpi-what?

Yes, an animal that we’ll be calling Koumpi for short. Named in 2015 by Steve Sweetman and colleagues (Sweetman et al. 2015), Koumpi is represented by a small, near-complete skull (about 11 cm long) in addition to an atlas vertebra and ten osteoderms.

**Caption:** *Koumpiodontosuchus*  had its time in the limelight back when it was first published in 2015. The original description (**Sweetman *et al*. 2015**) is open access at *Acta Palaeontologica Polonica*. The reconstruction here, created by Mark Witton and used in **Sweetman *et al*. (2015)**, shows *Koumpi* manipulating a viviparid gastropod, carcharodontosaurian theropods and damselflies in the background. Image: Mark Witton.

When alive, the whole animal would have been about 80 cm long. Skeletal fusions and other feature show that it was near mature, and thus small as an adult. Its polysyllabic name really isn’t all that bad once you know how to break it down: it’s a suchus, and it’s koumpio (button) donto (toothed). Nevertheless, it’s a complex name to be sure, and one that might, just might, have been created as a semi-spiteful act of retaliation in response to the 2011 naming of another Wealden crocodyliform, namely Anteophthalmosuchus Salisbury & Naish, 2011. That name, also, is not so bad once you get to know it.

Yes, Koumpi is from the Wealden – that famous, highly productive Lower Cretaceous succession of sandstones, mudstones and siltstones – and specifically from the Wessex Formation of the Isle of Wight. Many of the Wealden dinosaurs I’ve worked on, including Eotyrannus, Thecocoelurus, Aristosuchus, the spinosaurids Ceratosuchops and Riparovenator and the big sauropod ‘Angloposeidon’, are from the Wessex Formation too and hence would have lived alongside Koumpi. Incidentally, the Koumpi holotype was found at Yaverland on the island’s south-east coast, as of course was the controversial Yaverlandia, a second specimen of which was recently described (Naish & Sweetman 2026). More on Yaverlandia is coming in time.

**Caption:** a simplified representation of Wealden Supergroup stratigraphy and terminology, again. The Wessex Formation, mostly Barremian in age, yields the greatest number of Wealden fossil reptiles, and *Koumpi* is among them. Image: Darren Naish; the *Koumpi* skull is from **Sweetman *et al*. (2015)**.

Koumpi is heterodont, with caniniform anterior teeth and rounded, so-called tribodont, teeth at the rear of its jaws. The latter are suggestive of a diet that involved the crushing of shelled prey. As I keep saying whenever this comes up, such teeth do not indicate specialisation for durophagy (that is, a diet devoted to hard-shelled items), since living reptiles whose dentition includes at least some such teeth are generalists.

**Caption:** the Wealden crocodyliform assemblage includes an impressive number of goniopholidids, a group that is not the focus of interest here. These images show two of the best Wealden goniopholidids, *Hulkepholis willetii* at left, *Anteophthalmosuchus hooleyi* at right. Images from Salisbury & Naish (2011).

An important position in phylogeny. On the terminology in use for this group of animals, I’ll remind you that crocodylian (with a y, not an i) pertains only to the crown-group, Crocodylia. This is part of a much larger clade that includes numerous extinct groups, namely Crocodyliformes. This, in turn, is part of an even more inclusive clade – Crocodylomorpha – that includes lineages substantially different from crocodylians. A very unwise decision to disrupt and sabotage this taxonomy has been rebuffed (Brochu et al. 2009) and for good reason… for god’s sake don’t give up on it now!!

Within Crocodyliformes, crocodylians emerged close to the Jurassic-Cretaceous boundary from within Neosuchia, a group that includes lineages that, while undeniably crocodylian-like, lack the key specialisations of that group. These lineages – the goniopholidids, bernissartiids, susisuchids, hylaeochampsids and others – presage crocodylians in showing how their unusual palatal, vertebral and integumentary traits were assembled. Within Neosuchia, hylaeochampsids and some other groups are allied with crocodylians to form Eusuchia. I know that there are a lot of names to keep track of here, but what can I say… there are a lot of animals out there and the trees that depict their relationships are complex. There’s just no way round it.

**Caption:** a substantially simplified crocodylomorph cladogram, showing the memberships of Crocodylomorpha, Crocodyliformes, Neosuchia and Eusuchia. Bernissartiidae, according to the bulk of studies, is outside of Eusuchia and close to eusuchian ancestry. Image: Darren Naish.

When first described, it was immediately evident that Koumpi had much in common with the famous Bernissartia, another small Wealden neosuchian (albeit known also from several Upper Jurassic European units as well). Bernissartia has long served as a pivotal taxon is discussions of crocodylian origins. Sweetman et al. (2015) confirmed this via phylogenetic analysis, finding Koumpi to be part of Bernissartiidae. That looked pretty solid to me. So it was surprising when Groh et al. (2020, 2022) found that Koumpi might be closer to hylaeochampsids or to paralligatorids, or that Koumpi and certain other Mesozoic neosuchians might be within Eusuchia (Martin et al. 2020, Groh et al. 2020, 2022).

**Caption:** the mounted *Bernissartia fagesii* lectotype specimen on show at the IRSNB (Insitut Royal des Sciences Naturelles de Belgique, Brussels). It’s small, at around 60 cm in length, and is behind glass so hard to photograph without reflections. Crocodyliforms of this sort have a biserial compliment of dorsal osteoderms, not a tetraserial one like that typical of crocodylians (albeit not present in all of them). Image: Darren Naish.

A comprehensive anatomical atlas. In view of the importance of bernissartiids to hypotheses of crocodyliform evolution and the excellent preservation and intact nature of the holotype Koumpi skull (not to mention its availability to our research group), I and a team of colleagues led by Chris Barker and Neil Gostling at the University of Southampton arranged a full microCT scan of the specimen and the production of a detailed description that substantially updates the initial one provided by Sweetman et al. (2015).

**Caption:** images from **Barker *et al*. (2026)**, at left showing our retrodeformed, partly reconstructed model of the *Koumpiodontosuchus* cranium and mandible. At right, images showing blood vessels and nerves discovered during our analysis. Check the paper to see these images at full size. Images: **Barker *et al*. (2026)**.

Our study – it’s open access – has just been published (Barker et al. 2026) and it forms a substantial anatomical atlas of the animal’s known anatomy. The data is so comprehensive and substantial that Koumpi now ranks as one of the most thoroughly described non-crocodylian neosuchians. Our other authors are Ethan Tulloch, Mark Young, Lai-Cheuk Leung, and Kathryn Rankin of the University of Southampton’s MuVIS X-Ray Imaging Centre (Barker et al. 2026).

**Caption:** reconstructed brain, inner ear and associated blood vessels of *Koumpi* from **Barker *et al*. (2026)**. The olfactory lobes are obvious and what might be the optic lobes can be detected too. The overall shape of the brain recalls that of immature crocodylians but we think that this might be paedomorphic trait, and other *Koumpi* brain traits are characteristic of adults. Images: **Barker *et al*. (2026)**.

This new information enabled us to infer the hearing and neurosensory ranges of this animal, all indications being that it was similar in its abilities to extant crocodylians (Barker et al. 2026). This presumably means that there were ecological and behavioural similarities, which is what we’d predict in view of anatomical gestalt. We were also able to reconstruct the brain, the cranial nerves, the cerebral vasculature and the extremely complex paratympanic pneumatic system, this being a convoluted, multi-part arrangement of air-filled cavities and tubes connecting the middle ear and throat cavity (Barker et al. 2026).

**Caption:** more of our excellent anatomical imagery of the *Koumpi* cranial material. At left, the conjoined premaxillae, some of the images showing (in red) the neurovascular canals. At right, the dentaries, showing (in red) the ventral alveolar canal, the tooth roots and more. Those tribodont posterior tooth crowns are especially obvious here. Images: **Barker *et al*. (2026)**.

Phylogeny re-evaluated. We were also able to use our new anatomical data to look anew at the phylogenetic position of Koumpi. We evaluated those proposed affinities with hylaeochampsids and suggested placement within Eusuchia and did not find them supported (Barker et al. 2026). In contrast, we found strong support for the ‘conventional’ placement of Koumpi within Bernissartiidae, and for a placement of Bernissartiidae close to, but not within, Eusuchia (Barker et al. 2026).

Given that we incorporated a substantial amount of new anatomical data pertaining to the anatomy of the skull – over 240 new character states were coded for (Barker et al. 2026, supplementary data) – I’m going to make the argument that our assessment is much more data-rich than previous ones and therefore more likely to be correct. Koumpi is, once again, firmly back among the bernissartiids.

**Caption:** a strict consensus phylogeny resulting from our ‘equal weighing’ analysis, showing poorly resolved relationships among several lineages close to, and within, Neosuchia. The recovery of a clade that includes goniopholidids and tethysuchians (dyrosaurids and kin) is interesting, albeit not novel to our study. Note that *Koumpi* is a bernissartiid, and that bernissartiids are one step outside Eusuchia. Images: **Barker *et al*. (2026)**.

  **Caption:** a phylogeny using extended implied weights, and with a more resolved topology. The goniopholidid + tethysuchian clade is again present. Images: **Barker *et al*. (2026)**.

I haven’t used this article as an opportunity to discuss Wealden crocodyliforms in general, but it’s worth emphasising that Koumpi is but one of several, quite a few of which are also represented by excellent skull information (Salisbury & Naish 2011, Ristevski et al. 2018). Work similar to that done by our team on Koumpi could, one day, be done on those too and there are – as ever – more announcements on these animals yet to come…

Huge thanks and congrats to everyone involved in the Koumpi team, to everyone who assisted with this project at the University of Southampton and Dinosaur Isle Museum (where the fossil is housed), and to our phenomenally helpful and enthusiastic reviewers. The paper is OPEN ACCESS so please do check it out.

**Caption:** a montage of crocodylomorph-themed images, showing things that were published at Tet Zoo ver 2 and 3. There were articles there on thalattosuchians, recently extinct and extant crocodylians, and much more. Images: Tony Pyrakowski; Darren Naish.

Tet Zoo’s lament. A substantial amount of crocodylomorph material exists in the Tet Zoo archives but most of it is now only findable via the wayback machine and I haven’t yet added it to the WIP list, all of which means that I have to start from scratch and that virtually nothing is present here at ver 4, gah. Anyway, for previous articles at ver 4 on crocodylomorphs and Wealden animals in general, see…

Theropod Dinosaurs of the English Wealden, Some Questions (Part 1), March 2020
Dr Angela Milner and the Discovery of Baryonyx, August 2021
Two New Spinosaurid Dinosaurs from the English Cretaceous, August 2021
A Giant Spinosaurid Dinosaur from the Cretaceous of the Isle of Wight, June 2022
The Eotyrannus Monograph Is Finally Published, July 2022
A brain for Baryonyx: using CT-scanning to examine British spinosaurid brains, February 2023
Discovering 'Hidden' Diversity Within Wealden Spinosaurid Dinosaurs, May 2023
A New Theropod Dinosaur Assemblage from the Older Part of the English Wealden, December 2024
Crocodiles Attack Elephants Then, Now, and Still, August 2025

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

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Barker, C. T., Tulloch, E., Young, M. T., Naish, D., Leung, L.-C., Rankin, K. & Gostling, N. J. 2026. Re-evaluation of the Wealden crocodyliform Koumpiodontosuchus Sweetman et al., 2015: new osteological and neuroanatomical data from micro-computed tomography resolves a phylogenetic dispute. Zoological Journal of the Linnean Society 4, zlag035.

Brochu, A. C., Wagner, J. R., Jouve, S., Sumrall, C. D. & Densmore, L. D. 2009. A correction corrected: consensus over the meaning of Crocodylia and why it matters. Systematic Biology 58, 537-543.

Groh, S. S., Upchurch, P., Barrett, P. M. & Day, J. J. 2020. The phylogenetic relationships of neosuchian crocodiles and their implications for the convergent evolution of the longirostrine condition. Zoological Journal of the Linnean Society 188, 473-506.

Groh, S. S., Upchurch, P., Barrett, P. M. & Day, J. J. 2022. How to date a crocodile: estimation of neosuchian clade ages and a comparison of four time‐scaling methods. Palaeontology 65, e12589.

Martin, J. E., Smith, T., Salaviale, C., Adrien, J. & Delfino, M. 2020. Virtual reconstruction of the skull of Bernissartia fagesii and current understanding of the neosuchian–eusuchian transition. Journal of Systematic Palaeontology 18, 1079-1101.

Naish, D. & Sweetman, S. C. 2026. A second specimen of the enigmatic Wealden reptile Yaverlandia. Proceedings of the Geologists’ Association https://doi.org/10.1016/j.pgeola.2026.101179

Ristevski, J., Young, M. T., Brandalise de Andrade, M. & Hastings, A. K. 2018. A new species of Anteophthalmosuchus (Crocodylomorpha, Goniopholididae) from the Lower Cretaceous of the Isle of Wight, United Kingdom, and a review of the genus. Cretaceous Research 84, 340-383.

Salisbury, S. W. & Naish, D. 2011. Crocodilians. In Batten, D. J. (ed.) English Wealden Fossils. The Palaeontological Association (London), pp. 305-369.

Sweetman, S. C., Pedreira-Segade, U. & Vidovic, S. U. 2015. A new bernissartiid crocodyliform from the Lower Cretaceous Wessex Formation (Wealden Group, Barremian) of the Isle of Wight, southern England. Acta Palaeontologica Polonica 60, 257-268.

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Alien animals of any sort, by definition, cannot (in the phylogenetic sense) be tetrapods. To be sure, they can’t really be animals either, given that they’re not part of the Terran clade Animalia. But details details. I will discuss books and projects relating to tetrapod-like aliens if I deem them sufficiently worthy, and such it is with the book I’m going to discuss here…

For some years now, author, artist and retired neurology professor Gert van Dijk has been running and compiling the very excellent blog Furahan Biology and Allied Matters. This is required reading for anyone seriously interested in the designing of hypothetical organisms and Spec Bio (speculative biology or Dixoniana) in general. Blogs and other websites are all very well and good, but a project only has ‘weight’ and some greater amount of permanence when it spawns a book, and such it is with Gert’s Furahan thoughts. After teases and promotions extending over more than ten years, I’m pleased to report that *Wildlife on the Planet Furaha* now exists in hard, physical form, and it was very much worth the wait.

**Caption:** the current header for the Furahan blog, featuring two cursorial hexapod scalates, a bobbuck at left and prober at right. Both animals are mid-sized at close to 1.5 m in length. Image: Gert van Dijk.

The first thing to say is that this book is gorgeous: fantastically well designed, packed with illustrations of all sorts, and featuring spectacular full-page colour images. It might not be surprising in view of this that the author is a skilled artist who previously created cover illustrations for sci-fi works. Among the most interesting and compelling of books ever published are (in my opinion) those made by authors who have an artistic vision and a creative skill, since authors of this sort understand how to illustrate the concepts, objects and scenes they want to, in addition to knowing why this should be done in the first place.

People on Furaha. Furaha was discovered by Swahili-speaking spacefarers from East Africa and a backstory to its finding and the history of its study is discussed at the start of the book. Various key personnel, some pictured in the field or in their ceremonial academic garb, feature throughout and an entire history and academic genealogy of Furahan exploration and study, associated predominantly with the Institute of Furahan Biology, has been devised.

**Caption:** human explorers, travellers and scientists (including citizen scientists) feature throughout the book, and here are a happy pair adjacent to the bone-like internal elements of a brontorusp. Image: van Dijk (2025), used with permission.

It is a (sadly hypothetical) future to be proud of, where people have collaborated in honourable and scholarly pursuits, and where academic achievement is celebrated. We are reminded throughout the book that people have a tolerant, sensible view of the planet’s organisms, even deliberately avoiding the keeping of Furahan organisms as pets and use of the term ‘monster’ for those where it might be appropriate.

The organisms: plants and mixotrophs. The bulk of the book covers the main groups of organisms, all but two of the seven chapters being devoted to these. A complaint sometimes made about efforts to invent life on alien worlds is that plants, and plant-like organisms in general, are mostly ignored. That’s not entirely fair given that several authors who’ve written seriously about this topic have indeed devised alien plant life; I’m thinking of Lewis Dartnell, among others (e.g., Dartnell 2012). Furahan plants include giant, tree-like forms and many others, and not all are green.

**Caption:** a beautiful panorama showing red polypremnic trees alongside green and yellow grass-like ground plants. The animal is a Snafe *Factotum sequax*, a centauric hexapod belonging to a group endemic to the isolated continent of Meralgia. Image: van Dijk (2025), used with permission.

Fungi-like organisms have their own chapter. Like many of us, I have, in recent years, been surprised and impressed by the number and variety of slime-moulds, rusts and other mixotrophs that can be found in the area where I live. Furaha’s mixotrophs are a fascinating and, in part, terrifying bunch. Several are large and absorb nutrients from carcasses; others have evolved trap-like structures (like the dentated spheres of the Phalanx) that enable them (or, at least, enabled their ancestors) to capture and digest animals. Complex lifecycles and innovative means of dispersing larvae are explained.

**Caption:** a mixotroph montage, showing a stand of Nightsnare *Laqueus lentus* at left, and representatives of the different dispersal methods used by Purple flyfoam *Spumascansa damieni* at right. The frothy, methanous ‘rafts’ transport the ‘brochos’ larvae. Image: van Dijk (2025), used with permission.

The organisms: kwals and arthropod-like groups. Moving to animals, Furaha is alive with aquatic and terrestrial species. Its skies are inhabited by volant ones too. If it’s not already clear, Furaha is not home to creatures that look like alien copies of Terran vertebrates, though some amount of convergent evolution is inevitable, as we’ll see. Jelly-like marine organisms termed kwals occur in the seas. Though broadly similar to Terran jellies and with similar reproductive and trophic adaptations, they have evolved a suite of innovations and are substantially more complex.

**Caption:** a selection of kwals, showing (top to bottom) big yellow blob, purple flapper, milky petal pedal and sea wheel. Numerous kwal species exist and the text explains how at least 60 species occur in the small section of coast where these four were encountered. Image: van Dijk (2025), used with permission.

Arthropod-like organisms termed wadudu (singular mdudu) are abundant across the planet and include multi-limbed, terrestrial taxa that resemble spiders, millipedes and insects. Spidrids are superficially spider- or crab-like animals that have adapted to deserts, coasts, woodlands and other habitats, ranging in size from less than 1 mm to 30-40 cm in diameter. Unlike arthropods, they exhibit radial symmetry, with four eyes mounted on the top of the head. They hence don’t have a ‘front’ and can move equally well in any direction.

Explanations and tests. This brings me a key aspect of the book, and indeed the entire Furahan project. Van Dijk’s organisms don’t just do whatever they do based on the designer’s intuition. Rather, he has shown his working, explained his thinking and has even indulged in hypothesis testing when devising body shapes and patterns of locomotion. A detailed and well-illustrated guide to spidrid locomotion is provided (van Dijk 2025, pp. 82-83). If the insect-like organisms of Furaha possess wings and are capable of flight, how would flight occur given body plans built on radial symmetry? Tetrapter flight is explained in a devoted section (van Dijk 2025, pp. 88-89), and likewise for matters concerning locomotion in other organisms.

**Caption:** one of the many ‘technical’ illustrations that appears throughout the book, this one showing the field of vision of the Marblebill *Iaculator weismuelleri*, an arboreal predatory hexapod. Image: van Dijk (2025), used with permission.

Hypothesis testing does not always yield good news when inventing hypothetical organisms, and thus it is that the skies of Furaha are not filled with floating, balloon-like ‘ballont’ animals (“Unfortunately, physics said no to ballonts on an Earth-like planet, so all ballont paintings ended up in the archives”; van Dijk 2025, p. 152). I find this fascinating since it’s directly connected to a specific point I made in my recently published review of Dougal Dixon’s The New Dinosaurs (Naish 2026): in devising hypothetical or ‘alternative’ organisms, we might be providing an opportunity or opportunities to test certain possibilities that have otherwise gone unexplored. Well, here is an example of exactly that… albeit explored in the broader body of van Dijk’s writing online and not in this book.

This is all phenomenal stuff and those aiming to produce similar works should do likewise. I certainly intend to when I get round to creating a Squamozoic book.

**Caption:** two of the many cloakfishes illustrated and discussed in the book. At left, a Starfish *Tanypteryx archicus*; at right, a Common cloak *Tetrachlamus pycnus*, a species regarded as “the prototypical cloaky” (van Dijk 2025, p. 65). Image: van Dijk (2025), used with permission.

Furahan ‘fishes’: cloakfishes and aquatic scalates. Several aquatic Furahan groups look and act like ‘fishes’. Firstly, there are the cloakfishes, a group whose species differ fundamentally from Terran fishes in having four-sided symmetry and a method of propulsion based on the undulation of four longitudinally aligned membranes, these explaining the common name. There are over 10,000 cloakfish species in the Furahan seas and oceans and they feature a dazzling array of colours, patterns and lifestyles. The very biggest have a full span of 7 m. All cloakfishes are filter-feeders that, despite their alien shape, do have distinct dorsal and ventral surfaces and consequently have dorsoventral counter-shading.

More diverse in shape and ecology are the several ‘fish’ groups united as the scalates, the subgroups of which document an evolutionary transition. Archaic groups possess lateral swimming membranes and multiple jaw pairs, but indentations and a reduction of membrane extent ultimately led to the evolution of three sets of paired fins, the most posterior of which fused in one group. Changes in jaw configuration and respiratory anatomy occurred in step with these events. A cladogram depicts the relationships of these groups and where the relevant anatomical traits occurred.

**Caption:** the major Furahan clade of terrestrial scalates includes terrestrial and aquatic species, and fliers too. Some have four wings (the group Quadrialata) and others two (the group Dialata). The images here show korongoes *Dromodraco drungus*, in flight and in a diagrammatic takeoff sequence. This is a large dialate species with a wingspan of up to 5 m. Image: van Dijk (2025), used with permission.

Skeletons have evolved in scalates but the arrangement of the internal components is not like that of vertebrates, whereby the evolution of hard structures about a midline notochord was among the first of events. Scalates on Furaha started the evolution of their skeletons with the reinforcement of their lateral membranes, the result being a series of ‘rail bones’ (connected via ‘node bones’) that extend parallel to the long axis on the body’s outer edges. Rung bones then evolved, connecting left-side to right-side rail and node axes. The result is a ladder-like skeleton (hence the group name).

Terrestrial scalates termed hexapods (no connection to the Terran arthropod clade Hexapoda) include a diversity of heavy-bodied herbivores and omnivores, small arboreal forms, cursorial herbivores and predators, and large, long-limbed suspensory species like marblebills and berbies. Some terrestrial hexapods possess an elevated anterior section and their anterior-most limb pair is specialised for a role in disabling or killing prey, this configuration being termed centaurism. Some terrestrial hexapods have returned to life in water, such that there are superficially seal-like forms.

**Caption:** a Stickler *Perfixor artifex*, a predatory centauric hexapod of North Auralgia that undergoes colour change across the seasons. It is social and potentially dangerous to humans. The full illustration is taller and occupies an entire page. Image: van Dijk (2025), used with permission.

Rusps, including megarusps. Finally, hexapods are not the only large, terrestrial animals on Furaha. Rusps are long-bodied, multi-limbed animals that look something like armour-plated caterpillars, albeit equipped with an extensible mouth and, in some, a whip-like defensive organ attached to the head. The biggest – certain of the megarusps – are sauropod-sized. The existence of an apparently extinct megapredator that once posed a danger to these animals is hinted at.

**Caption:** the anterior end of *Brontocrambis brucus*, the Brontorusp, as it turns to face the viewer. Without the whip, this animal is 25 m long. The book features a double-page spread showing this species in its environment, plus another showing various megarusps to scale. Image: van Dijk (2025), used with permission.

This is a tremendously fun, charismatic and innovative menagerie. Many of the animals look great and I find their evolutionary backstories plausible. I should emphasise that my discussion here only scratches the surface and that numerous other creatures win coverage in the book. In addition, the book is not a complete list of all Furahan organisms but only a tour of various highlights. My only complaint is the absence of one or more big cladograms that show the relationships between the main groups, since it isn’t immediately clear how the main Furahan clades are related to one other.

As fun as it is to think that Furaha and its wildlife might be real… alas. An Appendix discusses the ‘prehistory’ of this project and discusses how the author got the whole thing off the ground, this section also featuring some of the art he created during its earliest stages. A glossary of Furahan terms follows, and there’s an index too.

**Caption:** proof that the idea of this book has been on the cards for a while. This photo was taken at the Spec Bio workshop at LonCon3 of August 2014, and the screen behind Gert (who is standing at left) shows the semi-hypothetical cover of an *Encyclopedia of Furahan Wildlife* (a double-page spread is visible at top). C. M. Kosemen is sitting at right, and you can see semi-hypothetical books devoted to Snaiad as well. Image: Darren Naish.

As is tradition ever since Dixon, Furahan organisms have binominal names. These feel real and are always worth checking, since they often include fun references to relevant people, among them Dougal Dixon and Johnny Weismueller. I have doubts about the species name in Pigritia sursumvergenspropterpenuriaponderis (the Righteous phelp) making it through peer review though. There are references to other Spec Bio projects here and there. Snaiad exists in the Furahan universe, for example (van Dijk 2025, p. 18). I spotted the Jaws reference but won’t spoil it.

**Caption:** the book features ‘range map’-type illustrations throughout, showing the known distribution of the species or group in question in red. Also featured throughout are scaled silhouettes showing the organisms with a selection of (often amusingly posed) humans. All of these diagrams are immaculate and wonderfully drafted. Image: van Dijk (2025), used with permission.

Wildlife on the Planet Furaha is a real tour-de-force that does its subject very, very proud. The book is beautifully designed, flawlessly edited, and tremendously fun to read. It is also highly affordable. Books on speculative biology are rare, and those few good ones that exist have – with exceptions – been discussed or even reviewed at Tet Zoo within recent years. It’s therefore very fitting to have the opportunity to discuss Wildlife on the Planet Furaha here.

In short, I adore this book and recommend it extremely highly. You must get hold of it if your interests overlap mine, and my congrats to the author and the publishers on a job very well done.

van Dijk, G. 2025. Wildlife on the Planet Furaha: A Speculative Biology Guide to Alien Life Forms. The Crowood Press Marlborough, UK. ISBN 978-0-7198-4571-0, softback, illustrations, pp. 160. £20.00/$29.99. Here from the publishers (but seemingly sold out). *Here from a giant international conglomerate.*

For previous Tet Zoo articles on Spec Bio and associated fields, see…

The Dougal Dixon After Man Event of September 2018, September 2018
Dougal Dixon’s After Man, the Initial Pitch Document, June 2020
Humanoid Dinosaurs Revisited Again: Russell and Séguin’s Dinosauroid at (Nearly) 40 Years Old, August 2021
Speculative Zoology and the World of After Man; an Interview With Dougal Dixon, April 2025
Speculative Zoology Grand and Photoreal: Boulay and Steyer's Demain, les Animaux du Futur, April 2025
Speculative Zoology’s 21st Century Redemption at the High Table: the 2025 Edition of Dougal Dixon’s The New Dinosaurs, March 2026

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Dartnell, L. 2012. Life under alien skies. Physics World 25, 26-30.

Naish, D. 2026. The New Dinosaurs: an alternative evolution. Historical Biology doi 10.1080/08912963.2026.2627439

van Dijk, G. 2025. Wildlife on the Planet Furaha: A Speculative Biology Guide to Alien Life Forms. The Crowood Press, Marlborough, UK.

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It’s time again to rescue another squamate-themed article from the Tet Zoo articles. This one – devoted to the treerunners, obviously – was first published at Tet Zoo ver 3 in December 2017 (the original Sci Am version is hosted here) and here it is again, with substantial updates…

**Caption:** at left, the head of a *Plica plica* specimen in dorsal view, showing scalation. Squamates vary in how symmetrical their head scales are, and minor asymmetry (as seen here) is common in many species. At right, a captive individual of what’s probably *Plica plica*. This individual lacks bright colours or a prominent nuchal crest, indicating that it’s a female or a juvenile male. Credits: Etheridge (1970); Darren Naish

What is Plica plica? It’s a strikingly proportioned, diurnal, arboreal iguanian lizard that can exceed 17 cm in total length and has a range that encompasses a huge part of northern South America (though, keep that in mind and read on). Etheridge (1970) described Plica plica and its close relative P. umbra as “among the most abundant, widespread, and earliest known lizards of South America” (p. 237). That “earliest known” always looked odd: what he meant is that P. plica is among the earliest of South American lizards to be scientifically recognised, since Linnaeus listed it (as ‘Lacerta plica’) in 1758.

**Caption:** captive *Plica* photographed in captivity in 2012, labelled as *P. plica* (but is it? Read on). Note the reddish head, light green or yellowish rings on the distal parts of the limbs, and the especially dark collar-like markings. Credit: Darren Naish.

I say that this lizard is “strikingly proportioned” because it’s shockingly suited for arboreal, trunk-clinging life, being highly gracile – almost spidery – and with a very slim tail, slender digits and hooked claws.

Plica is a tropidurine, a group of iguanian lizards that I thought I’d written about before. On checking, it turns out that I’ve written (more than once) about the liolaemines, another iguanian group sometimes confused or combined with tropidurines but nowadays thought to be quite distinct from them (Pyron et al. 2013). Indeed, a taxonomic system that recognises these major lizard groups as ‘families’ and separate from the super-inclusive Iguanidae of tradition probably best reflects their distinct nature and phylogenetic history. Tropidurines (Tropiduridae, if we go with that family-level thing) appear to be the sister-group to all other pleurodont iguanians, meaning that they’ve almost definitely been distinct since early in the Cenozoic if not before (Townsend et al. 2011). In addition to Plica, Tropiduridae includes Eurolophosaurus, Microlophus, Stenocercus, Strobilurus, Tropidurus, Uracentron and Uranoscodon. Some of those lizards are incredible and I really must cover them here some time.

**Caption:** much simplified, stripped-down cladogram for pleurodont iguanians. More complete, more complex versions of this tree will appear here in later articles (something I’ve been saying since 2017). It’s one of the hundreds of illustrations produced for my in-prep textbook on the vertebrate fossil record **on which go here**. Credit: Darren Naish

Anyway, those arboreal habits explain why the Plica species are sometimes called ‘tree runners’ or ‘treerunners’, though – frankly – this is a terribly vague name since I don’t think it’s clear that it refers to a group of lizards. The names Collared tree lizard, Collared tree runner and Harlequin racerunner are apparently also in use for P. plica. Calling it a ‘racerunner’ also seems less than brilliant given that the term ‘racerunner’ is more normally applied to a group of lacertids (see the article here); it’s also sometimes used for certain teiids. Thanks to Wikipedia, I know that a local name used for P. plica in Guyana is wakanama and I wish that this was in general use as its vernacular name.

Taxonomic revision and the post-2013 recognition of new species. An interesting thing long known about P. plica (in the traditional, inclusive sense of the name) is that it’s highly variable across its range: in size, the shape of the snout, the size and nature of the scales and neck spines, and in how much spotting there is on the throat. They’re sexually dimorphic, males having brighter, bolder colours and more prominent nuchal crest scales than females... and I’m going to assume here that discussions of variation within the species do take account of this dimorphism. Etheridge (1970) drew attention to geographical diversity in P. plica but noted that the poor preservation of many of the specimens used in his study didn’t allow this variation to be studied more carefully. It has long seemed plausible that various subspecies or even species might be involved.

**Caption:** a map from Etheridge’s 1970 study of *P. plica*, showing the reported localities he was aware of. As you can see, this species – as perceived by Etheridge – was meant to have an enormous range, and to occur across a fairly wide range of topographies and habitats. Credit: Etheridge (1970).

Murphy & Jowers (2013) used both morphological and molecular data to show that P. plica of tradition is indeed a complex of cryptic species. They named the Caribbean treerunner P. caribeana of the Eastern Coastal Range of Venezuela, Trinidad and Tobago, Kathleen’s treerunner P. kathleenae of Guyana, Medem’s treerunner P. medemi of Colombia, and Ray’s treerunner P. rayi of Venezuela and Colombia as new. P. plica proper is restricted to Surinam according to this view. Furthermore, this isn’t the end of it, since they noted that their study was not a revision of the entire species complex but merely an initial demonstration of ‘P. plica’s’ polytypic status. In another Plica-based study, Paula de Oliveira et al. (2016) found another species – P. umbra – to also be a species complex. A key point they made in that study is that: “If the observed diversity of lineages within the genus Plica is characteristic of squamate reptiles of the Amazon region, the diversity of squamates is grossly underestimated”. That’s very much a non-trivial observation.

**Caption:** a *Plica* montage. At top, a captive animal labelled as *P. plica* (the image has been rotated by 90°; it was actually clinging vertically from a trunk). At lower left, *P. umbra* photographed in the wild. At lower right, *P. rayi* in Puerto Ayacucho, Venezuela. Images: Darren Naish (top); Alessandro Catenazzi, CC BY-SA 2.5 (lower left; **original here**); Zelimir Cernelic, CC BY 3.0 (lower right; **original here**).

All of this leaves me wondering about the precise status of the captive animal or animals – labelled P. plica – shown in my photos here. The reddish head in the animal at the top of the article makes it look like one photographed in the Sierra de Lema of Venezuela, and figured by Murphy & Jowers (2013), though I can’t work out from their paper which species that specific specimen belongs to. Ho hum.

Behaviour and biology. Having mentioned sexual dimorphism, I should note the claim that dimorphism in these lizards is actually less pronounced than it is in other tropidurines (Vitt et al. 1997). That same study reported many interesting things about the ecology and biology of treerunners. They’re partly social, often occurring in pairs or groups. They’re ant specialists, a sample of 36 individuals revealing no other prey items whatsoever, even though other arthropod prey were apparently highly available. Vitt et al. (1997) wondered if the success of treerunners throughout South American lowland forests might be related to their exploitation of this resource, one otherwise not utilized by many other tree-climbing lizards across the region. You might be wondering what this means for maintaining them in captivity (where they do comparatively well). It turns out that they’ll accept crickets, weevils, cockroaches and fruit flies (Harding et al. 2016).

Egg-laying in Plica seems to occur throughout the wet season and during some of the dry months as well, and clutch size is small, averaging 2.9 in P. plica. This is presumably related to their compressed, gracile body shape (Vitt et al. 1997). They don’t nest in the arboreal environment but come down to the ground and bury the eggs among leaf litter (Harding et al. 2016). One final thing: despite living in environments where shade is common, treerunners do well at maintaining high temperatures and have generally been found to be a few degrees warmer than air and surface temperatures (Vitt et al. 1997).

**Caption:** *Plica plica* at left. At right, graph from Vitt *et al*. (1997) showing relationship between body temperature and substrate temperature in wild *P. umbra*. Images: DuSantos, CC BY 2.0 (**original here**); Vitt *et al*. (1997).

Iguanians might be over-represented (as lizards go) at Tet Zoo, in part because they’re kept more frequently in captivity than other lizard groups and hence are more likely to be photographed. One thing I aim to deal with when time allows: that whole Toxicofera thing!

For previous Tet Zoo articles on iguanians, see…

The Tet Zoo Guide to Mastigures, August 2018
The Remarkable Basilisks, May 2023
Leiosaurid Lizards: South America, Land of Iguanians, November 2024
What's With All These New Chameleon Names? Chameleons, Part 1, November 2024

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

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Etheridge, R. 1970. A review of the South American iguanid lizard genus Plica. Bulletin of the British Museum (Natural History) 19, 237-256.

Harding, L., Tapley, B., Gill, I., Kane, D., Servini, F., Januszczak, I. S., Capon-Doyle, J.-S. & Michaels, C. J. 2016. Captive husbandry and breeding of the tree-runner lizard (Plica plica) at ZSL London Zoo. The Herpetological Bulletin 138, 1-5.

Murphy, J. C. & Jowers, M. J. 2013. Treerunners, cryptic lizards of the Plica plica group (Squamata, Sauria, Tropiduridae) of northern South America. Zookeys 355, 49-77.

Paula de Oliveira, D., Tadeu de Carvalho, V. & Hrbek, T. 2016. Cryptic diversity in the lizard genus Plica (Squamata): phylogenetic diversity and Amazonian biogeography. Zoologica Scripta 45, 630-641.

Pyron, R. A., Burbrink, F. T. & Wiens, J. J. 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 2013, 13:93 doi:10.1186/1471-2148-13-93

Townsend, T. M., Mulcahy, D. G., Noonan, B. P., Sites, J. W., Kuczynski, C. A., Wiens, J. J. & Reeder, T. W. 2011. Phylogeny of iguanian lizards inferred from 29 nuclear loci, and a comparison of concatenated and species-tree approaches for an ancient, rapid radiation. Molecular Phylogenetics and Evolution 61, 363-380.

Vitt, L. J., Zani, P. A. & Avila-Pires, T. C. S. 1997. Ecology of the arboreal tropidurid lizard Tropidurus (=Plica) umbra in the Amazon region. Canadian Journal of Zoology 75, 1876-1882.

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Last year saw the appearance of the long-awaited second edition of Dougal Dixon’s The New Dinosaurs (Dixon 2025). If you’re a fan of the original, published in 1988, now is the time to get a second edition.

**Caption:** front and rear covers of the 2025 edition of Dougal Dixon’s *The New Dinosaurs: An Alternative Evolution*. The large theropod is a Cutlasstooth; to its right is a flightless pterosaur termed the Flarp. The rear cover shows (at top) a Sandle (at left, a fossorial theropod) and the head of a Gwanna, and (at bottom, left to right) a male Dingum (an Australasian coelurosaur), a Bricket (a Palaearctic lambeosaur) and a Nauger (a woodpecker-like arbrosaur).

I attended the London launch event, got hold of a review copy, and have just published a reasonably long article on the book in Historical Biology (Naish 2026). There is, of course, a huge amount of stuff to say about TND, some of which has been touched on here at Tet Zoo before (see the links below). But there are several additional points that haven’t been commented on at all, or – at least – not much, and I used the review as an opportunity to highlight them. I’ll summarise them here, meaning that this article is a summary of a review of a book, how meta. Let me know if you want the full pdf (of the review, not the 2025 edition of The New Dinosaurs. Buy it here).

**Caption:** parts of pages within the 2025 edition of *The New Dinosaurs*. A Malagasy megalodontosaur at left, a monocorn at right. Yup, that image on the left is by the legendary Denys Ovenden. The ones on the right are by Philip Hood… I own the original!

The New Dinosaurs is part of a 1980s ‘late renaissance’ wave in how Mesozoic archosaurs were depicted and described. Dougal Dixon is one of a list of authors who wrote about Mesozoic dinosaurs (and pterosaurs, and marine reptiles) from a ‘mainstream’, conservative view. There is, in fact, a ‘very British’, staid tradition of writing about these animals that extends from William Swinton in the 1930s to Alan Charig and Beverly Halstead in the 1970s and 80s, and to David Norman, Mike Benton and others during the 80s and beyond.

I mean no disrespect whatsoever here; it’s just the way it is. So when the American authors and artists Robert Bakker, Greg Paul, Mark Hallett and John McLoughlin (plus a few others) began publishing images of fuzzy and feathery ornithischians, thickly furred pterosaurs, bird-like theropods sprinting and leaping high off the ground, and sauropodomorphs rearing, tail-whipping and generally behaving more dynamically than had been the case before (as they did from 1975 onwards)… what to do?

**Caption:** it’s not true that fuzzy and feathery non-bird dinosaurs were ‘all new’ in the 1990s; they’d been kicking around in the literature for decades prior, as shown by this feathered *Syntarsus* from Bakker’s 1975 *Scientific American* article (upper right) and the feathered *Coelurus* (actually *Ornitholestes*) from McLoughlin’s 1979 *Archosauria* book. Fuzziness in pterosaurs was also well known by this time, and even Halstead endorsed this in his book of 1975 (art by Giovanni Caselli).

TND is about fictional animals of an alternative timeline. But a point that’s been missed or ignored until now is that it’s very much of its time, its animals reflecting views new in the 1980s. The pterosaurs are (mostly) capable bipeds whose hindlimbs are free of the wing membranes, a new-fangled idea advocated by Kevin Padian and picked up by artists including Paul and Hallett. The pterosaurs of TND are fuzzy, the theropods are too… but so are the ornithischians, and while there’s support for that view today (from Tianyulong and Kulindadromeus at least), it was elsewhere only being promoted in palaeoart like that by Paul.

**Caption:** new ideas on pterosaurs were appearing during the 80s, and one that had its time in the limelight proposed that these animals were agile, cursorial bipeds. This is an iconic image of the Jurassic pterosaur *Dimorphodon*, by J. Kevin Ramos, and it was used in several of Kevin Padian’s papers and articles published during the 1980s.

I put it that TND marks a departure from the ‘very British’ tradition of portraying these animals and hence should be considered part of this interesting phase of history. The old, conservative interpretation might be on the bonfire; it’s time to adopt a less familiar, stranger, perhaps more exciting view (Naish 2026). Importantly, it’s flat wrong to claim in seriousness than the Dinosaur Renaissance resulted in a negation of all that had gone before; things were far more nuanced (Naish 2021). But it’s also wrong to pretend that nothing happened.

Redemption. I’m hardly the first to point out that various of the animals featured in TND have been part-redeemed by recent discoveries. A giraffe-like, flightless pterosaur is consistent with the ‘terrestrial stalking’ interpretation of azhdarchids (Witton & Naish 2008), dwarf, island-dwelling macronarians are now a real-world thing, the long-faced, fish-eating Dip and superficially pangolin-like Pangaloon of the book are reminiscent of unenlagiines and alvarezsaurids, and ‘whelks’ might well have evolved had aristonectines not gone extinct (Naish 2026).

**Caption:** montage from Naish (2026). Several animals of *TND* pre-empt palaeontological discoveries and proposals. (A) The Lank of *TND* (illustration by Steve Holden; from Dixon (2025)) is a large flightless pterosaur that walks with a pacing gait and consumes grasses. (B) **Witton & Naish (2008)** proposed that azhdarchids were ‘terrestrial stalkers’, inviting comparisons between pterosaurs and giraffes. (C) The Dip of *TND* (illustration by Denys Ovenden; from Dixon (2025)), a long-faced, piscivorous coelurosaur. (D) Reconstruction of an unenlagiine, a long-faced maniraptoran with piscivorous adaptations. (E) The Pangaloon of TND (illustration by John Butler; from Dixon (2025)), a pangolin-like, terrestrial arbrosaur. (F) Reconstruction of an alvarezsaurid, a maniraptoran with pick-like forelimb and possible insect-eating adaptations.

Some TND animals are not realistic and never have been, among them snake-like and lizard-like theropods. Today I’m not so interested in heaping scorn on those, but think instead that we need to frame TND as the worthy successor to After Man that it is: the latter is a general introduction to the principles of evolution, the former specifically aims to teach its audience about zoogeography. It’s taken me a while to fully grasp the pre-eminence of zoogeography throughout TND, but today it’s obvious.

A tradition of reviewing. I’m already on record as explaining why, in my view, book reviews are important if you’re interested in charting the history of ideas on a subject. They allow authors – most notably, qualified and active researchers in the respective field – to muse, express personal opinions, and be critical or supportive in ways that aren’t usual for the technical literature (Naish 2025). Speculative Zoology and its cousins are generally not considered welcome at the high table. They don’t constitute real science, and who needs them anyway.

But is that true? If we survey our history, I don’t think it is. If it is, then why do Dougal Dixon’s books keep being reviewed in ‘mainstream’ science periodicals? Examples include Gee (1988), Tudge (1988), Paul (1990) and Unwin (1992). Oh, and Naish (2026). The fact is that Spec Zoo wins its place in the discourse because we know that it’s relevant to a great many of the broader questions we have. And on that note…

**Caption:** speculative dinosaurs of a world where the end-Cretaceous event never occurred, itself based on a scene featured in a previous review (Paul 1990) of *TND*, a scene that might now be criticised as too conservative. A giant, extravagant paravian culls out a dolicocephalic hadrosaurian grazer from a mixed herd that also includes large owlbear leptoceratopsians. A burrowing thescelosaur is visible at right and ducks flock overhead. The locale is the western grasslands of North America. From Naish (2026).

Relevance. Speculative Zoology mostly justifies its existence from its value as entertainment. It’s fun. But in asking questions about things that might have been, or might be at some point, are we doing something more? There’s a greater amount of this sort of thing in the literature than might be obvious: that is, where biologists and palaeontologists have asked questions about things that have seemingly never happened, or, if they have, have only happened once or twice, despite potential. Examples include articles on the distribution of viviparity across reptiles (including birds), body size limits in bats, the rarity of herbivorous aquatic tetrapods, and the supposed absence of suspension-feeding marine reptiles.

In fact, the increased realisation that the animals of the Mesozoic were very much guaranteed a future had a terrible event not occurred – they didn’t just slide into decline due to poor design – has been coupled with views on their diversity and adaptive scope to perhaps make speculations about things that might have been more acceptable. There are the very famous discussions of possible post-Cretaceous intelligence of course (Russell & Séguin 1982, Naish & Tattersdill 2021, Reiner 2023), but there are speculations on other topics too (see Naish 2026).

And thus it is that we want *The New Dinosaurs*. We might even need it. It might even be inevitable that such a book was destined to appear given what I just said about the primacy of ‘what ifs’ in discussions of evolutionary history. Could anyone else have done a better job than Dougal Dixon? Whatever, the fact is that they did not.

Let me know if you want a pdf of my review (not in the comments here: find me on social media or email). These are exciting times if you’re interested in Speculative Zoology, and it’s great news that this classic work – so long hard to get hold of if you missed it first time around – is back in print. Buy it here and tell your friends.

For previous Tet Zoo articles on speculative zoology, see...

The Dougal Dixon After Man Event of September 2018, September 2018
Dougal Dixon’s After Man, the Initial Pitch Document, June 2020
Humanoid Dinosaurs Revisited Again: Russell and Séguin’s Dinosauroid at (Nearly) 40 Years Old, August 2021
Speculative Zoology and the World of After Man; an Interview With Dougal Dixon, April 2025
Speculative Zoology Grand and Photoreal: Boulay and Steyer's Demain, les Animaux du Futur, April 2025

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Dixon, D. 2025. The New Dinosaurs: An Alternative Evolution. Breakdown Press, London.

Gee, H. 1988. Tales of future past. Nature 335, 505-506.

Naish, D. 2021. Dinopedia: A Brief Compendium of Dinosaur Lore. Princeton University Press, Princeton NJ.

Naish, D. 2025. Thoughts on reviewing books, from an incoming book review editor. Historical Biology 37, 2591.

Naish, D. 2026. The New Dinosaurs: an alternative evolution. Historical Biology doi 10.1080/08912963.2026.2627439

Naish, D. & Tattersdill, W. 2021. Art, anatomy and the stars: Russell and Séguin’s dinosauroid. Canadian Journal of Earth Sciences 58, 968-979.

Paul, G. S. 1990. An improbable view of Tertiary dinosaurs. Evolutionary Theory 9, 309-315.

Reiner A. 2023. Could theropod dinosaurs have evolved to a human level of intelligence? The Journal of Comparative Neurology 531, 976-1006.

Russell, D. A. & Séguin, R. 1982. Reconstruction of the small Cretaceous theropod Stenonychosaurus inequalis and a hypothetical dinosauroid. Syllogeus 37, 1-43.

Tudge, C. 1988. End points of an alternative evolution. New Scientist 120 (1641), 65-66.

Unwin, D. M. 1992. The New Dinosaurs: An Alternative Evolution (review). Historical Biology 6, 61-71.

Witton, M. P. & Naish, D. 2008. A reappraisal of azhdarchid pterosaur functional morphology and paleoecology. PLoS ONE 3 (5): e2271.

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It’s probably impossible to write about the history of fossil hominin discoveries in Africa and not discuss, or at least mention, the Leakeys…

**Caption:** Mary Leakey’s 1984 book, the topic of the article here, with a few other relevant works in the background. I have accrued a great many books on fossil hominins and am slowly working my way through them. Image: Darren Naish.

While Louis Leakey (1903-1972) is the most famous, if not notorious, member of this family, it’s the highly accomplished Mary (1913-1996), his wife, that we’ll be looking at here. In 1984, when she was in her early 70s, Mary penned Disclosing the Past: An Autobiography (Leakey 1984). As part of my slow-burn look at volumes devoted to palaeoanthropology, I’ve recently been obsessing over this book.

Mary notes early on that the writing of autobiographies is something of a Leakey trait, and it’s no trivial matter that Louis began this task when he was just 33 years old. Thanks both to her association with Louis – initially as an assistant, then through marriage – and her later status as an independent researcher with her own work programme, Mary was involved across much of her life with archaeology and has connections to a great many interesting discoveries and hypotheses. I looked forward to reading about these and wasn’t disappointed. The book is eminently readable and contains a vast amount of information.

**Caption:** Mary Leakey has had a small amount of fame, and has appeared on stamps, as a TV guest, and on the covers of various books. In 2013 - in celebration of the date of her birth - Google released a Doodle devoted to her work and life. You can read a bit about its backstory and creation **here**.

Early chapters dwell on Mary’s childhood, much of which involved London and part of it France. Her father’s occupation as a painter I think gave the family a bohemian, perpetually mobile life that must have involved financial fits and starts, though it’s hard from a modern perspective to understand how people claiming to have no money might be able to own homes in Kensington and France. As might be predicted for a person with such a background, Mary lacked a stable school life and was forever in and out of education, much of this Catholic. It’s also clear that she was a rebel, on occasion hiding in the boiler room to avoid lessons and on another causing an explosion in a chemistry class. A bold and adventurous streak is demonstrated by her adoption of gliding as a hobby.

Meeting Louis, and a career in archaeology. Mary (née Nicol) first met Louis Leakey in 1934 after being introduced to him by archaeologist Gertrude Caton-Thompson, best known at the time for her work on stone tools excavated at the Fayoum in Egypt. Caton-Thompson, Mary noted, “was the epitome of that remarkable breed of English ladies who for archaeology’s sake and by determinism, skill, expertise and endurance achieve discoveries of major and permanent importance” (Leakey 1984, p. 39). Mary had produced illustrations for Caton-Thompson’s book The Desert Fayoum and it was the quality of this work that resulted in the introduction, since Louis was in search of an illustrator for Adam’s Ancestors (Leakey L. 1934), a book that was to have numerous editions over the years. Louis was married (apparently unhappily) at this time to Frida, and Mary was taken by Louis to meet her in Cambridge.

**Caption:** this photo showing Louis and Mary working in the field, I think from the 1960s, obviously isn’t relevant to their early life together in the 1930s and 40s. Image: Smithsonian Institution, public domain (**original here**).

By this time (the early 1930s), Mary was already involved in archaeology of her own volition having attended courses in geology at University College, London and in archaeology at London Museum. Here, she attended lectures by R. E. M. Wheeler (later, Sir Mortimer Wheeler) and of interest is her comment that “I can’t help wondering whether many of the achievements in archaeological discovery and methodology that are so often attributed to him did not rightly belong to his charming and extremely able first wife, Tessa” (Leakey 1984, p. 37).

Mary soon assisted and even directed excavations at Meon Hill, Hembury, Swanscombe and Jaywick (near Clacton). Swanscombe’s famous hominin was found there by visiting amateur archaeologist A. T. Marston in 1934. The Jaywick excavations were successful too and Mary, working with Kenneth Oakley, published her results on this locality in 1937, this being her first publication (Leakey 1984, p. 49).

Louis was involved at some level in various of these excavations and in fact planned to study and resolve questions surrounding the ‘Clactonian’ industries of the Early Palaeolithic, though Mary reports that he never completed nor published this work (Leakey 1984, pp. 48-49). Louis was a remarkable man – I don’t just mean this in a positive sense – and a vast quantity has been written about him, much of it by himself. His relevance to Mary’s life of course means that he warrants at least some coverage in my article here, though I should note that I’m reading additional works on the Leakeys and will return to a discussion of Louis in time.

**Caption:** Clactonian sites yield distinctive Lower to Middle Palaeolithic tools, among them flint ‘choppers’ like this. Image: Sussex Archaeological Society, Stephanie Smith, CC BY-SA 4.0 (**original here**).

To Olduvai. It would have been during this English fieldwork that Louis first introduced the topic of Olduvai Gorge in Tanzania to Mary, a location with massive potential for Palaeolithic finds that might be important in global terms, not just East African ones (Leakey 1984, p. 49). Louis had been raising money to excavate there and wanted Mary to come along, and this she did, travelling there with her mother in 1935. In Zimbabwe, she had her first view of African rock art and described the memorable impression it had (Leakey 1984, p. 52).

Hominin remains identified as conspecific with the Asian species Homo erectus had only recently been found at Olduvai, as had the fossil antelope Phenacotragus recki (it’s Antidorcas recki today). A primate tooth found at Laetoli thought at the time to be that of a cercopithecid monkey later turned out to be that of a hominid. But, overall, the deposits weren’t as rich as hoped. Louis thought that the hippos of Ngorongoro didn’t belong to Hippopotamus amphibius but might be a surviving population of the especially big, ‘pop-eyed’ Pleistocene form H. gorgops. A long trek was made in order to discover a skull and thus test this possibility, but once one was found the ‘late-surviving H. gorgops hypothesis’ was no more.

**Caption:** hippos have a fascinating fossil record that involves Europe and Asia as well as Africa, and if you’ve been visiting Tet Zoo for long enough you might recall my article on this topic. At left, an Olduvai Gorge fossil of the giant extinct Pleistocene species *Hippopotamus gorgops* at the Natural History Museum, London. At right, a life reconstruction of this species. Images: Darren Naish.

Wildlife dangerous and not. Fieldwork in the African countryside of course involves the occasional encounter with wildlife, and tales of lions and other animals are peppered throughout the book. Some make for awkward reading, in particular one where a former student of Louis’s, now the local District officer, so disliked the presence of a local lion pride that he went and shot the whole lot. Mary expressed her disgust at this. There are also accounts of snakes being killed after entering premises. By the 1940s, Mary was a mother (first to Jonathan, then Richard, then Philip) and baby Jonathan was nearly killed by swarming army ants. Visiting geologist Robert Shackleton was bitten by a puff adder but survived after Mary ran two miles to the car to retrieve the serum they’d bought with them, leaving a very young Jonathan and her dog Janet with Shackleton to do this.

An aspect of Mary’s life that I can’t approve of concerns her habit of letting her pet dalmatians run off and do whatever they liked in the bush. At least some people saw these unfamiliar black and white beasts in the field and reported them to officials as a new species of predator (Leakey 1984, p. 205), not realising that they were exotic pets allowed to roam free. Like most pet dogs and cats given free roam, their time was of course mostly spent hunting, and Mary mentions more than once the fact that she and colleagues had to locate the dogs and bring them back to camp. This also, of course, presents substantial danger to the lives of the pets themselves and it’s unsurprising that Mary mentions the loss of dogs to snakebite and another to a lion.

**Caption:** Rusinga Island, Lake Victoria, Kenya, is a familiar location if you’re familiar with late Cenozoic African faunas. Here’s what it actually looks like, as seen in panoramic view from the south-east. Image: Küchenkraut, CC BY-SA 3.0 (**original here**).

Mary’s serious commitment to keeping and breeding Dalmatians is demonstrated by her and Louis’s founding, in 1949, of the Dalmatian Club of East Africa (Leakey 1984, p. 113). Mary and Louis were evidently very serious about keeping and showing these dogs and both had roles in the East African Kennel Club.

Also of interest is that the Leakeys kept hyraxes (both rock and tree hyraxes) as pets, basically by just capturing them in the wild and then getting them used to human (and domestic canine) company. A number of interesting anecdotes about the keeping of hyraxes are included. An eagle owl was also kept as a pet after being rescued as a chick and then there are the duikers and other antelopes, genets, bat-eared foxes and snakes.

**Caption:** hyraxes are fascinating animals, and they’ve recently become familiar to a massive, new audience thanks to the popularity of certain social media accounts (waa-waa!!). I plan to write about them at length and never have. The animal at left is a Bush hyrax *Heterohyrax brucei*; that on the right is a Rock hyrax *Procavia capensis*. Images: JaySef, CC BY-SA 3.0 (**original here**); Prosthetic Head, CC BY-SA 4.0 (**original here**).

Crocodiles presented a danger at the waterside Rusinga Island locations, and Mary describes Louis’s solution to this: “when we were all ready to bathe he would fire both barrels of a shotgun into the water” (Leakey 1984, p. 102).

African archaeology and phantom thumps. Discovery sites excavated during the 1930s and 40s include Hyrax Hill, a burial site and prehistoric shoreline, and Mount Olorgesailie in the Rift Valley, a site so rich in stone tools that it was made into an open-air museum. While working at Hyrax Hill, Mary felt a hard thump on her shoulder, caused by an unseen assailant, and interpreted this as displeasure from a local spirit. I’ve heard tell of various ghostly encounters like this. I’ve twice received prominent ‘phantom thumps’ myself, once on the shoulder and once on the back of my calf. The second one caused impressive bruising, and I assumed initially that someone had kicked me or hit me with a ball. No, these were strains caused by overloading an insufficiently warmed muscle. I reckon that most such events can be explained this way.

**Caption:** an image that gives a good idea of what the countryside near Mount Olorgesailie looks like, as of 1993. Image: Rossignol Benoît, CC BY-SA 3.0 (**original here**).

Another interesting character mentioned in passing is Joy Adamson, the famously popular author of Born Free and other works, whom Mary got to know through Joy’s marriage to Peter Bally at the Coryndon Museum (meaning that she was Joy Bally at this time). Joy was, apparently, “not a sympathetic personality, nor particularly easy to get on with”, but Mary did admire her genuine love of animals and hard work in raising funds for their benefit (Leakey 1984, p. 79).

Additional archaeological work of the 1940s led to Mary’s work in west Kenya on ‘dimple-based’ Iron Age pottery accrued by Archdeacon Owen, thought connected to a southward expansion of the Bantu. Similar pottery was later recovered from elsewhere in the continent (Leakey 1984, p. 86). Here’s your regular reminder that texts on the history of humanity and archaeology in general mostly do a poor job of explaining the events of Africa once you get to the evolution of modern humans, something that contributes to the idea that Africa is, and has been, a backwater where little to nothing happened. Archdeacon Owen was a champion of native rights and other causes who “would get down on his knees and pray for better luck”. He’s said to have deliberately buried a Proconsul skull after finding it. Its discovery, he feared, was at odds with the Old Testament story of creation (Leakey 1984, p. 86).

Rusinga Island’s fossils. The Miocene fossil sites of Rusinga Island in Lake Victoria became gradually better known during the 1940s. Among these, remains of the (supposed) hominoid Proconsul were considered the most exciting, a partial skull found by Mary in 1948 being among the best ever found (Leakey 1984, p. 98). Some of the fossils concerned here are no longer included in Proconsul but now belong to Ekembo, a taxon split from Proconsul by McNulty et al. (2015). Anyway, additional fragments, collected in 1947 but not studied until the 1960s by Martin Pickford, proved to be from the very same specimen.

**Caption:** several species have been included within *Proconsul* and the taxonomy of the genus is quite complicated. I haven’t taken the time to sort out which *Proconsul* taxon Mary was writing about, but the skull on the left is that of the type species *P. africanus*, this being a specimen on show at the American Museum of Natural History, New York. At right, a life reconstruction of *P. africanus* by Mauricio Antón, emphasising the superficially cercopithecid-like proportions of this animal. Image: Ryan Schwark, public domain (**original here**); Mauricio Antón, CC BY 4.0 (**original here**).

An important benefactor of the Leakey’s research in Rusinga was British-American businessman Charles Boise, whose name (I think pronounced boy-see) you’ll recognise due to the 1959 naming of Zinjanthropus boisei in his honour. While in France during the 1950s, the Leakeys had a private after-hours tour of the famous Périgord and Lascaux caves and were able to take Boise with them. This trip and others encouraged Boise to fund the Leakey’s work, and palaeoanthropology in general.

Such was international interest in Mary’s significant Proconsul specimen that she hand-carried it by way of a free flight offered by BOAC (the ancestor of British Airways) to England. Here, it was studied by William Le Gros Clark in Oxford, the great expert in primate evolution at the time and heavily involved in these Miocene investigations, before being loaned – yes, loaned – to the British Museum in London. The fate of this specimen is a long story, since Richard (the second of Mary’s three sons) later aimed to retain it for the National Museum in Kenya, only to be told by the British Museum that it hadn’t been loaned but gifted. This story does, however, end with the specimen back in Kenya in 1982 (Leakey 1984, pp. 100-101).

Later Rusinga finds included fossils of rhinos, rodents, arthropods, a tree ant nest and a remarkably complete lizard head that Mary described as preserved with its “tongue hanging out” (Leakey 1984, p. 103). I recognise this as a description of the gerrhosaurid described by Richard Estes in 1962 (Estes 1962).

**Caption:** the remarkably well-preserved gerrhosaurid lizard head and neck from Rusinga Island, as described by Estes (1962). Estes proposed that it should be referred to the extant *Gerrhosaurus* and perhaps even to the modern species *G. major* (he had it as ‘cf *G. major*), and authors since have supported this, so far as I know... though *G. major* is now *Broadleysaurus* and excluded from *Gerrhosaurus*. The scales, teeth, left eye and tongue are preserved, in part replaced by calcite. Images: Estes (1962).

Louis’s amazing productivity is documented by the fact that he wrote his second autobiography (White African) in 1936 in addition to another book – Stone Age Africa – in the same year. If the title White African seems odd, it’s related to the fact that Louis was recognised as a member of the Kikuyu people and very much saw himself as one of them. Included among the Kikuyu was the anti-colonial Mau Mau movement. Louis was a Mau Mau opponent, writing two critical books of 1952 and 54 on this subject, and working with the Keynan government to gather intelligence and broadcast propaganda. I, of course, know nothing proper of Kikuyu culture nor what it was like to be Louis Leakey so will withhold further comment. But for Mary and her family, this must have been concerning, since it put them at risk of terrorist attack.

Mary’s work on rock art. Mary’s interest in and study of African rock art is a constant throughout the book. Despite plans during the 1950s, neither Mary nor Louis were able to raise the funds needed to produce a large and lavish book of the sort the subject deserved, and such wasn’t achieved until 1983 with the publication of Africa’s Vanishing Art (Leakey 1983). There is today widespread awareness of the fact that climate change, vandalism and other factors are affecting the persistence of ancient rock art. Things weren’t quite as bad in the 1950s but there was awareness of rock art deterioration, and efforts were made to record and reproduce the art as well as protect it.

**Caption:** as is so typical of rock art, much of that studied and recorded by Mary involves astute and often brilliant observation… though some things are schematic or even symbolic and not meant to be technically accurate. At left, the ‘hartebeest frieze’ of the Kondoa region, Tanzania, showing animals grazing, feeding and ruminating. At right, ostriches and white rhinos at Kisese in Tanzania. Images: Leakey (1984).

How might people “record and reproduce” rock art? Tracings were made on large cellophane sheets, then transferred to drawing paper. The images were then reconstructed on moulded, artificial rock surfaces for museum display. As for protection, the local Warangi people had great interest in rock art and were paid to oversee it (Leakey 1984, p. 109). Mary mentions especially extravagant, very special art that existed at a rock shelter whose location was kept secret. She was never able to see it, let alone locate it, as is appropriate.

Across her recording efforts, 186 sites were catalogued, 43 of them with good art. Together, they featured over 1600 illustrated figures. As is the case for so much rock art across the ancient world, the images reveal a good understanding of form, anatomy and behaviour when it comes to non-human animals, combined with what are assumed to be exaggerations made for artistic effect. One notable example concerns a kudu with 11 horn spirals (Leakey 1984, p. 106).

**Caption:** some effort to date the rock art studied by Mary (and other scientists) since her primary contribution was published in the 1980s does exist, and Bwasiri & Smith (2015) explained the various difficulties inherent to this field. This image from their article shows Mary’s interpretation of a scene above, and their photo of the actual panel below. I think it’s clear that her illustration is highly accurate. How the scene should be interpreted is another matter (Bwasiri & Smith 2015).

‘Zinj’ is found. The 1950s were also the time when Mary and Louis made significant improvements in understanding the stratigraphy and chronology of the sediments of Olduvai Gorge, this work showing that the lower beds there were much older than previously thought.

**Caption:** at left, one of the many casts now in existence of the *Paranthropus boisei* holotype OH5, this one on show in the Springfield Science Museum, Massachusetts, USA. At right, life reconstruction of this species by Cicero Moraes and based on a 3D scan of the skull by Dr. Moacir Elias Santos. Images: Daderot, public domain (**original here**); Cicero Moraes, CC BY-SA 4.0 (**original here**).

While exploring a site there during July 1959, Mary noticed an unusually thick chunk of bone from the mastoid region of a hominid skull (Leakey 1984, pp. 120-121). This was the first discovery of the remarkable animal soon to be named Zinjanthropus, known today as Paranthropus boisei. I find it a bit remarkable that Louis “was sad that the skull was not of an early Homo, but he concealed his feeling well and expressed only mild disappointment” (Leakey 1984, p. 121). Louis believed that australopithecines were not part of the evolutionary lineage leading to Homo sapiens but a side branch, and he thus held out hope for the discovery of Homo-type animals as old as the oldest australopithecines (Leakey 1984, p. 123).

‘Zinj’ – published in Nature in 1959 by Louis alone (Leakey L. 1959) – inspired massive international interest and led to the winning of new financial support. Other palaeoanthropologists had the chance to examine the actual specimen when the Leakeys hand-carried it by plane, much as Mary had her famous Proconsul specimen, to the Third Pan-African Congress held at Kinshasa (then Léopoldville) in what is today Democratic Republic of the Congo. Doubts were already expressed about the placement of the specimen in its own genus, and Phillip Tobias – gifted by Louis the task of co-producing a more detailed study – was one of several to suggest allocation to Australopithecus. Mary says that Tobias was also responsible for the moniker ‘nutcracker man’ and first used it at the Kinshasa meeting (Leakey 1984, p. 125).

**Caption:** the discovery of the original *Paranthropus boisei* skull was deemed a sufficiently notable event that this special plaque was erected in Olduvai Gorge, Tanzania (this is the metal version; it had a predecessor made of what I assume is marble). As you see from this 2012 photo, people have taken to using it as a collection point for fossils, interesting rocks, bones and such. Image: George Lamson, CC BY-SA 2.0 (**original here**).

  **Caption:** Mary and Louis in 1962, and thus photographed at a point between the discovery of *Zinjanthropus* and the publication of *Homo habilis*. I’m not entirely sure which (probable) hominin fossil Louis is holding, but several photos showing Mary, Louis and this fossil were taken, so it was evidently newsworthy. Image: Smithsonian Institution Archives, public domain (**original here**).

Hello ‘Homo’ habilis. Additional remarkable finds were made at Olduvai during the early 1960s when parts of a more gracile, larger-brained but contemporary species were recovered. This was ‘handy man’, a creature that seemed to conform to the ‘early Homo’ that Louis hoped to discover, and of interest is that Louis apparently refrained from formally including it within Homo until he, Phillip Tobias and John Napier were ready to publish it in Nature in 1964 (Leakey 1984, pp. 127-128).

The name ‘Homo habilis’ was suggested by Raymond Dart but doubts about the inclusion of this animal within Homo – a well-known point of contention among specialists over recent decades – were expressed immediately by Le Gros (Leakey 1984, p. 128). Mary reflects on this point later in the book, her argument being that the larger brain size and association with diverse tools made inclusion within Homo more likely than an australopithecine suggestion. An interpretation I was previously unaware of is the argument (not attributed to a specific person) that H. habilis was “an advanced form of Australopithecus africanus” (Leakey 1984, p. 214). An additional notable find was made soon afterwards, this time of a skull identified as Homo erectus.

**Caption:** at left, a cast of the type juvenile individual of *Homo habilis* (OH-7), a mandible with dentition and parietal bones (an upper molar and hand bones were recovered as well), as displayed at Museo Arqueológico de Regional de Madrid. At right, the Koobi Fora (Kenya) specimen KNM-ER 1813 discovered in 1973, referred to *H. habilis* and integral to arguments that this taxon was sexually dimorphic. Images: Nachosan, CC BY-SA 3.0 (**original here**); Don Hitchcock, CC BY-SA 4.0 (**original here**).

The Pleistocene proves longer, and to have started earlier, than thought. One of many debates to occur among specialists across the time these discoveries were being made and interpreted was that concerning the age of the respective sediments. In 1960, geophysicist Jack Evernden from Berkeley visited Olduvai with plans to carry out potassium-argon dating, work that was then carried forward by Garniss Curtis. The results indicated that the Pleistocene was much longer in duration than the Leakeys had thought, the deeper parts of the Pleistocene being perhaps a million years older than considered beforehand (Leakey 1984, p. 131).

**Caption:** Olduvai Gorge as it looked in 2011. Exposed and actively eroding sedimentary layers are good for fossils, assuming of course that those layers contain a reasonable number of them. Almost 100 Pleistocene hominin fossils have been discovered at this location. Image: Mike Krüger, CC BY-SA 4.0 (**original here**).

Mary explains how this was controversial for a time, it meaning that the fossils that the Leakeys were working with were potentially far more distant, in evolutionary terms as well as chronological ones, from modern humans than they’d been thinking. Incidentally, the mention here of Evernden and Curtis ties the Olduvai story to that of another book I discussed here in 2025, Garniss Curtis, Carl Swisher and Roger Lewin’s 2000 Java Man: How Two Geologists Changed the Course of Human Evolution. You can read my article on that book here. That book explains how Louis so disliked the idea that Zinj might be almost two million years old than he “refused to believe dates that Garniss and Evernden produced for other fossils”, since “the dates did not jibe with what Louis wanted or believed” (Curtis et al. 2001, p. 19).

Louis’s decline. As is clear already, Louis Leakey was a major part of Mary’s life from the 1930s onwards. But the period extending from 1968 to 1972 marks one of a decline both in their relationship and in Louis’s health and faculties. Mary describes Louis’s growing competition with Richard, his own son, as well as his “disastrous” association with the Calico Hills discoveries of the Mojave Desert (Leakey 1984, p. 142). These involved numerous chert fragments, suggested to be human-made artefacts, and thought by Louis to perhaps be 80,000 years old. Mary disagreed not just with this interpretation but with everything about how the work was being carried out, and when Louis arranged a conference devoted to discussion of the site she felt that those who should have provided condemnation were holding back due to their sympathy for someone who, by now, was quite ill.

**Caption:** the Calico Hills story is fascinating and, in part, a case of people fooling themselves into thinking that the ‘evidence’ they had was far better than it was. At left, Calico Hills at Red Rock Canyon National Conservation Area. At right, a selection of the alleged hominin-made lithics from the site, from a 1938 publication but taken from Dempsey (2009). Images: Frank Schulenburg, CC BY-SA 4.0 (**original here**); Dempsey (2009).

Mary and Louis also disagreed on whether they should accept honorary degrees offered by the University of Witwatersrand in South Africa. Louis believed that these should be rejected in protest of apartheid. Mary thought that to do so would be a snub to a university that was doing what it could to fight against apartheid already, and to the colleague who had arranged these awards, Philip Tobias. Mary ultimately accepted the degree, and Louis was furious.

Towards the end, Louis was beset by health problems and continued to push himself too hard, never taking appropriate time to recover. For a while, he was cared for by Vanne Goodall, Jane’s mother, and he died in 1972.

**Caption:** the ‘site A’ hominin footprints of Laetoli, thought for a while to be those of a bear but now regarded once again as those of a hominin bigger and more robust than the taxon that made the majority of hominin footprints at the site. These show, at top, an image obtained via photogrammetry and, below, a contour map generated from a 3D surface scan. Image: **McNutt *et al*. (2021).**

The Laetoli tracks. Much else occurred in the 1970s, an event of international interest being the 1976 discovery of the Laetoli footprints. These included not just the famous ones made by hominins, but those made by numerous other animals too. Hadza hunters were used in the identification of these. An interesting controversy concerned the identification of the four ‘site A’ footprints that might have been created by a hominin but were suggested by some to be bear tracks, and here Mary takes time to remind readers that bears – not ordinarily associated with Africa bar the Atlas bear of the far north and, err, the semi-mythical Nandi bear I suppose – were a feature of the east African biota at this point in prehistory. Mary’s identification of the site A tracks as those of hominins has recently been vindicated, this work showing that ancient Laetoli was inhabited by two hominin taxa at least (McNutt et al. 2021).

By 1979, the more famous Laetoli hominin trackway, an impressive 24 m long and involving the footprints of three individuals, had been discovered. Such was the global importance of this site that plans were made to preserve it, initially via the creation of another open-air museum. This didn’t work out due to the remoteness of the site, so the decision was made to preserve the tracks and rebury the entire layer using river sand and plastic sheeting.

**Caption:** at left, Laetoli footprints as preserved at Site S, showing impressions left by hominins as well as horses, rhinos, giraffids and guineafowl. The grey areas at the rear of the hominin footpints are heel drag marks. At right, a photo of one of the most famous sections of the Site S trackway. Footprints made by small horses and other animals are on the same bedding plane. Images: **Masao *et al*. (2016)**, CC BY-SA 4.0.

Hadar, the ‘first family’, and Lucy. An especially interesting set of discoveries relative to Mary’s finds and research concerns the 1974 finding in Hadar, Ethiopia of Lucy by Donald Johanson, and the ‘first family’ in 1975. I was aware of some disagreement between Mary and Johanson, and Tim White too, over what happened next, but this was the first time I’ve read anything halfway lengthy about it.

Mary’s opinion was that the Hadar hominin fossils were substantially superior to those from Laetoli, and not from the same species. The Laetoli remains were not just found more than 1600 km away from Hadar but were around half a million years older. So she was unhappy with Johanson’s decision to use a Laetoli mandible – the LH4 specimen that Mary discovered in 1974 – as the holotype for Australopithecus afarensis, so much so that she asked to be removed from the authorship of the paper that named this taxon, this being Johanson et al. (1978) (Leakey 1984, p. 182). These comments aren’t unique to Mary’s book, by the way: there’s a complex to and fro in the technical literature on these matters (Johanson & White 1979, 1980, Day et al. 1980, Leakey & Walker 1980, McHenry & Corruccini 1980).

**Caption:** at left, a cast of the *Australopithecus afarensis* ‘Lucy’ skeleton Al 288-1 at Museum National d’Histoire Naturelle, Paris. At right, a cast of the LH4 *A. afarensis* holotype mandible from Laetoli, on show in a temporary exhibit at the Museo Arqueológico de Regional de Madrid. Images: 120, CC BY 2.5 (**original here**); Nachosan, CC BY-SA 3.0 (**original here**).

Mary also expressed her disagreement with the Johanson et al. view that A. afarensis held an ancestral position for later australopithecine species and Homo as well. She also took issue with comments made in Johanson and Maitland Edey’s 1981 book Lucy: the Beginnings of Humankind – she noted that her book is the only place where she can do it – and pointed out that quotes reported therein as historical events can only be fictional, since neither author was there to hear them. You may recall from earlier Louis Leakey’s apparent disappointment at Zinjanthropus being what it was, rather than the key Homo species he was hoping it might be. Johanson and Edey have it that Louis said that Zinj was “nothing but a god-damned robust australopithecine” on seeing the find. These words do have something of a Hollywood flair, but the fact is that they were never said (Leakey 1984, p. 183). Johanson and Edey’s book on Lucy is on my reading list, so I’ll be returning to this topic in time.

**Caption:** a 1981 edition of Donald Johanson and Maitland Edey’s *Lucy: The Beginnings of Humankind*. It’s an attractive book with a good number of photos and diagrams, but it contains points of fact that were contested by Mary Leakey.

20 million years of research. It goes without saying that the contribution of the Leakeys to palaeoanthropological research, and our understanding of prehistory in general, has been vast. Mary ended her autobiography by summarising the main discoveries she was involved in and what they mean for the big picture overall.

On that note, it’s almost unfathomable today to think that someone could perform the important and lasting studies she did with respect to such an incredible range of topics, these involving Miocene hominoids or near-hominoids like Proconsul, Pliocene and Pleistocene australopithecines, paranthropines and other stem-humans, and archaeological work on ancient humans, and on their art and artifacts in Africa as well as Europe.

Mary died in 1996, aged 83, and I think that this book does a good job of charting her life and work. I really enjoyed reading it and gaining so much insight into this important phase in palaeoanthropological history. There are, as stated earlier, other books on and by the Leakeys, and I’ll be writing about them too, in time.

**Caption:** certain of the books discussed and mentioned here, plus others connected to the life and work of the Leakeys. Yes, that includes work on living great apes, since the authors of those volumes have connections to Louis Leakey. Image: Darren Naish.

For previous Tet Zoo articles on hominins and other hominids, see…

Zihlman’s ‘pygmy chimpanzee hypothesis’, October 2012
Why Humans Are Important to Studies of Primate Diversity, July 2014
The Cautious Climber Hypothesis, March 2019
Piltdown Man and the Dualist Contention, March 2023
Curtis, Swisher and Lewin’s Java Man of 2000: Hominin-Themed Books, Part 1, March 2025

Huge thanks to those who help support this blog at patreon. You can see behind-the-scenes and in-prep stuff relevant to projects for as little as $1 per month.

Refs - -

Bwasiri, E. J. & Smith, B. W. 2015. The rock art of Kondoa District, Tanzania. Azania: Archaeological Research in Africa 50, 437-459.

Curtis, G., Swisher, C. & Lewin, R. 2001. Java Man: How Two Geologists Changed the History of Human Evolution. Little, Brown and Company, London.

Day, M. H., Leakey, M. D. & Olson, T. R. 1980. On the status of Australopithecus afarensis. Science 207, 1102-1103.

Dempsey, P. J. 2009. Calico – the Olduvai Gorge connection. Society for California Archaeology Proceedings 22, 1-13.

Estes, R. 1962. A fossil gerrhosaur from the Miocene of Kenya (Reptilia: Cordylidae). Breviora 158, 1-10.

Johanson, D. C. & White, T. D. 1979. A systematic assessment of early African hominids. Nature 203, 321-330.

Johanson, D. C. & White, T. D. 1980. On the status of Australopithecus afarensis. Science 207, 1104-1105.

Johanson, D. C., White, T. D. & Coppens, Y. 1978. A new species of the genus Australopithecus (Primates: Hominidae) from the Pliocene of Eastern Africa. Kirtlandia 28, 1-14.

Leakey, L. 1934. Adam's Ancestors: The Evolution of Man and His Culture. Metheun & Co., London.

Leakey, L. 1959. A new fossil skull from Olduvai. Nature 184, 491-493.

Leakey, M. 1983. Africa’s Vanishing Art: the Rock Paintings of Tanzania. Doubleday, Garden City, New York.

Leakey, M. 1984. Disclosing the Past: An Autobiography. Weidenfeld and Nicolson, London.

Leakey, R. E. F. & Walker, A. 1980. On the status of Australopithecus afarensis. Science 207, 1103.

Masao, F. T., Ichumbaki, E. B., Cherin, M., Barili, A., Boschian, G., Iurino, D. A., Menconero, S., Moggi-Cecchi, J. & Manzi, G. 2016. New footprints from Laetoli (Tanzania) provide evidence for marked body size variation in early hominins. eLife 5:e, 19568.

McHenry, H. M. & Corruccini, R. S. 1980. On the status of Australopithecus afarensis. Science 207, 1103-1104.

McNulty, K. P., Begun, D. R., Kelley, J., Manthi, F. K. & Mbua, E. N. 2015. A systematic revision of Proconsul with the description of a new genus of early Miocene hominoid. Journal of Human Evolution 84, 42-61.

McNutt, E. J., Hatala, K. G., Miller, C. Adams, J., Casana, J., Deane, A. S., Dominy, N. J., Fabian, K., Fannin, L. D., Gaughan, S., Gill, S. V., Gurtu, J., Gustafson, E., Hill, A. C., Johnson, C., Kallindo, S., Kilham, B., Kilham, P., Kim, E., Liutkus-Pierce, C., Maley, B., Prabhat, A., Reader, J., Rubin, S., Thompson, R., Thornburg, R., Williams-Hatala, E. M., Zimmer, B., Musiba, C. M. & DeSilva, J. M. 2021. Footprint evidence of early hominin locomotor diversity at Laetoli, Tanzania. Nature 600, 468-471.

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Foot deformities are ubiquitous in urban pigeons. Why?

**Caption:** poor toe-less or even mostly foot-less urban pigeons, one encountered in Lisbon in 2014, the other (on the right) seen close to Kew train station in September 2014. The right foot of the Kew bird is completely absent; the left one had at least one toe, curled round such that the bird was walking on the toe's lateral side. Image: Darren Naish.

Here’s a republished version of an article that originally appeared (here, at ver 3) back in 2022.

As you’ll know if you’ve spent any time watching the pigeons of towns and cities, something like one in every ten has missing or partial toes, or swollen toes, or other pedal deformities of some sort. And then there are really extreme individuals: the ones that are missing the majority of their toes altogether.

**Caption:** that bird from Lisbon again, showing how badly mangled its feet are. Image: Darren Naish.

I’m talking here about the domestic form of Columba livia, the so-called Rock pigeon or Rock dove. And, while I’ve mostly paid attention to deformed pigeons here in the UK, it certainly isn’t a UK-only thing, as continental European and North and South American people (and no doubt those of elsewhere in the world too) will confirm.

The Kew bird in the photos here was able to fly around and feed itself, and it might be broadly described as ‘healthy’. However, note that it’s in pretty poor condition. The feathers on its head and neck looked terrible and its rectrices (the big tail feathers) were frayed and shabby. I reckon this is partly due to an inability to groom and scratch itself: obviously, birds use their feet to reach parts that they can’t get to with the bill.

Looking nice, keeping clean. The impact of this is more than cosmetic, since birds with a poorly maintained or unrepaired plumage are disadvantaged in flight relative to tidier individuals, and less able to keep themselves warm and waterproofed. And a bird that isn’t able to groom parts of its plumage is also at risk of being unable to keep on top of parasites like ticks and feather mites. As demonstrated through various experiments, birds with shabby-looking plumage are less attractive as mates (Clayton 1990), so a bad-looking pigeon is likely to be a non-breeding pigeon. It’s also worth noting that (like many animals), pigeons preferentially use one foot more than the other as goes the way they land and perch and sit and so on (Fisher 1957). Individuals that lose or damage their preferred foot might, then, end up being doubly disadvantaged (imagine being right-handed, and then having to rely only on your left hand for evermore). I don’t know how concerned people are about the emotional well-being of animals like urban pigeons (or how far they’re prepared to go in admitting that non-human animals have feelings and states of mind), but I think we can be confident that the most severely deformed of these birds are - at least at times - miserable, unhappy and frustrated.

**Caption:** toe damage is very common in urban pigeons, but we mustn’t think that it’s ubiquitous. In this pigeon group in Bristol, photographed in 2023, I can only see one missing toe (on the right foot of the bird at extreme upper right). UPDATE: ok, I’ve now noticed a second pigeon with a mangled foot. The bird just left of the one at upper right has a club-like left foot. Image: Darren Naish.

The ‘tangled’ or ‘stringfoot’ hypothesis. Several ideas have been put forward to explain the many foot problems seen in urban pigeons. Some probably lose toes after getting them tangled in litter or anti-pigeon netting, or after they've received injuries from anti-pigeon spikes installed on signs and ledges. Fine wire, string, cotton thread and even human hair can all cause problems for birds when caught on or around digits, and some people say that interaction with fine thread and string and so on is the primary cause of pigeon foot damage. My impression is that this ‘tangled’ hypothesis, causing what’s increasingly known as stringfoot in birds, is the most likely explanation for the problem.

The ‘chemical injury’ hypothesis. It’s also sometimes suggested that the deformities result from infections received after standing on excrement, and that the birds become damaged through interaction with chemicals used on roofs and building stone. This ‘chemical injury’ idea is unlikely to be correct, since (a) exactly what sort of chemicals are we talking about here, and why have they been used on buildings in the first place?, and (b) a chemical would basically have to be a powerful acid or alkaline agent (hydrochloric acid, or a very strong bleach) before it could damage a bird’s feet. For completeness, note that hereditary deformities like those reported for some captive populations of other pigeon species (Flach & Cooper 1991) might also explain some abnormalities observed in urban pigeons.

**Caption:** pigeons are frequently encountered as roadkill here in the UK. The Common wood pigeon *Columba palumbus* at left was dead at the side of the road and its lack of rectrices show that it had undergone ‘terror moulting’ prior to death, though whether this happened literally moments before it expired or hours or days before is unknown to me. At right, a juvenile wood pigeon on a road. Its position implies that it fell from the tree above before being hit by a car. Images: Darren Naish.

  **Caption:** pigeons do have to deal with a list of predators, even in urban environments. Falcons and domestic cats are pigeon-killers of course, but so are gulls and corvids. This Lesser black-backed gull *Larus fuscus* at Bristol train station learnt how to kill feral pigeons and I watched it do this once. Image: Darren Naish.

And we’ll end things there. For previous TetZoo articles on pigeons, see…

The detachable tails of pigeons, September 2008
Are parrots actually pigeons?, May 2009
Giant club-winged pigeons and ninja ibises: clubs, spurs, spikes and claws on the hands of birds (part III), February 2011
Footless Urban Pigeons Suffer Untold Trials and Tribulations, September 2014
Voyeurism and Feral Pigeons, June 2015
The Asian Dove That Ate Europe Alive, January 2017
Pigeons and Doves 101, March 2018
The Dodo in Life, July 2020
Otidiphaps the Pheasant-Pigeons, April 2022

My writing and research is dependent on crowd-funded support. Thanks to those whose patronage made this article, and the others you read here, possible. Please consider assisting me if you can, thank you!

Refs - -

Clayton, D. H. 1990. Mate choice in experimentally parasitized rock doves: lousy males lose. American Zoologist 30, 251-262.

Fisher, H. I. 1957. Footedness in domestic pigeons. The Wilson Bulletin 69, 170-177.

Flach, E. J. & Cooper, J. E. 1991. Clinical and pathological findings in two Mauritian pink pigeons (Columba mayeri). Veterinary Record 129, 48-51.

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Once again it’s that time of year, by which I mean… spawnwatch season, of course.

**Caption:** peak spawning activity, occurring on 10th February. One important behavioural aspect to note is that activity occurs throughout daylight hours… just as it does throughout the night as well. I think that at least some frogs go for weeks without sleep during this period. Image: Darren Naish.

Yes, early February here in far southern England means that it’s once again that time when the lone amphibian species living in the grounds of Tet Zoo Towers – the Common frog Rana temporaria – gathers to breed. As ever, I’ve been keeping close tabs on things, so let’s see what happened this time round. There’s good news and bad news.

Some background. I’ve been maintaining ponds here at Tet Zoo Towers since 2006, initially with a rigid moulded pond, and more recently a large, pondliner-based one. Over that time, the number of adult frogs appearing in the ponds has increased from one or two to over 50, the number of spawn clutches increasing from one to a maximum of 31 (for 2024). In 2025, a cold weather spell caused an unusual break in breeding activity, and we never got above 18 clutches, which was surprising in view of the previous year’s high. I became worried that breeding females (which are typically the biggest frogs in the population and hence the most vulnerable to predation and maybe other things too) had disproportionally died off, and that the otherwise cumulative increase in our population was at an end, or even in reverse.

**Caption:** this is Plumpmot, a large and attractive female, here photographed in October 2023 with an earthworm. I didn’t see her during the breeding events of early 2026 and am not sure if she’s still around. Big females like this are especially important members of the population. Image: Darren Naish.

I should also add that the garden in which these ponds are sited is deliberately maintained for frogs and other animals. It’s a total mess and stands out markedly in contrast to the paved driveways and concrete death-yards otherwise so typical of the modern UK. There are tall hedges, piles of sticks, and any grassy areas are deliberately unkempt. In an age when people are, mostly, doing everything they can to obliterate every single scrap of greenery, I strongly believe that those of us who care should do everything we can to help the persistence of wildlife. We are fighting against those who want everything to be featureless concrete.

**Caption:** the look of pond 2 during early February following a couple weeks of constant, heavy rain. The water level is really high, and frogs are visible in parts of the pond that would be high and dry once the rain stopped. Note the vegetation and dead branches around the pond. These are there to provide refuge for frogs and deter pet cats. Image: Darren Naish.

As per usual, let me remind people in regions and nations that operate under what’s known as a continental climatic regime that, yes, the gathering and breeding of frogs at this time of year (early February) is normal for the mild, high latitude, maritime fringe archipelago in which I reside. A few decades ago, Common frogs here routinely started to gather and breed during the second week of February. Now, the first week is usual. Further to the west, in English counties like Cornwall, Common frogs regularly spawn in January, and late December spawnings are becoming increasingly common. I have a feeling that I’ve said all of this before, forgive me.

For a great overview of frog natural history in the UK you should consult Beebee and Griffiths (2000). More concise volumes include Inns (2009) and Beebee (2013). Cooke’s (2023) Tadpole Hunter is a very heavy and in-depth personal view of amphibian research in the UK that I’ve really enjoyed reading.

Things get underway. Male Common frogs are quite probably in a breeding pond throughout the whole of the winter, and hence ready to intercept arriving females and get things underway as early as possible. This year, a reasonable number of frogs (12) were readily visible on February 3rd, and two spawn clutches appeared on that day: a new earliest-ever record here (the previous was February 4th, for 2025). Constant heavy rain meant that the pond was at an all-time high, a fact that would have consequences once the rain actually died down…

**Caption:** daytime activity on February 7th, showing multiple spawn clutches in the shallow parts of the pond while frog combat and competition is very much underway. Image: Darren Naish.

  **Caption:** Common frogs are basically all recognisable as individuals on the basis of the dark markings on their dorsal surfaces and limbs. I’ve been trying to keep track of individuals but getting the right kind of photos is hard. Silviu Petrovan at the University of Cambridge has been leading a project that does this more properly. Image: Darren Naish.

Over 43 frogs were visible by February 5th, and by February 7th a substantial jump to 19 spawn clutches had occurred, putting us above the 2025 total count. A day later, and I counted just over 50 frogs, and over the next several days the clutch count increased to 31 by February 11th, this number being the previous all-time record of 2024. After that, a slow and incremental climb to the current (as of February 17th) total of 37 occurred, a new all-time record. As you can see from the cumulative graph here, we’re now (at the time of writing) at an approximate plateau, activity has mostly stopped and the frogs have mostly dispersed. I won’t be surprised if one or two additional clutches appear, but we’re essentially there, and with a very impressive new clutch record. I hope to see this increase next year!

**Caption:** cumulative graph for 2026, showing how we quickly increased from two clutches on February 3rd to over 20 by February 8th, peaking with 37 clutches as of right now.

  **Caption:** spawn clutches first appeared in our ponds here in 2010 (initially in an old plastic baby bath), but this graph shows how things improved markedly after 2020.

A rant on pet cats. As I’ve said ad nauseum before – forgive me – free-roaming pet cats are a perpetual, never-ending nuisance when you’re aiming to garden for wildlife. I live in constant dread of them killing various of the animals I cherish the most. Lone cats can and do wipe out entire families of fledgling birds and kill enough lizards, bats, frogs and other small animals to make a real difference to limited, isolated populations where there’s no migration from outside.

And a problem is not just that a lone cat from a neighbour’s garden will visit a wildlife-friendly garden ten, twenty, thirty, forty times a day, often staying there for hours at a time if allowed, it’s that this is often just one of many individuals that do the same. There are that many free-roaming pet cats. I’ve been using camera trapping in the garden for a while (see this article) and it’s disheartening to see how many pet cats – three, four, five – visit our property and hunt there during the day and night. This year, the camera trap let me down and only functioned for a single night (January 25th, days before spawning had started) before giving up, but it recorded three different pet cats on that one night.

**Caption:** pet cat 1, on the evening of 25th January.

  **Caption:** pet cat 2, photographed a few hours later during the evening of 25th January.

For 2026, I observed one attack on a frog by a pet cat myself, purely by chance when I looked out of a window. It had pulled the frog out of the pond before being chased away, but the frog was lucky and survived. Several of our regular frogs have damaged eyes, bear scars and one (called Three-point-five) is missing a hindfoot, and it’s likely that cat attacks are among the events that have caused these injuries. At one pond I used to monitor, a pet cat killed or mortally injured 12 frogs in a day; it was total carnage and extremely depressing. Seeing as you’re reading this blog, it’s likely that you already care about the plight of beleaguered wildlife and are doing what you can to help it persist. But do what you can to help make others aware of this problem. I wish that things would change.

**Caption:** Common frogs probably select shallow spawning places since these are warmest and allow for the fastest development of the spawn and tadpoles, but the massing of the spawn that occurs might also afford protection, both thermally and in terms of ‘safety in numbers’. Images: Darren Naish.

To the future. I’ve surely said previously at Tet Zoo that I made extensive modifications to the bigger, newer pond at the end of 2024 such that it was substantially improved as a frog breeding site. Common frogs preferentially spawn in water less than 10 cm deep and large areas of very shallow water are ideal, not just for the act of spawning, but for fast development of the spawn and tadpoles. This in mind, I now create ponds with very large, shallow areas. And such it is with the modified pond here. It has worked to perfection, all the activity and spawning occurring in the very shallow end.

**Caption:** frog activity at the shallow end of the pond on 7th Feburary. Water level is high and very new spawn clutches are visible left and right. When spawn is fresh, it’s tight, compact and rubbery. As it ages, it soaks up water, expands and flattens out. Common frog spawn is remarkably resilient and extremely good at protecting the embryos from low temperatures, ice, rain, hail and mechanical damage.

  **Caption:** we routinely have over 50 adult frogs massing in the pond during the breeding season now. When you combine this with the substantial number of juveniles that must be living in the same area, our population must be high. An aim of any successful conservation effort must be to have the population spread to appropriate habitat nearby. I hope that’s happening, but it’s not helped by the fact that we’re right next to a large and perpetually busy road. Image: Darren Naish.

But something is wrong. Round about February 8th, the water level started going down, and it did this so much that a few spawn clutches in the very shallowest area were ultimately left almost out of the water. This has undone a fair amount of my cleverly planned landscaping and means that I’ll have to renovate things in the future. I have no idea what’s caused this but it can only be a puncture, though how this came about I have no good idea. This isn’t an immediate disaster, since even the somewhat high-and-dry spawn is ok (Common frog spawn spreads out and there’s still enough water in that part of the pond for the embryos, once free of the jelly, to remain in water), but it is a pain. At the time of writing (February 17th), much of the shallow end is dry and I’ve had to do an emergency rescue of all the spawn. I’m going to have to do a rebuild. Again. The spawn and tadpoles will be ok though, don’t fear.

**Caption:** February 17th, and the water level has dropped a worrying amount. All the spawn has been (temporarily) placed in a large shallow tray and the pond will need rebuilding. These things happen.

I guess a pondkeeper’s pondwork is never done.

For previous articles on frogs, ponds, and spawning events, see…

A Love Letter to the Common Frog, August 2020
Live Spawnwatch Action From Pond 2 at Tet Zoo Towers, February 2024
News From the Pond's Edge: Spawnwatch 2025, February 2025
Suburban Camera Trapping, Week 1, May 2025

Tetrapod Zoology is dependent on funds raised at patreon. Please help support this blog if you consider it worthwhile and want to see me continue doing it, thanks!

Refs - -

Beebee, T. 2013. Amphibians and Reptiles. Pelagic Publishing, Exeter.

Beebee, T. & Griffiths, R. 2000. Amphibians and Reptiles. HarperCollins, London.

Cooke, A. 2023. Tadpole Hunter: A Personal History of Amphibian Conservation and Research. Pelagic Publishing, London.

Inns, H. 2009. Britain’s Reptiles and Amphibians. WILDGuides, Old Basing, Hampshire.

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With – as of January 2026 – an impressive and respectable two decades of history behind it, Tetrapod Zoology surely has a notable footprint in terms of academic achievement and pop culture influence, right? Whatever, let’s look at assorted articles worth talking about. A list of highlights, if you will.

**Caption:** images relevant to 20 years of Tet Zoo, a mix of dinosaurs, amphibians (the coloured *Platyhystrix* and banded newt), cryptids (the long-necked seal), speculative beasts (the demonic *Quetz* and flightless Miocene azhdarchid) and a plesiosaur skull. The demonic *Quetz* is by Bob Hersey and the *Platyhystrix* was coloured by Gareth Monger.

A quick history of Tetrapod Zoology. At this point in history, there are a reasonable number of Tet Zoo articles out there in the wild, unfortunately spread over four different sites. Tet Zoo ver 1 is at blogspot.com and the articles there are essentially still intact. There’s some ok content there, but I was a total newbie to blogging (as most of us were in those days) and did many things that aren’t acceptable today. I’m talking here about image use. Tet Zoo ver 2 happened at ScienceBlogs. I have a lot to say on the history and ultimate fate of ScienceBlogs but I’ll burn up too much space and time if I go into it. All you need to know is that things dissolved into a mess and hence Tet Zoo ver 3 arose from the ashes at the new and shiny Scientific American blogging network. Things here were good for a while (and I was even paid to blog), but ultimately it became clear that Sci Am weren’t interested in hosting blogs. Their policies on image use, reader comments, site design and more were completely at odds with my plans… and thus I left, and here we are at Tet Zoo ver 4.

**Caption:** a classic piece of Tet Zoo lore, specifically from ver 3 and April 2010 (**here**). The panbiogeography debate – which was attached to an article about ratites rather than bears – resulted in the longest comment thread in the whole of Tet Zoo history. I doubt if it’s findable or viewable now though, since Sci Am was absolutely not interested in the hosting of comments and appears to have removed all the comments sections from older articles. Images: Darren Naish.

Things wouldn’t be so bad if all the content published at ver 2 (ScienceBlogs) and ver 3 (Sci Am) was findable and available online. Alas, the enshittification of the internet has not been kind to it, since these days said content is variously paywalled, ruined or absent altogether. Over the past few years I’ve done what I can to find intact versions of all the relevant articles at wayback machine – the results are archived here – but there’s still tons to do and I’m way behind. Massive thanks to those who’ve assisted me with this task. It’s so disheartening to have to do this. One aim is to publish all of this material in books but that’s something else that I haven’t had time for, yet. Tetrapod Zoology Book One (Naish 2010), published in 2010, marks a start on this project.

My apologies to those of you who’ve heard much of this before.

**Caption:** over the years, there have been a reasonable number of Tet Zoo-themed spinoffs, some successful and with longevity, others less so. In 2014 and 2015, Ethan Kocak and I collaborated in the creation of the Tet Zoo Comic and the good news is that **it’s still online today**. Looking at these panels again right now, I still find them hilarious. Image: Kocak / Naish.

Tet Zoo’s Two Decades of Content. When it comes to seeing what was good, and sometimes not so good, about Tet Zoo’s two decades of operation, the good news is that my birthday articles prove useful in discussing those events that influenced or had attachment to the blog, even if they didn’t form the basis of articles per se. At least some of this material is pretty entertaining, at least to me.

**Caption:** relevant images from **the 10th birthday article from 2016**. Tet Zoo Podcats Top Trumps are hard to get these days. The Pterosaur Heresies image was created by Gareth Monger.

On Tet Zoo articles themselves, there have been several efforts to count and tabulate them. Cameron McCormick made major progress at the Tet Zoo Wiki, which I think is now sadly defunct... I just had a quick look online and can’t find it at all, wah. The main point I’m making here is that there isn’t a complete list of everything that I’ve published anywhere, this making it hard to keep track of what is or was especially notable. So let’s keep this intuitive: which articles do I consider highlights? Last time I asked this question – this was for the blog’s 5th anniversary – I invited friend and super-fan Albertonykus of Raptormaniacs to list personal highlights. The resulting article was great fun… but of course it was hosted at SciAm (I think) so now I can’t find it either [UPDATE: it’s here. Thanks to Albertonykus for finding it].

**Caption:** what was happening at Tet Zoo approximately ten years ago? By checking **part 1 of the 10th birthday series of articles**, I see that topics covered included taxonomic vandalism, the resurrection of *Brontosaurus*, the cult of turning live monitor lizards into giant geckos, and the scansoriopterygid *Yi qi*. I also published the notable April Fool’s article **Cetacean Heresies: How the Chromatic Truthometer Busts the Monochromatic Paradigm** in 2015 (try as I might, I cannot find an intact version, hence no link… UPDATE: **HERE IT IS**, thanks to David Gioia). This really nice illustration of a fancifully coloured *Peponocephala* (melon-headed whale) from that article is my line work, but all the colouring is by Gareth Monger. Image: Darren Naish and Gareth Monger.

Ok, enough preamble. The rest of this article is devoted to articles (grouped together by subject) that I consider among the highlights. It’s not complete nor especially thorough and I’d be interested to know if you, dear reader, have articles not listed here that you consider faves or highlights yourself. Ok, let’s get to it…

Frogs, salamanders, caecilians, oh my! It’s not all charismatic megafauna here and indeed never has been, since I’ve done everything I can to cover obscure small lizards, the incredibly diverse and abundant rodents, and the ever-fascinating and weird amphibians. Amphibians – the anurans (frogs and toads), the caudatans or caudates (salamanders) and the worm-like caecilians – have been a mainstay here, and I’ve made numerous efforts over the years to review the full diversity of these groups. These haven’t ever been fully successful, partly because doing this sort of thing takes forever, and partly because progress has been ruined by the fails of Tet Zoo ver 2 and 3, as discussed above. Indeed, I currently haven’t found intact versions of most ver 2 and 3 salamander- or frog-themed articles (a reminder that I’m compiling a list of archived articles here). Whatever, a lot of amphibian content has been published at Tet Zoo.

**Caption:** salamanders of several groups have been covered at Tet Zoo (albeit on an insufficient number of occasions), and I’ve released at least one article on giant salamanders, also known as cryptobranchids (**published 2010, it’s here**). Several species belonging to this group inhabited Europe during the geological past, among which is *Andrias scheuchzeri*, here shown in a 1964 illustration by Zdeněk Burian. At right, my cartoon representation of life during the giant salamander breeding season.

Let me say also that you can’t discuss amphibian diversity without focusing on conservation, the impacts of the climate crisis in global terms, and the impacts of habitat loss everywhere, this extending all the way down to what people are doing with the little bits of land near their houses (I live in the nature-depleted UK, where people are ceaselessly doing everything they can to get rid of every single scrap of greenery). On that last point, I’m doing what I can on a tiny local scale to help the one amphibian species that occurs where I live (the Common frog Rana temporaria) and have written about my local frog population quite a lot. I think that all of this is valuable content, and here are some of the articles that I feel are most interesting and impactful…

A Love Letter to the Common Frog, August 2020
Tiny Frogs and Giant Spiders: Best of Friends, August 2022
The Remarkably Weird Skeletons of Frogs, September 2022

Caption: over the past few years, pond 2 at Tet Zoo Towers has seen the laying of between 17 and 31 egg clutches. So it’s hard to believe that what you see in the photo here was the situation we had in 2012: a single clutch, and just two or three adult frogs. The pond shown in the photo here (pond 1) is not good for this species (the Common frog) being deep and choked with duckweed, but frogs will make do with what’s available. Image: Darren Naish.

I haven’t yet recovered intact versions of my 2009 articles on caecilians (these relating to a special event held at the Zoological Society of London) nor do I feel that I’ve given that particular group proper coverage. Yeah yeah, everyone knows about skin-eating and matrotrophy at this point; let’s talk instead about the dangerous poisonous species, the big aquatic ones that secrete themselves in submerged mud burrows, and the tiny species that have been evolving in isolation on Indian Ocean islands since, I dunno, the Cretaceous.

Azhdarchid Revolution. There’s no denying that azhdarchid pterosaurs – sometimes famously big, fantastically long-necked, and long-jawed too – have been a Tet Zoo mainstay since quite early on. A prominent event in my history as a publishing scientist is the 2008 publication of the initial ‘terrestrial stalking’ hypothesis with Mark Witton (Witton & Naish 2008), and that research is both related to earlier (and now mostly erroneous) musings shared at ver 1 as well as promotion that occurred once it was out.

It’s been extremely exciting to see our understanding of azhdarchids evolve and expand since then, both as we’ve learnt a lot more about azhdarchid anatomy and biology and as new taxa and specimens have been incorporated into phylogenetic studies. Here are links to my favourite, most notable azhdarchid-themed Tet Zoo articles…

Terrestrial stalking azhdarchids, the paper, May 2008
Quetzalcoatlus: the evil, pin-headed, toothy nightmare monster that wants to eat your soul, August 2013
Were azhdarchid pterosaurs really terrestrial stalkers? The evidence says yes, yes they (probably) were, December 2015
Some Azhdarchid Pterosaurs Were Robust-Necked Top-Tier Predators, February 2017
The Quetz Monograph Lives and Other News on Azhdarchid Pterosaurs, December 2021
Azhdarchid Progress, a Personal View, October 2024

Caption: azhdarchids (and related pterodactyloid pterosaur groups) have been covered a fair bit here over the years, though it’s been a long time since I’ve written about our changing views on their life appearance and behaviour. At left is an image from the 2000s depicting Quetzalcoatlus as stork-like, but as a wader adapted for grabbing aquatic invertebrates. At right is Richard Orr’s 1980s illustration of ‘demonic Quetzalcoatlus’.

Dinosaurs of non-feathered sort. As I often say, I don’t know whether I cover (non-bird) dinosaurs too often, or too…. not often. I tend not to cover dinosaur news stories at Tet Zoo, even though I find them exciting and sometimes have an insider perspective, since I feel that those stories are getting more than sufficient coverage elsewhere online. Look at the recent Nanotyrannus coverage: you couldn’t move for Nanotyrannus stories online for a couple of weeks.

The work that Mike Taylor, Matt Wedel and I published on neck posture in sauropods (Taylor et al. 2009) was great fun, associated with major interest from the media, and is connected to a substantial amount of research and discussion that’s been published since. The spinosaurid work that I’ve covered here – relating to the discovery of new baryonychine specimens and taxa in the Lower Cretaceous Wealden of southern England (Barker et al. 2021, 2022, 2023) – marks a highlight as does my recollections of the Dinosaurs Past and Present exhibition of the late 1980s and early 90s.

Sauropod dinosaurs held their necks in high, raised postures, May 2009
Recollections of Dinosaurs Past and Present, the 1980s Exhibition, February 2019
Two New Spinosaurid Dinosaurs from the English Cretaceous, September 2021

Caption: an argument might be made… might… that the concept of sexual sexual wasn’t sufficiently incorporated into studies of Mesozoic dinosaurs (and pterosaurs) prior to the 2010s. Among the several papers published during the opening salvos of 2011-2013 was Hone et al. (2011) on mutual sexual selection, a study discussed here at Tet Zoo ver 2 in 2012. Our initial thoughts on this topic were owed to our familiarity with work on such extant animals as wildfowl, seabirds and passerines. My Crested auklet Aethia cristatellus illustration at right featured in that paper.

Cassowaries and other ratites. Another of my favourite animal groups are the cassowaries, another extravagant and formidable group of ornithodirans (are you noticing a theme?). I’ve done my best to write about these incredible birds, and what we know and still don’t know about them, at Tet Zoo. This has led to technical research (Naish & Perron 2016) and also an association with cassowary expert and researcher Todd Green, this leading to my involvement in the once-in-a-lifetime opportunity to work with numerous live cassowaries kept in captivity in Florida. Our understanding of these birds is also connected to thoughts on the evolution of extravagant structures in dinosaurs in general and also to the mutual sexual selection work I published with Dave Hone and others back in 2011 (Hone et al. 2011), so that gets listed here too…

Did dinosaurs and pterosaurs practise mutual sexual selection?, January 2012
Inside the Cassowary's Casque, Updated for 2022, April 2022
The Great Florida Cassowary Relocation Event of 2023, April 2023

Caption: I’ve been lucky enough to spend time with cassowaries belonging to several different taxa (albeit not Dwarf or Bennett’s, so far), and have got to know several distinct individuals. These photos all show Piggy, a yellow-necked Northern or Single-wattled cassowary Casuarius unappendiculatus who did a number of quirky things during the time that I got to observe him. Images: Darren Naish.

We’re here for the esoterica 1, cryptozoology. Many people come to Tet Zoo because of what we can charitably call the weird stuff: cryptozoology, monster-themed content more broadly, and speculative zoology. Ok, you might be here because you like pterosaurs, unusual birds, obscure rodents or fossil mammals, but the fact is that the most-visited articles at Tet Zoo are on weird stuff. There are downsides to discussing these things, for sure… oh boy, do they attract problematic people… but maybe they can serve a purpose by showing how non-standard ideas can be traced back to real discoveries and hypotheses. Let’s start with articles on cryptozoology. Among those Tet Zoo articles worthy of considering highlights include…

Really: photos of the Loch Ness monster, December 2007
Pterosaurs alive in, like, the modern day!, December 2007
The 1972 Loch Ness Monster Flipper Photos, August 2020
The Lake Dakataua ‘Migo’ Lake Monster Footage of 1994, February 2021
What Was the Montauk Monster? A Look Back to 2008, October 2021

Caption: one of the most famous Loch Ness Monster encounters of all is the George and Mrs Spicer event of 1933. Their description of an amorphous, undulating mass crossing the road, a small head recalling that of a lamb or deer in its midst, culminated with this fantastic illustration by the legendary Gino D’Achille. The Spicer story is one of several accounts of ‘land nessies’, and I wrote about them back in the 2000s. Image: © Gino D’Achille.

The cryptozoological articles here are important personally because they’re connected to my involvement in TV projects on the Loch Ness Monster and such, and to the publication of Hunting Monsters (Naish 2016, 2017). An additional relevant book is currently in production.

**Caption:** in November 2015, Scottish tourist Harvey Robertson was in a boat off the coast of Corfu, Greece, when he photographed a peculiar grey object in the water. He didn’t notice it at the time and only later did he opt to pass the photo to the tabloid press. They, of course, promoted it as a possible sea monster, and the public went nuts for it too, terming it the Corfu Island Creature. Memo Kosemen had fun interpreting it as a possible marine mammal, hence the artistic reconstruction here. In reality, it’s very likely not a real animal at all and I suggested (as did other people at the time) that it might be a lost freeboard fender, like the one shown at top right here. Alas, many people interested in monsters much prefer to be led by their emotional sense of ‘rule of cool’ and any discussion of the case today will be dominated by those who think that ‘unknown sea monster off Corfu’ should be the preferred identity... I covered this case in my **annual round-up of Tet Zoo events published in January 2016**.

We’re here for the esoterica 2, speculative zoology. Again, I don’t know if I’ve covered SpecZoo too much or too little, but there’s been some good coverage of the field here over the years. My favourite articles include the following…

Aquatic proto-people and the theory hypothesis of initial bipedalism, March 2008
Dougal Dixon’s After Man, the Initial Pitch Document, June 2020
Humanoid Dinosaurs Revisited Again: Russell and Séguin’s Dinosauroid at (Nearly) 40 Years Old, August 2021

There are less opportunities to talk about SpecZoo away from this blog than there are for cryptozoology – like, no-one has ever approached me with the idea of doing a SpecZoo book or even a paid article – but maybe something further will happen in time, stay tuned. I have serious plans to get SpecZoo discussions into the technical literature, partly because there’s a lot of crossover already. I promise to come back to that point later in 2026.

**Caption:** certainly among the strangest of ideas I’ve covered is Initial Bipedalism, the fringe and little-known hypothesis in which it’s posited that marine humanoids, the direct descendants of vertically-oriented, lancelet-like animals, gave rise not only to us but to all other vertebrates. I wrote about Initial Bipedalism way back in March 2008 but an intact version of the article **isn’t showing for me right now**. These illustrations are by the late François de Sarre and artist Robert Dumont.

  **Caption:** more on Initial Bipedalism, though this time from my 2017 review of Bernard Heuvelmans’s *Neanderthal: Saga of the Minnesota Iceman* (**published here at ver 4 in 2023**). This illustration, by François de Sarre and from de Sarre (1997), shows how (*according to this model*) aquatic homunculus animals colonized the land and gave rise to animals like us, and how we then – via a process of de-hominization – begat primates increasingly specialized for quadrupedality. Image: François de Sarre.

We’re here for the esoterica 3, the cranks. In case you haven’t noticed, things are not great right now – I mean, in general, global terms – and among the many issues we have is that science is under siege. This is due both to the efforts of certain politicians and political parties to diminish the role of science across the board, and to a populist movement that’s leading to the spreading of pseudoscience and anti-science across popular media. It is right to fight back, though quite how we do this without making the relevant people even more entrenched is a good question. Anyway, I’ve written on cranks – on people who promote non-standard and poorly founded views and do so via a position of weird arrogance and elitism – a few times, the resulting articles being Tet Zoo classics…

Why the World Has to Ignore David Peters and ReptileEvolution.com, July 2020
Alan Feduccia’s Romancing the Birds and Dinosaurs: Forays in Postmodern Paleontology, October 2023
Ray Hoser, Number 1 Taxonomic Vandal, May 2024
Post-Truthism, Brian J Ford’s Aquatic Dinosaurs, and the Fate of ‘Too Big To Walk’, November 2024

And that’s where we’ll end things. Again, please tell me in the comments about those articles that you consider highlights, or maybe the podcast episodes too since I just realised that I totally forgot about the existence of the Tetrapod Zoology podcast until now.

**Caption:** additional illustrations created for the 2015 April Fool’s article **Cetacean Heresies: How the Chromatic Truthometer Busts the Monochromatic Paradigm** (an intact version of which is no longer findable, at least not for me. UPDATE: **it’s here**, thank you, David Gioia). Yes, this was created with the outlook of specific real-world cranks in mind: they’re weird, arrogant people who claim that only *they* are smart enough to see what the world is really like. It wasn’t created with **David Peters** in mind. Images: Darren Naish, colouring of whale image by Gareth Monger.

As ever, massive thanks for visiting and for being part of things here. This article is the second in my lookback at 20 years of blogging: for the first one do check out Two Full Decades of Tetrapod Zoologyif you haven’t already. Also as ever, do consider supporting the work here via the patreon if you can.

For previous TetZoo articles on birthdays and other landmarks, see…

Happy first birthday Tetrapod Zoology (part I), January 2007
Happy first birthday Tetrapod Zoology (part II), January 2007
Happy second birthday Tetrapod Zoology (part I), January 2008
Tetrapods of 2007 (happy birthday Tet Zoo part II), January 2008
Happy THIRD birthday Tet Zoo, January 2009
Tet Zoo = 4 years old today, January 2010
2009, a year of Tet Zooery, January 2010
Four years of Tet Zoo: to infinity... and beyond!, April 2010
It is with some dismay that I announce Tet Zoo's first hemi-decade, January 2011
Tet Zoo 5th birthday extravaganza, part II, January 2011
Happy Birthday Tetrapod Zoology: SIX YEARS of blogging, January 2012
Happy 6th Birthday, Tetrapod Zoology (part II), January 2012
Tetrapod Zoology enters its 8th year of operation, January 2013 (I cannot find any intact versions of this article, thanks SciAm)
Today marks NINE YEARS of Tetrapod Zoology, January 2015
Tetrapod Zoology 10th-Birthday Extravaganza, Part 1: 2015 in Review, January 2016
Tetrapod Zoology 10th Birthday Extravaganza, Part II: the Rest of 2015 Reviewed, January 2016
Tetrapod Zoology 10th-Birthday Extravaganza, Part 3: Tet Zoo's Tetrapod Treatment in 2015, January 2016
Today Is Tet Zoo's 11th Birthday, January 2017
The 12th Year of Tet Zoo, January 2018
The Tet Zoo 12th-Birthday Event, Part 2, January 2018
The Much Belated Final Part of the Tetrapod Zoology 12th Birthday Event, December 2018
Tetrapod Zoology Is A Teenager Now, January 2019
Tetrapod Zoology's 14th Year of Operation, 2019 in Review, January 2020
On Tetrapod Zoology’s 15th Birthday, the Year in Review, January 2021
Happy 16th Birthday, Tetrapod Zoology, February 2022
Four Years of Tetrapod Zoology ver 4, July 2022
The 17th Year of Tetrapod Zoology: 2022 in Review, January 2023
Five Years of Tetrapod Zoology ver 4, July 2023
Tetrapod Zoology Reaches 18 Years of Age, January 2024
It Was the 19th Year in the History of Tetrapod Zoology, January 2025
Two Full Decades of Tetrapod Zoology, January 2026

If you enjoyed this article and would like to see me do more, please consider supporting this blog (for as little as $1 per month) at patreon. The more support I receive, the more financially viable this project becomes and the more time and effort I can spend on it. Thank you :)

Refs - -

Barker, C. T., Hone, D. W. E., Naish, D., Cau, A., Lockwood, J. A. F., Forster, B., Clarkin, C. E., Schneider, P. & Gostling, N. J. 2021. New spinosaurs from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae. Scientific Reports 11: 19340.

Barker, C. T., Lockwood, J. A. F., Naish, D., Brown, S., Hart, A., Tulloch, E. & Gostling, N. J. 2022. A European giant: a large spinosaurid (Dinosauria: Theropoda) from the Vectis Formation (Wealden Group, Early Cretaceous), UK. PeerJ 10:e13543.

Barker, C. T., Naish, D. & Gostling, N. J. 2023. Isolated tooth reveals hidden spinosaurid dinosaur diversity in the British Wealden Supergroup (Lower Cretaceous). PeerJ 11: e15453.

de Sarre, F. 1997. Were aquatic pre-humans the first vertebrates to enter the land? In Downes, J. (ed) The CFZ Yearbook 1997. CFZ (Exeter), pp. 142-156.

Hone, D. W. E., Naish, D. & Cuthill, I. C. 2011. Does mutual sexual selection explain the evolution of head crests in pterosaurs and dinosaurs? Lethaia 45, 139-156.

Naish, D. 2010. Tetrapod Zoology: Book One. CFZ Press, Woolsery, Devon.

Naish, D. 2016. Hunting Monsters. Arcturus Books, London.

Naish, D. 2017. Hunting Monsters (hardcopy edition). Arcturus Books, London.

Naish, D. & Perron, R. 2016. Structure and function of the cassowary's casque and its implications for cassowary history, biology and evolution. Historical Biology 28, 507-518.

Taylor, M. P., Wedel, M. J. & Naish, D. 2009. Head and neck posture in sauropod dinosaurs inferred from extant animals. Acta Palaeontologica Polonica 54, 213-220.

Witton, M. P. & Naish, D. 2008. A reappraisal of azhdarchid pterosaur functional morphology and paleoecology. PLoS ONE 3 (5): e2271. doi:10.1371/journal.pone.0002271

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Today is the day….

**Caption:** a montage of animals that had relevance of one sort of another to my 2025…. proboscideans, varanids, phorusrhacids, marsupials, and cryptids like long-necked seals and tatzelwurms. Images: Darren Naish.

The blog Tetrapod Zoology – connected in some way to just about everything that’s happened in my professional life since the mid-2000s – has now been in operation for an absurd twenty years. Two. Decades. At the risk of lapsing into grotesque melodrama, I have the same feeling I get when attending a funeral… albeit without the melancholy… how did we get here already; how did so much time pass by?

2025 might have been the busiest year of my life; a lot happened. It was, however, primarily structured around four Big Events. The year was also punctuated by several publications, including the books Mesozoic Art II and the third edition of Ancient Sea Reptiles, as well as both popular and technical articles. I gave seven talks (in Bournemouth, Southampton, Portsmouth, Lyme Regis, Worthing, Aberdeen and Glasgow), travelled to Spain (by ship) and Scotland (by rail) but was otherwise mostly limited in my adventures to southern England.

**Caption:** Teddy the West Highland terrier is my companion on at least some excursions about the south of England. He’s a good boy but is afflicted by a list of ailments. As you’ll know if you’ve seen his comments on social media, he’s also a sage and vociferous commentator on the current sociopolitical state of the United Kingdom and the lamentable nature of its built environment. Images: Darren Naish.

Prologue. In this very long article – the first of at least two celebrating 20 years of Tet Zoo – I look back at 2025 as part of my regular annual reviews. As ever, the article could very well be seen as tremendously self-absorbed and self-congratulatory. But, hey, no-one else is gonna do it for me, so put up, shut up or go away, the internet is a big place.

In my 2024 review (published here in January 2025), I mentioned the existence of “[a] huge number of things … that I can’t talk about”. But now that 2025 has passed, I can talk about those things, and in fact they’re the main events relevant here. I should say to start with that 2025 began with me gainfully employed at the BBC’s Natural History Unit, specifically on the third season of Prehistoric Planet for Apple TV. More on this later.

**Caption:** 2025 included one of those milestone birthdays for myself, and here’s a view of some of the material accrued as a consequence of that event. It might be obvious how well known it is that I am clandestinely assembling a collection of animal figures, models and toys. Image: Darren Naish.

The start of 2025. 2025 was marked by a great many interesting discoveries concerning Mesozoic animals – yes yes, just as all the years are – and I had reason to write about some of these at Tet Zoo and elsewhere. My first article of this sort was on the unusual, long-handed Mexican ornithomimosaur Mexidracon and appeared at Discover Wildlife (it’s here). January also saw release of the Tet Zoo podcast episode that John Conway and I recorded at TetZooCon back in September 2024, that one being the last TetZooCon ever. It’s not a particularly good episode; we spoke about the suggested dicynodont identification of the La Belle France rock art image in South Africa (I totally don’t buy it) and the fact that Greg Paul wasn’t able to join us. On the subject of the Tet Zoo podcast – since we have a massive listenership, somewhere in the millions donchaknow – it’s not officially dead, it’s just that John and I have incompatible schedules. Also in January, I wrote an endorsement for C. M. Kösemen’s book All Tomorrows. I began consultancy work on Kevin Miguel’s Mesozoic Life Stories: River of Giants docuseries project.

**Caption:** yeah, you’re gonna be seeing a lot of figures, models and toys in this article. The *Spinosaurus* at left, named Seti, and the green *Carcharodontosaurus* at right, named Amon, are limited edition backer rewards for the **Mesozoic Life Stories: River of Giants project**. Image: Darren Naish.

A brief review of Neil Frost’s cryptozoological doorstop of a book (Fatfoot: Encounters With a Dooligahl) was published at Tet Zoo in February and I also published thoughts on Tama Zoological Park in Tokyo, a zoo I visited in Japan in 2024. I’m always literally years behind in my zoo review series and am perpetually aware of how much unpublished content I have on the zoos and wildlife parks I’ve visited.

Spawnwatch, Don Lessem, the CFZ Cryptozoology Summit. February is spawnwatch season here, so I again reported events as seen from the recently renovated big pond. I’m looking forward to seeing what happens in early 2026. Will there be more frogs and more clutches of spawn, or less, and will the ever-present free-roaming pet cats be a problem? We will see.

**Caption:** Common frogs *Rana temporaria* in the shallow end of pond 2 during early February 2025. We ended up with 17 clutches for that year and I will again be keeping track of things in 2026. Image: Darren Naish.

  **Caption:** Common frogs certainly benefit from large, very shallow areas and will preferentially spawn in water less than 10 cm deep. As is obvious from these images, the tadpoles pack into the shallows to absorb solar heat . These photos are from late March 2025. Images: Darren Naish.

Over the years I’ve worked on occasion with author, consultant and Dinosaur Society founder Don Lessem, and in January 2025 I had the opportunity to meet him while he was visiting London. Don signed books I threw in front of him and I also asked him about his time as consultant for Jurassic Park (the only movie in that franchise I care about or am interested in). More on the Jurassic films in a minute.

**Caption:** if you paid attention to dinosaur-themed books from the 1990s, 2000s and up to modern times, you’ll be very familiar with the work of Don Lessem. It was great to hang out in person during early 2025. Images: Darren Naish.

Regular readers of this blog will know of my ever-present interest in cryptozoology, and at this point I’m just as interested in the ideas promoted by the cryptozoologists themselves as I am in the putative items of cryptozoological focus (Naish 2016, 2022). In February I met up with British cryptozoologists Jon Downes and Richard Freeman at Jon’s residence – the CFZ or Centre for Fortean Zoology – in Woolfardiswarthy (pronounced Woolsery!) in north Devon. I needed info from both for a book project. We also spoke about giant fosa*, the status of cryptozoology, and rewilding in the UK. Some of our discussion is online here.

Following advice from colleagues who work in Madagascar, I’m switching from the spelling ‘fossa’ to ‘fosa’.

Caption: let me say, as I so often do, that you can be super interested in mystery animal research without being a ‘believer’. I cherish the time I spend with other people involved in cryptozoology, wherever they are on the ‘believer’-’sceptical’ spectrum. Here I am at the CFZ in Devon with Richard Freeman (holding his 2024 book Creatures That Eat People) and (seated) Jon Downes. At right, a view within the old CFZ office, various relevant items on show. Images: Graham Inglis, Darren Naish.

Tom Jackson’s Dorling Kindersley book Eyewitness Animal (Jackson 2025) arrived at Tet Zoo Towers in February. It warrants mention because the extinct animals section includes a short bit about me and a photo of me sat next to the Bone Clones Gigantopithecus skull. Finally, some recognition.

**Caption:** the 2025 DK *Eyewitness Animal*. The image at right is featured therein; it was taken at the Society of Vertebrate Paleontology meeting in Brisbane in October 2019, but I regret that I don’t have a record of who took it. This wasn’t, of course, the only *Gigantopithecus*-relevant event of 2025…

Goodbye BBC, goodbye Bristol. March was my final month at the BBC’s Natural History Unit (NHU). I made a special homage to Bristol Museum and Art Gallery: if I’m not working in Bristol anymore, I won’t be visiting it so often… folks, make the most of your local museums and other visitor attractions while you can. Being based at the NHU – and working in particular on big-budget shows about prehistoric animals – has been a dream job, the most insane privilege. Having this excellent job has been a major obstacle to other things, since I’ve had to decline or delay numerous projects and other jobs. But that’s a very niche complaint, and finishing there was truly the end of an era. I’ve had the most fantastic time at the NHU since starting there in 2017 and have met the most incredible set of people, many of whom are now lifetime friends.

**Caption:** Bristol Museum and Art Gallery is home to a massive number of historically interesting items. The vintage dinosaur models here, made by Alan Braddock, are familiar to many as they were featured on postcards released commercially during the 1970s. The *Archaeopteryx* model at far left has appeared in several books and on TV, as has the stop-motion Aardman Animations stop-motion *Dimorphodon* at right. A replica azhdarchid humerus is visible at extreme right. Image: Darren Naish.

I also have vast nostalgia attached to my time at the ramshackle Broadcasting House in Whiteladies Road (during my tenure, we moved out and went to the new, more sterile premises in Bridgewater House). In addition, I’m happy that I had the chance to see people, places, facilities and even archive film associated with NHU projects that had a major impact on my younger self… key episodes of Wildlife on One, the David Attenborough 1970s series Fabulous Animals, The Velvet Claw… I could write a whole book on the behind-the-scenes work involved in the making of all three seasons of Prehistoric Planet. Maybe that opportunity will arise one day, maybe it won’t.

**Caption:** the older I get, the more interested I am in documenting and discussing the things that inspired me when I was young. Back in the 1970s, 80s and 90s, the BBC Natural History Unit produced some of the most excellent, influential pieces of media on those subjects I’m interested in. The images here relate to Sir David’s *Fabulous Animals*, and an early outing of stork-faced *Quetzalcoatlus* from the Wildlife on One film *Pterodactyls Alive!* At right is my cherished VHS set of *The Velvet Claw*.

Tet Zoo on Monster Hunter. March also saw the release of a few videos I made during January for PC Gamer on Monster Hunter. This was all part of a ‘reality check’ series PC Gamer does where a given expert watches game footage and then says what they like, what they don’t like, and how things do or don’t align with their knowledge. I’m not a gamer – I don’t want to spend more time on computers than I already do – but I’m at least aware of this franchise due to its appearance in pop culture, plus I went to the trouble of sitting through the 2020 movie.

**Caption:** PC Gamer created a nice little desk-top display for my appearance on ‘reality check’, and I even had genuine opportunity to use that model *T. rex* skull. Image: PC Gamer.

The creatures of the Monster Hunter games represent a mashup of fossil species (dinosaurs especially), living animals, and mythical dragons and other beasts. A huge number of things could be said about each and every one of them. Watch the PC Gamer review here on YouTube. A second video about Elden Ring – a game less relevant to my interests – was recorded at the same time but not released until April (it’s here on YouTube). Elden Ring is based more around high fantasy and magic and less on creatures, but there are dragons, snake monsters and so on.

Also in March, the digital version of my article on reviewing books appeared in Historical Biology (where I’m now book reviews editor); the hardcopy version appeared in December (Naish 2025a). DinoCon 2025 tickets went on sale during the month. At some point during the early months of 2025, Arturo García, a Chilean artist based in Santiago who does great palaeoart, asked me an interesting question: if all my model animals were melted down and turned into a new one, what would it be? Answer: a hypothetical giant flightless bat. And thus…

**Caption:** a giant flightless bat as depicted by Arturo García, with one standard Naish unit for scale. Image: Arturo García.

In pursuit of newts. Observational work involving wild amphibians occurred in March when Phil Budd of the Southampton Natural History Society, Helian and I again visited the ponds in the north of our city to monitor the situation. Remember that the UK has a tiny number of native amphibian species, this being just three newts, two toads and two frogs. A very active area of research concerns the occurrence, distribution and spread of invasives. Do we have the fast-spreading, ecologically flexible Alpine newt Mesotriton alpestris in the Southampton area? We couldn’t find them in 2024. But this time… success. They’re here, they’re easy to find, and it looks like they’re becoming more abundant. We will continue to monitor the situation.

**Caption:** a shallow pond in the north of Southampton, surrounded by marsh and heavily vegetated land and hence good for amphibians. A single male Common toad *Bufo bufo* is visible here, as well as numerous male and female Smooth newts *Lissotriton vulgaris*. Image: Darren Naish.

  **Caption:** and here’s one of the animal we were looking for, a male Alpine newt. This animal has a complex taxonomic history and, after being named *Ichthyosaura* for a while (a very annoying name), is – as of 2025 – back in *Mesotriton*. It doesn’t appear especially close to the other newts we have in the UK (*Lissotriton* and *Triturus*), instead being close to the group that includes fire-belly *Cynops* newts and kin. Image: Darren Naish.

The Palaeontology in Public book launch event, organised by Chris Manias of University College London, happened in London in late March; there were talks on fossil plants, Crystal Palace, the history of palaeoart and more. I was part of a panel that also featured Natalie Lawrence (author of the 2024 book Enchanted Creatures: Our Monsters and Their Meanings), Pleistocene expert Victoria Herridge, and world-renowned palaeontologist Professor Michael Benton. I got my first look at Mark Witton’s very impressive Princeton University Press book King Tyrant.

**Caption:** a scene from the Palaeontology in Public Panel Discussion of March 2025. Left to right: Michael Benton, Tori Herridge, Darren Naish, Natalie Lawrence. Image: Elliott Edwards, used with permission.

  **Caption:** a March 2025 scene from Titchfield Haven, a SSSI on the banks of The Solent, southern Hampshire. I really like the composition of this scene. We can see Black-headed gull *Chroicocephalus ridibundus*, Ruddy turnstone *Arenaria interpres* and Mallard *Anas platyrhynchos*. Image: Darren Naish.

I published another Discover Wildlife article in March, this one on the two-fingered therizinosaur Duonychus (it’s here). I also visited the Bournemouth Natural Sciences Society to talk on Mesozoic marine reptiles, all part of my continuing promotion for Ancient Sea Reptiles.

**Caption:** my marine reptile talk at the Bournemouth Natural Science Society (BNSS) in March 2025 resulted in some interesting things being encountered. Prior to that time I didn’t know that Dougal Dixon had published a series of books made for Walking With Dinosaurs 1999 (thanks to Brandon Mason for bringing them along). At right, a model Great auk *Pinguinus impennis* on show at BNSS at that time. Images: Darren Naish.

Mokele-mbembandwagon, redux. I was also quoted in a Popular Mechanics article on mokele-mbembe, this one titled ‘‘Dinosaur’ sightings are on the rise in the Congo. Could this legendary creature realty exist?’ and written by Jordan Smith. Therein, my words were used alongside those of Eddie Guimont and Loren Coleman. A longer, more authoritative article, again quoting me, appeared in New Lines Magazine during April; this one is titled The Congo’s Dinosaur of Discord and is by Ryan Biller. In both cases, I repeat the same stuff about this phenomenon I’ve been saying for a while: that the idea of ‘living sauropods in the Congo’ is related to an erroneous and romanticised late 1800s and early 1900s notion of ‘backwater prehistoric Africa’ mixed with adventure ‘lost world’ fiction and the cultural impact of sauropod skeletons in museums (Naish 2016).

In addition, there isn’t much good reason to think that people in the relevant regions have seen ‘living sauropods’ at all. Sure, they have myths and stories about giant dangerous animals, but Europeans have erred in connecting these with sauropods. If people have seen weird animals, I like reminding people that very large softshell turtles and big pythons are endorsed as real and findable within the cryptozoological literature.

**Caption:** you can’t modernize the Westernized, cryptozoological concept of the mokele-mbembe, not that there’s been much effort to do so, because it is, and always will be, linked to an anachronistic cartoon view of what the ‘Dark Heart of Africa’ was meant to be like more than a century ago. The animal at rear here is a modern view of a saltasaurid titanosaur; at right is the mokele-mbembe as generally imagined, right out of a 1901 encyclopaedia. Image: Darren Naish.

The whole mokele-mbembe thing is now self-perpetuating both as Congolese people work as canny guides and assistants to foreign wannabe adventurers and while wannabe adventurers and self-styled explorers – who never do work of any value bar writing popular books and doing podcast interviews – larp their way through the region in an effort to confirm what they already think is true. This has been going on since the 1980s and the subject is now so tarnished that we’ll probably never understand what the phenomenon was based on originally... assuming, of course, that it was based on anything. What’s weird is that there’s currently a resurgence of interest in the idea that mokele-mbembe is (a) real and (b) worth searching for, and very similar articles keep appearing. Sharon Hill wrote about this during March 2025 and used the term ‘mokele-mbembandwagon’ for this new bout of interest.

**Caption:** there are a great many books on fossil hominids, and I’m gradually working through the ones that I own. Due to be covered here next are Mary Leakey’s 1984 biography, and Harry Shapiro’s *Peking Man* of 1975. Curtis *et al*.’s *Java Man* was discussed **here** at Tet Zoo. Images: Darren Naish.

On fossil hominids, sea monsters and Prehistoric Planet megathreads. Moving to something wholly different, I’ve been meaning for years to start writing about the massive number of palaeoanthropological books I’ve accrued, many of which cover the same discoveries, discussions and hypotheses albeit from different perspectives. I made a start on this in March with my article on Curtis et al.’s Java Man of 2000. Other Tet Zoo articles are coming in the series. I’m working my way through books on the Leakeys (of which there are many) as well as others on the supposed presence of Homo erectus in Africa. Remember that H. erectus was first named from Java… how did we arrive at the view that it was present throughout virtually the whole of the Old World, persisting for millions of years?

A review of a book more directly relevant to my own research – Adrian Shine’s A Natural History of Sea Serpents – was also published in March, as was a recycled article on the Southern sea lion Otaria byronia/Otaria flavescens, one of my favourite pinnipeds. I also published (both on the increasingly problematic site known as Twitter and at the Tet Zoo patreon) long-form thoughts on the Oceans episode of Prehistoric Planet 2. Better late than never (the series aired in May 2023, approximately two years earlier).

**Caption:** I’m a simple man, and for long have I hoped to own my very own life-sized plastic heron. That dream was realized in April 2025. I took the heron with me on various adventures about the place, and here it in in the car, and near the shore in Swanwick, near the River Hamble. Images: Darren Naish.

SpecZoo, marine reptiles, the Heuvelmans project, a dinosaur book. Ever aiming to rescue old material from the previous versions of Tetrapod Zoology (the ScienceBlogs and Scientific American years), I revamped and republished two articles on speculative zoology during April, one my 2013 interview with Dougal Dixon and one my review of the 2015 book Demain, les Animaux du Futur.

Luke Muscutt and I hung out with marine reptile fossils (and replicas of them) in London, this being relevant to the next step in our work on plesiosaur swimming behaviour. Like all academic projects, this is slow-burn and taking forever, but I promise we’ll return to it in time. On the subject of slow-burn projects, progress continued on my Cryptids of Bernard Heuvelmans project, the results again appearing at patreon. Increasing workload during the year meant that I had to temporarily abandon that project. I cannot express how frustrating this is. Maybe I’m trying to do too much.

**Caption:** images from a visit to Holly Hill Woodland Park (Fareham, Hampshire) during April 2025. Pictures like this do a god job of reminding people who don’t live in the UK how incredibly mild and warm our climate is despite our far northern latitude. Note the (non-native) tree ferns. Images: Darren Naish.

  **Caption:** I visited Blenheim Palace during April 2025, and it turned out to be home to several things of TetZooniverous interest, among them this antler display. These mostly belong to Red deer *Cervus elaphus* but there’s Moose *Alces alces* here too. The big surprise, though, is the *Megaloceros* set in the middle. Image: Darren Naish.

TikTok and taxonomic trolling. And so to May… the month in which I started a new venture, namely a TikTok account (TetZooTowers_collection) devoted to model and toy animals. We started with Star Wars creatures for May 4th but later began going through Spinosaurus toys in rough chronological order. These do a good job of charting changing views on thoughts about this animal, but to hear the full story on that you’ll need to watch the series… which currently exists as 47 separate episodes and is still incomplete! I gave more talks, firstly at God’s House Tower in Southampton and given to accompany the Hidden in Stone exhibition organised by my colleague Neil Gostling, and then for the University of Portsmouth’s Palaeo Society. Both talks were on dinosaurs and given to accompany a signing of Dinopedia, my Princeton University Press book (buy it direct from me here).

**Caption:** a montage of screengrabs (showing Instagram stories) that give some idea of the content online at the TetZooTowers\_collection TikTok, everything here being related to spinosaur figures.

  **Caption:** the studio area… yeah, let’s call it a studio area… that we work with for the TetZooTowers\_collection TikToks. I continue to hold out hope that I might one day obtain a larger house and hence eventually have the collection on proper display. There’s no hope of that today and most of it is in storage boxes. Image: Darren Naish.

  **Caption:** scenes from the opening of Hidden in Stone at God’s House Tower, Southampton, during April. Part of the exhibition focuses on the spinosaurid work published by Chris Barker, Neil Gostling, myself and others, so at left we see a reconstructed baryonychine skull together with casts of the *Ceratosuchops* holotype. At right, myself and Karen Fawcett with her brilliant Crystal Palace *Megalosaurus* model.

A niche topic covered here at Tet Zoo on a few occasions concerns taxonomic vandalism, the event in which rogue researchers take to publishing their own scientific names for populations or specimens they consider distinct. I’ve contributed to multi-authored articles on the issue before (Rhodin et al. 2015) but have never published a stand-alone piece on it… until May, since the new issue of The Biologist, released that month, includes my article on this topic (Naish 2025b). The online version is here. Palaeornithologist Hanneke Meijer visited the University of Bournemouth here in southern England and we caught up, giving me an opportunity to get my SVP dodo monograph signed. Only another three editors to go.

**Caption:** mediocre bird photos taken on a trip to Fishlake Meadows Nature Reserve during May 2025. Top row, left to right: Sedge warbler *Acrocephalus shoenobaenus*, Bullfinch *Pyrrhula pyrrhula*, Common kestrel *Falco tinninculus*. Greylag goose *Anser anser* and Egyptian goose *Alopochen aegyptiaca* are having an altercation in the photo below. The Egyptian goose isn’t a goose at all, but a tadornine duck. Images: Darren Naish.

  **Caption:** Eurasian jay *Garrulus glandarius* photographed in the New Forest during June 2025. Jays are on that list of bird species that were like mythical animals to me as a young person… animals that I knew from books but had no hope of ever seeing in real life. Today I see them on regular occasion and know that they’re findable wherever the habitat is right. Woodpeckers and nuthatches are also on that list. Image: Darren Naish.

More cryptozoology-relevant content was released in June when I participated in an interview for Barnaby Jones’s Monsters on the Edge podcast, the episode being titled ‘It’s a Dinosaurs World’. My main points will be familiar to those who’ve read my stuff: we covered the prehistoric survivor paradigm, claimed sightings of many-humped sea monsters and more. It’s here on YouTube.

**Caption:** Flame the bearded dragon remains a prominent fixture of my domestic life, and I do what I can to give her interesting experiences and opportunities. Here she is among the undergrowth in our front garden during the summer. Image: Darren Naish.

The Lyme Regis Fossil Festival 2025. June’s big event, however, was the Lyme Regis Fossil Festival. As I’ve said on previous occasion, this has increased over recent years in size and complexity such that it’s now the UK’s premier palaeontology-themed event. There was a time when the Fossil Festival basically involved a single, small marquee, occupied by fossil dealers, and nothing else. Ok, there were a couple of evening social events but that was it. That contrasts massively with the situation of today: events span most of the length of the town, tens of vendors are present in giant marquees and various buildings, and numerous authors, scientists, artists, film-makers and others give talks and presentations.

**Caption:** promotional imagery created for the 2025 Fossil Festival, featuring art by me (at left) and by Lee Brown (of leebrownpaleoart) at right. They have done such a great job of making this an enormous, and enormously fun, annual event.

  **Caption:** view of the Lyme Regis promenade, looking east, and giving some loose idea of the huge number of people that visit and attend during the course of the weekend. You should be able to see Nathan and a DinoCon sign in the middle of the image. Image: Darren Naish.

  **Caption:** oh dear; me being subtle and low-key in demonstrating enthusiasm for *Spinosaurus*. This replica skull was brought to the Fossil Festival by the University of Portsmouth team. Note the little promo card for the *Spinosaurus*-themed episode of *Walking With Dinosaurs 2025*.

I gave my Dinopedia-related dinosaur talk again (‘Reconstructing the Lives of Dinosaurs’) but my efforts to sell books were thwarted by my phone and SumUp machine refusing to talk to each other. I’ve learnt from this mistake and now only use the SumUp phone app. The Fossil Festival was the ideal event for DinoCon promotion, so Nathan, Annie, Mike and I (the core DinoCon team) handed out leaflets, waved placards about and spoke to potential stall-holders.

**Caption:** Darren and Nathan holding the DinoCon placard, with (at far right) Lyme Regis palaeontologist and museum worker Kieran Satchell in attendance too.

In London, I finally made time to look at the palaeontological garden at the Natural History Museum (NHM). My main reason for visiting was to catch up with Steve Zhao at Sandbox VR, since a job I had from early in the year concerned consultation for the brand-new Age of Dinosaurs VR experience, created in partnership with the NHM and due to go live in early 2026. I also travelled to Worthing to talk to the West Sussex Geological Society about dinosaurs and sell copies of Dinopedia again, and met up at the Grant Museum of Zoology with Michael Mills (aka Professor Flint). In late June, Natalia Jagielska and I spoke about Natalia’s section in Mesozoic Art II in an open-to-everyone zoom event. The book wasn’t officially out until September but I and others had received our copies by then.

**Caption:** the **Sandbox VR** immersive experience *Age of Dinosaurs* goes live in early 2026. We've done what we can to make it technically right, but it's tremendous fun too and we can't wait for you to see it. We’re pleased to have the support of NHM London in promotion. Image: (c) Sandbox VR.

  **Caption:** Fern the bronze *Diplodocus*, on show in the gardens at NHM London, is a beautiful and spectacular thing. The gardens as a whole are a great addition to what’s already one of London’s best visitor attractions. Images: Darren Naish.

  **Caption:** the Grant Museum in London is notable for many things, but one of them is its jar of moles. It’s so famous that the museum sells cuddly moles and a ‘moles in a jar’ badge. At right, myself and Mike Mills with said famous far. Images: Darren Naish.

A diversion on Jurassic World Rebirth. A weird thing happened at the start of July when Toni (my wife) and I attended, by invitation, a Jurassic World Rebirth event at The Barbican in London. The event itself wasn’t great, basically a series of team tasks that had only the flimsiest of connections to the movie and its animals, but I much enjoyed exploring the Barbican Conservatory, a huge tropical house surrounded by brutalist architecture. Getting to meet the brilliant Dolores Aquilops puppet was a highlight, as was my first look at the new toys released to accompany the movie. They’re pretty fun, even if they have nothing to do with science or real Mesozoic animals at this point.

**Caption:** the brilliant Delores puppet at left, with human companion too. At right, my efforts to form an affectionate bond with a rancor monster didn’t end well, I guess I need to cut down on the snickers bars. Images: Darren Naish; Toni Naish.

  **Caption:** scenes from the *Jurassic World Rebirth* event at the Barbican. I was later to obtain one of those tail-thrashing *Spinosaurus* figures for myself (a 2025 birthday present). Images: Darren Naish.

Jurassic World Rebirth includes a sequence showing a T. rex swimming, itself based on content from the book Jurassic Park (and planned for a time to be included in Jurassic Park the original film). Season 1 of the Apple TV series Prehistoric Planet also featured a swimming T. rex, this one most certainly not connected to Jurassic Park but merely to the fact that animals of all sorts are good swimmers (I can say “most certainly” because Paul Stewart and I devised this sequence during the making of the series). A consequence is that a few articles on the ‘Could T. rex really swim?’ issue appeared… one of which involved input from myself. This is interesting for people who haven’t thought about this idea before, but familiar in view of previous discussions on dinosaur behaviour.

Big Event Number 1: Discovering Dinosaurs at Lightroom London, King’s Cross. And so we come to the first of 2025’s Big Events, namely the July 7th opening of the Discovering Dinosaurs show at Lightroom London. Lightroom is an immersive visitor attraction, somewhat like a planetarium but where you enter a giant square exhibition space and experience a movie-length show as visuals and film is projected on the walls (and floor) around you.

**Caption:** Lightroom London gives you more than one opportunity to see the *Prehistoric Planet* dinosaurs at life-size, both outside (as here) and as part of the event. At right, me with actor Damien Lewis, the show’s narrator. Damien was one of several celebs at the opening event in July. Billie Piper was there too but I didn’t talk to her. Images: Darren Naish; Mary Gunton.

When I joined the team in April, Lightroom was showing Moonwalkers, an event devoted to lunar discovery and exploration narrated by Tom Hanks. I was blown away by the trouble the team had gone to in obtaining visuals, info and segments of film, and oh boy did it deliver in terms of immersion. Well, a similarly grand and immersive event was constructed for Discovering Dinosaurs, the whole thing being a spin-off of the Apple TV / BBC Studios show Prehistoric Planet.

**Caption:** scenes from one of several Lightroom London events I went to, these from July 2025. The show features scenes and stories from *Prehistoric Planet* seasons 1 and 2 but a large quantity of novel material was produced specially for it, including entirely new sequences and fantastic art. Images: Darren Naish.

If you have any interest in being surrounded by life-sized dinosaurs (and Cretaceous marine reptiles and pterosaurs) in a brilliant and educational event, or if you’re a Prehistoric Planet super-fan, you just have to visit. Involvement with Discovering Dinosaurs occurred throughout 2025 and I’m thrilled with how it turned out. The Discovering Dinosaurs official brochure features an article I wrote (Naish 2025c).

**Caption:** the Lightroom London entrance at Lewis Cubitt Square, King’s Cross. At right, literature that accompanies the event. Learnt about Lightroom London and Discovering Dinosaurs **at their website here**. Images: Darren Naish.

Cetaceans, tapejarids, sea reptiles. I participated in another ORCA sea safari across the Bay of Biscay in July. As ever, we saw a good amount of wildlife, cetaceans and seabirds in particular. For a detailed account see the Tet Zoo article here. The discovery of phytoliths in a Cretaceous tapejarid pterosaur was a big deal for those of us interested in pterosaur biology and I covered it at Tet Zoo during July 2025, in part because some of the commentary echoed points I made in an unpublished article I wrote on this group back in 2010.

**Caption:** the ORCA sea safaris are all about cetaceans, of course. But we see other animals too, and here are a number of Eurasian spoonbills *Platalea leucorodia* seen near the Spanish coast, and a poor photo of a tuna hunting out at sea. Images: Darren Naish.

The Japanese language edition of Ancient Sea Reptiles appeared in late July, the first non-English version to see print. I’ve said before what a big personal deal it is when I see versions of my books published in other languages: the sad fact is that this is something that authors have zero control over. You basically wait for publishers in other countries to approach your own publisher. Japan always comes through on this front, and I admire the Japanese here not just for their interest in the science but also in the attention to detail with respect to design.

**Caption:** the 2025, Japanese edition of my *Ancient Sea Reptiles*. The fact that art appears on the hardboards beneath the dustjacket is a great feature. The art here is by Jaime Chirinos (the plesiosaur-themed image) and Davide Bonadonna (the thalattosuchian-themed image). Image: Darren Naish.

A cluster of events occurred at the end of July. Apple TV announced, at last, the existence of a third season of Prehistoric Planet, one devoted to the Pleistocene. Tet Zoo ver 4 – the version you’re visiting now – celebrated its seventh birthday on July 31st, this being a mini-introspective of adventures in these here parts. And I appeared as a guest on another podcast, this time Things Visible and Invisible. It was pitched as a general discussion of cryptozoology and (when released in September) was titled ‘Can Science Explain Legendary Creatures?’. It’s here at YouTube.

At The New Dinosaurs book launch. August started with my attending the launch of the second, 2025 edition of Dougal Dixon’s The New Dinosaurs, hosted at Waterstones at Gower Street, London. Dougal spoke about the book’s backstory and contents with Ross MacFarlane, a research development specialist at the Wellcome Collection. Several people interested in both dinosaurs and SpecZoo attended and here’s where I learnt that models depicting various creatures from TND and other Dixonian projects were in exist and soon to be on sale.

**Caption:** scenes from the August 2005 launch of the second edition of Dougal Dixon’s *The New Dinosaurs* at Gower Street, London. The Cutlasstooth model was made by **EXEtinct**. The gentleman leading the event with Dougal is Ross MacFarlane of the Wellcome Collection. Images: Darren Naish.

My review of Graham Weedon and Sandra Chapman’s 2022 book Ichthyosaurs From the Early Jurassic of Britain was published in Historical Biology (Naish 2025d).

**Caption:** an August 2025 trip to the New Forest, during which deer, birds and Viviparous lizards *Zootoca vivipara* were seen. But the most surprising wildlife observation? At the edge of a pool (a drying tributary of Mill Lawn Brook), I saw a small, eel-like fish. I thought it was an elver but it wasn't translucent, and the obvious eye, dark dorsum, and gill slits show that it was a European brook lamprey *Lampetra planeri*. It was tiny, only about 6 cm long. Images: Darren Naish.

Big Event Number 2: DinoCon 2025. A big part of my life over the past 11-ish years has been the running of a big, London-based zoology convention – TetZooCon – that is itself a spinoff of this blog. As mentioned above, TetZooCon came to an end in 2024 and plans for its descendant – DinoCon – were announced at the last TetZooCon. To cut straight to the chase, the first ever DinoCon happened at the University of Exeter during August, and it was a massive and heartening success. The publication of my write-up of what went down was delayed for a few months and didn’t appear until October: it’s here. It should be obvious that DinoCon 2025 went well enough that we’re doing it again, the 2026 meeting happening at Hilton Birmingham Metropole on July 25th and 26th. Tickets are due to go on sale very very soon, so keep an eye on our social media accounts and our website.

**Caption:** just a little of the Mesozoic-themed art I picked up at DinoCon 2025. An excellent range of stickers and miniature pieces of art. I hope you can see the ‘Big Bony Club’ ankylosaur sticker by Speed Thief (aka Sean Hennessy), a hit bit of merch. Image: Darren Naish.

Reviews of DinoCon 2025 appeared at Geek Ireland, SV-POW!, The Inquisitive Biologist, Furahan Biology And Allied Matters, EXEtinct, Raptormaniacs and other places too. Tom Fishenden created a brilliant video on the event which you can view here at YouTube.

**Caption:** an anonymous person’s DinoCon 2025 haul. Look at all those sweet sweet acquisitions. Image: Darren Naish.

Snake Summer and Monsters of the Deep in Scotland. Regular readers will be aware of Monsters of the Deep (MotD), a museum exhibition I co-curated for the National Maritime Museum Cornwall at Falmouth, and which revolves around European myths about sea monsters and our scientific discovery of the deep sea. After finishing its run in Cornwall, MotD went to Chatham Historic Dockyard in Kent for 2023, but things there came to an end in November of that year. What next? Scotland! Yes, Scotland, specifically Aberdeen Art Gallery.

**Caption:** while in Glasgow, I met up with David Armsby (at right) and successfully obtained these amazing *Stegosaurus* and *Carnotaurus* models.

And thus, during late August and early September, I travelled about Scotland, my first stop being Glasgow city centre where I had errands to run. One was catching up with film-maker and artist David Armsby and here’s how I came to possess two of the giant dinosaur sculpts that David makes as part of his creative process. Words cannot convey how pleased I was to obtain these, but I knew that getting them north to Aberdeen and then aaaaall the way back south to Tet Zoo Towers – a distance of around 1100 km – would be quite the chore. I took big suitcases and a ton of packaging. Both survived the trip with no breakages except for a single hoof tip on the stegosaur.

**Caption:** the Armsby *Carnotaurus* and *Stegosaurus* safe in their new home, Tet Zoo Towers. You might just be able to see the broken hoof on the left hand of the stegosaur (since repaired). I also picked up a Greyfriars Bobby model, because how could I not. The actual statue (which is in Edinburgh, not Glasgow) has been featured at Tet Zoo in the past. Image: Darren Naish.

At the Hunterian Zoology Museum, Glasgow, I attended the Snake Summer event organised by Will Tattersdill and Jordan Kisler, this occurring due to 2025 being Year of the Snake. The museum is also home to a famous alleged snake specimen, the holotype of Bothrodon pridii. I gave a talk linked to my Heuvelmans project and discussed giant snakes, cryptids like the African crowing crested cobra, and ultimately the Japanese tsuchinoko and winged and flying snakes. It’s incredible how many snake-themed myths and stories there are, a fact relating to the impact snakes have had on human culture and tradition. A write-up, summarising my thoughts on mythical and mystery snakes in general, was published in Fortean Times in December (Naish 2025e). Snake Summer explains why I republished my Crowing crested cobra article, first released at Tet Zoo back in 2011, here in September.

**Caption:** a view across the main hall of the Hunterian Zoology Museum, a trove of wonders. It includes notable and historically interesting arthropod, fish, reptile, bird and mammal specimens and owes its existence to the bequest of Dr William Hunter’s collection to the University of Glasgow in 1807. Image: Darren Naish.

  **Caption:** the *Bothrodon pridii* display cabinet at the Hunterian Zoology Museum. *B. pridii* was named by John Graham Kerr in 1926 for a giant fang 6.5 cm long, identified as that of back-fanged snake around 20 m long. You can see the specimen at middle right here. Alas, it turned out to be one of the finger-like outgrowths of a Chiragra spider conch *Harpago chiragra*. Image: Darren Naish.

  **Caption:** cover slide for my snakes talk, featuring several of the cryptids I’ve illustrated for the Heuvelmans cryptids project…. plus an antiquarian image of a European flying snake.

I then travelled to Aberdeen to see Monsters of the Deep in its new home. It’s a good fit at Aberdeen Art Gallery and it’s always interesting to see how a venue has modified an exhibition to make it work within their space. I gave another talk – Sea Monsters Past and Present – before doing an Ancient Sea Reptiles book signing. An article discussing the Monsters of the Deep exhibition is in review at a publication.

**Caption:** Aberdeen Art Gallery is a grand, spectacular building with really impressive indoor spaces and an amazing view from the roof balcony (where the cafe is). Aberdeen itself is a fascinating city with a lot of grey, blocky architecture that almost has an eastern European vibe. Image: Darren Naish.

  **Caption:** part of the Monsters of the Deep exhibition on show at Aberdeen Art Gallery in September 2025, this being segments of the part devoted to historical manuscripts and the sea monsters of Olaus Magnus’s *Carta Marina* of 1539. Images: Darren Naish.

  **Caption:** do enough public speaking, and you *will* get to deliver orations from spectacular spaces. My September 2025 talk at Aberdeen Art Gallery occurred in Cowdray Hall, home of an enormous upright chamber organ. Image: Darren Naish.

An article that took a good long while to piece together – one on the speculative possible existence of hybrid Mesozoic dinosaurs – was published here in September. The remarkable Early Cretaceous pachycephalosaur Zavacephale was out by September, and I published a Discover Wildlife article on it… it’s here. And the new megaraptoran Joaquinraptor was covered by me there as well.

**Caption:** the main books I had out in 2025. *Mesozoic Art II*, and the third and Japanese editions of *Ancient Sea Reptiles*. Image: Darren Naish.

Big Event Number 3: Mesozoic Art II. The big, beautiful, art-themed book Mesozoic Art, published by Bloomsbury in 2022, was enough of a success that the decision was made to do a follow-up. And so it was that part of 2025 was taken up with the assembling and writing of the bigger sequel volume Mesozoic Art II (White & Naish 2025), release of which happened in late September. I wrote about it here. We’re showcasing technical and artistic excellence in modern palaeoart, but are also doing what we can to represent the creators of this work with respect to geography and diversity.

Doing these books is not easy and the main push to get them done always coincides with maximum workload at my end. Whatever, we work hard and we get things done. Steve and I spoke at private and public events and I was also able to donate an advance copy for the DinoCon auction. A Love in the Time of Chasmosaurs episode on the book (ep 44) was released in October and an online discussion event organised and hosted by Karim Zanaty can be seen here on YouTube.

**Caption:** cover slide from one of the several events on *Mesozoic Art II*. The art on the cover of the book is by the amazing Anthony Hutchings, who I’ve now worked with on several occasions (the results of a new collab are due out soon).

Also on publicity, I appeared as a guest on the Terrible Lizards podcast hosted by Dave Hone and Iszi Lawrence, this time on Mesozoic marine reptiles as part of my promotion of the third edition of Ancient Sea Reptiles (Naish 2025f). That book was out by this time but I didn’t get to announce it at Tet Zoo until December. I screwed up on a few technical things in the Terrible Lizards episode, wrongly stating that the only ichthyosaur group to persist from the Triassic into the Jurassic were the thunnosaurs… I meant the parvipelvians, the group that includes temnodontosaurs in addition to the thunnosaurs. We all make mistakes when discussing things on the fly. More Ancient Sea Reptiles promotion happened in October when I spoke about the same topic for the Portsmouth Palaeo Society. The hardcopy version of my paper on Brian Ford’s Too Big To Walk appeared in October (Naish 2025g), meaning that this should now be cited as a 2025 paper, not a 2024 one (the paper one replaces the original digital release).

The Colour of Dinosaurs. Also in October, I attended The Colour of Dinosaurs at MAST Mayflower Studios in Southampton, a made-for-kids blend of science advocacy and theatre, with tons of music. It revolves around the work of palaeontologist Jakob Vinther, who plays himself, but also emphasises the diversity of its human cast and different approaches to colour and thinking. I really enjoyed it and found it equally fun, moving and informative. Fossils star too, as does the brilliant Psittacosaurus model made by Bob Nicholls. I liked the show so much that I purchased the soundtrack through Bandcamp, and I owe thanks to Jacob Vinther for bringing the event to my attention.

**Caption:** a scene from The Colour of Dinosaurs, taken in October 2025 and during the Southampton part of its run, in which Jacob presents the Bob Nicholls *Psittacosaurus* model to his musician colleagues. Image: Darren Naish.

Dark Folklore. Another event revolving around Ancient Sea Reptiles then happened as Toni and I travelled to Porthtowan in north Cornwall for the first ever Dark Folklore Festival, organized by Rob Vickery. I spoke about Sea Monsters Past and Present (the same talk I’d given at Aberdeen) and sold more copies of Ancient Sea Reptiles. The whole thing was a brilliant mix of West Country weirdness, storytelling, live music and monstrous shenanigans. I enjoyed it immensely and am very much planning to attend the next one. We used the trip as an excuse to visit Bodmin, Mount Hawke and other places of cryptozoological relevance.

**Caption:** yes, another ‘sea monsters’ talk, this time at the Dark Folklore Festival in Porthtowan, Cornwall. The talk combines discussion of cryptozoology and the ‘prehistoric survivors’ idea with what we know about relevant fossil animals, in particular Mesozoic marine reptiles and basilosaurid whales. Image: Darren Naish.

  **Caption:** select scenes from the 2025 Dark Folklore Festival. I attended a ritual, camped on site, listened to fantastic live music, and had a great time. Definitely planning to attend the next one if I can. Images: Darren Naish.

  **Caption:** the UK has some incredibly exciting and picturesque coastline, and Porthtowan on the north coast of Devon is notable for its ferociously rocky scenery. Here’s an image from our October 2025 trip. Image: Darren Naish.

Also on monsters, late October saw the release of the IFLS (I ****ing Love Science) YouTube movie on the Loch Ness Monster. I was interviewed for this – I’ve lost count now of how many Nessie-themed interviews I’ve done for film and TV – and I do quite like the final product, which you can watch here.

Palaeoart in Birmingham. Something really fun happened in early November, specifically as part of the 85th annual meeting of SVP (the Society of Vertebrate Paleontology), held at the International Convention Centre (or ICC) in Birmingham (UK). It’s rare for SVP meetings to be held in the UK (I think this is only the second time) so I tried my best to attend. Speaking as someone who’s been running conventions for more than ten years, I know how irksome and unfair it can be when people complain about prices… but, my god, the price was unfortunately prohibitive. However, preceding the event was the palaeoart workshop organized by Mark Witton, titled 'Paleoart [sic] Past and Present: the View from the UK’, and I’m pleased to say that I was able to attend that at least.

**Caption:** Liam Elward (the taller individual) and your humble author at Mark Witton’s Paleoart Past and Present: the View from the UK workshop at SVP 2025. I left the event with a small pile of stuff, among which are stickers and prints created by Liam. Image: Julianne Zelda Kiely.

The event involved talks, panel discussions and show-and-tell sessions and was hugely positive. I gave the talk ‘Conway et al.'s All Yesterdays and the Case for Speculative Palaeoart’. It was great to meet a good number of interested people, palaeoartists among them, for the first time as well as others I’ve known for a while. I owe thanks to Mark for organizing it and SVP for assisting my attendance. We talk often about the fact that the UK is a bit of a powerhouse in the production of palaeoart, but an argument can be made that those of us here at ground zero maybe don’t take sufficient advantage of this. Then again, we do have the Popularizing Palaeontology meetings, numerous book launches, the Lyme Regis Fossil Festival and DinoCon, so….

Big Event Number 4: Prehistoric Planet Ice Age! The big event of November, and Big Event Number 4 of 2025, was the release on Apple TV, late in the month, of Prehistoric Planet Ice Age. As per seasons 1 and 2, it was of course years in the making. There was no Los Angeles red carpet event this time, alas, but we did do various press events and big-screen showings in London, including one at Lightroom. Here’s where I got to meet geneticist, author and science communicator Adam Rutherford.

**Caption:** Lightroom London hosted a special screening of *Prehistoric Planet Ice Age* during late November 2025. Seeing these shows projected at size is a phenomenal experience that I’ll always make time for if I can. A number of other people in the UK palaeo-scene were able to attend this showing and we hung out afterwards. Image: Darren Naish.

Apple are – with all due respect to my friends and colleagues there – an enigma in terms of when and how they do publicity, and it was quite frustrating for those of us at the BBC end of things to see the months go by with nothing, absolutely nothing, released in terms of promotion. The trailer was finally out on November 6th and the series itself was out in time for Thanksgiving (which was November 27th for 2025). In preparation for the release of the series I published articles at Tet Zoo on sloths, giant fosas, Woolly mammoths, glyptodonts, Thylacoleo and diprotodontians in general, but I didn’t get through as much as I hoped I would. More Pleistocene-themed content is yet to come at Tet Zoo.

Another technical paper – this one being the next instalment in the dinosaur cognition wars – appeared in December (Caspar et al. 2025) and the Tet Zoo article on it is of course still fresh at the time of writing. A spinosaurid paper appeared in the month too (Barker et al. 2025) but I haven’t yet had time to discuss it here. Also fresh and relevant to things covered in this article is the completely unexpected and shocking passing in January of long-time friend and colleague Richard Forrest, covered here. Not a good start to 2026, and something that will make the year’s Lyme Regis Fossil Festival, and other events, very different in tone.

**Caption:** the top shelf of one of my cabinets, rearranged for 2025. There are some real treasures here if you’re interested in model and toy animals. Image: Darren Naish.

Assessing 2025’s Tet Zoo coverage. This is a good point at which to stop and do what I do in every annual recap: assess Tet Zoo coverage across the year to see which animal groups, which subjects, won coverage and which did not. We start with a list of the year’s articles (remembering that a Tet Zoo Year extends from Jan 21st to Jan 21st), and we then view things on a graph.

SpecBio

Speculative Zoology and the World of After Man; an Interview With Dougal Dixon, April 2025
Speculative Zoology Grand and Photoreal: Boulay and Steyer's Demain, les Animaux du Futur, April 2025

Cryptozoology

Yowies and the Marsupial Hominoid Hypothesis – Neil Frost’s Fatfoot: Encounters With A Dooligahl, February 2025
Adrian Shine's A Natural History of Sea Serpents, March 2025
The Crowing Crested Cobra Once More, September 2025

Non-bird dinosaurs

In Quest of Hybrid (Non-Bird) Dinosaurs, September 2025
The Continuing Debate on Dinosaur Cognition, January 2026

Pterosaurs

Tapejarid Palaeobiology and the Concept of the ‘Giraffized’ Azhdarchid, July 2025

Turtles

World Turtle Day 2025!, May 2025

Crocodile-group archosaurs

Crocodiles Attack Elephants Then, Now, and Still, August 2025

Permian and Mesozoic swimming reptiles

Ancient Sea Reptiles Goes to Third Edition!, December 2025

Squamates

Racerunner Lizards of the World Unite, New for 2025!, April 2025
Record-Holding Champion Giant Slow-Worms, July 2025

Mammals

Curtis, Swisher and Lewin’s Java Man of 2000: Hominin-Themed Books, Part 1, March 2025
Otaria, the Southern Sea Lion, March 2025
The Fate of the Woolly Long-Nosed Armadillo of Peru, May 2025
Of Zaedyus, the Pichi, May 2025
Armadillo Empire, Part 1: of Euphractines and Eutatines, June 2025
Armadillo Empire, Part 2: Fairies, Tolypeutines, and Where Glyptodonts Go, June 2025
Whales and Dolphins Around the Coasts of Europe, 2025, July 2025
Sloth World, 2025 (Part 1), October 2025
The Amazing Giant Black Fossa, October 2025
A Woolly Mammoth Primer, November 2025
The Life Appearance of Glyptodonts, November 2025
Thylacoleo, the Incredible Marsupial Lion, November 2025
Diprotodontians Forever: the Vombatiform Radiation, Part I, November 2025

Amphibians

News From the Pond's Edge: Spawnwatch 2025, February 2025

Miscellaneous musings

Tet Zoo Reviews Zoos: Tama Zoological Park, Tokyo, February 2025
Announcing DinoCon … Tickets Now on Sale!, February 2025
Three Months to DinoCon!, May 2025
Suburban Camera Trapping, Week 1, May 2025
Tetrapod Zoology ver 4’s 7th Birthday, July 2025
Mesozoic Art II, Palaeoart Portfolio for the 2020s, September 2025
DinoCon 2025 Has Left the Building, October 2025
British Palaeontologist Richard Forrest, a Brief Obituary, January 2026

As ever, the results are both frustrating and interesting. The good number of articles relating to miscellaneous notices and announcements (concerning DinoCon and such things as a book announcement and a zoo review) obviously loom large, but if 2025 was anything at Tet Zoo it was mammal heavy. Virtually all of those mammal articles exist as a consequence of Prehistoric Planet Ice Age. Otherwise, coverage looks moderately balanced (that being the thing I ultimately strive for), the low number of articles in the bulk of categories being an inevitable consequence of my inability to do more than I already do. The total lack of birds is a surprise. Remember that I would blog so much more if only I could put more time into it. Hint hint.

Caption: ok, so I didn’t write about birds at Tet Zoo during 2025 but I did spend a lot of time looking at live ones. These photos are from a Birdworld trip made in September 2025. They show Australian pelican Pelecanus conspicillatus, Greater flamingo Phoenicopterus roseus and Magpie goose Anseranas semipalmata. Images: Darren Naish.

So far I’ve been talking about things as if 21st Jan 2026 is a normal birthday for Tet Zoo. But it isn’t: this particular birthday marks 20 years, if my maths works out. That means, my friends, that even more introspection is required. That’s what’s happening in the next article: a look back at 20 years of Tetrapod Zoology. Thanks, as ever, for visiting Tet Zoo – especially so if you’ve been with me since the beginning in 2006 – and also for leaving comments and for assisting my work and efforts to remain solvent.

**Caption:** in celebration of this landmark in Tet Zoo history we’re releasing a new lot of t-shirt designs. **They’re available at our SumUp shop** from 10.30am (GMT) on Wed 21st January.

For previous TetZoo articles on birthdays and other landmarks, see…

Happy first birthday Tetrapod Zoology (part I), January 2007
Happy first birthday Tetrapod Zoology (part II), January 2007
Happy second birthday Tetrapod Zoology (part I), January 2008
Tetrapods of 2007 (happy birthday Tet Zoo part II), January 2008
Happy THIRD birthday Tet Zoo, January 2009
Tet Zoo = 4 years old today, January 2010
2009, a year of Tet Zooery, January 2010
Four years of Tet Zoo: to infinity... and beyond!, April 2010
It is with some dismay that I announce Tet Zoo's first hemi-decade, January 2011
Tet Zoo 5th birthday extravaganza, part II, January 2011
Happy Birthday Tetrapod Zoology: SIX YEARS of blogging, January 2012
Happy 6th Birthday, Tetrapod Zoology (part II), January 2012
Tetrapod Zoology enters its 8th year of operation, January 2013 (I cannot find any intact versions of this article, thanks SciAm)
Today marks NINE YEARS of Tetrapod Zoology, January 2015
Tetrapod Zoology 10th-Birthday Extravaganza, Part 1: 2015 in Review, January 2016
Tetrapod Zoology 10th Birthday Extravaganza, Part II: the Rest of 2015 Reviewed, January 2016
Tetrapod Zoology 10th-Birthday Extravaganza, Part 3: Tet Zoo's Tetrapod Treatment in 2015, January 2016
Today Is Tet Zoo's 11th Birthday, January 2017
The 12th Year of Tet Zoo, January 2018
The Tet Zoo 12th-Birthday Event, Part 2, January 2018
The Much Belated Final Part of the Tetrapod Zoology 12th Birthday Event, December 2018
Tetrapod Zoology Is A Teenager Now, January 2019
Tetrapod Zoology's 14th Year of Operation, 2019 in Review, January 2020
On Tetrapod Zoology’s 15th Birthday, the Year in Review, January 2021
Happy 16th Birthday, Tetrapod Zoology, February 2022
Four Years of Tetrapod Zoology ver 4, July 2022
The 17th Year of Tetrapod Zoology: 2022 in Review, January 2023
Five Years of Tetrapod Zoology ver 4, July 2023
Tetrapod Zoology Reaches 18 Years of Age, January 2024
It Was the 19th Year in the History of Tetrapod Zoology, January 2025

Refs - -

Barker, C. T., Naish, D. & Gostling, N. J. 2025. Insufficient evidence for spinosaurid survival into the latest Cretaceous: a comment on Olmedo-Romaña et al. (2025). Ameghiniana 62, 572-580.

Caspar, K., Gutiérrez-Ibáñez, C., Hady, G., Holtz, T. R., Naish, D. & Hurlburt, G. R. 2025. Endothermy, neuron counts, and other issues: Further remarks on neurocognitive evolution in fossil vertebrates. The Anatomical Record doi.org/10.1002/ar.70113

Jackson, T. 2025. Eyewitness Animal. Dorling Kindersley, London.

Naish, D. 2016. Hunting Monsters. Arcturus Books, London.

Naish, D. 2022. A cultural phenomenon. The Biologist 69 (3), 16-21.

Naish, D. 2025a. Thoughts on reviewing books, from an incoming book review editor. Historical Biology, 1. https://doi.org/10.1080/08912963.2025.2471237

Naish, D. 2025b. Taxonomic trolling. The Biologist 72 (2), 26-29.

Naish, D. 2025c. Dinosaurs are not extinct. In Tucker, L. (ed) Discovering Dinosaurs. Lightroom, London, pp. 16-27.

Naish, D. 2025d. Ichthyosaurs from the Early Jurassic of Britain: by Graham P. Weedon and Sandra D. Chapman, 2022, Siri Scientific Press, 448 pp., £99.99 (Hardcover), ISBN 978-1-8381528-6-4. Historical Biology 10.1080/08912963.2025.2541788

Naish, D. 2025e. Snake summer at the Hunterian. Fortean Times 464, 52-53.

Naish, D. 2025f. Ancient Sea Reptiles (Third Edition). Natural History Museum, London.

Naish, D. 2025g. The response to and rejection of Brian Ford’s Too Big to Walk, a 21st century effort to reinstate the aquatic dinosaur hypothesis. Historical Biology 37, 2147-2156.

White, S. & Naish, D. 2025. Mesozoic Art II: Dinosaurs and Other Ancient Animals in Art. Bloomsbury Wildlife, London.

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Those of you specially interested in the behaviour and biology of Mesozoic dinosaurs will be well aware of the dinosaur cognition debate that’s been going on in the animal cognition literature…

**Caption:** a wholly speculative effort to portray potential cognition in an animal like a non-bird theropod. Which of these dotted lines is more realistic… *or*, is it totally misleading, perhaps wrong, to imagine or portray things this way? Images: Darren Naish.

As discussed at Tet Zoo back in 2024 (go here), things started when neuroscientist Suzana Herculano-Houzel argued that such extinct dinosaurs as Tyrannosaurus might have had primate-like numbers of neurons, and thus might have been similar to anthropoids in behavioural traits and overall intelligence (Herculano-Houzel 2023). I don’t need to say that these conclusions were surprising, the consensus view among those working on extinct dinosaurs otherwise thinking they were potentially similar in intelligence to extant squamates, crocs, and perhaps to such birds as ratites and gallinaceous birds.

I’m part of a team that pushed back on the details that Herculano-Houzel used to build her case: Kai Caspar, Cristian Gutiérrez-Ibáñez, Grant Hurlburt, myself and others showed that Herculano-Houzel’s brain size estimates were likely incorrect, that her neuron count estimates were unreliable and based on erroneous assumptions, and that the neuron counts she estimated weren’t clearly linked to anthropoid-like cognitive abilities anyway (Caspar et al. 2024). We published in The Anatomical Record and our article is open access. Concepts like ‘intelligence’ and ‘overall cognitive complexity’ are still difficult to quantify and describe in any meaningful way, so our takehome as expressed in popular sources (like this blog) was that animals like tyrannosaurs were more likely ‘reptile smart’ than ‘primate smart’… It is important to note, however, that it’s very hard to find us stating exactly this in our article. Consider the following statements from Caspar et al. (2024)…

“Cognitive traits identified exclusively in birds or crocodiles cannot simply be extrapolated to Mesozoic dinosaurs with any degree of certainty since they might represent crown group apomorphies. Although it might be appealing to hypothesize that cognitive patterns found among modern palaeognaths are representative for their maniraptoriform forerunners … this idea is (in most cases) not testable and should hence not be disseminated uncritically.”

“…. the extant archosaurian groups leave us in a rather suboptimal position to infer cognitive traits in non-avian dinosaurs.”

Our article (Caspar et al. 2024) was, of course, far from the end of the discussion. It turns out that people working elsewhere in animal cognition have also been following the argument, and – yes – they have thoughts.

**Caption:** ongoing studies of extant non-bird reptiles show that there’s a lot going on here in cognitive terms, and at least some are explorative, investigative creatures that are good at learning. It’s increasingly well known that monitor lizards (like the Komodo dragon at left) and iguanians like bearded dragons (like the pet one at right) pass numerous tests relating to 'intelligence’. Images: Darren Naish.

Jensen et al. (2025) respond. Thomas Rejsenhus Jensen of Lund University’s Cognitive Zoology Group, plus colleagues, responded; while not agreeing with Herculano-Houzel (2023) outright, part of their argument was that Caspar et al. (2024) might be criticized for creating the impression that a typological view holds sway in animal cognition; that reptiles (like non-bird dinosaurs) can only be ‘reptile-like’, for example. Instead, they point out, cognition involves multiple shades of grey in addition to a plasticity that relates to the specific lifestyle and adaptations of a species (Jensen et al. 2025)… a position, by the way, that we agree with. T. rex should not, then, be considered either ‘baboon-like’ or ‘crocodile-like’ but ‘T. rex-like’ and it’s misleading to use such terms as “smartness” “intelligence”, “baboon-like cognition” and “crocodile-like cognition” (Jensen et al. 2025).

**Caption:** “these animals are like x”, “no, they’re like y”, “no, they’re like what they’re like”. The articles shown here are from a debate that occurred in the journal *Evolution* between 1970 and 1974.

In addition, they argued that we’d ignored the ‘endothermic brain hypothesis’ and also suggested the presence of a ‘cognitive arms race’ as a potential explanation for the evolution of convergent neurocognitive traits in mammal-line and bird-line animals in the Mesozoic (Jensen et al. 2025).

There’s a lot to consider here, and late last year another article was published on this topic, this one being a response to Jensen et al. (2025) from Kai Caspar, Cristian Gutiérrez-Ibáñez, Grant Hurlburt, myself and others (Caspar et al. 2025). We took issue with several of Jensen et al. (2025)’s key points. Here, I summarise these and explain how we responded. Our article was again published in The Anatomical Record and is again open access. We owe substantial thanks to the editorial team at The Anatomical Record for swift and efficient handling, plus to our reviewers and colleagues.

**Caption:** the topic being covered here is of broad interest and *will* get covered in the popular and semi-technical press. In view of this, what language do we use and which comparisons do we make when discussing the subject? The articles here are from **CBBC** and Tet Zoo; I can no longer find ‘Scientists change their minds about *Tyrannosaurus rex* yet again’ (a similar article by the same author **is here**).

On typology and non-bird dinosaurs as ‘reptile-brained’. As noted above, Jensen et al. (2025) accused us (Caspar et al. 2024) of simplifying or underselling the issue of dinosaur cognition. But this is complicated by the fact that the discussion as a whole is occurring in two locations: (1) the technical literature, and (2) the popular sphere (blog articles like the one you’re reading now, and in the popular press, since journalists enjoy covering this topic). Nowhere in our technical article did we (Caspar et al. 2024) use the terms that might be associated with a typological view: yes, we drew attention to the fact that extinct dinosaurs are often more ‘crocodylian-like’ in brain form and proportional size than birds, but we stated a nuanced position when it comes to what this might mean for cognition, as is clear from the quotes from Caspar et al. (2024) provided above.

When it comes to summarising the situation in popular sources, I don’t see an issue with using terms like ‘smartness’ and ‘intelligence’ so long as caveats, or quote marks (as here), are used. It remains difficult, if not impossible, to quantify or precisely measure overall cognitive complexity, meaning that approximate comparisons to living taxa are about the best way we have of describing things. Saying that dinosaurs like tyrannosaurs might have been approximately similar in intelligence to certain lizards or crocodylians, for example, is reasonable based on what we know (and, as I’ve emphasized here, not ‘a bad thing’ in terms of what it might mean for behavioural complexity, adaptability and so on). Jensen et al. (2025) appeared under the impression that the notion of ‘reptile-brained’ (non-bird) dinosaurs had been abandoned by scientists working on these animals, but that’s not so, as we demonstrated via quotes and citations from recent work on dinosaur biology (Caspar et al. 2025).

**Caption:** there’s a popular idea that palaeognaths (which include ratites, like this rhea) are not real birds and don’t do birdy things. Having spent a lot of time with them (cassowaries especially)… no. This captive rhea spent a good while engaging in what sure looked like object play with a feather. An argument has been made that some extinct dinosaurs were similar to ratites in cognitive terms. Images: Darren Naish.

An aside on nomenclature. As someone who sometimes publishes on the fossil record and diversity of crocodile-line archosaurs, I always make a point of using the specific terms that have been established in the technical phylogenetic literature. The term crocodile applies to one specific group of these animals and people – and this applies especially to palaeontologists – really, really need to wean themselves off the childish idea that it can be used indiscriminately.

The crown-group – the one that includes alligators, crocodiles and their close relatives – is Crocodylia, which we use to avoid confusion with the old ‘Crocodilia’, a term used in the past for the entire group (meaning Crocodylomorpha, and incorporating a massive diversity of extinct archosaurs that are wholly different in shape and behaviour from crocodylians). In short, modern crocodiles, alligators are kin should be referred to collectively as crocodylians. Back to the main issue at hand…

**Caption:** a substantially simplified cladogram showing relationships within crocodylomorphs. There is still a tendency within the natural history and zoological literature for the crown-group to be termed the ‘crocodilians’ but the name ‘crocodylian’ is more firmly established within phylogenetic literature. The incorrect tradition of referring to any and all of these animals as ‘crocodiles’ also needs to be put to bed. Crocodiles are a specific group of crocodylians. Image: Darren Naish.

Is the ‘endothermic brain hypothesis’ valid for Mesozoic vertebrates? The main reason we hadn’t mentioned the endothermic brain hypothesis or EBH in our initial article (Caspar et al. 2024) is that it hadn’t been proposed in print when we were writing. Its first published outing was in Osvath et al. (2024), which appeared in November 2024: Caspar et al. (2024) appeared in April 2024. Simply put, the EBH argues that whole-body tachymetabolic endotherms have evolved unusual cognitive traits (like larger brain size and higher neuron counts) in step with their need to find greater amounts of fuel for their elevated metabolisms, and that they operate via an advanced ‘model-based’ system of optimized foraging that requires greater cognitive processing than that present in other animals (Osvath et al. 2024).

Our main point of response (Caspar et al. 2025) is that robust evidence for a link between the evolution of endothermy and a marked change in brain anatomy (especially one measurable in fossils) is lacking. Fossils simply don’t show an anatomical ‘upgrading’ in brain form or complexity occurring in step with the hypothesised development of endothermy among the relevant groups. In addition, the idea that a modified (as in, proportionally big or especially neuron-dense) brain is associated with endothermy is countered by fossil pseudosuchians (which apparently combined endothermy with proportionally small brains) and big-brained, neuronally dense but ectothermic fishes (Caspar et al. 2025).

**Captions:** these images are here to remind you that there are actinopterygians (ray-finned fishes) and chondrichthyans (cartilaginous fishes) with proportionally enormous brains and extremely high neuron densities. How do these fit into patterns proposed for terrestrial animals like archosaurs? Images: opencage, CC BY-SA 2.5 (original **here**); Shiyam ElkCloner, CC BY-SA 3.0 (original **here**).

Was there a Mesozoic ‘cognitive arms race’? Jensen et al. (2025) proposed that the evolution of endothermy in synapsids and archosaurs might have been engaged in a “cognitive competition for food” and that this “could have boosted success compared to less cognitively apt competitors”. They were inspired by Benton’s (2021) very useful overview on the evidence for endothermy in Permo-Triassic synapsids and archosaurs, in which a ‘Triassic arms race’ relating to postural and locomotor changes was suggested.

**Caption:** mammal-line and bird-line animals interacted continuously throughout the Mesozoic, as they do today. But is there any evidence that anything like a cognitive ‘arms race’ was underway? This image, from the Apple TV series *Prehistoric Planet*, shows an interaction between a troodontid theropod and a multituberculate. Image: © Apple / BBC Studios.

There may or may not be merit to this idea. But even if there is, the fossil record does not show a concomitant cognitive ‘up-grade’ occurring in either group at the right time. “The proposal is thus implausible to us” (Caspar et al. 2025). See the article for the full discussion and relevant data.

Neuron counts and relative brain size again. Finally, we end our article by discussing the fact that neuron counts are too ‘noisy’ a metric to be especially reliable when it comes to making inferences about cognition. Jensen et al. (2025) criticized us for apparently questioning the idea that neuron counts are linked to cognitive ability, but we didn’t exactly do that: our point instead was that neuron counts are one among many things that need to be considered, and that they can’t be used alone as a proxy in making conclusions (as they were by Herculano-Houzel) (Caspar et al. 2024, 2025).

In fact, despite the argument from Jensen et al. (2025) that a clear link between raw neuron count and cognition exists, it’s still difficult to pin down the tightness of this correlation and determine whether absolute or relative neuron count is more important. For birds, most work indicates that relative brain size holds sway. Big members of animal groups have high neuron counts relative to small members of those groups because high counts correlate with overall size and don’t clearly grant cognitive advantages (Caspar et al. 2025).

**Caption:** graphs from **Caspar *et al*. (2025)** showing neuron count in the telencephalon plotted against body mass in mammals (at left) and birds (at right). The line in the mammal graph shows the regression line for primates, showing that *Homo sapiens* has a neuron count expected for its body mass. The line in the bird graph shows likewise for Psittacopasserae, here showing that *Corvus corax* demonstrates the same thing within its clade. Image: **Caspar *et al*. (2025)**.

Clearly, much remains to be learnt about the anatomy of dinosaur brains and what that anatomy does, or does not, mean for behaviour and cognition. The notion that we can make simple inferences about cognition based on overall physiology (the ‘endothermic brain hypothesis’) or that brain enlargement and hence behavioural complexity occurred as the consequence of a Mesozoic ‘arms race’ are both, however, so flawed that they should probably be considered wrong, and sections of our new article are devoted to discussing and refuting these ideas (Caspar et al. 2025).

As should be clear by now, this is very much an active area of debate and consideration. It’s difficult not to be intrigued and even excited by claims that Mesozoic dinosaurs possessed “largely unknown neurocognitive functions approaching those seen in birds” (Jensen et al. 2025). But on dinosaur cognition overall, these animals quite probably overlapped with turtles, squamates and crocodylians as well as birds (Caspar et al. 2024, 2025), the great caveat being that our understanding of cognitive traits in so many of these animals remain very much rudimentary. Again, this mustn’t be interpreted to mean that the extinct animals were deficient or poor in performance given what we currently think about learning, complexity and memory in those living animals.

Acknowledgements. I thank Kai Caspar for checking the text and providing useful suggestions and corrections.

For previous Tetrapod Zoology articles on dinosaur brains, biology and connected issues, see…

The Sensitive Face of a Big Predatory Dinosaur, June 2017
Could We Domesticate (Non-Bird) Dinosaurs?, August 2018
Postcranial Palaeoneurology and the Lifestyles of Pterosaurs, August 2018
Alternative Timeline Dinosaurs, the View From 2019 (Part 3): the Dinosauroid and its Chums, December 2019
Did Dinosaurs and Pterosaurs 'Glow'? Extinct Archosaurs and the Capacity for Photoluminescent Visual Displays, March 2020
Humanoid Dinosaurs Revisited Again: Russell and Séguin’s Dinosauroid at (Nearly) 40 Years Old, August 2021
A brain for Baryonyx: using CT-scanning to examine British spinosaurid brains, February 2023
T. rex the Tool Maker -- Testing Controversial Claims About Dinosaur Smarts, April 2024

You can support this blog – and my work in general – at patreon for as little as $1 per month. Do that, and you also get to see behind-the-scenes and in-prep material I’m working on. Huge thanks to everyone who helps.

Refs - -

Benton, M. J. 2021. The origin of endothermy in synapsids and archosaurs and arms races in the Triassic. Gondwana Research 100, 261-289.

Caspar, K., Gutiérrez-Ibáñez, C., Ornella, B. C., Carr, T., Colbourne, J., Erb, A., Hady, G., Holtz, T. R., Naish, D., Wylie, D. R. & Hurlburt, G. R. 2024. How smart was T. rex? Testing claims of exceptional cognition in dinosaurs and the application of neuron count estimates in palaeontological research. The Anatomical Record 2024, doi 10.1002/ar.25459.

Herculano-Houzel, S. 2023. Theropod dinosaurs had primate-like numbers of telencephalic neurons. Journal of Comparative Neurology 531, 962-974.

Jensen, T. R., Jacobs, I., Kverková, K., Lalic, L., Polonyiová, A., Stehlík, P., Reber, S. A., & Osvath, M. 2025. T. rex cognition was T. rex-like – A critical outlook on diverging views of the neurocognitive evolution in dinosaurs. The Anatomical Record doi.org/10.1002/ar.70074

Osvath, M., Němec, P., Brusatte, S. L., & Witmer, L. M. 2024. Thought for food: the endothermic brain hypothesis. Trends in Cognitive Sciences 28, 998-1010.

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Today (Friday 9th January 2026) comes the sad announcement that UK-based marine reptile worker Richard Forrest is no longer with us, this information having been passed to me from Sally Hollingworth, Mammoth Project Leader for the Cerney Wick excavations, and Neil Gostling of the University of Southampton. This ultimately comes from Sue Forrest, Richard’s wife.

**Caption:** there are, unsurprisingly, a great many photos online showing Richard with plesiosaur fossils. In this one he’s examining bones of the Scunthorpe pliosaur at North Lincolnshire Museum. Image: (c) North Lincolnshire Museum.

This is a huge shock and a major loss. Richard is a notable person in the British Mesozoic research community, known in particular for his work on plesiosaurs (and other marine reptiles) (e.g., Benson et al. 2013, Forrest 1998, 2000, 2003a, b, c, 2005, Forrest & Oliver 2003) and his significant role in running, organising and archiving conferences and meetings, in particular SVPCA (the Symposium on Vertebrate Palaeontology and Comparative Anatomy). Richard hosted the website for that meeting and was responsible for archiving images and recollections from years past. A brief celebration of Richard’s interests and work formed the focus of a Guardian article published in 2024.

**Caption:** some reasonably bad photos featuring Richard, and taken at various of the SVPCA meetings. At left, an especially bad one from 2001, taken at the Square Tower in Portsmouth (Darren Naish at left, Richard Forrest at right). At right, Richard with noted palaeoartist Bob Nicholls at SVPCA Oxford, September 2012.

I've known Richard since the late 1990s and he was a mentor, advisor and friend to many of us. At this point in his life and career, he was – whether he liked it or not – regarded as a pillar of the British Mesozoic marine scene and a wise old Gandalfian holder of skill and knowledge. Trained as an architect and an expert at compiling databases, Richard did a massive amount to build collaborations, helped with excavations across the UK, and was also great at passing on his advice and enthusiasm for the subject. He was warm and welcoming and promoted inclusion, collaboration and the role of amateurs in science. In short, he was exactly the sort of person you want in a field like palaeontology: passionate and interested, careful and always aiming to improve his skill and knowledge, respectful of data and detail but always prepared to consider outside and left-field ideas, keen to work with professionals and institutions as well as amateurs and people wholly new to the field, and appropriately supportive of everyone he was connected to. He was an expert preparator as well as a publishing researcher.

**Caption:** Richard (in the middle, with blue t-shirt) was one of many people who worked hard at Jeff Liston’s 2002 Star Pit excavation of Ariston the *Leedsichthys* specimen. None of my photos are good but it should be obvious from this one that torrential rain put an end to the day’s work on this occasion. Image: Darren Naish.

We were actively working together on a very exciting project and he also attended TetZooCon on several occasions; he gave a talk at the 2023 event (which included a Mesozoic marine reptiles selection of talks).

**Caption:** Richard Forrest (RIP Jan 2026) at right, with Darren (left) and Neil Gostling (centre) during July 2025. Photo taken near Neil’s office at the University of Southampton.

Richard leaves behind a substantial family; condolences to everyone there. I only know his wife Sue but am aware that he had many children and grandchildren. My photos are a random mix from the 2000s and 2010s (the oldest is from 2001) plus others that are more recent. I'm sure that longer and more adept obits will appear in time... farewell old friend, you will be much missed.

Please add thoughts and recollections below, and thank you for doing so.

Refs - -

Benson, R. B. J., Evans, M., Smith, A. S., Sassoon, J., Moore-Faye, S., Ketchum, H. F. & Forrest, R. 2013. A giant pliosaurid skull from the Late Jurassic of England. PLOS ONE 8,e65989.

Forrest, R. 1998. A possible early elasmosaurian plesiosaur from the Triassic/Jurassic boundary of Nottinghamshire. Mercian Geologist 14, 135-143.

Forrest, R. 2000. A large rhomaleosaurid pliosaur from the Upper Lias of Rutland. Mercian Geologist 15, 37-40.

Forrest, R. 2003a. Evidence for scavenging by the marine crocodile Metriorhynchus on the carcass of a plesiosaur. Proceedings of the Geologists’ Association 114, 363-366.

Forrest, R. 2003b. Notes on a specimen of the plesiosaur Plesiosaurus dolichodeirus (Reptilia; Plesiosauria) from the Lower Lias of Charmouth. Proceedings of the Dorset Natural History and Archaeological Society 125, 101-104.

Forrest, R. 2003c. Taphonomic distortion of cervical vertebrae of a specimen of Plesiosaurus dolichodeirus (Reptilia; Plesiosauria) from the Lower Lias of Charmouth. Proceedings of the Dorset Natural History and Archaeological Society 125, 105-108.

Forrest, R. 2005. The application of multivariate analysis in the reconstruction of the skeleton of a specimen of Muraenosaurus cf. leedsi. The Quarterly Journal of the Dinosaur Society 4 (3), 22-29.

Forrest, R. & Oliver, N. 2003. Ichthyosaurs and plesiosaurs from the Lower Spilsby Sandstone Member (Upper Jurassic), north Lincolnshire. Proceedings of the Yorkshire Geological Society 54, 269-275.

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Yes … it’s time again to talk about the sloths, how I love them so. By the way, over recent months I’ve had to train myself to pronounce ‘sloth’ in the American way, not ‘sloathe’…

**Caption:** there are two very basic things to remember about sloths... that there are small, tree-climbing ones, and a substantial diversity of mostly larger or MUCH larger, extinct ones. Here are two exemplars of those facts: a captive *Choloepus* at left, and a museum skeleton (a cast) of the South American giant *Megatherium* (at the Natural History Museum, London) at right. Images: Darren Naish.

Following the armadillo-themed articles that appeared here within recent months, it only seems fair to give sloths some coverage too. And such a thing is well overdue, since I haven’t written properly about sloths for years now. Indeed, a sloth-themed review article I published back in 2005 (Naish 2005) is now chronically out of date with respect to what it says about sloth phylogeny, diversity and history.

Sloths are a big group and there’s a lot to say about them, so my aim here is to focus on a few select highlights. I can say now that – yes – sloths are a group I’ve been considering at length within recent years thanks to Prehistoric Planet Ice Age, due out in late November 2025. More on that in due time.

**Caption:** there’s a fair bit about sloths in the Tet Zoo archives, but it’s now hard to find and much of it has been ruined by removal of images and such. Here are two of the relevant articles, one from 2007, one from 2010. Of relevance here is that I’m gradually building a new archive of Tet Zoo articles (**it’s here**), aiming where possible to find intact versions at the internet archive.

Sloths of the Caribbean. We’ve known since the 1860s that sloths inhabited various of the Caribbean islands, since that’s when Megalocnus rodens – initially deemed notable because of its rodent-like anterior dentition – was described from the Pleistocene of Cuba. Megalocnus was similar in size to an average bear and seems from limb proportions and foot shape to have been a terrestrial, ground-feeding animal.

Later finds showed that similar, related sloths occurred in Puerto Rico (Acratocnus) and Hispaniola (other species of Acratocnus, plus Parocnus and Neocnus); additional taxa lived in Cuba too (Imagocnus, and other species of Acratocnus and Neocnus). As many as six species lived on certain of these islands (Viñola-Lopez et al. 2021). These were generally smaller than Megalocnus (the Neocnus species were tiny; smaller than most living tree sloths at less than 15 kg) and some (like Acratocnus) were especially short-snouted. Excepting Imagocnus (which is Miocene), these are animals of the Late Pleistocene and Holocene, and remains of Megalocnus and Parocnus (plus of indeterminate species) show that some were alive 6000-5000 years ago (Steadman et al. 2005). They’re modern and recently extinct, not prehistoric.

**Caption:** megalocnoid skulls from Hispaniola as illustrated by **MacPhee *et al*. (2000)**. (A) *Acratocnus ye*, (B) *Neocnus dousman* and (C) *N. toupiti*. Note the scale bars: the skulls of these animals are (relatively!) tiny (it’s 9.7 cm in *N. toupiti*). The skulls of these animals are also narrow across the snout. Image: **MacPhee *et al*. (2000)**.

All the sloths mentioned here share traits and have traditionally been grouped together in Megalocnidae. A popular idea on their phylogenetic placement has been that they’re part of Megalonychidae – the group anchored on Megalonyx, a particular famous North American ground sloth – and, within that group, close to Choloepus, the extant two-fingered sloths (e.g., Gaudin 1995, 2004, Pujos et al. 2007). This seems logical in view of superficial similarities in skull form and the small size and probable semi-arboreal lifestyle of Caribbean taxa like Neocnus*.

I’ve always disliked the terms ‘two-toed’ and ‘three-toed’ for extant sloth species, given that we’re talking about fingers, not toes… so I welcome new efforts to change the common names of these animals.

Caption: a simplified version of the morphology-based view of sloth phylogeny generally thought correct prior to the 2010s. Bradypus was regarded as the earliest diverging sloth lineage (in which case it has a suspiciously long ghost lineage), nothrotheres and megatheriids were considered close kin, and Choloepus was a megalocnid and hence part of one of the youngest sloth groups. Images: Darren Naish.

However, molecular results indicate that this is completely wrong. Instead, it seems that the Caribbean sloths represent an entirely distinct lineage that diverged early on within sloth history, way back in the Eocene (Delsuc et al. 2019, Presslee et al. 2019, Viñola-Lopez et al. 2021). They’re thus not close to Megalonyx or kin, nor to Choloepus, and their early colonization of the Caribbean has required a rethink on how they became distributed. In addition, an Oligocene divergence of the Aractocnus and Parocnus lineages (it seemingly occurred around 30 million years ago) has led to a revised taxonomy where Acratocnidae and Parocnidae are regarded as the main constituent groups within an ‘up-ranked’ Megalocnoidea (Delsuc et al. 2019).

**Caption:** molecular data indicates that the pattern of sloth evolution was rather different from what we thought based on anatomy. Caribbean sloths are the sister-group to the rest of the clade and *Choloepus* is not close to this group (instead being closer to mylodontids); *Bradypus* does not belong to an especially archaic lineage but is instead within the clade that includes megatheres and nothrotheres. Image: Darren Naish.

Big megatheres: furry, naked or both or neither? Within recent years, the view that extinct sloths of (essentially) all sorts should be imagined as shaggily-pelted has come into question. We have direct evidence that at least some mid-sized, extinct sloths (namely Mylodon and Nothrotheriops) were like this, since of course we have preserved segments of their pelt. But was it true for all of them, in particular the very big ones that lived (in parts of their ranges) in tropical climes?

**Caption:** one of the famous fragments of *Mylodon* skin (with pelt) from Cueva del Milodón Natural Monument in Chilean Patagonia, specifically the one collected by Francisco P. Moreno in 1897. Despite claims that sloth fur is coarse and stiff, you’ll note that the hair here actually has a soft and lustrous look. Small bony nodules are embedded on the skin’s inner surface. This specimen is in the collections of the Natural History Museum, London. Some dung from the same location is visible at back. Image: Darren Naish.

On a few occasions, experts have argued that giant taxa – in particular the megatheres Megatherium and its close relative Eremotherium – were more sparsely haired, perhaps recalling extant elephants or domestic pigs in hair coverage. Supposedly, they would have benefited from this, since a thick pelt would result in heat stress (Fariña 2002). However, sloths have a relatively low metabolism compared to elephants and pigs (and this is apparently true even if we accept arguments about extinct sloths having higher metabolisms than conventionally assumed; Varela et al. 2024), so… are we sure that it’s ‘one integumentary rule applies to all’ when it comes to physiological arguments of this sort?

Exactly this was examined by Deak et al. (2025). By simulating the climatic regimes that would have been experienced by megatheres when they were alive and combining this with a metabolic rate, Deak et al. (2025) found that megatheres would have suffered from cold stress across large swathes of their range if they had sparse or absent fur. A dense, long coat would likely have caused heat stress in tropical places, but a dense and thick pelage was otherwise needed even for these giants. In view of this, Megatherium and Eremotherium might even have been variable in integument across their ranges (Deak et al. 2025).

**Caption:** at left, a graph from **Deak *et al*. (2025)** showing the temperature regimes experienced by select extinct sloths. A famous ‘tropical’ sloth like the gigantic *Eremotherium* experienced warm temperatures in part of its range, but it lived in places that were cool and cold as well. At right, *Eremotherium laurillardi* skeleton at Houston Museum of Nature and Science. Images: **Deak *et al*. (2025)**; Kamraman, CC BY-SA 2.5 (original **here**).

Marine sloths. If you’re at all similar in age to myself, you’ll remember how exciting it was to see the 1990s discovery of Thalassocnus natans, a coastal South American sloth, initially described from the Upper Miocene of Peru, that possesses adaptations throughout the skeleton for swimming (de Muizon & McDonald 1995). Additional, geologically younger Thalassocnus species were discovered later on, some of which look to be more specialized for swimming than T. natans (de Muizon et al. 2003, 2004a).

**Caption:** the 1995 publication of *Thalassocnus natans* from Peru caused quite the sensation. As you see from this articulated skeleton (on show at Muséum National d'Histoire Naturelle, Paris), it’s not tremendously modified relative to other sloths: adaptations in the tail, hindlimb and skull are consistent with a wading, aquatic feeding lifestyle. However, the discovery since 1995 of other thalassocnine species show that members of this lineage became increasingly modified for marine life over time. Image: FunkMonk, CC BY-SA 3.0 (original **here**).

All species are mid-sized as sloths go, with body length estimated at being between 2.5 and 3 m long. In the youngest species – T. yaucensis from the Late Pliocene – the bones at the front of the skull are elongate and narrow and indicative of flexible lips specialized for marine feeding. T. yaucensis also lacks microwear indicative of sand ingestion and hence was apparently feeding in deeper water than the less specialized species (de Muizon et al. 2004b). In the rest of the skeleton, the radius superficially recalls that of pinnipeds (de Muizon et al. 2003). As tempting as it is to imagine these last members of the group as pelagic ‘seal-sloths’, if you like, it might be that they were more adapted for shallow-water floating and walking and not under evolutionary pressure to evolve a fusiform shape. They’re not unique to Peru by the way, later finds demonstrating a presence in Argentina and Chile.

This lineage – grouped together as Thalassocninae – was originally included within the nothrotheres (read on for more on that group), and some studies do still support this view (Pujos et al. 2016). However, other studies place them within Megatheriidae (Amson et al. 2017), the sloth group best known for including the giant megatheres discussed above. Incidentally, suggestions have been made that other extinct sloths were adapted for life in water. Lestodon – a robust-skulled mylodontid with thick and prominent caniniform teeth – has been compared to hippos on occasion and hence imagined as an amphibious animal. I’d like to see this formally tested though.

**Caption:** *Lestodon* of the Pliocene, Pleistocene and Holocene of Brazil, Paraguay, Uruguay and Argentina is a large and hilariously chunky mylodontid sloth with a notably broad, robust rostrum. The suggestion has been made several times that it might have been hippo-like in some aspects of behaviour and ecology. Images: skeletal reconstruction modified from **Tomassini *et al*. (2020)**; the skull illustration is from a paper by Greg McDonald but I’m unsure of its original provenance, CC BY 4.0 (original **here**).

Shasta ground sloths and other nothrotheres. Excluding the thalassocnines, nothrotheres – or nothrotheriids to be more formal – are a group of small- and mid-sized, mostly terrestrial sloths that lack the specializations typical of other groups and share relatively narrow cheek teeth, a prominent and asymmetrical vomerine keel on the bony surface of the pharyngeal cavity (Gaudin 2004), bulbous mastoid processes at the back of the skull and a list of other traits in the skull, forelimb and ankle. They have slender, more elongate arm bones than most other extinct sloth groups. Genetic data shows that they aren’t archaic sloths, but a relatively young group akin to megalonychids and the extant three-fingered Bradypus (Delsuc et al. 2019).

**Caption:** skeleton of the familiar nothrothere *Nothrotheriops*, aka Shasta ground sloth, as displayed at Peabody Museum. At right, a 1938 photo of *Nothrotheriops* dung from Rampart Cave in Arizona. This dried dung is simply old and dry and has never been buried, so I don’t think people have ever referred to it as ‘fossil’. Images: public domain (originals **here** and **here**).

The history of thoughts on nothrothere taxonomy and phylogeny is complex and involves more taxa than I have time to talk about here. A number of Miocene taxa appear to be early-diverging members of the group outside a Pliocene-Holocene clade (Nothrotheriinae) that includes Nothrotherium itself, a well-studied animal from the Pleistocene and Holocene of South America. Coprolites and isotopic data show that this sloth inhabited parkland-like forests and ate dry-adapted fruits and leaves. Similar things can be said for the related Nothrotheriops, one species of which (N. shastensis) is known as the Shasta ground sloth following its early 20th century discovery in Shasta County, California. Copious air-dried dung belonging to N. shastensis is known from the American south-west and shows that these animals used caves and rocky overhangs as refuge or resting sites. Related nothrotheres include Aymaratherium from the Pliocene of Bolivia, Pronothrotherium from the Miocene and Pliocene of Brazil and Mionothropus from the Miocene of Peru (De Iuliis et al. 2011, Pujos et al. 2016).

**Caption:** cladogram and map showing nothrothere evolution, from **Pujos *et al*. (2016)**. This work shows nothrotheres as a mostly Miocene radiation of non-equatorial South America, one lineage of which migrated into North America. It remains controversial whether thalassocnines are nothrotheres, since they also share traits with megatheres. Image: **Pujos *et al*. (2016)**.

That was a very brief look at just a few aspects of extinct sloth diversity and biology. There is so much more to say and I absolutely must come back to them in time. Watch this space.

For previous Tet Zoo articles on xenarthrans, see…

Five things you didn’t know about armadillos, June 2007
What was that skull? (on glyptodonts), May 2008
I, Priodontes, the tatuasu, September 2008
Predation and Corpse-Eating in Armadillos, September 2010
The Fate of the Woolly Long-Nosed Armadillo of Peru, April 2024
Of Zaedyus, the Pichi, May 2025
Armadillo Empire, Part 1: of Euphractines and Eutatines, June 2025
Armadillo Empire, Part 2: Fairies, Tolypeutines, and Where Glyptodonts Go (July 2025)

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for Tet Zoo and the more productive I can be on those long-overdue book projects. Thanks!

Refs - -

Amson, E., Muizon, C. de & Gaudin, T. J. 2017. A reappraisal of the phylogeny of the Megatheria (Mammalia: Tardigrada), with an emphasis on the relationships of the Thalassocninae, the marine sloths. Zoological Journal of the Linnean Society 179, 217-236.

Deak, M. D., Porter, W. P., Mathewson, P. D., Lovelace, D. M., Flores, R. J., Tripati, A. K., Eagle, R. A., Schwartz, D. M. & Butcher, M. T. 2025. Metabolic skinflint or spendthrift? Insights into ground sloth integument and thermophysiology revealed by biophysical modeling and clumped isotope paleothermometry. Journal of Mammalian Evolution 32, 1.

De Iuliis, G., Gaudin, T. J. & Vicars, M. J. 2011. A new genus and species of nothrotheriid sloth (Xenarthra, Tardigrada, Nothrotheriidae) from the Late Miocene (Huayquerian) of Peru. Palaeontology 54, 171-205.

Delsuc, F., Kuch, M., Gibb, G. C., Billet, G., Hautier, L. & Poinar, H. N. 2019. Ancient mitogenomes reveal the evolutionary history and biogeography of sloths. Current Biology 29, P2031-2042.

Fariña, R. A. 2002 Megatherium, the hairless: appearance of the great Quaternary sloths (Mammalia; Xenarthra). Ameghiniana 39, 241-244.

Gaudin, T. J. 1995. The ear region of edentates and the phylogeny of the Tardigrada (Mammalia, Xenarthra). Journal of Vertebrate Paleontology 15, 672-705.

Gaudin, T. J. 2004. Phylogenetic relationships among sloths (Mammalia, Xenarthra, Tardigrada): the craniodental evidence. Zoological Journal of the Linnean Society 140, 255-305.

MacPhee, R. D. E., White, J. L. & Woods, C. A. 2000. New megalonychid sloths (Phyllophaga, Xenarthra) from the Quaternary of Hispaniola. American Museum Novitates 3303, 1-32.

Muizon, C. de & McDonald, H. G. 1995. An aquatic sloth from the Pliocene of Peru. Nature 375, 224-227.

Muizon, C. de, McDonald, H. G., Salas, R. & Urbina, M. 2003. A new early species of the aquatic sloth Thalassocnus (Mammalia, Xenarthra) from the Late Miocene of Peru. Journal of Vertebrate Paleontology 23, 886-894.

Muizon, C. de, McDonald, H. G., Salas, R. & Urbina, M. 2004a. The youngest species of the aquatic sloth Thalassocnus and a reassessment of the relationships of the nothrothere sloths (Mammalia: Xenarthra). Journal of Vertebrate Paleontology 24, 387-397.

Muizon, C. de, McDonald, H. G., Salas, R. & Urbina, M. 2004b. The evolution of feeding adaptations of the aquatic sloth Thalassocnus. Journal of Vertebrate Paleontology 24, 398-410.

Naish, D. 2005. Fossils explained 51: sloths. Geology Today 21 (6), 232-238.

Presslee, P., Slater, G. J., Pujos, F., Forasiepi, A. M., Fischer, R., Molloy, K., Mackie, M., Olsen, J. V., Kramarz, A., Taglioretti, M., Scaglia, F., Lezcano, M., Southon, J., Feranec, R., Bloch, J., Hajduk, A., Martin, F. M., Gismondi, R. S., Reguero, M., Muizon, C. de, Greenwood, A., Chait, B. T., Penkman, K. & MacPhee, R. D. E. 2019. Palaeoproteomics resolves sloth relationships. Nature Ecology & Evolution 3, 1121-1130.

Pujos, F., De Iuliis, G., Quispe, B. M., Adnet, S., Flores, R. A., Billet, G., Fernández-Monescillo, M., Marivaux, L., Münch, P., Prámparo, M. B. & Antoine, P.-O. 2016. A new nothrotheriid xenarthran from the early Pliocene of Pomata-Ayte (Bolivia): new insights into the caniniform–molariform transition in sloths. Zoological Journal of the Linnean Society 178, 679-712.

Pujos, F., de Iuliis, G., Argot, C. & Lars, W. 2007. A peculiar climbing Megalonychidae from the Pleistocene of Peru and its implication for sloth history. Zoological Journal of the Linnean Society 149, 179-235.

Steadman, D. W., Martin, P. S., MacPhee, R. D. E., Jull, A. J. T., McDonald, H. G., Woods, C. A., Iturralde-Vinent, M. & Hodgins, G. W. L. 2005. Asynchronous extinction of late Quaternary sloths on continents and islands. Proceedings of the National Academic of Sciences 102, 11763-11768.

Tomassini, R. L., Montalvo, C. I., Garrone, M. C., Domingo, L., Ferigolo, J., Cruz, L. E., Sanz-Pérez, D., Fernández-Jalvo, Y. & Cerda, I. A. 2020. Gregariousness in the giant sloth Lestodon (Xenarthra): multi-proxy approach of a bonebed from the Last Maximum Glacial of Argentine Pampas. Scientific Reports 10,10955.

Varela, L., Tambusso, S. & Fariña, R. 2024. Femora nutrient foramina and aerobic capacity in giant extinct xenarthrans. PeerJ 12: e17815.

Viñola-Lopez, L. W., Suárez, E. E. C., Vélez-Juarbe, J., Milan, J. N. A. & Bloch, J. I. 2021. The oldest known record of a ground sloth (Mammalia, Xenarthra, Folivora) from Hispaniola: evolutionary and paleobiogeographical implications. Journal of Paleontology 96, 684-691.

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Among the most successful of books I’ve been involved in are those devoted to palaeoart. In particular, I’m thinking here of 2022’s Mesozoic Art, edited by Steve White and myself and published by Bloomsbury UK…

Well received critically and commercially, *Mesozoic Art* has done, and still does, a good job of showcasing modern palaeoart (White & Naish 2022). Maybe it always was inevitable that a second volume would appear, one bigger and (arguably) better than the first.

**Caption:** our cover image is by Anthony Hutchings, this one depicting a social interaction between two psittacosaurs. At right, social media avatars of your two humble author-editors (mine is by Ethan Kocak).

This week sees the publication (in the UK!) of just such a successor, namely Mesozoic Art II (MAII from hereon), again edited by Steve and myself, again published by Bloomsbury (White & Naish 2025). Here, we take a quick look at this lavish, riotously colourful, spectacular volume that showcases the state of modern palaeoart, one that we hope audiences will enjoy. For all its size and quality, MAII is also highly affordable: please buy it here. Its size means that shipping costs will add substantially to the cost, alas, but we hope that our friends and colleagues around the world will be able to obtain it.

Mesozoic Art II, the contents. MAII is large (31 x 28 cm), and includes a foreword kindly provided by conservationist, TV presenter, photographer and author Chris Packham in addition to an introduction penned by Steve and myself (White & Naish 2025). MAII is, however, very much devoted to its 25 separate artist portfolios. One of the main tasks that Steve and I (and the Bloomsbury team) face in compiling these volumes is selecting artists who both produce the sort of art we want and have a working portfolio.

**Caption:** contents page from *MAII*, showing thumbnails for our 25 amazing artists.

Included among these for MAII are established professionals whose work is well known due to its inclusion in books and museum exhibitions, Andrey Atuchin, Brian Engh, Beth Zaiken and Bob Nicholls among them. We also feature the work of technically qualified palaeontologists who also produce incredible art, like Henry Sharpe and Natalia Jagielska. Then there are artists who’ve come to us from outside of conventional palaeoart, like concept artist Gaëlle Seguillon and comic artist Lewis Larosa. And notable too is our inclusion of portfolios by people who are getting big-time exposure in print for perhaps the first time, among them DJ Washington, Haider Jaffri, Ramón M. González and Mattia Yuri Messina. We’re talking 256 pages of large-format, high standard, full-colour palaeoart excellence.

**Caption:** producing captions for these books is quite the challenge, since a delicate balance has to be found. How much technical content do we share about the organisms in the art, how much do we say about the actual art and its backstory, and are we pitching this for a lay-audience or a more technical one? We reach a compromise but there’s still room for improvement. Image: Ramón M. González, from White & Naish (2025).

What do we mean by ‘Mesozoic Art’ anyway? As I’ve stated several times (including in both MA and MAII), the term ‘Mesozoic Art’ doesn’t relate specifically to our inclusion of art showing organisms and environments from the Mesozoic (the section of geological time that extended from 252 to 66 million years ago and includes the Triassic, Jurassic and Cretaceous periods).

**Caption:** here, I’m going to repeat the caption I used back in 2022 when I published an article on *Mesozoic Art*, the first book. Animals of today are animals of the past when imagined or seen in the right way; similarly, the animals of the past can often be transposed to the present if environments and conditions are right. *I know* I’m not the only one who looks at animals and environments in this way. Images: Darren Naish.

Rather, it relates to the idea that many of us use a sort of ‘prehistoric gaze’ when viewing scenes of any age. If I look at a deer in a meadow, am I experiencing a vista that also existed 20 million years ago? What if it was 100 million years ago, in which case the animal sure wasn’t a deer? What was similar, what was different? The term ‘Mesozoic Art’ is our best effort to capture that idea. While it’s true that Mesozoic dinosaurs and their contemporaries dominate the book, we also include portfolios where mammals are very much on show (Beth Zaiken) and where Cenozoic birds are overwhelmingly obvious (Simone Giovanardi).

As you’d expect, the quality of the work we feature is staggering. It would be wrong to describe a work featuring 25 artists as a ‘who’s who’; nevertheless, MAII can be described as representing a good overview of the state and health of palaeoart in 2025.

**Caption:** sample pages from the jaw-dropping Brian Engh section, showing Late Jurassic and Late Cretaceous scenes and an outstandingly complex marine image depicting Late Cretaceous Western Interior Sea life. Images: Brian Engh, from White & Naish (2025).

  **Caption:** sample pages from the section showing Edyta Felcyn-Kowalska’s work. Some incredible stuff here; I especially like that juxtaposing Mesozoic archosaurs belonging to highly disparate lineages. Images: Edyta Felcyn-Kowalska, from White & Naish (2025).

This leads me to another point. It’s standard for palaeoart to involve a style that (in talks) I’ve been terming technical hyper-realism; for most people (palaeontologists among them) palaeoart is ‘meant’ to be like this. As such, it’s generally not always that ‘arty’, functioning instead as scientific reconstruction work that might decorate the pages of a technical academic paper more than a gallery wall.

But a topic very relevant to MAII concerns diversity, both with respect to our increasingly global cadre of contributing artists and the styles and forms of palaeoart we showcase. Volumes devoted to palaeoart – whatever we mean by that term (and that itself is a topic for discussion) – have generally been very ‘European’, focusing very much on English and North American artists. That’s still true to a degree, but it’s also true that there’s a more global feel to the work that’s visible. Palaeoartists of multiple nations worldwide have, of course, been contributing to the canon for decades, but it’s increasingly the case that this work is visible from afar. MAII, we hope, goes some way in terms of highlighting the global nature of the field.

**Caption:** there’s a lot of good palaeoart out there online, much of it highly similar in quality to that we regard as professional. For a time recently, I gathered the work of (mostly) unpublished palaeoartists and shared it via a social media ‘artboost’ project. These screengrabs show just a few of these. I had to give up due to other demands on my time. What’s clear is that there is tons of great work by great people that we have yet to see shared in print.

Yes, we do need cubist dinosaurs. Of similar interest is that we’re moving to a world where art styles very different from technical hyper-realism are increasingly evident. Back in 2012 – when John Conway, Memo Kösemen and I were promoting the then-new All Yesterdays (Conway et al. 2012) – John drew attention to palaeoart-themed commentary from an academic where the statement “after all, there’s no need for a cubist dinosaur” appeared in print. John’s response was: oh yeah, who says?

**Caption:** this article isn’t about *All Yesterdays*, but here are images relevant to the 2012 publication of that book (including a scene from the launch event; from left to right, John Conway, Memo Kösemen, Darren Naish).

Fact is, someone denying potential interest in a cubist dinosaur is promoting disinterest in the idea that extinct dinosaurs might be depicted in artistically novel, non-standard ways. If you’re interested in extinct dinosaurs, or other fossil animals, for visual, aesthetic reasons – as many of us are – why shouldn’t we have cubist dinosaurs? We’re not just interested in extinct animals, or animals of any sort, for dry, academic reasons.

With that in mind, a notion floated at the 2012 publication of All Yesterdays was that we need more artistic experimentation in palaeoart. With MAII, an argument can be made that things are going well on that front, and exhibits A and B are the portfolios of Natalia Jagielska and DJ Washington (White & Naish 2025). Both are creating wonderful pieces of art that tick many boxes in terms of technical accuracy, but many others in terms of artful style. I predict that we’re going to see much more work of this sort, and indeed my impression from the art on show and on sale at DinoCon 2025 – the giant dinosaur-themed meeting I and others recently hosted in Exeter, UK – is that work like this is increasing in popularity.

**Caption:** select images from the Natalia Jagielska and DJ Washington pages included in *MAII*. Our sections on these two artists have proved stand-out favourites among at least some people who’ve looked at the book. Images: Natalia Jagielska, DJ Washington, from White & Naish (2025).

For now, that will do. We hope that people like Mesozoic Art II and buy it. Steve and I (working with various of the contributing artists where possible) are hoping to do various additional promotional events (we’ve done a few already); pay attention to what we say on social media, and watch also for announcements at my website. And huge thanks to everyone who helped this milestone book come together, not least of those being our 25 amazing contributing artists! As we say in the intro… here’s to Mesozoic Art III!

For previous TetZoo articles on other palaeoart books, see

The Great Dinosaur Art Event of 2012, November 2012
All Yesterdays: Unique and Speculative Views of Dinosaurs and Other Prehistoric Animals – the book and the launch event, December 2012
Artistic Depictions of Dinosaurs Have Undergone Two Revolutions, September 2014
Announcing Mesozoic Art, a Lavish New Volume on Modern Palaeoart, October 2022

If you enjoyed this article and want to assist me in what I do — and see behind-the-scenes stuff in development — please support me at patreon. Click anywhere on this text in bold.

Refs - -

Conway, J., Kosemen, C. M. & Naish, D. 2012. All Yesterdays: Unique and Speculative Views of Dinosaurs and Other Prehistoric Animals. Irregular Books.

White, S. & Naish, D. 2022. Mesozoic Art: Dinosaurs and Other Ancient Animals in Art. Bloomsbury Wildlife, London.

White, S. & Naish, D. 2025. Mesozoic Art II: Dinosaurs and Other Ancient Animals in Art. Bloomsbury Wildlife, London.

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If you know anything about animals, you’ll know that hybridization – here meaning the crossing of species – is commonly reported among wild, free-living animals, and appears ubiquitous in some species pairs.

**Caption:** the *Jurassic World* universe has, over time, included more and and more hybrid dinosaurs, some very silly. 'Stegoceratops' – a genetic hybrid between *Stegosaurus* and some sort of ceratopsian (not necessarily *Triceratops*, maybe *Nasutoceratops*) – has been a mainstay of the games and expanded universe for a while. Image: *Jurassic Park* wiki (**here**).

We’ve known this for centuries, mostly because it’s obvious from the anatomy of the hybrid offspring but also because we’ve observed interspecies pairing – and its results – in action. Late 20th century advances in genetics have shown that hybridization is even more common than previously thought, and that whole populations, or sections of populations, have a hybrid ancestry.

We also know, thanks to centuries of keeping animals in captivity, that hybrids of most remarkable sort can occur when conditions allow, certain animal groups possessing behavioural, anatomical and genetic traits that permit an ‘anything goes’ approach to crossing (e.g., Allen & Short (1997) on equids, Arantes et al. (2020), Vilaça et al. (2021) on sea turtles, Káldy et al. (2020) on sturgeon x paddlefish hybrids). In cases, the species concerned are not especially closely related and have been going their separate evolutionary ways for 10, 20, or 30 or more million years.

**Caption:** hybrids that, by now, we all know and love. At left, Russian sturgeon *Acipenser gueldenstaedtii* (a) and American paddlefish *Polyodon spathula* (d), and hybrids (sturddlefish) between the two. These are captive hybrids and couldn’t have come about in the wild. At right, juvenile hybrid sea turtle captured off Florida in 2016. It looks like a Green turtle *Chelonia mydas* but has a few traits characteristic of Loggerhead *Caretta caretta*. These species belong to lineages that separated between 35 and 46 million years ago [UPDATE: see comments. Those claims of very old divergences in extant sea turtles are probably very wrong]. Images: **Káldy *et al*. (2020)**; Shamblin *et al*. (2018).

Because the hybrid animals that have received the greatest amount of discussion are big mammals – like canids, bears, zebras and cetaceans – you might have the impression that hybridization in the wild is a mostly mammals, perhaps even mammals-only, thing. It totally isn’t. We know of cases in which related bird species hybridize in the wild, examples including ducks, grouse, gulls, hummingbirds and corvids. Among non-bird reptiles, wild hybrids are known among sea turtles (Arantes et al. 2020, Vilaça et al. 2021), Galapagos tortoises (Miller et al. 2017), crocodiles (Milián-García et al. 2011, 2015) and numerous lizards (Rassman et al. 1997, Reeder et al. 2002, Jančúchová-Lásková et al. 2015). In amphibians, certain western Palaearctic water frogs (Spolsky & Uzzell 1986) and ambystomatid salamanders famously hybridize (Spolsky et al. 1992). Wild-living fish hybrids are well known among salmon and carp.

**Captions:** if you look at gulls a lot (the white-headed *Larus* species especially), you’ll know that hybrids are moderately common in some populations, and often difficult to identify precisely. At left, a gull photographed in Lisbon in 2014 and which seems to combine traits of Lesser black-backed *L. fuscus* and Herring gulls *L. argentatus*. At right, an Olympic or Puget Sound gull photographed in Oregon in 2022. This sort of gull appears to be a hybrid between Western *L. occidentalis* and Glaucous-winged gulls *L. glaucescens*. Images: Darren Naish.

Why animals hybridize. Why animals hybridize is a complex question with several answers. Certain animals recognize the vocal, acoustic and visual signals of close relatives as ‘similar enough’ to those of their own species that they function as appropriate stimuli when the time is right. In those cases, hybridization might be described as opportunistic.

Incidentally, an idea popular in the dinosaur literature – that the extravagant horns, frills, crests and so on of dinosaurs helped the species recognise their own and avoid mating with the members of other species – is surely mostly erroneous (Hone & Naish 2013). They don’t function that way in living species. It was more likely a combination of factors, involving pigmentation, behaviour and olfactory and vocal signals, that ‘controlled’ courtship and mating events.

**Caption:** there are numerous dinosaur communities where some or several close relatives lived alongside one another; the animals shown here weren’t all sympatric but are representatives of certain of the relevant groups (lambeosaurine hadrosaurs and both chasmosaurine and centrosaurine ceratopsids). These animals often exhibit extravagant structures but – even so – we might speculate that hybridization occurred here and there. This illustration is looking very dated. Image: Darren Naish.

I digress. In some cases, it’s hypothesized that hybridization occurs because individuals fail to find mates of their own species. From an evolutionary perspective, a ‘bad mating’ is better than no mating at all: there’s an advantage in using opportunities to pass on genes, species boundaries be damned. Species that are rare (due to human hunting, habitat loss or deterioration and so on) may therefore have little option other than to hybridize. At least some of the hybridization we see in the modern world might, then, not have occurred – or, at least, not have occurred at regular frequency – in the geological past. Having said that, a world devoid of human pressures could still result in situations where species were forced to hybridize with related species, and we would do well to keep this in mind when looking at fossils from ecosystems thought to have been under environmental stress.

A second possible impact of humans on hybridization might run the other way: that is, that the declines we’ve caused to species and populations might reduce the opportunity for hybridization, since we’ve lowered the number of cases where species are sympatric and thereby lowered the opportunities for hybridization. The fossil record shows, in general, that sympatry between related species was more common in the past than it is today. Indeed, many ancient ecosystems look packed in terms of related species relative to modern ones. Don’t forget that the world today is depauperate when it comes to megafaunal diversity and distribution.

**Caption:** it’s well known that *Panthera* species can hybridize, with ligers (*Panthera leo* x *P. tigris*) being probably the most familiar. Ligers have been known to science since the late 1700s and can be (relatively) easily created in captivity. Notably, the two parent species here have, in the past, occurred sympatrically and there are unconfirmed accounts of their hybrids existing in the wild. Image: Camphora, public domain (**original here**).

Are hybrids really ‘common’? Not in birds. Hybridization in some animal groups is, statistically speaking, rare and statements implying that it’s common or ubiquitous need to be seen in context. Yes, hybrids between species x and species y might often get reported (indeed, hybrids are deemed so interesting that they’re almost certainly over-reported), but they’re rare in terms of frequency.

Ernst Mayr, one of the most influential and famous figures in our understanding of bird evolution, suggested on the basis of museum specimens that hybrids occurred on the order of 1 in 60,000, meaning that about 0.001% of individuals were hybrids (Mayr 1963, p. 114). More recently, Justyn et al. (2020) used eBird data for the United States collected between 2010 and 2018 and found that hybrids accounted for 0.064% of records. That’s a higher percentage than Mayr’s, this perhaps explained by differing views on species boundaries (the gulls and ducks that modern workers regard as distinct species were not treated as such by Mayr). When very ‘hybrid-heavy’ species were excluded (like Mallard Anas platyrhynchos and Mexican duck A. diazi), hybridization frequency was 0.009% (Justyn et al. 2020).

**Caption:** two ducks that engage in a lot of hybridizing. Mexican duck at left, Mallard at right. Images: **ALAN SCHMIERER**, CC0 (**original here**); Darren Naish.

In addition, nearly 86% of all bird hybrids in the dataset compiled by Justyn et al. (2020) involved just ten species pairs, all of which were ducks, geese, gulls and chickadees. In fact, nearly 83% of all US hybridization events involved ducks and gulls alone. Huge swathes of the avian tree of life (at least, based on US data) have no meaningful hybridization going on at all (Justyn et al. 2020).

Crocodylians, turtles and squamates. The take-home from this perspective is that hybridization is rare overall, is aberrant and remarkable across big swathes of the avian tree and is only ‘expected’ or ‘usual’ if you’re dealing with a very particular, very small set of unusual species. It’s not clear how applicable this observation is when it comes to other animal groups though, and it should be obvious from the title of this article that I’m especially interested here in possible hybridization among non-bird dinosaurs. In view of that, the animals most relevant after birds are crocodylians, turtles and squamates. How common, or rare, is hybridization in those animals?

For crocodylians, cases of hybridization among wild animals have been well recorded between certain species pairs, namely American crocodiles Crocodylus acutus and Morelet’s crocodile C. moreletii in the Mexican Caribbean (e.g., Machkour-M’Rabet et al. 2009), and Cuban crocodiles C. rhombifer and a population historically (but probably incorrectly) identified as American crocodiles in the Zapata Swamp of Cuba (e.g., Milián-García et al. 2011, 2015). In the Zapata Swamp case, as many as 49% of the wild-living individuals had a hybrid ancestry (Milián-García et al. 2015) and a repeated claim made in the relevant publications is that hybridization among crocodile species is common.

**Caption:** Cuban crocodile, a species that has hybridized in the wild with the related crocodile species. Whole populations in parts of Cuba appear to have a hybrid ancestry *but* human intervention may have a role in contributing to this. The Cuban crocodile is a fantastic animal, highly capable on land, good at grabbing animals from overhead branches (by leaping from water), and with great pigmentation. Image: Zanbog, CC BY-SA 2.0 (**original here**).

However… we’re referring here to declining, relictual populations of species whose ranges have been substantially reduced relative to the historical norm. Indeed, it’s been suggested that this hybridization might have been “enhanced by anthropogenic pressures”, to quote Cuban crocodile expert Yoamel Milián-García. Is hybridization in crocodiles really ‘common’? I would challenge that contention.

Among lizards, hybridization in the wild is known among around 70 species, including assorted iguanas (Rassman et al. 1997, Vuillaume et al. 2015, Moss et al. 2018), fence lizards, anoles and other iguanians (Jančúchová-Lásková et al. 2015), whiptails (Reeder et al. 2002), certain snakes, lacertids and several gecko groups (Jančúchová-Lásková et al. 2015). Some whiptail species appear to owe their origins to hybridization.

**Caption:** very useful phylogenetic tree from Jančúchová-Lásková *et al*. (2015) showing prevalence of hybridization within lizards. It's widespread enough that we should regard it as ubiquitous across the group, but note that there are numerous groups where it basically hasn't been reported.

This indicates that an ability to hybridize is widespread among lizards, but “[r]eliable records of hybridization are scarce” (Jančúchová-Lásková et al. 2015, p. 169) and, again, the numbers of species we’re dealing with are low relative to the number of lizards there are in total. In addition, at least some examples (those involving Iguana species) are due to human introduction. Yes, hybridization occurs between some species pairs, but it shouldn’t be regarded as widespread across the animals as a whole, nor ‘normal’ or ‘ubiquitous’.

Very similar things could be said about turtles (which now appear to be on the archosaur lineage and thus closer to dinosaurs than lizards). Assorted rare hybridizations have been reported from the wild (Miller et al. 2017, Arantes et al. 2020) and at least some of these, as well as some from captivity (Buskirk et al. 2005), are consistent with the ‘anything goes’ tendency noted earlier. Again, some and maybe even most or all of the recorded wild hybridizations are linked to human-caused factors.

In quest of hybrid (non-bird) dinosaurs. Here we come to the raison d’âtre of this article. If hybridization occurs among animals of so many sorts today, we absolutely can and should assume that it occurred among the animals of the past. We could discuss this issue as it pertains to fossil hominins, elephants, crocodylians or frogs, but we’re here because of the non-bird dinosaurs.

**Caption:** the very famous *Triceratops* specimen AMNH 5116, on show at the American Museum of Natural History in New York. It's a really famous *Triceratops* but also a deeply odd one. Image: Kabacchi, CC BY 2.0 (original **here**).

My primary contention (or assumption) is that hybrid non-bird dinosaurs did exist as live animals. Based, however, on the points we’ve already seen, we should be working on the assumption that hybridization was rare, if not extremely so. It remains possible that there were certain populations or species – like those duck and gull species noted above – where hybridization was frequent.

While we don’t have a firm idea of which species pairs might have been ‘gull- or duck-like’ in their propensity to hybridize, an argument can be made that we can predict which species pairs were most likely to do so. Obviously, we might start with closely related, anatomically similar, sympatric animals. Late Cretaceous lambeosaurines and chasmosaurines come to mind, as do Morrison Formation diplodocids, Wessex Formation iguanodontians, and the Late Cretaceous east Asian ceratopsians grouped together as protoceratopsids or bagaceratopids (yes, different group name endings there), for starters.

**Caption:** at left, the ceratopsian specimen MPC-D 100/551B from Ömnögovi, Mongolia, identified by Czepiński (2020) as ‘cf *Bagaceratops* sp’ and suggested to perhaps be a hybrid. At right: opinions differ among experts as to how many taxa should be recognized among Late Cretaceous east Asian ceratopsians (*Magnirostris*, shown here, is generally thought today to be synonymous with *Bagaceratops*). If any of these animals were sympatric, it is at least plausible that hybridization occurred. Images: Czepiński (2020); Darren Naish.

A suggestion already exists that a Bagaceratops-like ceratopsian from the Gobi Desert might be a hybrid between Protoceratops andrewsi and B. rozhdestvenskyi (Czepiński 2020) but alternative explanations for its anatomy – that it represents a transitional evolutionary form between the two or is simply a new species – is on the cards too. Various Triceratops specimens look anatomically ‘intermediate’ between the older T. horridus and the younger, shorter-snouted T. prorsus (Scannella et al. 2014). The favoured explanation is that these are, again, evolutionary intermediates, consistent with the view that the younger species evolved directly from the older one, and likewise for other ceratopsids that look like ‘intermediates’ (Fowler & Freedman Fowler 2020). Comments have occasionally been made about the Triceratops on show in New York (AMNH 5116), since (while heavily reconstructed in the frill especially) it looks something like an intermediate between Triceratops and Torosaurus, this at least raising the possibility that it might be a hybrid between the two.

**Caption:** several *Triceratops* specimens (like UCMP 113697 and MOR 3027, shown here in the middle part of the tree) were found by **Scannella *et al*. (2014)** to be 'intermediates' between the two recognized species. Within their hypothesis of anagenetic evolution, such animals must have existed. But the possibility that some of these specimens might be hybrids is on the cards. Image: **Scannella *et al*. (2014)**.

  **Caption:** the famous Yoshi’s Trike, MOR 3027, at left, on show at the Museum of the Rockies, Montana. This animal is anatomically intermediate between the *Triceratops* species *T. prorsus* and *T. horridus*... could it be a hybrid? That’s a fun idea, since its ridiculous supraorbital horns could then be imagined (I said *imagined*) as a consequence of hybrid vigour. Image: WernerG2011, CC BY-SA 2.0 (**original here**).

Another point: even if we do find two species that might have engaged in hybridization, we have no way of knowing how typical they were of their broader groups. Stated another way, just because two species in a group of 50 regularly hybridize, it doesn’t follow that regular hybridization should be assumed for the remaining 48 within the group.

Can we test for hybridization in fossil dinosaurs? Even if hybridization did occur, regularly or not, the big question is… will we ever be able to do more than speculate about it? Will we ever be able to demonstrate it? We don’t exactly have large datasets for the majority of fossil dinosaur species. Most are known from singletons.

In living animals, we can spot hybrids because they combine physical traits belonging to both parent species. And if we measure the physical traits of hybrid individuals, we often find that they’re anatomically intermediate between their parent species. Could we determine, via morphometrics, that a given fossil dinosaur specimen is a hybrid? A sample size of between 20 and 30 individuals is considered exceptionally good for fossil dinosaurs; we’re virtually always well short of the few hundred you need to be confident about the distribution of shape and size within a population. If we do find a specimen that’s unusual in morphometric terms (that is, it’s intermediate between individuals of species x and species y, or has proportions that make it partly overlap either or both x and y), our default and most conservative hypothesis is that we’ve found a new species.

**Caption:** Rackelhuhn, a naturally occurring and well known European hybrid between Western capercaillie and Black grouse. Image: F. C. Robiller, CC BY-SA 3.0 (**original here**).

One group of dinosaurs is represented by numerically sufficient samples to test for hybrids, and here I’m referring to Pleistocene and Holocene birds. Based on the prevalence of hybrids between three grouse species pairs – the Western capercaillie Tetrao urogallus and Black grouse T. tetrix, Western capercaillie and Willow ptarmigan Lagopus lagopus and Black grouse and Willow ptarmigan – Bocheński & Tomek (2000) looked at the bones of taxiderm hybrid specimens to see if their hybrid nature could be detected via osteology alone. They found that some animals could be identified as hybrids based on size and anatomical detail but that not all could, and they concluded that “it is probably only a matter of time to prove the presence of galliform hybrids in fossil and archaeological materials” (Bocheński & Tomek 2000, p. 698). This, and cases like it, are not exactly applicable to extinct species, however, since this is a case where we know that hybridization occurs, and have then gone looking for it in fossils. If we only had fossil grouse to go on, would we know that specimens with ‘intermediate’ measurements and details represent hybrids? That’s impossible to answer, but the answer is probably no.

**Caption:** Bocheński & Tomek (2000) showed that hybrids between extant galliform species could be identified osteologically, if you know exactly what to look for. As shown in these and other diagrams, known hybrids possess small osteological details in the sternum, coracoid, humerus and elsewhere in the skeleton that are intermediate between the conditions of the two parent species. Images: Bocheński & Tomek (2000).

Despite all of these caveats, excuses and handwringing: yes, there are specimens that look like they might be hybrids. The bad news is that we have no real way of testing, let alone confirming, a potential hybrid origin. In addition, the statistical rarity of hybridization relative to other potential explanations for an unusual anatomical configuration – in particular the possibility that the fossil concerned represents a new taxon of conventional, non-hybrid origin, or an evolutionary intermediate – renders any suggestions of a hybrid origin speculative in the extreme.

**Caption:** palaeoartists have indulged in a modicum of speculation on the sorts of hybrids that might have existed in some Mesozoic dinosaur communities. This hypothetical ‘Toroceratops’ represents a *Triceratops* x *Torosaurus* hybrid. Image: **FALC Paleoart**, used with permission.

‘Impossible’ hybrids that are not impossible. I want to end this article by playing with even more speculation, because where else can you do this if not on a blog. All the ideas about the possible presence of hybridization discussed above concern closely related taxa. But more exciting, and even more speculative, is the idea – again, inspired by hybrids in the modern world – that species only distantly related might hybridize too.

It’s highly unlikely that species from far-flung branches on the dinosaur family tree might have got it on with one another (no, for example, thyreophoran x ceratopsian crosses), but… based on hypothesized hybridization events that seem to exist across lineages within deer (Pitra et al. 2004), turtles (Buskirk et al. 2005, Arantes et al. 2020) and such birds as woodpeckers (Fuchs et al. 2013), it’s possible that extinct dinosaurs that were part of the same ‘family-level’ clade, but were not necessarily all that close, could still produce viable crosses. So, imagine a cross between a Parasaurolophus-like lambeosaurine and a Corythosaurus-like one, or between a Chasmosaurus-like chasmosaurine and something like Anchiceratops or Regaliceratops.

**Caption:** anatomy and genetics indicate that several extant animal species or groups probably do owe their origins to hybridization between species that are not especially closely related. As a consequence they do look weird and have often been hard to place in phylogenetic terms. Examples include the Milu or Père david’s deer *Cervus davidianus* (captive individual at left), and the campephiline woodpeckers (Crimson-crested woodpecker *Campephilus melanoleucas* shown here). I wrote about the ancestry of the Milu **here**. Images: Darren Naish; Bernard Dupont, CC BY-SA 2.0 (**original here**).

Even more extreme crosses – occurring between lineages distinct for over 20 or 30 million years – are also at least plausible based on what some living animals have done. Of course, we can’t say anything about the real probability of this given our lack of knowledge on their genetics. But it does mean that (arguably) ridiculous things are conceivable, like hybrids between centrosaurine and chasmosaurine horned dinosaurs, between apatosaurine and diplodocine sauropods, between microraptorines and velociraptorines within maniraptorans, or, shock horror, between baryonychines and spinosaurines within megalosauroids.

**Caption:** another hypothetical Late Cretaceous hybrid, this time one between members of the *Parasaurolophus* and *Lambeosaurus* lineages within lambeosaurines. Would such an animal succeed in winning mates itself? Would it be a loner, would it live with one or either parent species, and would its unique anatomy be reflected in unique acoustics? Questions we will likely never have answered… should such an animal have existed. Image: Hodari Nundu, used with permission.

To sum up this article in the most concise way possible… Could hybrid (non-bird) dinosaurs exists: yes, of course, they surely did, but they were likely vanishingly rare, so rare that they’re unlikely to be represented in the current palaeontological sample. If they are represented, it will be among species known from a good number of individuals. Can we ever identify hybrids, even among such species? Based on our current knowledge and analytical techniques, almost certainly not, but we should remain open-minded to the possibility that they might be both findable and identifiable.

For previous articles on Mesozoic dinosaur biology relevant to the topics covered here, see…

The Sensitive Face of a Big Predatory Dinosaur, June 2017
Could We Domesticate (Non-Bird) Dinosaurs?, August 2018
Postcranial Palaeoneurology and the Lifestyles of Pterosaurs, August 2018
Alternative Timeline Dinosaurs, the View From 2019 (Part 3): the Dinosauroid and its Chums, December 2019
Did Dinosaurs and Pterosaurs 'Glow'? Extinct Archosaurs and the Capacity for Photoluminescent Visual Displays, March 2020
Pete Dunne and Kevin T. Karlson’s Gulls Simplified: A Comparative Approach to Identification, October 2021
A brain for Baryonyx: using CT-scanning to examine British spinosaurid brains, February 2023
T. rex the Tool Maker -- Testing Controversial Claims About Dinosaur Smarts, April 2024

Refs - -

Allen, W. R. & Short, R. V. 1997. Interspecific and extraspecific pregnancies in equids: anything goes. The Journal of Heredity 88, 384-392.

Arantes, L. S., Ferreira, L. C. L., Driller, M., Filho, F. P. M. R., Mazzoni, C. J. & Santos, F. R. 2020. Genomic evidence of recent hybridization between sea turtles at Abrolhos Archipelago and its association to low reproductive output. Scientific Reports 10, 12847.

Bocheński, Z. & Tomek, T. 2000. Identification of bones of galliform hybrids. Journal of Archaeological Science 27, 691-698.

Buskirk, J. R., Parham, J. F. & Feldman, C. R. 2005. On the hybridisation between two distantly related Asian turtles (Testudines: Sacalia × Mauremys). Salamandra 41, 21-26.

Czepiński, Ł. 2020. New protoceratopsid specimens improve the age correlation of the Upper Cretaceous Gobi Desert strata. Acta Palaeontologica Polonica 65, 481-497.

Fowler, D. W. & Freedman Fowler, E. A. 2020. Transitional evolutionary forms in chasmosaurine ceratopsid dinosaurs: evidence from the Campanian of New Mexico. PeerJ 8: e9251.

Fuchs, J., Pons, J.-M., Liu, L., Ericson, P. G. P., Couloux, A. & Pasquet, E. 2013. A multi-locus phylogeny suggests an ancient hybridization event between Campephilus and melanerpine woodpeckers (Aves: Picidae). Molecular Phylogenetics and Evolution 67, 578-588.

Hone, D. W. E. & Naish, D. 2013. The ‘species recognition hypothesis’ does not explain the presence and evolution of exaggerated structures in non-avialan dinosaurs. Journal of Zoology 290, 172-180.

Káldy, J., Mozsár, A., Fazekas, G., Farkas, M., Fazekas, D. L., Fazekas, G. L., Goda, K., Gyöngy, Z., Kovács, B., Semmens, K., Bercsényi, M., Molnár, M. & Patakiné Várkonyi, E. 2020. Hybridization of Russian sturgeon (Huso gueldenstaedtii, Brandt and Ratzeberg, 1833) and American paddlefish (Polyodon spathula, Walbaum 1792) and evaluation of their progeny. Genes 11 (7), 753.

Jančúchová-Lásková, J., Landova, E. & Frynta, D. 2015. Are genetically distinct lizard species able to hybridize? A review. Current Zoology 61, 155-180.

Justyn, N. M., Callaghan, C. T. & Hill, G. E. 2020. Birds rarely hybridize: a citizen science approach to estimating rates of hybridization in the wild. Evolution 74, 1216-1223.

Machkour-M’Rabet, S., Henaut, Y., Charruau, P., Gevrey, M., Winterton, P. & Legal, L. 2009. Between introgression events and fragmentation, islands are the last refuge for the American crocodile in Caribbean Mexico. Marine Biology156, 1321-1333.

Mayr, E. 1963. Animal Species and Evolution. Harvard University Press, Cambridge, MA.

Milián-García, Y., Ramos-Targarona, R., Pérez-Fleitas, E., Sosa-Rodríguez, G., Guerra-Manchena, L., Alonson-Tabet, M., Espinosa-López, G. & Russello, M. A. 2015. Genetic evidence of hybridization between the critically endangered Cuban crocodile and the American crocodile: implications for population history and in situ/ex situ conservation. Heredity 114, 272-280.

Milián-García, Y., Venegas-Anaya, M., Frías-Soler, R., Crawford, A. J., Ramos-Targarona, R., Rodríguez-Soberón, R., Alonso-Tabet, T. J., Sanjur, O, Espinosa-López, G. & Beringham, E. 2011. Evolutionary history of Cuban Crocodiles Crocodylus rhombifer and Crocodylus acutus inferred from multilocus markers. Journal of Experimental Zoology 315, 358-375.

Miller, J. M., Quinzin, M. C., Poulakakis, N., Gibbs, J. P., Beheregarey, L. B., Garrick, R. C., Russello, M. A., Ciofi, C., Edwards, D. L., Hunter, E. A., Tapia, W., Rueda, D., Carrión, J., Valdivieso, A. A. & Caccone, A. 2017. Identification of genetically important individuals of the rediscovered Floreana Galápagos giant tortoise (Chelonoidis elephantopus) provides founders for species restoration program. Scientific Reports 7, 11471.

Moss, J. B., Welch, M. E., Burton, F. J., Vallee, M. V., Houlcroft, E. W., Laaser, T. & Gerber, G. P. 2018. First evidence for crossbreeding between invasive Iguana iguana and the native rock iguana (Genus Cyclura) on Little Cayman Island. Biological Invasions 20, 817-823.

Pitra, C., Fickel, J., Meijaard, E. & Groves, P. C. 2004. Evolution and phylogeny of old world deer. Molecular Phylogenetics and Evolution 33, 880-895.

Rassman, K., Trillmich, F. & Tautz, D. 1997. Hybridisation between the Galapagos land and marine iguana (Conolophus subcristatus and Amblyrhynchus cristatus) on Plaza Sur. Journal of Zoology 242, 729-739.

Reeder, T. W., Cole, C. J. & Dessauer, H. C. 2002. Phylogenetic relationships of whiptail lizards of the genus Cnemidophorus (Squamata: Teiidae): a test of monophyly, reevaluation of karyotypic evolution, and review of hybrid origins. American Museum Novitates 3365, 1-61.

Scannella, J. B., Fowler, D. W., Goodwin, M. B. & Horner, J. R. 2014. Evolutionary trends in Triceratops from the Hell Creek Formation, Montana. Proceedings of the National Academy of Sciences 111, 10245-10250.

Shamblin, B. M., Mansfield, K. L., Seney, E. E., Long, C. A., Bagley, D. A. & Nairn, C. J. 2018. Brazilian origin of a neritic juvenile hybrid loggerhead x green turtle foraging in Florida. Marine Turtle Newsletter 155, 4-7.

Spolsky, C., Phillips, C. A. & Uzzell, T. 1992. Gynogenetic reproduction in hybrid mole salamanders (genus Ambystoma). Evolution 46, 1935-1944.

Spolsky, C. & Uzzell, T. 1986. Evolutionary history of the hybridogenetic hybrid frog Rana esculenta as deduced from mtNDA analyses. Molecular Biology and Evolution 3, 44-56.

Vilaça, S. T., Piccinno, R., Rota-Stabelli, O., Gabrielli, M., Benazzo, A., Matschiner, M., Soares, L. S., Bolten, A. B., Bjorndal, K. A. & Bertorelle, G. 2021. Divergence and hybridization in sea turtles: inferences from genome data show evidence of ancient gene flow between species. Molecular Ecology 30, 6178-6192.

Vuillaume, B., Valette, V., Lepais, O., Grandjean, F. & Breuil, M. 2015. Genetic evidence of hybridization between the endangered native species Iguana delicatissima and the invasive Iguana iguana (Reptilia, Iguanidae) in the Lesser Antilles: management implications. PLoS ONE 10 (6), e0127575.

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In which I once more rescue an article from the broken Tet Zoo archives, this time from ver 3 at Sci Am, and specifically from November 2011…

**Caption:** at left, cover of the 1979 *Sun, Sand & Snakes*, a valuable source of info on the Crowing crested cobra and African snakes in general. At right, reconstruction of Crowing crested cobra by Karl Shuker, used with permission.

Yes, here’s an article that appeared here in the past: the original is here at the internet archive, but I can only find ruined versions lacking images. I’m republishing this article for two reasons, one of which will become clear very soon, and one of which is due to my continuing efforts to republish squamate-themed articles here at ver 4. To business…

I’ve recently been reading Stephen Spawls’s Sun, Sand & Snakes, a 1979 volume that charts Spawls’s childhood interest in snakes and other reptiles and recounts his numerous japes and scrapes with local, east African herpetofauna. Today, Spawls is a well-known herpetologist, co-author of the excellent The Dangerous Snakes of Africa (Spawls & Branch 1995) and A Field Guide to the Reptiles of East Africa (Spawls et al. 2002). Sun, Sand & Snakes contains many interesting anecdotes about east African snakes but it’s his brief section on the ‘feathered serpent’ that inspired me to write this article.

**Caption:** I’m a big fan of the herpetological books of Stephen Spawls and his co-authors and own these two (plus the *Sun, Sand & Snakes* one shown above).

Well known to both those who know snakes and those who know the literature on animal myths and anecdotes, the ‘feathered serpent’ is more often known as the Crowing crested cobra. “In some versions the snake has the head of a chicken, complete with combs and wattles, in others it has merely a crest of feathers. This fabulous serpent is, of course, highly poisonous [sic]. In most cases it is believed to have the ability to kill its human victims merely by looking at them. It lives on human flesh and can be detected in the areas where it lives by its offensive smell and the strange and frightening noises it makes at night” (Spawls 1979, p. 95).

Stories and sightings that supposedly pertain to the Crowing crested cobra come from South Africa, Malawi, Mozambique, Zimbabwe, Zambia and Tanzania, and apparently as far north as the Central African Republic. I’m somewhat sceptical of the idea that the various mystery snake-like entities written about across this enormous area really do pertain to the exact same kind of animal, but it’s said that the many named used for the Crowing crested cobra across this range include bubu, inkhomi, hongo, songo and mbobo (Shuker 1991).

**Caption:** at left, the original (non-coloured) illustration of the Crowing crested cobra from Karl Shuker’s 1991 *Extraordinary Animals Worldwide*, also published in the revised 2007 volume *Extraordinary Animals Revisited*, shown at right. These books are of direct interest to me for several reasons, one being that they feature Karl’s thoughts on *Ameranthropoides*… as hinted by the cover of the 2007 work. Image: Karl Shuker, used with permission.

The literature on this semi-mythical creature is fairly widely scattered in the arcane literature. However, most of it was gathered together by Bernard Heuvelmans for his 1978 Les Dernier Dragons d’Afrique and again by Karl Shuker for his 1991 Extraordinary Animals Worldwide (and his 2007 Extraordinary Animals Revisited). Karl’s chapter also includes what seems to be the only decent, published reconstruction of the Crowing crested cobra’s purported appearance. That picture (penned by Karl himself) is reproduced here (with permission: © Karl Shuker). UPDATE: for my project on The Cryptids of Bernard Heuvelmans (see progress at my patreon), I’ve illustrated a Crowing crested cobra myself.

**Caption:** reconstruction of Crowing crested cobra, based very obviously on Karl’s version, and created for my in-prep *The Cryptids of Bernard Heuvelmans* (**progress on which is shared at patreon**). Remember that this snake is meant to be very large, around 6 m long. Image: Darren Naish.

Clearly, the Crowing crested cobra is fairly ridiculous in appearance. Overall, it’s said to be brownish or greyish, but it’s scarlet about the face and in possession of a forward-projecting, serrated cockscomb. Additionally, males supposedly possess paired, chicken-like wattles on either side of the face.

A list of superlatives and super powers. All of this would be radical enough, but the Crowing crested cobra is supposedly in possession of a number of additional remarkable traits. It’s apparently very large – something like six metres long – and males supposedly make a loud crowing noise, like a rooster, while females make a chicken-like clucking. So, it’s a crowing, crested snake, which kind of explains the name. It’s said to be arboreal and to strike downwards at the heads of unsuspecting people who pass beneath. And it’s said in some stories that, on killing an animal (by either striking and injecting it with venom, or by spraying venom in spitting cobra fashion*), it doesn’t eat it but waits for flies to lay their eggs and then feeds on the maggots. Other stories say that it kills prey by merely looking at them. I should add, however, that there are also more plausible accounts of it preying on hyraxes.

Real spitting cobras only ‘spit’ (= spray) in self-defence; they do not and cannot kill prey by spraying venom.

I’m reminded of the fact that a fair number of other mystery animals are similarly imbued with – not one – but a whole string of incredible superlative traits. The Brazilian Mapinguari, for example, suggested by some investigators to be a surviving ground sloth, isn’t just a sloth-like mammal with big claws… it also roars like a jet engine, secretes an noxious gas through a mouth-like opening in its belly, and is impervious to bullets. Some of the original stories about the Puerto Rican version of the Chupacabra said how it had wings, could change colour, had kangaroo-like hindlimbs, spikes on its back, a giant fang projecting from its mouth and glowing red eyes, and made the air vibrate with a low-pitched throbbing. It also had psychic powers, liked astrology, was deeply religious, and possessed strong right-wing views about immigration and social welfare.

**Caption:** the chupacabra/chupacabras has a complex history, and what it’s imagined to be like has changed substantially over the years. Prior to it being thought of as a hairless, dog-like animal, it was supposed to be a spiky-backed vampiric humanoid, possibly of alien origin, as shown in the image at right. The illustration at left, by famed palaeoartist John Sibbick, was done for the cover of *Fortean Times* and reflects an effort by the artist to ‘rationalize’ the entity (rather than show it in the manner that would have been more accurate according to canon). Images: © John Sibbick / *Fortean Times*; LeCire (**original here**), public domain.

As is sometimes the case with reports of incredible mystery animals, there are a few accounts that refer to the discovery of Crowing crested cobra specimens. Shuker (1991) discussed the 1944 review compiled by J. O. Shircore in which Shircore (1944) described what was supposedly the partial skeleton of a Crowing crested cobra’s cockscomb. The specimen mostly consisted of a lanceolate plate of bone, marked on its sides with presumed muscle attachment scars, and connected on its upper and lower regions to a section of reddish skin and another fragment of dark, wrinkled head skin. Unfortunately it isn’t possible to make much sense or use of Shircore’s description and it isn’t clear to me how a ‘lanceolate plate’ of bone can be linked with the supposedly chicken-like comb of a Crowing crested cobra. It would be interesting to know what happened to this specimen, or if it’s illustrated anywhere. Shircore also claimed to be in possession of a few other Crowing crested cobra remains (Shircore 1944, Shuker 1991), but these were just vertebrae and skin fragments and I can’t see how he could determine that they were from a possibly novel species of snake.

There’s also a case from 1959 where John Knott was driving through the Kariba area of Zimbabwe (then Southern Rhodesia) when he ran over and mortally wounded a large, jet-black snake about 1.8 m long. It possessed a distinct, symmetrical head crest that possessed five internal prop-like struts. Seemingly, these allowed the crest to be raised at will (Shuker 1991). Shuker noted how the ‘struts’ sounded reminiscent of the rods that help the Frill-necked lizard Chlamydosaurus kingii erect its neckfrill, but we don’t know what became of this snake, nor did Knott illustrate or photograph it.

**Caption:** vintage 1827 illustration of deceased Frill-necked lizard, published in Phillip Parker King’s *Narrative of a Survey* Volume 2, the first outing of this species in the scientific literature and the place where it was officially named and described. The illustration (an engraving) is by Mr Curtis and was based on a drawing by H. C. Field… I assume the ‘Mr Curtis’ is John Curtis (1791-1862). Image in public domain.

Reports like Shircore’s and Knott’s sound somewhat plausible since they seem to make an anecdotal animal more real by imparting it with apparently genuine remains, but the reports are unfortunately also anecdotal and thus not all that useful.

Needless to say, the existence of the Crowing crested cobra is not accepted by mainstream science, nor is there any good evidence that might support its existence. Nor does it seem likely that a very large, arboreal, elaborately crested, highly vocal snake (inhabiting an area that – comparatively speaking – is well trodden by herpetologists) might really exist. It would be great to be wrong about this, of course.

Possible origins. So – how might belief in this remarkable serpent have arisen? Several ideas have already been suggested and I don’t have any new ones. One possibility is that people have seen ‘crested’ and/or gaudily coloured snakes, and combined them with other stories of spectacularly dangerous, frighteningly big snakes.

Things sometimes go wrong when snakes shed their skin: sections of skin can get stuck and fail to slough off completely, in cases creating peculiar ‘ruffs’, ‘crests’ or ‘frills’ about the head or neck. Apparently a few Black mambas Dendroaspis polylepis – originally identified as ‘crested snakes’ – have been captured and examined in which incompletely sloughed skin was at first mistakenly interpreted as a crest (Spawls 1979, Shuker 1991). Some authors have even said that raised patches of old skin are common on the heads of Black mambas, though I’m not sure that this is true.

**Caption:** it might be that part of the Crowing crested cobra legend is based on encounters with Black mambas, perhaps individuals with a ‘frill’ form of sloughed skin. The Black mamba is so named because its mouth interior is black. Image: TimVickers, public domain (**original here**); Tad Arensmeier, CC BY-SA 3.0 (**original here**).

Some of the other traits ascribed to the Crowing crested cobra are common motifs in snake stories and legends. Big, scary snake-like animals are discussed in cultures worldwide – I’m thinking of the Naga of southeast Asia, the Gambian Ninki-Nanka and the various stories of a giant snake (sometimes called the Taguerga) from Algeria – and what’s interesting is that they’re often described as being crested, or having wattles or dewlaps or other cranial adornments. Tales of a Caribbean mystery snake remarkably similar to the Crowing crested cobra were reported by Philip Gosse in his 1867 The Romance of Natural History (and, incidentally, the accounts did not all come from people of recent African ancestry), and tales of a Chinese ‘rooster-crested’ snake that crowed like a chicken have also been recalled by some (Shuker 2007). Comparisons with the European myths of the Cockatrice and Basilisk are of course irresistible.

**Caption:** wooden sculpt of a Naga at Keraton Yogyakarta, Java. Note the crown and lappet-like structure on the throat. Image: © CEphoto, Uwe Aranas (**original here**), CC BY-SA 3.0.

Noise-making snakes. The idea that the Crowing crested cobra might kill animals in order to later eat the maggots on the carcass is also seen elsewhere in snake folklore. As for the idea that the Crowing crested cobra might be highly vocal and capable of making bird-like noises: again, the ability to make bird-like calls is a peculiar but oft-recounted folkloric ability of certain snakes.

Shuker (1991, 2007) recounted a long series of anecdotes in which people claimed to hear snakes of diverse species make goat-like bleats, duck-like quacks, cat-like purrs, bell-like noises, shrill calls and bird-like notes. The mainstream explanation is that these were mistakes and that people had heard other noises and wrongly associated them with the snakes. Alternatively, some people have suggested that the noises were actually made by prey animals that the snakes were swallowing (not theoretically impossible in some cases), while others have proposed that the noises were mechanical – we know that some snakes and other reptiles can stridulate by rubbing their body or tail scales together.

For the most part, I think that the majority of ‘snake calls’ were indeed the result of confusion on the part of the witnesses, but the possibility remains that just a few weird snake calls really do reflect otherwise unappreciated vocal prowess. We now know, after all, that Bornean cave racers Orthriophis taeniurus can make miaowing-like noises. But, whatever, the alleged chicken-like clucking and crowing of the Crowing crested cobra just seems to be a mythical add-on to an already mythical creature. Known species like gaboon vipers (Bitis gabonica and B. rhinoceros) are claimed in some regions to be “crested snakes which crow”, again showing that bird-like crests and bird-like calls are traits sometimes superimposed onto snakes in myth and anecdote.

**Caption:** a speculative effort to reconstruct the Crowing crested cobra as if it were a real elapid snake, with the sort of cephalic scalation expected for cobras and their kin. Image: Marcus Bühler, used with permission.

As might be clear by now, I’m not holding out much hope that the Crowing crested cobra ever existed. If it did, it would be fun to wonder what sort of snake it could be – presumably an elapid related to, or within, the cobra radiation. Tthe illustration above – by Markus Bühler – is an imaginative attempt to reconstruct the Crowing crested cobra as if it were a real, biologically plausible elapid. However, it could just be a ‘mythified’ version of the Black mamba (Spawls 1979). It would be awesome to be completely wrong about this. But, alas, the animal almost certainly represents a curious amalgam of myth, superstition and faulty observation.

For previous Tet Zoo posts on other squamates see…

In Quest of Anguids, May 2006
Cambodia: now with dibamids!, May 2011
The Tet Zoo Guide to Mastigures, August 2018
The Remarkable Basilisks, May 2023
Do Lizards Really Have ‘Mite Pockets’?, March 2024
Meeting Lake Zacapu’s Garter Snake, March 2024
Ray Hoser, Number 1 Taxonomic Vandal, May 2024
The Mysterious Dibamids, May 2024
Side-Stabbing Stiletto Snakes, Redux for 2024, May 2024
The Rehabilitation of Günther’s Black Cameroonian Snake, May 2024
Ikaheka and Other ‘Palatine Draggers’, Cryptozoic Elapid Snakes of Melanesia, June 2024
Arboreal Alligator Lizards of Mesoamerica... and Beyond!, July 2024
Grayiids: Africa Has Water Snakes Too, July 2024
Record-Holding Champion Giant Slow-Worms, July 2025

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

Refs - -

Shircore, J. O. 1944. Two notes on the Crowing crested cobra of Africa. African Affairs 43, 183-186.

Shuker, K. P. N. Extraordinary Animals Worldwide. Robert Hale, London.

Shuker, K. P. N. 2007. Extraordinary Animals Revisited. CFZ Press, Woolsery, Devon.

Spawls, S. 1979. Sun, Sand & Snakes. Collins and Harvill Press, London.

Spawls, S. & Branch, W. 1995. The Dangerous Snakes of Africa. Blandford, London.

Spawls, S., Howell, K., Drewes, R. & Ashe, J. 2002. A Field Guide to the Reptiles of East Africa. Academic Press, San Diego.

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Regular readers here will be familiar with my lamentations about the old, archived material from ver 2 (ScienceBlogs) and ver 3 (Scientific American). It’s been lost, destroyed, vandalized, paywalled, or some combination of those things. Today, something happened which has inspired me to rescue one of those articles from ver 3, specifically from 2013 (here’s the original). What inspired it, huh? Well, THIS DID…

**Caption:** *these screengrabs are obviously not the best* but… yes, that’s a large Saltwater crocodile *Crocodylus porosus* attacking a small Asian elephant *Elephas maximus*. I’m very confident that the footage is real and absolutely not AI. The croc can be identified as *C. porosus* on the basis of its scute-free dorsal neck surface, not size alone. The footage is credited to © Sufri.Johny and was shared on Facebook by © **Wildman Adventures**.

You can watch the footage yourself since it’s visible here on Facebook where it’s been shared globally (as in, with the entire internet), meaning that you don’t need Facebook membership or login for access. Ok, the elephant is one of the small ones from the Bornean Sabah population, but it’s still an elephant, apparently an adult one (on balance, it might be a ‘sub-adult’, or or teenager; I can’t tell), and the interaction is still rad as hell. It would appear that the crocodile’s behaviour is opportunistic… and also unsuccessful. On that note, the elephant is not only very much alive at the end of the encounter, but also not seemingly injured at all. I can’t even see any perforations or lacerations on the skin.

**Caption:** another screengrab, here showing the second lunge from the crocodile, and showing it biting the elephant right across the upper right side of the shoulder region. The footage is credited to © Sufri.Johny and was shared on Facebook by © **Wildman Adventures**.

Here's where we come to the aforementioned 2013 article. This, you see, is far from the first time that a crocodile has been filmed attacking an elephant. Ok, the other cases we’re aware of – or, I’m of, anyway – are not so impressive and dynamic. But exist they do.

Way back in November 2010 a remarkable photo appeared online, this time showing an adult Nile crocodile Crocodylus niloticus biting the trunk of an adult female African bush elephant Loxodonta africana (a plague upon those bloggers and others who identified the crocodylian as an... alligator. Come on). You've almost certainly seen the photo already: it was widely features in newspapers, magazines, blogs and such. My initial plan was to assume that everybody's seen it, but that would be frustrating to those of you that haven't so, whatever, here it is...

**Caption:** one of the best of the relevant photos. They were used uncredited quite a bit, but should be considered © Ben Campbell.

Above we see the moment where the mother elephant is raising her trunk, the croc clamped around the end and still in the water. But a whole sequence of events was photographed: the elephant actually pulled the croc right out of the water entirely, at which point the croc let go. The baby elephant tripped over the crocodile, but all three animals survived. In fact, both elephants were definitely ok since they were seen later in the day at the same river (the Luangwa River). This incident happened in Zambia's South Luangwa National Park; the photographer was Swiss tourist Martin Nyfeler. Here's the whole sequence, as it appeared here on the blog Ben in Zambia...

**Caption:** a full, detailed account of what happened and was photographed is present on Ben Campbell’s blog where it was published in September 2010. The blog is no longer active, but the photos should be considered © Ben Campbell.

Remarkably, this isn't the only time a crocodile has been photographed grabbing an elephant's trunk. During October 2010, Johan Opperman photographed an incident in Kruger National Park, this time featuring a baby Bush elephant. The baby was part of a group that waded across a small river, the surface totally covered in floating plants. As you can see, the crocodile grabbed the baby's trunk. The other elephants rallied to the baby's defence and the crocodile released its grip and disappeared. Here's one of the several images...

**Caption:** Johan Opperman’s 2010 photo of a crocodile pulling the trunk of a juvenile elephant was published in several places; **here’s an appearance in *The Telegraph***. Image: © Johan Opperman.

But it gets even better. Did you know that croc-attacks-elephant-trunk incidents had been FILMED? Not once, but at least twice. Yup, here's number one... it’s titled Crocodile bites elephant on the trunk and was uploaded by YouTube user 92BMW318is, if that is their real name. Here’s a screengrab, sorry for the terrible low quality…

I can't pretend to have done any research on this specific incident and don't know anything about it, other than it looks like it was filmed in the wild, somewhere in Africa. It's clearly genuine. The elephant that gets attacked appears to be another adult female. After recovering from the shock of "OMG there's a crocodile on my trunk", her reaction seems to involve aggression and retaliation, since she seems to charge into the water in pursuit. And here's number two, though be aware that it’s blurry as hell... (the main incident happens at 0:16-0:17; this version is a shortened version of what was originally a longer video). Again, here are screengrabs of the main moments of interest…

**Caption:** this attack occurred in moderately deep water (for a river), and a few moments show that the crocodile grabbed the trunk’s tip before letting go. Thanks to RoryD for bringing this piece of footage to my attention. It happened in the Chobe River in Botswana and was uploaded in April 2012 by YouTube user jwakf.

Here's the thing that immediately strikes me about these incidents. They're been photographed or filmed at least four times*. As you'll know if you're an expert on elephant and/or crocodile anecdotes, there are allusions in the literature to at least a few other cases of the same sort of thing. I recall one incident in particular where a crocodile attacked a juvenile elephant; an adult elephant grabbed the crocodile, pulled it on to the bank, trampled it, and threw its corpse into a tree. Anyone recall this as well?

Five times. Here’s another case from 2015, this time from Sabi Sands Nature Reserve in South Africa.

Anyway, if crocodiles have been seen to attack elephant trunks on at least four five separate occasions, it seems safe to assume that this has actually happened on more occasions than this, since there must be occasions where (1) people didn't film/photograph it, and (2) people weren't there to see it happening. In the grand scheme of things, crocodile attacks on elephant trunks are probably pretty rare… and probably always have been, even if we think of a time when there were far more crocodiles and far more elephants, but... it does make you wonder.

**Caption:** I’ve never read the Rudyard Kipling stories, but I do know that there’s one called *The Elephant’s Child*, and that it includes a segment where the elephant gains its trunk thanks to the actions of a crocodile. I believe that this illustration was created by Kipling in 1902; image in the public domain.

None of the attacks mentioned here resulted in the death of an elephant. Indeed, it may be that the crocodiles involved in these incidents made mistakes, thinking that the object that they were grabbing was attached to something far more manageable. However, note that a damaged trunk can prove fatal for an elephant: they can literally be disabled by trunk blockages and amputations, since an inability to forage, drink or breathe can result in declining health and eventual death. This at least makes it plausible that a crocodile could result in an adult elephant's death. On the other hand, elephants can survive with mutilated trunks. The photo below shows an individual photographed by Marlon Du Toit: it had a mutilated trunk and had to learnt to squirt water into its mouth from its damaged trunk tip.

**Caption:** the full story behind this specific elephant used to be findable online, but I can no longer locate any trace of it, not even at wayback machine. There are plenty other images online of elephants with damaged – *sometimes even mostly missing* – trunks; it’s clearly a not uncommon problem, linked in part to the widespread use of snares. Image: © Marlon Du Toit.

One final thing. Having said that these incidents are probably very rare, I wonder if they might have happened on enough occasions that elephant behaviour has been modified accordingly. I mean: are there places with high crocodile densities that elephants deliberately avoid as drinking spots? Or are there places where elephants are especially careful, or where they do unusual things to 'test' for the initial presence of lurking crocs? And let us not forget the crocodylians and elephants (and other proboscideans) of the geological past. Forgive me, I couldn't help but knock up the following quick illustration...

**Caption:** the famous extinct proboscidean (*not an elephant!*) *Platybelodon* has an unpleasant encounter with the big, long-jawed fossil crocodylian *Euthecodon* somewhere in Miocene or Pliocene east Africa. *Platybelodon* has traditionally been reconstructed as having a broad, flattened structure where elephants have a trunk, but it’s now widely thought that this is wrong, and that it and its relatives had trunks. They weren’t the wading ‘shovel-tuskers’ of tradition, but parkland-dwelling herbivores. Image: Darren Naish.

And everything you read here was inspired by a conversation I had with Mike P. Taylor and Matt Wedel.

For previous Tet Zoo articles on elephants, see...

RIP Yeheskel Shoshani, May 2008
How do you masturbate an elephant?, June 2008
The tangled mammoths, November 2008
The Domes of Wisdom: an Asian Elephant Tale, August 2022

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

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Today is July 31st 2025… do you know what this means?

**Caption:** a montage of things somehow relevant to the history of Tet Zoo ver 4. From left to right: **the 2022 publication of *Ancient Sea Reptiles*** is deeply relevant to the history of this blog (it saw release as a second edition in 2023); my interest in cassowaries led to **field collaboration with Todd Green in 2023** (that’s not Todd in the photo, but Liberace the red-necked northern cassowary); finally, the tenure of Tet Zoo ver 4 overlapped with the final days of **Dinosaurs in the Wild**, a travelling exhibition connected to various things that happened before and since. Images: Darren Naish.

It means, dear reader, that Tetrapod Zoology ver 4 – the version of the blog you’re consulting right now – began life on this very day, all but seven years ago, in 2018. And thus: happy 7th birthday Tet Zoo ver 4!

As a regular reader, you’ll know that Tet Zoo’s proper official birthday is on January 21st, and the next of those is a big one, marking 20 YEARS OF TETRAPOD ZOOLOGY. I feel like I should mark that event specially in some way, but quite how is not yet clear. Maybe a special line of merch should be released or something like that. The ver 4 birthday isn’t such a big deal, but it’s still one worth marking, and thus here we are.

**Caption:** TetZooCon – the annual Tetrapod Zoology Convention – has been a mainstay throughout the duration of Tet Zoo ver 4. The 5th and 6th of those events happened in 2018 and 2019, respectively, but a global pandemic meant that we switched to zoom-based versions in 2020 and 2021. The 9th event (2022) was in-person again, and things were back to normal for the 10th (2023) and 11th (2024). And that 2024 TetZooCon was the last one ever. Thanks to those who made it what it was.

The usual backstory stuff. As explained in the very first ver 4 article from July 2018, ver 4 started life as I packed my bags and said goodbye to Scientific American’s so-called blogging network in order to set up home here, a web location already created thanks to the podcast that John Conway and I record on an intermittent basis. Yes, the podcast is still a viable enterprise, it’s just that John and I have wholly incompatible schedules. We have had for years now.

**Caption:** John Conway and I have been working closely together for what feels like an eternity at this point. We talk about palaeoart a lot; at left, John is pondering different versions of a Greg Paul illustration. **The podcast back-catalogue is here.** John insists on wearing sunglasses indoors, that’s how cool he is (kidding, he actually doesn’t). Images: Darren Naish.

Here’s your reminder that the podcast back catalogue is here; many of the episodes are trainwreck garbage but at least some are good listening. My favourite is ep 72, the Loch Ness Monster special. I thank the people who say kind words about the whole podcast endeavour; it’s really appreciated.

Blog content, some thoughts. Anyway, the podcast aside, the blog itself has gone from strength to strength and there’s now a good amount of content covering most of the major tetrapod-themed topics I’m interested in or qualified to write about. There is, as ever, so much more to do, and I’m forever driven furious by the fact that I just can’t make time to write and publish more than I do already. Support me at Patreon dammit.

**Caption:** a good chunk of my time over the years since 2016 has been taken up by the making of TV shows that have a sort of connection to the seminal *Walking With Dinosaurs* of 1999. Partly for that reason, I thought it appropriate to publish my thoughts on that series: check out **Part 1** and **Part 2**, both published in 2021. The image at left shows Mike Milne in an Impossible Picture office (RIP Mike; he died in 2024); the image at right is the intro from Haines (2000).

One of my aims is to rescue material from ver 2 (the ScienceBlogs years) and ver 3 (the Sci Am years) and re-publish it here at ver 4, since those respective hosters either no longer really exist, or (in the case of Sci Am) have ruined the material they host by removing its images or paywalling it, or both. You’ll know that I’ve rescued some material but there’s still so much more to get to. I also aim to have this older content published in book form, but that’s something I just can’t make time for either.

Massive, heartfelt thanks to the wonderful people who support me at patreon and assist in the perpetuation of this blog and the support of these planned publication projects. Please consider assisting me if you don’t already; there’s tons of unique, secret content there that’s never been shared anywhere else.

For the remainder of this article I want to bring attention to a hand-picked, annotated list of Tet Zoo ver 4 articles that I’m especially fond of, grouped together by subject. My aim in part is that you might be inspired to look at them again.

**Caption:** my workload has constantly thwarted my ability to publish as many books as I might like, but at least a few have seen release while ver 4 has been active. Among them are the third edition of *Dinosaurs: How They Lived and Evolved* (2023; with Paul Barrett), *Dinopedia* (2021) and *Mesozoic Art* (2022; with Steve White). Other books are currently in production or due to see print soon.

Mesozoic dinosaurs are a mainstay here at Tet Zoo and always have been, which is understandable both because I publish technical and popular research on these animals (my publications are here) and because they are – sorry other tetrapod groups – undeniably among the most awesome and fascinating animals of them all. Having said that, I’m still not sure whether I cover non-bird dinosaurs too much, or too little. Or do I cover them… just right? Among my favourite ver 4 articles are…

Could We Domesticate (Non-Bird) Dinosaurs?, August 2018
Recollections of Dinosaurs Past and Present, the 1980s Exhibition, February 2019
Two New Spinosaurid Dinosaurs from the English Cretaceous, September 2021
The Eotyrannus Monograph Is Finally Published, July 2022
Robert Plot’s Lost Dinosaur Bone, December 2022
Post-Truthism, Brian J Ford’s Aquatic Dinosaurs, and the Fate of ‘Too Big To Walk’, November 2024
My Weird 2000 Paper on Tree-Climbing Dinosaurs, December 2024
Heptasteornis, My Beloved; Alvarezsaurids in Europe, the Backstory, January 2025

Caption: this was intended as a dinosaur-themed montage, but it says a lot that it’s ended up as, specifically, a theropod-themed one. Clockwise from top left: tree-climbing juvenile dromaeosaurids, from here; Eotyrannus in life, by Loana Riboli (copyright, used with permission), from here; a friendly ornithomimid with its human companion, by Mike Skrepnick (used with permission), from here; and British baryonychine spinosaurids new to science as of 2021, by Anthony Hutchings, from here.

Speculative Zoology – SpecZoo – is another topic that’s always been a Tet Zoo mainstay. My approach to this subject has been piecemeal so far, largely taking the approach of looking at one book or one idea, even, at a time. There’s call for big, synthetic articles that combine and contrast the writings of various contributors. Regular readers of Tet Zoo will know my argument (albeit not original to me) that SpecZoo has significant overlap with cryptozoology, a matter I aim to explore later this year in an academic manuscript.

The Dougal Dixon After Man Event of September 2018, September 2018
Dougal Dixon’s After Man, the Initial Pitch Document, June 2020
Humanoid Dinosaurs Revisited Again: Russell and Séguin’s Dinosauroid at (Nearly) 40 Years Old, August 2021
The Snouters or Rhinogradentians, a Brief Review, October 2021
Alternative Timeline Dinosaurs, the View From 2019 (Part 1), November 2019
Alternative Timeline Dinosaurs, the View From 2019 (Part 2), December 2019
Alternative Timeline Dinosaurs, the View From 2019 (Part 3): the Dinosauroid and its Chums, December 2019
Speculative Zoology and the World of After Man; an Interview With Dougal Dixon, April 2025
Speculative Zoology Grand and Photoreal: Boulay and Steyer's Demain, les Animaux du Futur, April 2025

Caption: ‘dinosauroids’ of two very different sorts. At left, the Russell-Séguin version (as illustrated for my 2021 book Dinopedia), a humanoid invented to show the inevitability of the humanoid form (see the full article here). At right, a more ‘realistic’ dinosauroid (in my view, of course): a smart maniraptoran theropod that kills mammals for a living. Images: Darren Naish.

Megamammals. I’m more of a reptile guy than a mammal one but I really do like mammals too, and among those I keep coming back to are the big ‘ungulates’, both of the land and of the water. Perissodactyls have had a fair bit of coverage here (and more is set to come), artiodactyls too… and that includes whales, which are artiodactyls: down with the pointless and redundant ‘Cetartiodactyla’ (Prothero et al. 2021). Proboscideans are under-represented here, as are fossil megamammals.

Whatever Happened to the Kabomani Tapir?, October 2018
Remembering Lyall Watson’s Whales of the World, June 2019
Extreme Cetaceans, Part 1, September 2019
Kogia, Shark-Mouthed Horror, March 2022
Hartebeest, Long-Faced Antelopes of Many Forms, June 2022
The Domes of Wisdom: an Asian Elephant Tale, August 2022
The Slightly Surprising Diversity of Zebras, Part 1, October 2022
The Discovery of the Okapi, Part 1, April 2023
The Discovery of the Okapi, Part 2, April 2023
The Slightly Surprising Diversity of Zebras, Part 2, July 2023

Caption: come on, who doesn’t love tapirs? Tapirs have been mentioned or discussed so many times within the TetZooniverse that they might be unofficial emblems of the whole enterprise, as implied in this illustration (left) by Patrick Murphy. At right, this cladogram (from Ruiz-García et al.’s (2015) study of South American tapirs) shows how Kabomani tapirs (green) appear to be a lineage within Tapirus terrestris (red). For more, see the 2018 Kabomani tapir article here.

Cryptozoology. I’m a sceptic and emphatically not (at this point in my life) a ‘believer’ when it comes to Bigfoot, Nessie, sea monsters, living thylacines and so on (Naish 2017). But what seems to confuddle some people beyond all measure – yes, I do have a specific Wikipedia editor in mind here – is my belief that cryptozoological claims, data and hypotheses should be evaluated fairly in case they represent descriptions of real observations or encounters, not rejected out of hand. There’s been a fair bit of content here that approaches the subject in that way, but… let’s face it: it’s also really interesting to write about the beliefs and pet (sometimes wayward) hypothesis of cryptozoological authors.

Sea Monster Sightings and the ‘Plesiosaur Effect’, April 2019
Books on the Loch Ness Monster 3: The Man Who Filmed Nessie: Tim Dinsdale and the Enigma of Loch Ness, August 2019
Lore of the Loveland Frog, January 2020

Caption: cryptozoology includes the classic famous cryptids – Bigfoot, Nessie and the like – but what about the fringe creatures at the edges? Among my favourite is the Loveland frog of the 1950s and 70s, a creature I covered here in 2020. Images: alleged original eyewitness sketch, and a later redrawing by Ron Schaffner.
The 1972 Loch Ness Monster Flipper Photos, August 2020
The Case of the Cadborosaurus Carcass: a Review, November 2020
The Lake Dakataua ‘Migo’ Lake Monster Footage of 1994, February 2021
eDNA, Footprints and the Biological Bigfoot: Comments on an Interview With Jeff Meldrum, April 2021
Legend of the Black Dog, August 2022
Werewolves in America; the Tale of Dogman, July 2024

Caption: water monsters are great value for money, and doubly fascinating because many (albeit not all) of the accounts do involve real sightings of real animals. At left, stills from the Migo footage of 1994, covered here. Middle: it sure would be nice if Nessie sightings like this actually occurred, but they really didn’t. This book is by Tim Dinsdale, and for more on him go here. At far right, the Shiels Nessie, which I’ve written about at length but not necessarily here at Tet Zoo.

Caption: recalling what I said above the Loveland frog, are semi-mythical creatures like the Black dog (a special kind of phantom, not just a dog that’s black) relevant to conventional zoology? Well, I’m sufficiently interested in them that I have written about them (go here).

Debates in bird evolution. I have a vested interest in what we politely term non-standard hypotheses, and birds – for whatever reason – are afflicted by these. You can’t work on Mesozoic dinosaur or fossil birds and be unaware of the ‘Birds Are Not Dinosaurs’ (BAND) crowd, and – for my sins – I’ve covered that topic more than once. But birds of more modern sort (the neornithines) have an association with non-standard hypotheses too, and what’s interesting and telling is that the people who promote these are… the exact same set of contrarian nay-sayers, the BAND guys…

Heilmann, Thompson, Beebe, Tetrapteryx and the Proavian, January 2019
Controversies in Ratite and Tinamou Evolution (Part I), July 2023
Alan Feduccia’s Romancing the Birds and Dinosaurs: Forays in Postmodern Paleontology, October 2023

Caption: Alan Feduccia has had more than his fair share of time in the limelight, but his style of argumentation really needs to be called out for the contrarian pseudointellectualism that it is. At right, a rough consensus cladogram showing relationships within Archosauria. Feduccia argues in his latest book that we should abandon this (relatively) resolved tree and instead throw our arms in the air and embrace something more chaotic. Images: Darren Naish.

Frogs, salamanders, caecilians. I’m a massive amphibian nerd and have done what I can to give them fair credit here: there’s a ton of fascinating diversity in their biology and history, their conservation and global decline needs adequate coverage, and I’ve invested massively in assisting the lone species that occurs in the overbuilt, suburban bit of little England where I live. I haven’t covered amphibians enough, for sure, but there’s still some good stuff here…

A Love Letter to the Common Frog, August 2020
Allodapanura, the Biggest Frog Group You’ve Never Heard Of (Part 1), October 2020
Tiny Frogs and Giant Spiders: Best of Friends, August 2022
The Remarkably Weird Skeletons of Frogs, September 2022
The Amazing Caecilians, October 2022
What, If Anything, Is A Strabomantid?, January 2024
Small Obscure African Toads; Toads Part 14, January 2024
Live Spawnwatch Action From Pond 2 at Tet Zoo Towers, February 2024

Caption: my efforts to help Common frogs Rana temporaria and create more, and better, ponds, have been regular points of discussion here. I’ve built (and helped manage) several ponds during ver 4’s duration but still need to write at length about pond construction and design. Images: Darren Naish.

Squamates. I mentioned earlier my hopes that I might be able to rescue old content from previous versions of the blog. But this is such a daunting task that I can only make progress if I stick to articles on a specific animal group. For that reason, I’ve been rescuing and republishing article on squamates. Here are some of my favourites (with some others that were published here at ver 4 for the first time).

The Remarkable Basilisks, May 2023
Do Lizards Really Have ‘Mite Pockets’?, March 2024
The Mysterious Dibamids, May 2024
Side-Stabbing Stiletto Snakes, Redux for 2024, May 2024
The Rehabilitation of Günther’s Black Cameroonian Snake, May 2024
Ikaheka and Other ‘Palatine Draggers’, Cryptozoic Elapid Snakes of Melanesia, June 2024

Caption: squamates have had some amount of coverage here, but it’s still not enough. For whatever reason, iguanians are over-represented (probably because they’re among the squamates most frequently encountered in captivity here in the UK). At left, captive Omani spiny-tailed lizard or Thomas’s mastigure U. thomasi from this 2018 article. At right, a skink cladogram from this 2020 article.
```
 **Caption:** a montage which helps show how many squamate-themed articles were published at Tet Zoo back in the olden times. Virtually all of these articles are now only findable at wayback machine, and even then not in intact version. I’ve been doing what I can to rehabilitate them here at ver 4.
```

Finally, I should add that there are a number of additional articles that I also regard as useful and worthy but which don’t fit into the above categories. We can regard them as random favourites. Among them is my article my take on the internet culture of the potoo (February 2019), that on the life appearance of the Dodo Raphus cucullatus (July 2020), a compilation of surprising and odd, animal-themed Palaeolithic rock art (July 2019), and my retrospective on the 2001 Dorling Kindersley book Encyclopedia of Dinosaurs and Prehistoric Life (January 2022). And I haven’t rounded up articles on several other topics that have been revisited here several times, among them Mesozoic marine reptiles, my birdwatching efforts, the Tet Zoo Reviews Zoos series, and my writings on azhdarchoids (and other pterosaurs).

**Caption:** ‘Our Lord and savior potoo bird’. This image is by dragongirl222, who does **a whole range of potoo-themed merchandise at redbubble**. Image: (c) **dragongirl222**.

And hello DinoCon. Ok, that’s where we’ll end. Thank you for joining me in this 7th birthday celebration; special thanks again to those who help support Tetrapod Zoology. The proper birthday – in which I review the activities and events of the year that’s passed (and, oh boy… 2025…) – will be a far lengthier, more complex article, but we have some way to go yet before that time comes.

As I write, I and others are of course busy getting ready for another major event in the Tetrapod Zoology calendar: the first ever DinoCon (go here for info and tickets), the descendant of TetZooCon, this year held at the University of Exeter (UK) on the weekend of August 16th and 17th. The gamble has paid off and DinoCon is set to be huge, way bigger than even the biggest TetZooCon. This basically means that DinoCon is already set to be a regular fixture and can only get bigger and better with time. Do come along if you’re not already planning to do so.

**Caption:** the promotional art for DinoCon (which will be available on merchandise) was created by Natalia Jagielska.

And that’s that. Thanks for joining me here… more coming soon! For previous Tet Zoo birthday articles, see…

Happy first birthday Tetrapod Zoology (part I), January 2007
Happy first birthday Tetrapod Zoology (part II), January 2007
Happy second birthday Tetrapod Zoology (part I), January 2008
Tetrapods of 2007 (happy birthday Tet Zoo part II), January 2008
Happy THIRD birthday Tet Zoo, January 2009
Tet Zoo = 4 years old today, January 2010
2009, a year of Tet Zooery, January 2010
Four years of Tet Zoo: to infinity... and beyond!, April 2010
It is with some dismay that I announce Tet Zoo's first hemi-decade, January 2011
Tet Zoo 5th birthday extravaganza, part II, January 2011
Happy Birthday Tetrapod Zoology: SIX YEARS of blogging, January 2012
Happy 6th Birthday, Tetrapod Zoology (part II), January 2012
Tetrapod Zoology enters its 8th year of operation, January 2013 (I cannot find any intact versions of this article, thanks SciAm)
Today marks NINE YEARS of Tetrapod Zoology, January 2015
Tetrapod Zoology 10th-Birthday Extravaganza, Part 1: 2015 in Review, January 2016
Tetrapod Zoology 10th Birthday Extravaganza, Part II: the Rest of 2015 Reviewed, January 2016
Tetrapod Zoology 10th-Birthday Extravaganza, Part 3: Tet Zoo's Tetrapod Treatment in 2015, January 2016
Today Is Tet Zoo's 11th Birthday, January 2017
The 12th Year of Tet Zoo, January 2018
The Tet Zoo 12th-Birthday Event, Part 2, January 2018
The Much Belated Final Part of the Tetrapod Zoology 12th Birthday Event, December 2018
Tetrapod Zoology Is A Teenager Now, January 2019
Tetrapod Zoology's 14th Year of Operation, 2019 in Review, January 2020
On Tetrapod Zoology’s 15th Birthday, the Year in Review, January 2021
Happy 16th Birthday, Tetrapod Zoology, February 2022
Four Years of Tetrapod Zoology ver 4, July 2022
The 17th Year of Tetrapod Zoology: 2022 in Review, January 2023
Five Years of Tetrapod Zoology ver 4, July 2023
Tetrapod Zoology Reaches 18 Years of Age, January 2024
It Was the 19th Year in the History of Tetrapod Zoology, January 2025

Refs - -

Haines, T. 2000. J’accuse: Tim Haines. The Dinosaur Society UK Quarterly Magazine 3 (4), 8-9.

Naish, D. 2017. Hunting Monsters. Arcturus Books, London.

Prothero, D. R., Domning, D., Fordyce, R. E., Foss, S., Janis, C., Lucas, S., Marriott, K. L., Metais, G., Naish, D., Padian, K., Rössner, G., Solounias, N., Spaulding, M., Stucky, R. M., Theodor, J. & Uhen, M. 2021. On the unnecessary and misleading taxon “Cetartiodactyla”. Journal of Mammalian Evolution 29, 93-97.

Ruiz-García, M., Castellanos, A., Agueda Bernal, L., Navas, D., Pinedo-Castro, M. & Mark Shostell, J. 2015. Mitochondrial gene diversity of the mega-herbivorous species of the genus Tapirus (Tapiridae, Perissodactyla) in South America and some insights on their genetic conservation, systematics and the Pleistocene influence on their genetic characteristics. Advances in Genetic Research 14, 1-51.

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Once again, I’m back from time spent in the North Atlantic looking at wild cetaceans, specifically on a Bay of Biscay trip (a journey made between Plymouth in England and Santander in Spain) organised by the wildlife charity ORCA…

**Caption:** ORCA does its whale surveying aboard various vessels that cover numerous routes. So far, I’ve only done the southern England to northern Spain route; here’s our vessel in Santander, Spain. Images: ORCA; Darren Naish.

Over 520 individual cetaceans were recorded on this one trip, belonging to at least six species. This article mostly serves as an excuse to share photos taken by myself and by my associate Alex Srdic, but let’s also use it as an opportunity to look at news pertaining to western Europe’s cetacean species.

Porpoises, dolphins, and bycatch. A lot of the news about cetaceans around Europe is, of course, bad and concerns deaths caused by bycatch and entanglement. A 2025 report compiled by EIA (the Environmental Investigation Agency) states that over 35000 (not a typo) Harbour porpoises Phocoena phocoena are trapped each year in European waters as bycatch (mostly by gillnets), and that Common dolphin Delphinus delphis numbers killed by bycatch have increased markedly since 2020. The porpoise numbers are ridiculous and we clearly have a major problem. We’re talking about multiple individuals being killed every single day in every single sea and coastal inlet of our region, with the North Sea, Celtic Sea and English Channel being among the areas with highest porpoise mortality. Here’s the UK government’s assessment of this situation if you want more detailed breakdown.

**Caption:** a Common dolphin montage, showing individuals leaping as well as a family group moving at a more sedate pace. Images: Alex Srdic; lower one Darren Naish.

15 Harbour porpoise were seen on our trip and over 330 Common dolphins, these numbers indicating that both species are abundant and easy to find, despite these concerning declines. It also appears that animals are unable to make a living due to ecological disruption caused by both climate change and industrial fishing. In recent years, reports have increased of dolphins that are in poor nutritional condition, with low fat reserves and even obvious emaciation (e.g., Levesque et al. 2021 for Irish coast; Albrecht et al. 2021 for Celtic Sea). I do not have a positive view at all of the fishing industry but at least it’s becoming better known how bad things are. If you really must eat seafood, make sure that it’s from sustainable fisheries… though even that doesn’t seem to be curtailing things.

**Caption:** leaping Common dolphin. This species is highly variable in pattern and colour (and there’s ecomorphological variety in beak length as well); this one is interesting in having irregular pale patches on its dorsal cape. Image: Alex Srdic.

A diversion on Stenella dolphins. Having mentioned abundant dolphin species, we also saw around 10 individuals of the mostly deepwater Striped dolphin Stenella coeruleoalba. They tend to be highly acrobatic and prone to leaping high, and their lateral stripes and mostly grey palette are usually obvious.

**Caption:** leaping Striped dolphin showing eponymous markings. Image: Alex Srdic.

One of the most interesting things about Striped dolphins (to me) is that it’s recently been argued that a related species – the Clymene or Euphrosyne dolphin S. clymene – is a naturally occurring hybrid between the Striped and the Spinner S. longirostris (Amaral et al. 2014), though it seems (for now) that experts are still happy to regard it as a species. This is not, however, as clear-cut as generally implied since not all studies unambiguously support hybrid status for S. clymene (Faria et al. 2022). Yes, the Stenella dolphins do hybridize (including with Common dolphins as well), and some hybrids do backcross with the parent species, but not all studies find all Clymene dolphins to be hybrids!

**Caption:** a pair of leaping Striped dolphins. Spray and very visible splashing is typical of Striped dolphin leaps. Image: Alex Srdic.

Bottlenose and Risso’s dolphin: all change. On ecological disruption and climate change, at least some cetacean species around Europe’s shores are changing their distributions and perhaps even their habits as a consequence. Sighting records of Bottlenose dolphins Tursiops truncatus around the UK reveal some considerable change. This species is mobile enough that individuals recorded off Scotland have later been seen off The Netherlands (Hoekendijk et al. 2021); however, static and semi-resident groups are also present off Wales and eastern Scotland. Over the past ten or so years, Bottlenose dolphins around the UK have been more frequently reported off the eastern coast of England and are seemingly increasing their range southwards (e.g., Aynsley 2017). Along England’s southern coast, they appear to be moving eastwards, and data from some years suggests reduced presence (which could mean a declining population) in England’s south-west. Anyway, we didn’t definitely see any Bottlenose dolphins on this trip (a contrast to previous years) but… what is this?

**Caption:** this dolphin (the two images show the same individual) was seen and photographed by several people but remained ambiguous. It was mostly grey with a pale belly and robust rostrum. It looks something like a bottlenose dolphin but also doesn’t quite match one. Images: Darren Naish.

At the same time as the Bottlenose has moved south and east, the remarkable Risso’s dolphin Grampus griseus has been moving north over the last three decades such that it’s now regularly seen off Scotland and the northern isles and is now a relatively abundant animal in European shelf waters, including those around the coasts of the UK (Hodgins et al. 2024). It also seems to be becoming more coastal in habits overall, previous records revealing it to be mostly a deepwater whale. This is presumably linked to changing prey distribution but isn’t well understood.

Prior to this year, I’ve only fleetingly seen a Risso’s dolphin once, and this was a dorsal fin alone. For 2025, we saw a group of nine off the southern English coast, and I also saw what I think was a very white one at close range off the coast of Brittany. Incidentally, Risso’s dolphin is likely not a close relative of ‘typical’ dolphins (like Tursiops and Delphinus): it instead seems allied to the globicephalines – the pilot whales and such (LeDuc et al. 1999, Caballero et al. 2008, McGowen 2011, Geisler et al. 2011). I wrote an article about this in 2012 at Tet Zoo ver 3.

**Caption:** two of a group of nine Risso’s dolphin seen on our trip. The blunt head, extensive pale scarring and big, slender dorsal fin are all characteristic. Image: Alex Srdic.

  **Caption:** characteristic dorsal fins of Risso’s dolphins. Image: Alex Srdic.

Beaked whales and rorquals. No Bay of Biscay trip would be complete without at least one Cuvier’s beaked whale Ziphius cavirostris sighting, and two were seen on our trip (albeit only distantly and fleetingly). Yes, there’s a bit of a push to get this animal’s name changed to the alternative ‘Goose-beaked whale’ but that certainly hasn’t caught on among people who study whales or survey them. This species dives to 2.9 km and thus holds the current deep-dive record over elephant seals and sperm whales. As you’ll recall if you’re a regular reader, I’ve seen numerous beaked whales (both Ziphius and Mesoplodon) on these trips before and had high hopes. Alas.

**Caption:** a mid-sized whale identified as a Northern minke, though the field sign used to support this identification are not immediately clear from this photo. Minke surface briefly once or twice and then disappear. Image: Alex Srdic.

  **Caption:** a second Northern minke individual seen closer to the UK, this time giving us a great view of its distinctive dorsal fin. Image: Alex Srdic.

Finally, we saw numerous (11) Northern minke whales Balaenoptera acutorostrata. We really should call them ‘Northern minke’ rather than just ‘Minke’ since a second species – the Antarctic minke B. bonaerensis – has been reported from the North Atlantic (a lone one was killed off Jan Mayen in 1996 and hybrids are known for the region too) (Glover et al. 2010) and there are suspicions that they might be present in the region more than realized so far. We saw no Fin whales B. physalus this time, which is weird given the more than 30 seen on 2024’s trip.

The unidentified ones. As typical for sea trips that involve looking at cetaceans, numerous animals were seen for which definitive identifications couldn’t be achieved. A big, unidentified whale was seen moving swiftly away from the ship’s wake at one point, an unidentified beaked whale was seen, and some number of indeterminate dolphins were also recorded.

During the time of our survey, two Killer whales Orcinus orca were reported from off the Scilly Isles and recognised on the basis of their markings as individuals from an Iberian population. Needless to say, we had vague hopes that we might get to see them… but nope.

**Caption:** I like seeing animals, but landscapes, seascapes and the sky are all great as well. We had fantastic views of the full moon over various coastal seascapes, like this one – I think off the coast of France. Image: Darren Naish.

That’s where I’ll end. The trip was massive fun and I’m thrilled with what I saw. Consider supporting ORCA and joining their trips if you can. Massive thanks to Alex for use of his photos, and thanks too to my travel companions Matthew Harrop and Richard Hing.

**Caption:** these trips are one of the few occasions on which I get to see unobstructed views of both sunrise and sunset. This sunrise is from the morning of 11th July 2025. Image: Darren Naish.

Cetaceans have been covered a few times at Tet Zoo before, see…

Remembering Lyall Watson’s Whales of the World, June 2019
Whale Watching in the Bay of Biscay, August 2019
Extreme Cetaceans, Part 1, September 2019
Extreme Cetaceans, Part 2, September 2019
Extreme Cetaceans, Part 3, December 2019
Kogia, Shark-Mouthed Horror, May 2023
In Memory of Richard Ellis, Influential Artist and Author, June 2024

You can support this blog and my other projects here at Patreon

Refs - -

Albrecht, S., Minto, C., Rogan, E., Deaville, R., O’Donovan, J., Daly, M., Levesque, S., Berrow, S., Brownlow, A. Davison, N. J., Slattery, O., Mirimin, L. & Murphy, S. 2024. Emaciated enigma: decline in body conditions of common dolphins in the Celtic Seas ecoregion. Ecology and Evolution 14, e70325.

Amaral, A. R., Lovewell, G., Coelho, M. M., Amato, G. & Rosenbaum, H. C. 2014. Hybrid speciation in a marine mammal: the clymene dolphin (Stenella clymene). PLoS ONE 2014; 9: e83645.

Aynsley, C. L. 2017. Bottlenose Dolphins (Tursiops truncatus) in North-East England: a Preliminary Investigation into a Population Beyond the Southern Extreme of its Range. MSc thesis, Newcastle University, Newcastle, UK.

Caballero, S., Jackson, J., Mignucci-Giannoni, A. A., Barrios-Garrido, H., Beltrán-Pedreros, S., Montiel-Villalobos, M. G., Robertson, K. M., Baker, C. S. 2008. Molecular systematics of South American dolphins Sotalia: sister taxa determination and phylogenetic relationships, with insights into a multilocus phylogeny of the Delphinidae. Molecular Phylogenetics and Evolution 46, 252-268.

Faria, D. M., Steel, D., Baker, C. S., da Silva, J. M., de Meirelles, A. C. O., Souto, L. R. A., Siciliano, S., Barbosa, L. A., Secchi, E., Couto Di Tullio, J., de Oliveira, L. R., Ott, P. H. & Farro, A. P. C. 2022. Mitochondrial diversity and inter-specific phylogeny among dolphins of the genus Stenella in the Southwest Atlantic Ocean. PLoS ONE 17: e0270690.

Geisler, J. H., McGowen, M. R., Yang, G. & Gatesy, J. 2011. A supermatrix analysis of genomic, morphological, and paleontological data from crown Cetacea. BMC Evolutionary Biology 2011, 11:112

Glover, K. A., Kanda, N., Haug, T., Pastene, L. A., Øien, N., Goto, M., Seliussen, B. B. & Skaug, H. J. 2010. Migration of Antarctic minke whales to the Arctic. PLOS ONE 5 (12), e15197.

Hodgins, N. K., Steel, E. M., Dyke, K., Walters, A. E. M., Dolman, S. J., Hall, K., Neave-Webb, E., Evans, P. G. H., Bird, C., Robinson, K. P., Marwood, E. M., Foubister, R., Harrop, H., Knight, A. & Munro, K. 2024. Using citizen science to better understand Risso’s dolphin (Grampus griseus) presence in northeast Scotland and the Northern Isles. Frontiers in Conservation Science 5, 1366064.

Hoekendijk, J., Leopold, M. & Cheney, B. 2021. Bottlenose dolphins in the Netherlands come from two sides: across the North Sea and through the English Channel. Journal of the Marine Biological Association of the United Kingdom 101, 853-859.

LeDuc, R. G., Perrin, W. F. & Dizon, A. E. 1999. Phylogenetic relationships among delphinid cetaceans based on full cytochrome b sequences. Marine Mammal Science 15, 619-648.

Levesque, S., O'Donovan, J., Daly, M., Murphy, S., O'Connell, M., Jepson, P., Deaville, R., Barnett, J., & Berrow, S. D. 2021. Supply of Vertebrate Necropsy and Sample Recovery Services Merged Final Reports. Marine Institute.

McGowen, M. R. 2011. Toward the resolution of an explosive radiation – a multilocus phylogeny of oceanic dolphins (Delphinidae). Molecular Phylogenetics and Evolution 60, 345-357.

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The fossil record is a cruel and fickle mistress, and there are a vast many fossil animals for which key data on lifestyle and biology is simply not preserved, or – at least – not known. Yet…

**Caption:** images that depict tapejarids as herbivores or frugivores are few and far between, but they do exist. Inspired by my reading of Peter Wellnhofer’s 1991 book on pterosaurs, I once chose to depict *Tapejara* as a leaf eater; the image at right – from issue 95 of the Orbis *Dinosaurs!* partwork (published 1994) – shows *Tapejara* as a frugivorous fruitbat/hornbill mashup. Images: Darren Naish; © Robin Bouttell.

How did the toothless azhdarchoid pterosaurs of the Cretaceous make a living? Azhdarchoids include mid-sized, relatively short-jawed groups (‘mid-sized’ = wingspans of 1.5-2m) – I’m thinking of the tapejarids – as well as much longer-jawed, mostly larger (wingspans 4-10m) groups, like the fantastic and rightfully famous azhdarchids. It is mandatory to your understanding of this article that you pay attention to the difference between azhdarchids and azhdarchoids, by the way.

**Caption:** this reconstructed (model) skeleton of the Brazilian tapejarid *Tupandactylus imperator*, formerly on show at the Museu Nacional, Rio de Janeiro (tragically ruined by fire in 2018), shows the deep rostrum, pointed jaw tips and remarkable bony crests (on the snout, rear of the skull, and lower jaw) typical of this azhdarchoid pterosaur group. Image: Darren Naish.

Tapejarids as omnivorous ‘hornbill pterosaurs’. Almost immediately after their 1989 recognition, experts drew attention to an approximate similarity between tapejarids and hornbills. And thus was born the hypothesis that tapejarids might have been frugivorous (Wellnhofer & Kellner 1991). Later authors built on this, and by the time I wrote a book chapter dedicated to this pterosaur group in 2010, the hypothesis of tapejarid frugivory or herbivory was widely considered a reasonable one.

Yes, a book chapter; one that to this day remains unpublished. Back in the 2000s, a team of editors who shall remain nameless made efforts to get a giant, pterosaur-themed textbook off the ground. Several authors – myself among them – submitted manuscripts. But alas, the book was destined never to appear. Months of time and effort were wasted. This happens a lot in academic publishing and it (partly) explains why some of us are so weirdly reticent about committing to new projects, though there are other reasons too, for sure. I’m just lazy, for example. Here, shared publicly for the first time, is an edited version of part of the tapejarid palaeobiology section from my aforementioned and ill-fated, unpublished book chapter…

**Caption:** once upon a time, this was going to happen. Even the publishers (Cambridge University Press) were taking it seriously enough to put this ad online.

At the time of writing, no detailed or empirical work on tapejarid palaeobiology has been published, with the exception of an abstract by Meijer et al. (2007). Several different ecological roles have been imagined for tapejarids. Wellnhofer and Kellner (1991) regarded it as most likely that tapejarids were frugivores: the short, narrow head might, they speculated, have been well suited for arboreal foraging, and they noted that pointed bill tips appear “a perfect tool for picking or plucking fleshy fruits” (Wellnhofer and Kellner 1991, p. 103). A superficial similarity with the (mostly) frugivorous hornbills was also suggested to support frugivory, as was the relatively small wingspan and body size of Tapejara wellnhoferi. Wellnhofer and Kellner (1991) noted that Fleming and Lips (1991) had hypothesized the existence of frugivory in pterosaurs given the apparent ecological requirement for dispersers of angiosperm fruits and seeds during the Cretaceous. The hypothesis of frugivory for tapejarids was also mentioned by Unwin (2006, p. 99).

**Caption:** tapejarid skulls aren’t quite like those of any living animal group, but they resemble those of hornbills more than those of anything else. In the two skulls here (*Tupandactylus navigans* at left, *Tu. imperator* at right), the orbit (eye socket) is the small, subcircular opening at the rear, way outsized by the massive antorbital fenestra. Both of these specimens show how the bony crests were continuous with giant, sheet-like soft tissue extensions. Images: Mark Witton; Darren Naish.

Morphometric and comparative work on tapejarid crania lend support to Wellnhofer and Kellner’s (1991) hypothesis that these pterosaurs were terrestrial omnivores or herbivores. Meijer et al. (2007) used multivariate morphometric techniques to analyse jaw structure in Tapejara wellnhoferi and concluded that it could not generate enough force to crack seeds or other hard objects; however, other kinds of herbivory could not be excluded. As described above [in a descriptive section not included here], the retroarticular process on the tapejarid mandible is unusual in being proportionally large, and in having a sloping dorsal surface. Among birds, a similarly shaped retroarticular process is present in some hornbills. Contrary to expectations, this does not support an unusual large depressor mandibulae compared to that of related birds (hoopoes and wood-hoopoes); instead, the ventral part of the pterygoideus ventralis lateralis, associated with the posteroventral part of the jaw, is particularly bulky in hornbills (Burton 1984). This is combined with a gently decurved, laterally compressed rostrum and a heavily ossified palate (the so-called ‘doubly desmognathous’ condition of Beddard (1898)) where the palatines and other elements form a robust, unfenestrated sheet anterior to the orbits. While the sequence of transformations leading to the evolution of the tapejarid palate within Pterosauria were altogether different from those that culminated in the doubly desmognathous palate of hornbills, the unfenestrated palatal sheet present in Tapejara wellnhoferi (and probably other tapejarids as well) is superficially much like a hornbill palate.

**Caption:** I’ve been bitten by several hornbill species in my time, albeit never with malicious intent. This is a captive Trumpeter hornbill *Bycanistes bucinator* photographed in 2012, a mid-sized African species. Hornbills like this differ from tapejarids in having a curved bill, but this isn’t the case in all of them; the two groups are similar in skull proportions and in being highly pneumatic in the rostrum. Image: Darren Naish.

In conclusion, tapejarid skull and lower jaw shape, and inferred wing loadings and wing shape, suggest an ecology and lifestyle most similar to that of scansorial, omnivorous birds such as hornbills, as proposed by Wellnhofer and Kellner (1991).

A different generic name was used in the book chapter, but we don’t need to worry about that.

So, there we have it: Naish (unpublished) supported frugivory – or, at least, omnivory involving frugivory – in tapejarids. And here’s where this becomes newsworthy, because an interesting publication just out in the pterosaur literature reports the discovery of phytoliths (microscopic mineral grains that some plants embed within their tissues) and gastroliths (swallowed stones used to aid digestion) within the gut of the Chinese tapejarid Sinopterus (Jiang et al. 2025). This is (so far as I can tell from what’s been released so far) good evidence for herbivory of some sort in Sinopterus. Wow – hypothesis confirmed!

**Caption:** montage from Jiang *et al*. (2025) showing preserved stomach contents of the tapejarid *Sinopterus*, revealing gastroliths and phytoliths, and thus evidence for herbivory of some sort. I can’t say much more at this stage since the paper has only been released as a preprint that I can’t access. Image: Jiang *et al*. (2025).

Azhdarchids as ‘giraffized’ herbivores. Inspired by this discovery, the very excellent artist and purveyor of… interesting ideas Hodari Nundu recently posed the following interesting question on social media: “what evidence do we have to confidently say azhdarchids were predators and not a sort of giraffized version of vegetarian tapejarids?”. Oh ho ho, a challenge! I will note to start with that the whimsical linking of azhdarchids with giraffes has happened before, and I must avoid elaborating on it here.

**Caption:** it’s a most curious and interesting thing that azhdarchids (and similar azhdarchoids) acquired superficially artiodactyl-like proportions, albeit with wings and a very different skull. And thus, we find justification for the flightless, superficially giraffe-like pterosaurs of Spec Zoo, right? Follow the links below if you’re interested. Images: © Steve Holden/Dougal Dixon; Darren Naish.

That aside, here are my thoughts on this matter. Imagine we live in a world where any and all direct data on azhdarchoid diet is absent… as we did until a few weeks ago (Jiang et al. 2025). At that time, the only thing we had to go on when it comes to azhdarchoid ecology and diet is form-function correlation: the idea that we can use the shape of an animal and its body parts to infer its natural history, mostly via analogy with living animals.

**Caption:** views have differed on whether hornbills were ancestrally mostly frugivorous and then gave rise to the mostly predatory ground hornbills, or whether mostly predatory ancestors gave rise to the mostly-frugivores. Either way, the differences between the two sorts aren’t that profound: they’re certainly less different than azhdarchids are from tapejarids. Here are captive Southern ground hornbills *Bucorvus leadbeateri*. Images: Darren Naish.

As discussed above, tapejarids are sufficiently similar in jaw and skull shape for us to hypothesize that they might have been omnivorous and partly or mostly frugivorous, and perhaps similar in feeding habits to birds like hornbills. Azhdarchids are different from tapejarids in having radically long, narrow, spear-like jaws. If we’re playing that ‘form-function correlation’ game, azhdarchids most recall storks and ground hornbills (Witton & Naish 2008). Ergo, we should – in the absence of evidence to the contrary – hypothesize that they were similar to storks and/or ground hornbills in feeding habits and diet.

**Caption:** there will come a time when I’ll stop talking about the impact of, and publicity associated with, the Witton & Naish 2008 ‘terrestrial stalking’ paper, but that time is not today. Here are screengrabs showing a few of the treatments the story was given by the great British press. Cor blimey, wot a whopper, *Gary Lineker’s woke pet donkey stole my garden shed*, and so on.

Readers aware of the literature on azhdarchids will know that this is exactly what we have been hypothesizing at least since Mark Witton and I published our initial ‘terrestrial stalking’ paper back in the 2000s (Witton & Naish 2008). While we mostly view these animals as predatory – as faunivorous or animalivorous (yes, these terms really are in use and preferable to ‘carnivorous’ when it comes to a diet involving small animals) – we’ve noted that a generalist, omnivorous diet is conceivable. Here’s what Mark and I said in a 2015 study…

We interpreted the elongate necks, hypertrophied, stork-like rostra, and distinctive limbs of azhdarchids as adaptations to “terrestrial stalking”, a lifestyle akin to that of modern ground hornbills and large stork species in which small foodstuffs (e.g., fruit, carrion, invertebrates, and small vertebrates) are procured during sustained periods of terrestrial foraging. (Witton & Naish 2015, pp. 651-652)

**Caption:** numerous aspects of azhdarchid anatomy provide support for the view that these animals were terrestrial stalkers that picked up animal prey from the ground; the hypothesis is different from most of those proposed for pterosaurs in that it isn’t based on just one perceived trait or adaptation. Image: **Witton & Naish (2015)**.

Note the mention of fruit. Yes, it’s conceivable that a hungry azhdarchid encountering a tree laden with fruit could consume some quantity of it, just as some mostly predatory animals will do today. But, otherwise: we think that mostly predatory is the lifestyle we should be imagining. Here are some additional points that might show why this view is preferable to a more herbivorous one…

The long, pointed azhdarchid rostrum does not appear suiting to the snipping, breaking or processing of plant parts. More study is needed, but a specialized herbivorous lifestyle would surely involve the evolution of shorter jaws with a shorter level arm, as is the case in tapejarids.
Azhdarchids appear to have been habitat generalists, with taxa known from deserts and semideserts as well as regions dominated by arid savannah, parkland, open woodland and subtropical riparian forest, and there are taxa of coastal habitats too. Given that they were likely good at covering distance (I’m not convinced by the argument that they were burst-fliers and more like bustards than storks), it might be that their presence in any one habitat does not necessarily require that they had to make a living there. However, the impression I get from their distribution is more consistent with that of generalist predators, able to exploit animal prey of diverse sort. They aren’t consistently associated with places where we have good evidence for ecologically ‘reliable’ vegetation.

Caption: variation in neck proportions and skull shape in particular show that not all azhdarchids were alike, a point made by numerous authors at this point. The remarkably long and slender neck of Arambourgiania, shown here, suggests a lifestyle, ecology and diet quite different from that of azhdarchids with a shorter, more robust neck and skull, for example. Image: © Mark Witton.
Specialized herbivores have relatively large guts and other adaptations for the processing and fermenting of plant material. Azhdarchids have relatively small bodies for their size and all indications are that their carrying capacity for food was low, which is more in keeping with carnivory than a diet regularly involving plants. I and others are working on a project directly relevant to this issue right now.

My main conclusion on azhdarchids is that they were indeed predominantly predatory, and sufficient generalized that they could take advantage of all kinds of small animals, from arthropods and fish to small dinosaurs, other pterosaurs and other reptiles (Witton & Naish 2008, 2015, Naish & Witton 2017). As noted earlier, the idea is already out there (Witton & Naish 2015) that they may well have exploited carrion and some plant foods in the same way that their extant analogues do. We should also always keep in mind that this is a large enough group for there to be all kinds of species-level peculiarities that may well counter the trend otherwise typical for the group. But that, I argue, is where we’re at right now. Study relevant to the issues covered here is underway, and – as ever – we await more exciting news on pterosaur biology and lifestyle.

For previous TetZoo articles on azhdarchids and other pterosaurs, see…

Tiny pterosaurs and pac-man frogs from hell, March 2008
Come back Lank, (nearly) all is forgiven, September 2008
Daisy’s Isle of Wight Dragon and why China has what Europe does not, March 2013
In Rio for the 2013 International Symposium on Pterosaurs, May 2013
Quetzalcoatlus: the evil, pin-headed, toothy nightmare monster that wants to eat your soul, August 2013
Were azhdarchid pterosaurs really terrestrial stalkers? The evidence says yes, yes they (probably) were, December 2013
Some Azhdarchid Pterosaurs Were Robust-Necked Top-Tier Predators, February 2017
Postcranial Palaeoneurology and the Lifestyles of Pterosaurs, August 2018
Did Dinosaurs and Pterosaurs 'Glow'? Extinct Archosaurs and the Capacity for Photoluminescent Visual Displays, March 2020
New Perspectives on Pterosaur Palaeobiology, the TetZoo Review, September 2020
Baby Pterosaurs Were Excellent Fliers and Occupied Different Niches From Their Parents, July 2021
The Quetz Monograph Lives and Other News on Azhdarchid Pterosaurs, December 2021
Azhdarchid Progress, a Personal View, October 2024

If you enjoyed this article and want to assist me in what I do — and see behind-the-scenes stuff in development — please support me at patreon. Click anywhere on this text in bold.

Refs - -

Beddard, F. E. 1898. The Structure and Classification of Birds. Longmans, Green, London.

Burton, P. J. K. 1984. Anatomy and evolution of the feeding apparatus in the avian orders Coraciiformes and Piciformes. Bulletin of the British Museum of Natural History (Zoology) 47, 331-443.

Fleming, T. H. & Lipps, K. R. 1991. Angiosperm endozoochory: were pterosaurs Cretaceous seed dispersers? The American Naturalist 138, 1058-1065.

Jiang, S., Zhang, X., Wu, Y., Zheng, M., Kellner, A. W. A. & Wang, X. 2025. First occurrence of phytoliths in pterosaurs—evidence for herbivory. Science Bulletin doi:10.1016/j.scib.2025.06.040

Meijer, H. J. M., Van der Meij, M. M. E., van Waveren, I. & Veldmeijer, A. J. 2007. Linking skull morphology to feeding in Tapejaridae: adaptations to frugivory in Tapejara wellnhoferi. In Hone, D. (ed.) Flugsaurier: the Wellnhofer Pterosaur Meeting. Bavarian State Collection for Palaeontology, unpaginated.

Naish, D. & Witton, M. P. 2017. Neck biomechanics indicate that giant Transylvanian azhdarchid pterosaurs were short-necked arch predators. PeerJ 5: e2908.

Unwin, D. M. 2006. The Pterosaurs From Deep Time. Pi Press, New York.

Wellnhofer, P. & Kellner, A. W. A. 1991. The skull of Tapejara wellnhoferi Kellner (Reptilia, Pterosauria) from the Lower Cretaceous Santana Formation of the Araripe Basin, northeastern Brazil. Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historische Geologie 31, 89-106.

Witton, M. P. & Naish, D. 2008. A reappraisal of azhdarchid pterosaur functional morphology and paleoecology. PLoS ONE 3 (5): e2271.

Witton, M. P. & Naish, D. 2015. Azhdarchid pterosaurs: water-trawling pelican mimics or “terrestrial stalkers”? Acta Palaeontologica Polonica 60, 651-660.

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As a regular reader of Tetrapod Zoology, you will know I’m sure that I made some effort in 2024 to rescue ruined squamate-themed articles from vers 2 and 3 of the blog…

**Caption:** I photograph all the slow-worms I see, and here are a few encountered here in southern England during the 2010s and 20s. The deceased individual at left, found in a churchyard while I was on a bat survey, was a big, robust male, and when complete would have been quite large. How large? I can’t reliably say seeing as its posterior half was missing. Images: Darren Naish.

For 2025, events have not allowed much progress, and I’m way behind on my plans, as I am at all times. Here’s your regular reminder that you can help me spend more time generating content for the blog by supporting me at patreon. But today I want to rescue another old squamate-themed Tet Zoo article, this time on one of my favourite animals: Anguis fragilis. The article first appeared here in 2007 and has been updated where appropriate.

The slow-worms. I’ve returned several times on Tetrapod Zoology to the European common slow-worm Anguis fragilis, a legless anguid lizard traditionally thought (read on!) to occur across Europe and Asia as far east as western Siberia. I find slow-worms very charismatic animals. Part of the appeal might be that they are easy to find in the places where I’ve lived; part of it might be that we Brits have such a poor reptile fauna that we hold those few species we do have in special regard; and part of it might be that they’re really cute and cool to look at.

**Caption:** slow-worms are part of the anguimorph lizard group Anguidae, and indeed are the ‘type’ genus for this group. Here’s a montage of extant anguids, using the brilliant 1983 paintings of Alan Male (I’ve used this same montage in previous Tet Zoo anguid articles). The animals shown here are Sheltopusik *Pseudopus apodus* (at top), then the brightly coloured galliwasp *Diploglossus*, the alligator lizard *Gerrhonotus* at lower left, and *Anguis* itself at lower right. Images: Alan Male, from Whitfield (1983).

Slow-worms, and their close relatives within Anguidae, are relatively intelligent by lizard standards and captive individuals exhibit behaviours suggesting that they learn to recognise their keepers. They’re long-lived: a captive slow-worm at Copenhagen lived for 54 years. Until recently, experts recognised just two slow-worm species: A. fragilis and A. cephallonica of the Peloponnese and Ionian islands.

However, 21st century molecular studies have led to the recognition of divergences within this group that occurred way back… as in, during the Miocene (so, around 25 million years ago or more). Lineages that old, in tetrapods, are surely separate species, and these discoveries are mostly responsible for the view that multiple species are present, hence the recent recognition of the Eastern slow-worm A. colchica of Russia and Georgia, Greek slow-worm A. graeca and Italian slow-worm A. veronensis (Gvoždík et al. 2010, 2013). Thus, we now have five extant slow-worm species. Populations from the more eastern part of slow-worm range are relatively understudied, so it’s possible that east Eurasian animals currently labelled as A. fragilis might turn out to be something else, but let’s just see.

**Caption:** slow-worm phylogeny, from Gvoždík *et al*. (2013), showing the revised taxonomy they suggest based on this phylogenetic structure. Peloponnese and Italian slow-worms form a clade that’s the sister-group to the remaining taxa. The divergences between the main lineages here appear to have occurred 25 million years ago or more. Image: Gvoždík *et al*. (2013).

  **Caption:** map showing the geographic origins of the slow-worms analysed by Gvoždík *et al*. (2013) for their phylogenetic study. The assumption until recently was that slow-worms from throughout the range of these animals (the brown area) belong to *Anguis fragilis*, but there’s now doubt about this: *A. fragilis* is widespread in the west, but what about all unsampled regions from the east? The Scandinavian animals should be *A. fragilis*, and some authors have shown all the animals in the east to be *A. colchica* (Speybroeck *et al*. 2016), albeit sharing a fuzzy boundary (running north-south through Poland, and also through Czechia, Lithuania, Latvia and Estonia) with *A. fragilis*. Ireland is shown here as devoid of slow-worms but we now know that that’s not right: they’re present, and seem to owe their presence to introduction by people. Image: Gvoždík *et al*. (2013).

An aside worth mentioning here is that these multiple species are very hard (albeit not impossible, based on scale counts and so on) to distinguish morphologically, and you basically need to know their geographic origins in order to work out which species they might be (e.g., Renet et al. 2018). Some people think that this is fine and that morphology isn’t destiny (especially so in a cryptozoic group of lizards where olfaction is more important than what you look like).

And some other people think that we’ve gone too far in splitting lineages like this and that we don’t need to be giving different species names to each new genetic lineage we document. This is, as you likely know, one of the biggest areas of dispute within 21st century biology and is usually framed as a debate about taxonomy. It’s not about taxonomy (the naming of things) alone though, since it also concerns how we interpret phylogeny, and how we interpret molecular data.

Another aside: as is so often the case with vernacular names, there’s considerable ambiguity on how the common name of this group of lizards should be written. Is it ‘slow worm’, ‘slow-worm’ or ‘slowworm’? Given that these animals aren’t, actually, worms, my thinking is that ‘slow worm’ is wrong. Of the other two options, I dislike both; the hyphen is ugly, but so is the ‘double w’ in ‘slowworm’. Herpetologists who’ve written about these animals have preferred ‘slow worm’ (e.g., Arnold et al. 1978, Inns 2009, Speybroeck et al. 2016) or ‘slow-worm’ (e.g., Knight 1965, Laňka & Vít 1986, Langton 1989, Beebee & Griffiths 2000, Beebee 2013), with the latter winning out by number (I’ve only seen one published use of ‘slowworm’: Hvass 1972). For that reason, slow-worm is the term I’m using here (and will do so in future, when I remember).

**Caption:** many parts of the world are home to reptile species that are but covered perhaps once, maybe twice, in the literature. Western Europe’s reptile species have been written about on numerous occasions; here are a selection of books that cover slow-worms, albeit often relatively fleetingly. Image: Darren Naish.

Some slow-worms are big. One thing that I find particularly interesting about slow-worms is the question of how big they get. I find this subject interesting because – of all the slow-worms I’ve seen (and I’m now talking only of A. fragilis) – none have ever been longer than 25 cm. Other British people with experience of A. fragilis talk of individuals of 25-30 cm as exceptionally big, so I’m not alone. Despite this, some slow-worms well exceed these lengths: most texts on these lizards list the world record as a male 48.9 cm, discovered in Portsmouth, southern England (Fairfax 1965) but it now seems that the true record-holders are from continental Europe. In 2012, Wolfgang Böhme reported a monstrous German specimen 57.5 cm long (Böhme 2012), but even this was exceeded by a Croatian giant 60.7 cm long (Zadravec & Golub 2018).

Particularly big slow-worms have been reported from Flat Holm and Steep Holm, two small islands in the Bristol Channel. In 1975, British herpetologist Tony Phelps discovered a Steep Holm slow-worm that was 45 cm long, estimated by him to be between 60 and 70 years old. In 1984, an individual 39 cm long was photographed on Steep Holm by slow-worm expert Nick Smith but, because it had a regenerated tail, its original length (i.e., with its intact original tail) would have been about 53 cm.

**Caption:** it might be obvious how Steep Holm got its name. Muntjac deer used to live on it (maybe they still do) and some died by falling off the sides. I’ve never visited, but I’ve viewed it from afar on several occasions. This photo was taken from Weston-super-Mare in August 2020. Image: Darren Naish.

Back when the biggest slow-worms on record were English, it seemed that an ‘island effect’ was at play in the development of large individuals (Portsmouth is technically an island, albeit one only weakly differentiated from the English mainland). But now that we have very large individuals from widely separated places in continental Europe, it seems that these animals have the potential to become large wherever they occur.

Giant slow-worms are Sheltopusik-like. As documented more than once at his blog – Bestiarium – my friend Markus Bühler discovered a particularly big male slow-worm in Germany. It was dead, perhaps as a result of ingesting poisoned prey, and measured 48 cm.

**Caption:** extremely large, and sadly deceased, male slow-worm found in Germany by Markus Bühler in 2006. This specimen was preserved and properly measured, and found to be 48 cm in total length. Unlike snakes, legless anguids have long tails: with a bit of imagination you can work out where the body-tail junction is, and note how slender the tail is relative to the thicker part of the body. Image: Markus Bühler, used with permission.

What’s weird is that some of these especially big slow-worms look quite different from the smaller animals I’m used to. Superficially, what with their bulkier proportions and more clearly demarcated scales, they look rather more like the European glass lizard or Sheltopusik Pseudopus apodus than do typical British slow-worms. The Sheltopusik sometimes gets described as looking like a particularly big slow-worm, the biggest individuals reaching 1.4 m. It occurs in shrubland and steppe habitats in south-east Europe, part of the Middle East and western Asia as far east as Kazakhstan. In the past, it was often included in Ophisaurus*, a genus whose numerous species occur across Asia as well as North America (though… there are competing views on the taxonomy of these animals. I’ll come back to this matter in time).

Slow-worms and glass lizards don’t just possess scales: they’re encased in subrectangular osteoderms. These are highly distinctive as fossils and help explain why the group has such a good fossil record.

Caption: the Sheltopusik does look somewhat like a gigantic slow-worm, but it has a pluck and charm all its own too. Their eyes in particular are striking and make them look very different from snakes (something we’ve discussed here at Tet Zoo before). A consequence of large size is that the skin has more of a ‘plated’ look than it does in slow-worms, the osteoderms appearing more defined and obvious. These photos were taken in Corfu. Images: Markus Bühler, used with permission.

What makes the similarity between big slow-worms and the Sheltopusik especially interesting – in my mind – is that some phylogenetic studies find the Sheltopusik to be the sister-taxon to Anguis (Macey et al. 1999). This makes me wonder if (a) slow-worms (maybe of all species) have the potential to become big and gnarly in appearance like the Sheltopusik, but that this opportunity ordinarily doesn’t allow, or (b) that the relatively small, relatively smooth-scaled and dainty slow-worms of places like the UK represent an unusual, presumably geologically young, clade, with the big, Sheltopusik-like ones being more typical, or phylogenetically ‘basal’, within the slow-worm clade. But that can’t be right in view of the slow-worm phylogeny shown above, where small, dainty British slow-worms are not obviously geologically younger or ‘more derived’ (whatever that means these days…) than other Anguis lineages.

I’ll finish by saying that we still know surprisingly little about slow-worms, despite their familiarity in a scientifically well-studied region. Questions remain about their diet and ecology (they seem to have a tight association with ants. Are they predators of ants or their larvae, or is something else going on here?), and what, exactly, is going on with their distribution (they seem widely distributed in suburban and urban environments across their range – are they good at dispersing? If so, how?).

**Caption:** I had the impression that slow-worms have been covered on an unusually high number of occasions here, but it turns out that I’ve only written about them three times or so. Given that Tet Zoo is approaching its 20th year of operation, it shouldn’t be surprising that certain familiar species have been covered more than once. These two articles are from ver 1 (2006) and 2 (2007), respectively.

We will return to them in time, I’m sure. Plus, I still have a great many other anguid lizards to write about. My thanks to Markus Bühler and Tobias Möser for their kind assistance with this article.

For previous Tet Zoo articles on squamates, see…

In Quest of Anguids, May 2006
Pompey and Steepo, the World-Record-Holding Champion Slow-Worms, May 2007
Cambodia: now with dibamids!, May 2011
The Tet Zoo Guide to Mastigures, August 2018
The Remarkable Basilisks, May 2023
Do Lizards Really Have ‘Mite Pockets’?, March 2024
Meeting Lake Zacapu’s Garter Snake, March 2024
Ray Hoser, Number 1 Taxonomic Vandal, May 2024
The Mysterious Dibamids, May 2024
Side-Stabbing Stiletto Snakes, Redux for 2024, May 2024
The Rehabilitation of Günther’s Black Cameroonian Snake, May 2024
Ikaheka and Other ‘Palatine Draggers’, Cryptozoic Elapid Snakes of Melanesia, June 2024
Arboreal Alligator Lizards of Mesoamerica... and Beyond!, July 2024
Grayiids: Africa Has Water Snakes Too, July 2024
Smaug, Ouroborus, Flat Lizards and More, August 2024
Leiosaurid Lizards: South America, Land of Iguanians, November 2024
What's With All These New Chameleon Names? Chameleons, Part 1, November 2024
Racerunner Lizards of the World Unite, New for 2025!, April 2025

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for Tet Zoo and the more productive I can be on those long-overdue book projects. Thanks!

Refs – -

Arnold, E. N. A., Burton, J. A. & Ovenden, D. W. 1992. Reptiles and Amphibians of Britain and Europe. Collins, London.

Beebee, T. J. C. 2013. Amphibians and Reptiles. Pelagic Publishing, Exeter.

Beebee, T. & Griffiths, R. 2000. Amphibians and Reptiles. HarperCollins, London.

Böhme, W. 2012. A record-sized specimen of the western slow worm (Anguis fragilis). Zeitschrift für Feldherpetologie 19, 117-118.

Fairfax, R. A. 1965. Very large English slow-worm. British Journal of Herpetology 2, 229.

Gvoždík, V., Benkovský, N., Crottini, A., Bellati, A., Moravec, J. Romano, A., Sacchi, R. & Jandzik, D. 2013. An ancient lineage of slow worms, genus Anguis (Squamata: Anguidae), survived in the Italian Peninsula. Molecular Phylogenetics and Evolution 69, 1077-1092.

Gvoždík, V., Jandzik, D., Lymberakis, P., Jablonski, D. & Moravec, J. 2010. Slow worm, Anguis fragilis (Reptilia: Anguidae) as a species complex: genetic structure reveals deep divergences. Molecular Phylogenetics and Evolution 55, 460-472.

Hvass, H. 1972. Reptiles and Amphibians in Colour. Blandford Press, London.

Inns, H. 2009. Britain’s Reptiles and Amphibians. WILDGuides Ltd, Old Basing, Hampshire.

Knight, M. 1965. Reptiles in Britain. Brockhampton Press, Leicester.

Langton, T. 1989. Snakes & Lizards. Whittet Books, London.

Laňka, V. & Vít, Z. 1986. Amphibians and Reptiles. Hamlyn, Twickenham.

Macey, J. R., Schulte, J. A., Larson, A., Tuniyev, B. S., Orlov, N. & Papenfuss, T. J. 1999. Molecular phylogenetics, tRNA evolution and historical biogeography in anguid lizards and related taxonomic families. Molecular Phylogenetics and Evolution 12, 250-272.

Renet, J., Lucente, D., Delaugerre, M., Gerriet, O., Grégory, D., Abbat-Tista, C. & Cimmaruta, R. 2018. Discovery of an Italian slow worm (Anguis veronensis Pollini, 1818) population on a Western Mediterranean Island confirmed by genetic analysis. Acta Herpetologica 13, 165-169.

Speybroeck, J., Beukema, W., Bok, B. & Van Der Voort, J. 2016. Field Guide to the Amphibians & Reptiles of Britain and Europe. Bloomsbury, London.

Whitfield, P. 1983. Reptiles and Amphibians: An Authoritative and Illustrated Guide. Longman Group Ltd, Harlow, UK.

Zadravec, M. & Golub, A. 2018. Longer than the longest - two record-breaking specimens of Anguis fragilis from north-western Croatia. Zeitschrift Für Feldherpetologie 25, 105-107.

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Once more, it’s time to look at armadillos, both at their diversity and at some aspects of their evolutionary history…

**Caption:** the evolution of tiny burrowing forms (look for the fairy armadillo at lower left) and giant, heavily armoured, herbivorous forms (like the giant glyptodont *Doedicurus*) means that armadillos exhibit a really profound, and rarely commented on, variation in size. Images: James Dana, Hermann Burmeister; both in public domain.

If you read the previous article – which you really should have done by now, come on – you’ll have noticed my repeated emphasis of the fact that the mostly large, mostly herbivorous, entirely extinct glyptodonts (and a few related groups of large-bodied, armoured xenarthrans) are deeply nested within the armadillo radiation. They are armadillos, not a separate group. That’s quite the surprise relative to convention, because the ‘standard view’ – repeated in all classic and even some modern texts on fossil mammal diversity (e.g., Prothero 2017) – is that glyptodonts and armadillos share a long-ago ancestry, and that the two groups are close… not that one is nested within the other.

If you pay attention to discoveries relevant to the world of fossil mammals, you might have heard that the placing of glyptodonts within armadillos is a discovery we owe to molecular phylogenetics. In 2016, Frédéric Delsuc and colleagues published the results of a molecular analysis of armadillo phylogeny, incorporating palaeogenomic data from the Pleistocene-Holocene giant glyptodont Doedicurus (Delsuc et al. 2016). Their primary take-home was that glyptodonts “are not … a sister group [to armadillos], but a subfamily”, “firmly within the family tree of modern armadillos”, according to this article at BBC News. More recent molecular studies have provided additional support for this view (Brambilla et al. 2025).

Don’t get me wrong: none of the discussion or reporting surrounding this study has been technically wrong, nor notably misleading. But it’s certainly created the impression that glyptodonts were universally regarded as the ‘armadillo sister group’ prior to this gene-based work.

**Caption:** armadillo skulls are highly variable, and contain a ton of anatomical information that can be analysed within a phylogenetic context. If you do analyse this data and include sufficient taxa… what do you find? Not enough people have paid attention to the article shown at left (Gaudin & Wible 2006); read on. The skull diagrams here, from Gaudin & Wible (2006), are (clockwise from upper left) *Dasypus*, *Tolypeutes*, *Proeutatus*, *Cabassous*.

We need to push back on this a bit: the finding that glyptodonts are crown-armadillos, surrounded by extant lineages, doesn’t ‘belong’ to the world of molecular phylogenetics, and I say this based on statements made by palaeontologists interested in anatomy (and not working on palaeogenomics). George Engelmann (1978, 1985) was first to imply this, initially in his 1978 PhD thesis, since he argued that the extinct eutatines (conventionally regarded as euphractines, and hence close to the living hairy armadillos and kin: see the previous article) were closer to glyptodonts than to other armadillos… a discovery which implied paraphyly among ‘conventional’ armadillos.

Building on this work, Gaudin & Wible (2006) went further, and argued that craniodental data showed that glyptodonts (and the also extinct pampatheres) were closer to living euphractines than to both eutatines and the living long-nosed armadillos. In their 2011 study on the Late Oligocene tolypeutine Kuntinaru of Bolivia, Billet et al. (2011) reported similar findings.

**Caption:** an armadillo cladogram published by Tim Gaudin and John Wible in 2006, and emphasizing the fact that phylogenetic hypotheses whereby glyptodonts (and pampatheres and their kin) are nested within crown-armadillos are not ‘owned’ by molecular biologists. Anatomists discovered this first! Images: three-banded armadillo in the public domain (**original here**); *Eutatus* from Krmpotic *et al*. (2009); *Macroeuphractus* from Vizcaíno & De Iuliis (2003); *Euphractus* and glyptodont by Darren Naish.

I remember reading Gaudin & Wible (2006) when it was new and thinking it so heterodox that it surely couldn’t be right. I mean: we know that glyptodonts are the sister-group to living armadillos, right? Nope... glyptodonts have been considered nested within crown-armadillos since the 2000s. All of which means that it’s wrong to frame the nesting of glyptodonts within crown-armadillos as a surprising, molecular-led discovery.

**Caption:** I’ve mentioned a few times how pleased I am to own the museum-quality model of the Pliocene-Pleistocene-Holocene glyptodont *Neosclerocalyptus* shown here, created by Santiago Druetta in Córdoba, Argentina. Thanks to Rebecca Groom for printing and painting. At right, a *Glyptodon clavipes* skull photographed at the Oxford University Museum of Natural History. Note the massive jugal flange and deep and broad nose. Images: Darren Naish.

On the need for an expanded taxonomy of armadillos. I have one final thing to say on the phylogenetic position of glyptodonts, and it concerns taxonomy. In those phylogenetic schemes where glyptodonts are the sister-group to other armadillos, they’ve traditionally been given their own ‘higher order’ name, this variously being Glyptodontia, Glyptodontoidea or Glyptodonta (McKenna & Bell 1997, Vizcaíno & Bargo 1998, Gaudin & Wible 2006).

But the discovery that glyptodonts belong within crown-armadillos – and, specifically, are surrounded by euphractines and tolypeutines (Delsuc et al. 2016) – has led to the implication that they should be absorbed into Chlamyphoridae (see the discussion in this Tet Zoo article), a group formulated as if it’s a Linnaean ‘family’. How inclusive or otherwise a ‘family-level’ clade should be is wholly subjective, and there might well be some specialist workers entirely happy with the idea that all the lineages concerned (eutatines, euphractines, glyptodonts, pampatheres, fairy armadillos and tolypeutines) should indeed be lumped into a super-inclusive Chlamyphoridae.

**Caption:** a molecular armadillo phylogeny as published by Delsuc *et al*. (2016). The phylogeny is broadly consistent with other studies and other data (though don’t go thinking that all phylogenetic studies on armadillos are in agreement — oh ho ho!), but the issue I have with what’s shown here is the taxonomy… should we really be including all those lineages within a super-inclusive Chlamyphoridae? Image: **Delsuc *et al*. (2016)**.

But I’m not happy with it. I think that glyptodonts are absolutely ‘distinct enough’ (and diverse enough, and geologically old enough) to warrant ‘higher-order’ taxonomic distinction, and I think it’s helpful to have them differentiated from pampatheres (Pampatheriidae) and from the clade that includes fairy armadillos and tolypeutines. Those last two could be united within Chlamyphoridae. If the extinct eutatines and the extant euphractines are outside the glyptodont + chlamyphorid clade, they need ‘higher-order’ names too, so here’s a push to have us recognise Eutatidae and Euphractidae…. if the phylogenetic hypotheses I’m using here as a framework remain robust.

Fairy armadillos or Pichiceigos. Arguably the most specialized and unusual armadillos are the fairy armadillos, generally regarded these days as a clade (Chlamyphorinae) that shares a Late Eocene or Oligocene ancestor with tolypeutines (e.g., Billet et al. 2011, Delsuc et al. 2012, 2016, Gibb et al. 2016). Anatomical evidence previously suggested that fairy armadillos belong with euphractines – this group was termed Euphracta by Engelmann (1978, 1985) – since all are alike in the bony anatomy of the ear region, most memorably in possessing an elongate and tubular external auditory meatus (Gaudin & Wible 2006). Some modern phylogenetic work still finds this result (Herrera et al. 2017).

**Caption:** a Pink fairy armadillo taxiderm specimen and mounted skeleton, both on display at the Grant Museum of Zoology, London. It should be obvious that there’s a whole lot of weird going on here. Just about everything in the skeleton is remarkable. Check out how the rounded shield at the rear looks fused to the pelvis, the strongly reduced tail and the giant foreclaws. As for the skull… I have no idea what’s going on with those two rounded bosses that look like horns. Rad. Images: Darren Naish.

There are two extant fairy armadillo species: the Pink fairy armadillo Chlamyphorus truncatus of central Argentina, and the Greater or Chacoan fairy armadillo Chaetophractus retusus of Bolivia, Paraguay and northern Argentina. The second species has sometimes been included in Chlamyphorus and the generic name Burmeisteria Gray, 1865 has also been used for it; the two extant species belong to lineages that appear to have been separate since the Oligocene, so distinct generic status is appropriate (Delsuc et al. 2012). These animals are tiny, some adults of C. truncatus being only just over 8 cm in head and body length. They inhabit dry grasslands and sandy plains and are rapid burrowers that apparently use the subcircular armour plate covering the rear end as a shield or plug to block their burrow (Nowak 1999).

**Caption:** it turns out that the two fairy armadillo lineages are highly distinct, more so anatomically than most non-specialists realize (Greater or Chacoan fairy armadillo illustrations at right, from Hermann Burmeister’s description of the holotype). And molecular phylogenetics shows that they’ve been distinct for around 17 million years, as shown by the phylogeny (from Delsuc *et al*. 2012) at left. Images: Delsuc *et al*. 2012; public domain.

Indeed, they’ve converged on the ‘mole’ ecotype that evolved at least four times elsewhere within mammals (marsupial moles, golden moles, true moles, palaenodonts). They have reduced eyes, enlarged claws on their forelimbs, a silky pelt, a strongly reduced and mostly immobile tail, and a streamlined and flexible armour covering that includes the aforementioned posterior plate. Little is known of their ecology or behaviour for the obvious reason that they’re rarely seen alive. A lot more could be said about them but this will do for now. A few years ago, there was a non-serious effort to get them renamed ‘battle hamsters’ but it didn’t go anywhere… which is good, because it sure wouldn’t be helpful to have a group of xenarthans with the word ‘hamster’ in the name.

Remaining tolypeutine armadillos. Molecular studies find remaining tolypeutines to be a clade, with the remarkable Giant armadillo Priodontes maximus being the sister-taxon to the three-banded armadillos (Tolypeutes) and the naked-tailed armadillos (Cabassous). All three genera are united by some authors within Priodontini (Delsuc et al. 2003). Priodontes is a superstar armadillo, well known for its comparatively large size: it reaches 30 kg at least (note that all those mentions of specimens exceeding 60 kg are apparently based on overweight captive individuals). It’s also notable for its amazing hand claws and digging prowess, it probably being the most proficient and specialized digger among armadillos after the fairy armadillos (Vizcaino et al. 1999, Vizcaino & Milne 2002). The good news is that I’ve written about this species before (albeit at Tet Zoo ver 2): go here for that article.

**Caption:** this, obviously, is an illustration from a children’s book (specifically, Peter Spier’s 1971 *Gobble Growl Grunt*). I include it to emphasize the fact that the semi-bipedal tendencies and ‘claw-supported walking’ abilities of certain armadillos are widely known, and long have been, yet rarely mentioned outside of specialized literature. Image: (c) Peter Spier.

Three-banded armadillos are very special in being the only armadillos that can properly ‘enroll’ to form a sphere. The two species – Tolypeutes tricinctus of Brazil and the more widespread T. matacus – possess shell-like anterior and posterior carapacial segments that allow the limbs, head and tail to be tucked within. So effective, tight and rapid-action is this defence that they can even hop a short distance off the ground before snapping the shell shut before hitting the ground.

Like some other tolypeutines, their forelimb claws are so long and big relative to the rest of the hand that they’re ‘claw walkers’. Three-banded armadillos don’t excavate burrows but will use shelters constructed by other animals, like anteaters [UPDATE: see comments. It’s now know that they can dig their own burrows too]. They’re mostly denizens of grasslands and marshes; ants, termites and beetle larvae form most of their diet but they also eat fruit (Nowak 1999).

**Caption:** how, exactly, do three-banded armadillos enroll? It might be that their vertebral anatomy isn’t tremendously unusual relative to that of other xenarthrans but that the intricate, interlocking joints of the dermal skeleton are the key innovation here. Maybe I should have checked before writing this. This photo was taken at the Museum of Osteology, Oklahoma, USA. Image: Mathew Wedel, SV-POW!, **original here**.

  **Caption:** three-banded armadillos are very unusual little animals. They’re partly cute, but also partly quite disturbing. Features to look for include the intricate sculpting of the scutes, the scaliness of the ears, and the massive length of the foreclaws. Images: *T. tricinctus* at left by ChrisStubbs, CC BY-SA 3.0 (**original here**); *T. matacus* at right by Hedwig Storch, CC BY-SA 3.0 (**original here**).

Finally, naked-tailed armadillos are among the most obscure armadillos of all. They superficially resemble Priodontes but are mostly 30-40 cm in head and body length (the tail adds an extra 9-20 cm) and generally weigh between 3 and 6 kg. They’re short-snouted, have relatively big ears, and possess 10-13 mobile bands across the middle of the body. As suggested by the common name, they’re unique in having a mostly unarmoured tail. It’s long and slender and decorated with a reduced number of small, widely spaced scales.

Naked-tailed armadillos – four extant species are recognized – are among the most generalized of armadillos, variously occurring in lowlands and uplands, grassland, and riverside woodland and marsh, and they’re good at burrowing and swimming as well as running. They appear to mostly rely on ants and termites.

**Caption:** there aren’t many good photos of naked-tailed armadillos in the wild. This is a Southern naked-tailed armadillo *Cabassous unicinctus*, a species that occurs across most of Brazil and in adjacent countries across northern continental South America. Image: Ben P, CC BY 4.0 (**original here**).

And that’s where we’ll end things. This is absolutely not a thorough look at fossil armadillos, since there are several fascinating taxa – interesting anatomically and in phylogenetic position – that I’ve deliberately ignored here. So, we have to come back to these animals in time. The main function of this article was to show how glyptodonts (and other big armadillos known only as fossils) fit within armadillo phylogeny and diversity as a whole. When we revisit these animals – which we will do, reasonably soon – it will be to discuss glyptodonts alone…

For previous Tet Zoo articles on armadillos and other xenarthrans, see…

Five things you didn’t know about armadillos, June 2007
What was that skull? (on glyptodonts), May 2008
I, Priodontes, the tatuasu, September 2008
Predation and Corpse-Eating in Armadillos, September 2010
The Fate of the Woolly Long-Nosed Armadillo of Peru, April 2024
Of Zaedyus, the Pichi, May 2025
Armadillo Empire, Part 1: of Euphractines and Eutatines, June 2025

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for Tet Zoo and the more productive I can be on those long-overdue book projects. Thanks!

Refs - -

Brambilla, L., Ibarra, D. A., Barboza, M. C., Bresso, E. G., Rosano, G., Pérez, G., Straccia, P., Scian, R. D. & Brun, L. R. 2025. Mitochondrial genome of Neuryurus rudis (Xenarthra, Cingulata); contribution to phylogeny and origin of glyptodonts. Gene 936, 149059.

Delsuc, F., Gibb, G. C., Kuch, M., Billet, G., Hautier, L., Southon, J., Rouillard, J.-M., Fernicola, J. C., Vizcaíno, S. F., MacPhee, R. D. E. & Poinar, H. N. 2016. The phylogenetic affinities of the extinct glyptodonts. Current Biology 26, R155-R156.

Delsuc, F., Superina, M., Tilak, M.-K., Dousery, E. & Hassanin, A. 2012. Molecular phylogenetics unveils the ancient evolutionary origins of the enigmatic fairy armadillos. Molecular Phylogenetics and Evolution 62, 673-680.

Delsuc, F., Stanhope, M. J. & Douzery, E. J. P. 2003. Molecular systematics of armadillos (Xenarthra, Dasypodidae): contribution of maximum likelihood and Bayesian analyses of mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 28, 261-275.

Engelmann, G. 1978. The Logic of Phylogenetic Analysis and the Phylogeny of the Xenarthra. Ph.D. dissertation, Columbia University, New York.

Engelmann, G.1985. The phylogeny of the Xenarthra. In Montgomery, G. G. (ed) The Ecology and Evolution of Armadillos, Sloths, and Vermilinguas. Smithsonian Institution Press, Washington, DC, pp. 51-64.

Gaudin, T. J. & Wible, J. R. 2006. The phylogeny of living and extinct armadillos (Mammalia, Xenarthra, Cingulata): a craniodental analysis. In Carrano, M. T., Gaudin, T. J., Blob, R. W. & Wible, J. R. (eds) Amniote Paleobiology: Perspectives on the Evolution of Mammals, Birds, and Reptiles. University of Chicago Press, pp. 153-198.

Gibb, G. C., Condamine, F. L., Kuch, M., Enk, J., Moraes-Barros, N., Superina, M., Poinar, H. N. & Delsuc, F. 2016. Shotgun mitogenomics provides a reference phylogenetic framework and timescale for living xenarthrans. Molecular Biology and Evolution 33, 621-642.

Herrera, C. M. R., Powell, J. E., Esteban, G. I. & del Papa, C. 2017. A new Eocene dasypodid with caniniforms (Mammalia, Xenarthra, Cingulata) from northwest Argentina. Journal of Mammalian Evolution 24, 275-288.

Krmpotic, C. M., Carlini, A. A. & Scillato-Yané, G. J. 2009. The species of Eutatus (Mammalia, Xenarthra): assessment, morphology and climate. Quaternary International 210, 66-75.

McKenna, M. C. & Bell, S. K. 1997. Classification of Mammals: Above the Species Level. Columbia University Press, New York.

Nowak, R. M. 1999. Walker’s Mammals of the World, Sixth Edition. The Johns Hopkins University Press, Baltimore and London.

Prothero, D. R. 2017. The Princeton Field Guide of Prehistoric Mammals. Princeton University Press, Princeton and Oxford.

Vizcaíno, S. F. & Bargo, M. S. 1998. The masticatory apparatus of the armadillo Eutatus (Mammalia, Cingulata) and some allied genera: paleobiology and evolution. Paleobiology 24, 371-383.

Vizcaíno, S. F. & De Iuliis, G. 2003. Evidence for advanced carnivory in fossil armadillos (Mammalia: Xenarthra: Dasypodidae). Paleobiology 29, 123-138.

Vizcaíno, S. F., Fariña, R. A. & Mazzetta, G. 1999. Ulnar dimensions and fossoriality in armadillos and other South American mammals. Acta Theriologica 44, 309-320.

Vizcaíno, S. F. & Milne, N. 2002. Structure and function in armadillo limbs (Mammalia: Xenarthra: Dasypodidae). Journal of Zoology 257, 117-127.

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If you’ve read recent articles here, you’ll have seen the coverage I’ve been giving to armadillos…

**Caption:** a euphractine montage… though read on for ideas about what ‘euphractine’ does or does not mean. At left, skeleton of a hairy armadillo (*Chaetophractus*). At right, a Six-banded armadillo *Euphractus sexcinctus* demonstrating how easy it is for armadillos of this sort to stand bipedally. This one was photographed at Edinburgh Zoo in 2010. Images: Darren Naish.

As usual, my initial plan wasn’t to embark on some grand tour of total armadillo diversity but, rather, to cover select facts about the group, my primary aim being to discuss where glyptodonts – the mostly enormous, entirely extinct, primarily herbivorous armadillos of the American fossil record – fit within the armadillo radiation. Follow me as I cover more of armadillo diversity, all the while aiming to get in the end to the glyptodonts and what they are…

We saw in the previous article that armadillos are today considered to consist of two primary clades, namely long-nosed armadillos (Dasypodidae) and all the others (Chlamyphoridae). Current thinking, based mostly on molecular phylogenetics (read on), has it that Chlamyphoridae consists of euphractine armadillos, fairy armadillos, and tolypeutine armadillos (Delsuc et al. 2003, Gibb et al. 2016, Barasoain et al. 2020). I must add – without getting bogged down in a discussion of armadillo taxonomic history – that armadillo specialists have historically preferred to include all armadillos within a lone family (Dasypodidae), and to recognise within it two subfamilies, Dasypodinae and Euphractinae. This was still being used as of 2024 (e.g., Carlini et al. 2009, Klimeck et al. 2024). In the taxonomic system where non-dasypodid armadillos are united within Chlamyphoridae, rather than Euphractinae, the term ‘euphractine’ is used for the Euphractes lineage alone, not for the whole of the non-dasypodid clade. Gaudin & Wible (2006) used the name Euphracta for the Euphractes clade.

**Caption:** a very simplified cladogram depicting the relationships of some of the animals discussed here. Eutatines, the living six-banded, hairy armadillos and kin, *and* *Macroeuphractus* and similar forms are conventionally allied within Euphractinae. However, some of these animals appear to be closer to the pampathere + glyptodont clade than others. Images: three-banded armadillo in the public domain (**original here**); *Eutatus* from Krmpotic *et al*. (2009); *Macroeuphractus* from Vizcaíno & De Iuliis (2003); *Euphractus*, pampathere and glyptodont by Darren Naish.

With this taxonomic framework in mind… we looked previously at one of the euphractines, the Pichi Zaedyus pichiy of southern South America. It’s now time to look at remaining euphractines, and to start getting through remaining chlamyphorids in general. As we’ll see (in the next article), this ‘two family’ taxonomy is a bit problematic, and I think it should be revised.

Remaining euphractines. Ok, with the Pichi out of the way, the other extant euphractines are the Six-banded armadillo Euphractus sexcinctus and the hairy armadillos (Chaetophractus). All (including the Pichi) are robust ecological generalists that eat just about anything, are good diggers and burrowers, and occur mostly in grasslands and semi-deserts. They vary in their degree of hairiness – as you might guess, the several Chaetophractus species are the hairiest – and all have pointed ears, a head that’s subtriangular when seen from above, and a slender, fully armoured tail.

**Caption:** six-banded and hairy armadillos are omnivore-carnivores that eat carrion, arthropods of all sorts, and also catch and kill vertebrates, sometimes of surprising size. For more on this issue see the 2020 article **Predation and Corpse-Eating in Armadillos**. Images: original video that yielded screengrab no longer online; Darren Naish.

Current convention is to regard all three of the lineages here as distinct genera, but they’re sufficiently alike that some authors have combined all of them in one genus (Euphractus Wagler, 1830 wins out). At least some molecular studies suggest that the Pichi might be nested within Chaetophractus, rather than be its sister-taxon (Delsuc et al. 2016). I was interested to learn (thanks to David Howlett in the comments of the previous Tet Zoo armadillo article) that supposed Pichi specimens kept in captivity have sometimes turned out to be misidentified Screaming hairy armadillos C. vellerosus, which goes to show that even people with hands-on experience of live armadillos can confuse Zaedyus with the Chaetophractus species.

Fossil euphractines in name only, perhaps. A substantial number of fossil taxa have been regarded as euphractines. The oldest of these date to the Eocene or Early Oligocene (like Isutaetus, Parutaetus, Meteutatus and Anutaetus), a few others are from younger parts of the Oligocene (Hemiutaetus, Amblytatus and others), and others are Miocene and/or Pliocene (Vetelia, Proeuphractus, Paleuphractus and so on).

**Caption:** Early Oligocene armadillos from the Tinguiririca Fauna of Chile, both of which would conventionally have been included within the inclusive version of Euphractinae. At left, *Meteutatus tinguiririquensis*; at right, *Parutaetus chilensis*. This very nice illustration is from Carlini *et al*. (2009): no artist is credited for this illustration in the paper (unless I’ve missed it), so I assume it’s one of the authors.

These taxa are all very obscure, at least some were named for osteoderms alone, and they’re hardly ever discussed outside of the technical literature except for the very large Macroeuphractus of Pliocene Argentina. I previously covered this very interesting animal in this 2020 article devoted to carrion-eating and predation in armadillos. Macroeuphractus and a few similar taxa are sufficiently well represented in anatomical terms to win inclusion in phylogenetic studies (Gaudin & Wible 2006, Herrera et al. 2017), and results mostly indicate that they’re likely not euphractines after all, by which I mean that they’re not close relatives of Euphractus and its extant kin. Paleuphractus, Proeuphractus and Macroeuphractus were all found by Gaudin & Wible (2006) to be close to the pampathere + glyptodont clade, the main characters responsible for this position being a proportionally short snout and the detailed anatomy of the ectotympanic bulla (the bulging bony region that forms the underside of the ear region). Barasoain et al. (2021), in contrast, did find certain of these taxa to be close to Euphractus and kin.

**Caption:** a *Macroeuphractus* montage, based on the work of Vizcaíno & De Iuliis (2003). At left, the big, robust and heavily built skull of this animal (the scale bar is probably 100 mm: the paper accidentally fails to provide its length), as illustrated by Richard Lydekker in 1894. At right, speculative art provided by Vizcaíno & De Iuliis (2003) and showing *Macroeuphractus* about to break into a burrow inhabited by the chinchillid rodent *Lagostomus*. This will not end well for the rodents.

If these fossil, Euphractus-like taxa are not close relatives of Euphractus after all, it means that the features conventionally used to unite ‘euphractines’ are widespread within this section of the armadillo family tree. This wouldn’t affect the monophyly of euphractines proper. It would also mean that a series of taxa – these ‘former euphractines’, the pampatheres, and other taxa too – form a series of intermediates between glyptodonts and extant armadillos.

Eutatines and others. If it’s not already obvious, numerous fossil armadillos are known. A number are grouped together as the eutatines – yes, a word annoyingly similar to euphractines – after the relatively well-represented Eutatus of the Pliocene, Pleistocene and Holocene of Argentina, several species of which are known. Making things additionally complex is the fact that eutatines are considered within euphractines in those taxonomic systems where Euphractinae is used in the old, inclusive sense (so, certain fossil ‘euphractines’ listed above – like Meteutatus of the Eocene – are also eutatines). Eutatus was large, with a skull 26 cm long. This suggests an overall size about similar to the living Giant armadillo Priodontes maximus or maybe bigger.

**Caption:** *Eutatus* is known from very good specimens, some of which have the bulk of their armour preserved in full articulation. This skeleton of *Eutatus punctatus* is on show at Museu de Ciències Naturals de València, Spain. Image: Joanbanjo, CC BY-SA 3.0 (**original here**).

  **Caption:** ok, so a lot of fossil armadillos are known from isolated osteoderms alone. It figures that these tough, super numerous bones have high preservation potential and thus might be over-represented in the fossil record relative to other parts of the skeleton. However, there are also quite a few fossil armadillos that are extremely well preserved, with mostly intact armour. Obviously, much of the armour on this *Eutatus* specimen is reconstructed, but at least some of it is articulated. Note the long, triangular form of the head shield. Again, this is *E. punctatus* on show at Museu de Ciències Naturals de València, Spain. Image: Joanbanjo, CC BY-SA 3.0 (**original here**).

Eutatus was long-snouted, with a relatively slender lower jaw and toothless premaxilla. Its teeth have flattened crowns that are elliptical in cross-section and the jaw joint resembles that of ruminants.

All of these features indicate that Eutatus was a high-fibre herbivore, presumably eating leaves, buds and maybe grasses too (Vizcaíno & Bargo 1998). A few older armadillos conventionally classified alongside Eutatus within Eutatini – they include Proeutatus and Stenotatus of the Miocene, and Doellotatus and Ringueletia of the Pliocene – are superficially similar to Eutatus but smaller and with less prominent muscle insertion points on the jaw and cranium, so they might have been omnivorous and perhaps intermediate in ecology between armadillos like Euphractus and the strongly herbivorous Eutatus (Vizcaíno & Bargo 1998).

**Caption:** a eutatine/eutatin montage drawing on images compiled by Vizcaíno & Bargo (1998), and with the caveat that these animals don’t group together in current phylogenetic analyses. *Eutatus* is obviously bigger, long-snouted and longer-jawed than the others shown here, and with a longer toothless section at the front of the upper and lower jaws.

Variation in osteoderm anatomy present across the Eutatus species indicates variation in terms of how hairy they were, and also in how adapted they were for aridity and low or high temperature (Krmpotic et al. 2009). Yes, you can make inferences about that sort of thing from osteoderm anatomy. It’s a subject I can’t do justice here but will in another article.

If you know anything about fossil armadillos, you might be thinking from the description of Eutatus given above that it sounds superficially like a pampathere, these being a group of even larger, fully herbivorous extinct armadillos (indisputable species of which are known from the Miocene to the Holocene, and from North America as well as South America). Pampatheres have often been considered close to glyptodonts.

**Caption:** pampatheres (like *Holmesina*, here on the left) and glyptodonts (like *Glyptodon*, on the right) are going to be discussed a bit more in the next armadillo-themed article. The fact that both groups are nested *within* armadillos is now well known and today we’re in the habit of giving credit to molecular researchers for discovering this, but… nope, people working on anatomy discovered this first! That’s a non-trivial point that I’ll be bigging up in the next article. This photo was taken at the Royal Ontario Museum, Toronto. Image: Darren Naish.

Engelmann (1978, 1985) suggested that all three groups (eutatines or eutatins, pampatheres, glyptodonts) formed a clade, with eutatins being closer to glyptodonts than pampatheres. However, Gaudin & Wible (2006), Billet et al. (2011) and Herrera et al. (2017) all found so-called eutatins to be non-monophyletic: as previously noted by Vizcaíno & Bargo (1998), it turns out that Proeutatus shares unusual anatomical details with pampatheres and glyptodonts (including teeth where the crowns are worn flat and have a resistant orthodentine core), whereas Eutatus itself does not (Gaudin & Wible 2006, Billet et al. 2011). In fact, the distribution of anatomical features means that Eutatus is outside the clade that unites euphractines (in the strict, restrictive sense) with pampatheres and glyptodonts. That’s bad news if you’re a eutatin superfan, but not for everyone else because what this means is that armadillos gave rise to big-bodied, specialized herbivores at least twice.

Ok, I need to stop there… to be continued…

For previous Tet Zoo articles on armadillos and othr xenarthrans, see…

Five things you didn’t know about armadillos, June 2007
What was that skull? (on glyptodonts), May 2008
I, Priodontes, the tatuasu, September 2008
Predation and Corpse-Eating in Armadillos, September 2010
The Fate of the Woolly Long-Nosed Armadillo of Peru, April 2024
Of Zaedyus, the Pichi, May 2025

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for Tet Zoo and the more productive I can be on those long-overdue book projects. Thanks!

Refs - -

Barasoain, D., Contreras, V. H., Tomassini, R. L. & Zurita, A. E. 2020. A new pygmy armadillo (Cingulata, Euphractinae) from the late Miocene of Andean Argentina reveals an unexpected evolutionary history of the singular Prozaedyus lineage. Journal of South American Earth Sciences 100, 102589.

Barasoain, D., González Ruiz, L. R., Tomassini, R. L., Zurita, A. E., Contreras, V. H. & Montalvo, C. I. 2021. First phylogenetic analysis of the Miocene armadillo Vetelia reveals novel affinities with Tolypeutinae. Acta Palaeontologica Polonica 66, S31-S46.

Billet, G., Hautier, L., de Muizon, C. & Valentin, X. 2011. Oldest cingulate skulls provide congruence between morphological and molecular scenarios of armadillo evolution. Proceedings of the Royal Society B 278, 2791-2797.

Carlini, A. A., Ciancio, M. R., Flynn, J. J., Scillato-Yané, G. J. & Wyss, A. R. 2009. The phylogenetic and biostratigraphic significance of new armadillos (Mammalia, Xenarthra, Dasypodidae, Euphractinae) from the Tinguirirican (Early Oligocene) of Chile. Journal of Systematic Palaeontology 7, 489-503.

Engelmann, G. 1978. The Logic of Phylogenetic Analysis and the Phylogeny of the Xenarthra. Ph.D. dissertation, Columbia University, New York.

Klimeck, T. D. F., Ciancio, M. R., Sedor, F. A. & Kerber, L. 2024. Revealing the diversity of Paleogene cingulates from Brazil: a new species of Parutaetus (Euphractinae) in the Guabirotuba Formation (middle-late Eocene). Journal of Vertebrate Paleontology e2403581.

Krmpotic, C. M., Carlini, A. A. & Scillato-Yané, G. J. 2009. The species of Eutatus (Mammalia, Xenarthra): assessment, morphology and climate. Quaternary International 210, 66-75.

Vizcaíno, S. F. & Bargo, M. S. 1998. The masticatory apparatus of the armadillo Eutatus (Mammalia, Cingulata) and some allied genera: paleobiology and evolution. Paleobiology 24, 371-383.

Vizcaíno, S. F. & De Iuliis, G. 2003. Evidence for advanced carnivory in fossil armadillos (Mammalia: Xenarthra: Dasypodidae). Paleobiology 29, 123-138.

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It’s World Turtle Day, and what kind of person doesn’t love and admire turtles?!

**Caption:** I sometimes consider it kinda odd that a group of reptiles built the way they are have been so strongly associated with amphibious and aquatic habits across their history, but here we are. At left, an unusual close-up view of a captive Green turtle *Chelonia mydas*. At right, a *Pseudemys* turtle (maybe a Florida red-bellied *P. nelsoni*?) observed in the wild in Florida, in a swimming pool of all places. Images: Dave Hone, used with permission; Darren Naish.

  **Caption:** aquatic turtles at very different ends of the size spectrum. At left, a pleurodire, specifically a Roti Island snake-necked turtles *Chelodina mccordi* observed at Brighton Aquarium (sorry the snout is cut off: these animals move far too much to enable nice, clear photos). At right, a reconstruction of the gigantic Late Cretaceous North African sea turtle *Ocepechelon*. Images: Darren Naish; Joschua Knüppe.

Because I’m drowning in work, I have nothing ready to go, so here are random turtle-themed thoughts relating either to material published in the past here at Tet Zoo, or connected to content that’s been amassed for my in-prep textbook… which, yes, is still a thing and due to be finished one day. Oh, this isn’t the first time that World Turtle Day has snuck up on me by surprise – ha, the irony – since I see from Tet Zoo articles of the dim and distant past that exactly the same thing has happened before, as evidenced by…

Turtles I Have Recently Seen, June 2015

Pleurodires or side-necked turtles occur today in South America, Africa, Madagascar and Australasia and are less diverse and less widespread than their sister-group, the cryptodires. There’s sometimes the implication that pleurodires are ‘more archaic’ than cryptodires (possibly due to Northern Hemisphere chauvinism) but there’s no reason to adopt this given that both lineages are of equal age: it’s true that pleurodires are less diverse than cryptodires (in ecology, morphology, size and habitat choice) but this is less of an issue when extinct taxa are accounted for.

**Caption:** the matamata(s) is taken for granted as an odd pleurodire turtle, but it’s something very, very special. Just peer into the eyes of this surreal, highly aquatic, flanged, cryptic predator with a proboscis and remind yourself that it’s a turtle. The suggested use of plural there is a reference to the fact that there might be more than one matamata species. The photos of the captive individual appear here courtesy of Mark Hollowell; the drawing is by me. Images: Mark Hollowell (used with permission); Darren Naish.

The fossil record reveals that pleurodires were once far more widespread, exhibited more anatomical disparity, and occurred in a greater variety of habitats than they do today, especially during the Cretaceous and Paleogene. I haven’t written about pleurodire diversity in much detail, to my shame, but I did publish a whole series of articles on matamatas at Tet Zoo ver 2 back in 2010…

Matamata: turtle-y awesome to the extreme, June 2010
The familiar Matamata, known to us all since the 1700s, and its long, fat neck (matamatas part II), July 2010
“Adaptation perfected” (possibly) in a turtle’s head (matamatas part III), July 2010
Turtles that suck, turtles that blow (matamatas part IV), August 2010
Giant fossil matamata turtles (matamatas part V), April 2011

Caption: another matamata montage, showing a familiar photo of a captive one ‘snorkeling’, and captive individuals observed in various UK collections. Images: Francis Miller, from the Time-Life International 1963 volume The Reptiles; Darren Naish.

Softshell turtles or softshells (Trionychidae) are one of my favourite turtle groups and are a widely distributed lot of aquatic cryptodires. They possess a flexible proboscis (a feature we might not predict based on their osteology alone) and a soft dermal covering to the carapace that extends well beyond the shell’s bony margins. Key features include the presence of a wrinkled shell bone texture and the loss of virtually all carapacial scales and scutes.

**Caption:** at left, a surprisingly big African softshell *Trionyx triunguis* photographed at the mouth of the Congo River; this is one of a series of photos shared in 2009 at the SA Reptiles discussion board. At right, skull of an Indian narrow-headed softshell *Chitra indica*, another large member of the group. Note how elongate and (yes) narrow that skull is, and how the orbits are aaaaaalll the way at the front. Images: © Herphabitat (**originals here**); Darren Naish.

Softshells are typically associated with tropical and subtropical swamps, lakes and pools in North America, Africa and Asia but also occur in estuarine and marine environments. I’ve found them especially exciting over the years due to their ecological flexibility, the relatively large sizes reached by individuals within some species, and their sometimes ferocious predatory prowess. Plus they often look very, very weird.

In combination, these factors help explain why they’re one of the few turtle groups that I’ve covered here at Tet Zoo on repeat occasion, though I haven’t given them due credit because boy do they need more coverage. Check out…

The goat-eating hot water bottle turtles, November 2007
Giant African softshells – wow!, January 2010
In case you forget, softshell turtles are insanely weird, December 2011

Caption: a Florida softshell Apalone ferox that I photographed in captivity in 2011. The red hue is partly caused by the colour of its basking lamp. Either way, the turtle still looks very, very strange. Image: Darren Naish.

Leatherbacks or leatherback sea turtles (Dermochelyidae) are named for the extant Dermochelys coriacea, a remarkable marine turtle well known for its substantially reduced carapace encased in leathery skin, enormous front flippers, deep head, prominent pseudoteeth, and size (the carapace can be 1.8 m long and the flipper span can be 2.7 m).

Dermochelys is a predator of cnidarians (jellies, siphonophores) that dives as deep as 1.5 km. It used to be considered a specialist predator of these animals but there are some indications that it might be better characterized as a generalist, since we now know that it also eats algae. It’s fast growing (reaching maturity in less than a decade) and near-globally distributed. There’s tons to say about it. I have covered the fossil history of leatherbacks in my 2023 book Ancient Sea Reptiles but still need to write about the behaviour, anatomy and ecology of the extant species.

Ancient Sea Reptiles Is Out Now, February 2023
News from the World of Ancient Sea Reptiles, May 2023

Caption: at left, this blog’s author with a life-sized model of the Harlech leatherback at Anglesey Sea Zoo. At right, one of the original photos of the actual animal, which stranded on the Welsh coast (after drowning in fishing gear) in 1989. It was huge, weighing 916kg. Images: Toni Naish; © Western Mail & Echo, Wales.

Snapping turtles (Chelydridae) are excusively American today and are represented by two genera: the common snappers (Chelydra) and alligator snappers (Macrochelys; it is termed Macroclemys in some of the older literature). These are strongly aquatic, omnivorous turtles with large heads, renowned for their powerful, hooked beaks and extremely rapid strikes. Molecular studies indicate that chelydrids are close relatives of mud and musk turtles. I love snapping turtles and have had reason to write about them a few times at Tet Zoo…

They bite, they grow to huge sizes, they locate human corpses: the snapping turtles, part I, February 2006
Snapping turtles, part II: hyperexcitability, supercooling, and the inevitable recolonisation of Europe in the Anthropocene, February 2006
Snapping turtles, part III: bite, lunge, lure and snap, May 2006

Caption: an unusually pale Common snapping turtle on show at Bluereef Aquarium, Portsmouth, UK. Images: Darren Naish.

Tortoises (Testudinidae) are the most terrestrial of testudines, their terrestrial features including a strongly domed carapace, large (even gigantic) size, compact hands and feet and hoof-like or nail-like unguals. An external dermal covering of large, sometimes plate-like or spiny, scales often covers the limbs and the snout is typically deep, broad and short. This is another group that I’ve covered here quite a few times. Check out…

Gilbert White’s pet tortoise, and what is ‘grey literature’ anyway?, October 2008
Turtles that eat bone, rocks and soil, and turtles that mine, April 2014
Tortoises as Consumers of Carrion, March 2021
Tortoises as Consumers of Carrion, Part 2, June 2021
Leopard Tortoises, January 2022
Megalochelys, Truly a Giant Tortoise, January 2024

Caption: Megalochelys, a giant tortoise of prehistoric Eurasia, included giant specimens where the CCL (curved carapace length) was 2 m long. Here, I’ve followed the long tradition of showing this animal to scale with a modern human: the one used here has a standing height (without hat) of 1.6 m. Image: Darren Naish.

That will have to do for now. How I’ve love to write about turtles at length. But I have to stop there. Thanks to LOLAMARINA – if that is her real name – for reminding me that it’s World Turtle Day once again, and let us all remember that so many turtle species worldwide require conservation and compassion if they are to persist. That means helping to maintain the existence of wetlands, woodlands, natural deserts and coasts, preventing pollution, and objecting to their exploitation in the pet and restaurant trade. We love turtles; the world will be so much poorer without them.

If you enjoy Tetrapod Zoology, please consider supporting its persistence at patreon. Thanks to those who keep things going already, you are the finest of humans.

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Let’s experiment with camera trapping!

Some months ago, I was very graciously loaned two full HD hunting cameras (aka trail cameras or remote cameras) by friend and colleague Rick Minter, specifically for use in Central Asian fieldwork (and if you want to know more about that, go here and here). For assorted practical reasons they were never used as planned. But in my keenness to use them – and ever hoping that I might catch a wayward British bigfoot or panther on camera – I recently installed one of them in the small area of overgrown, designed-for-wildlife space I own, and here we’re going to look at the results. Let’s do some camera-trapping in tiny little English suburbia.

The camera. I’ll say to start with that the camera – I’m hazy on its specifications but think it’s a Brown A280 with infrared motion sensor – is outstandingly good at its job and has performed exactly as I hoped. I initially had it on high sensitivity, but this resulted in hundreds of photos where it had been triggered by moving vegetation. I set it up to overlook part of one of our ponds. In that location, it isn’t really concealed but merely strapped to a metal post.

**Caption:** what sort of tetrapods might you expect to see in a small, suburban area in the south of England? These three are average, and even they are less abundant that you might like. Clockwise from left: Common frog *Rana temporaria*, Dunnock *Prunella modularis*, Eurasian blackbird Turdus merula. The blackbird is currently undergoing decline in the UK (by about 50% in some areas), apparently as a consequence of Usutu virus. Indeed, there are none living in our immediate vicinity this year… which is concerning. Images: Darren Naish.

The UK is unlike most other parts of the world. In discussing the sort of wildlife you might photograph with a remote camera in the UK, I feel the need to provide some background that might be necessary to readers overseas. To those of you outside the UK, I need to emphasize how little wildlife there is here, the south-east in particular.

The UK is – you have to believe me here – embarrassingly bereft of wildlife, and given this situation it’s simply unbelievable that successive governmental administrations have each brought in new plans to reduce what denuded little scraps of fauna and flora remain. Our current government has just (May 2025) announced a plan to allow development of infrastructure projects without public consultation at the pre-application stage, which basically means that companies can see a patch of land they want to develop and proceed without there being a need for people or organizations to raise objections. Remember that we humans massively benefit from the existence of green space: the argument that wild spaces are required isn’t ‘just’ about wildlife.

**Caption:** the green area in front of our house. It’s fringed by tall hedges and there are two ponds on the right. Image: Darren Naish.

When people say to me such things as “How many species of amphibians are there where you live?”, “Might snakes visit your pond to catch frogs?”, “Could shrews have eaten those half-eaten beetles you found?”, “Could it have been a wildcat or marten near your house?”, I’m both entertained and saddened by the naivety. One species of amphibian. No snakes. No shrews. No wildcats. No martens, no mustelids of any sort. And so on. I say it again: the UK is embarrassingly bereft of wildlife, such that the area where I live (Southampton in Hampshire) essentially has no more than a handful of species, and those species we do have are the hardiest survivors within their respective groups, the ones that occur right across Eurasia.

On that note, the area where the camera is set up is tiny, enclosed, fringed by suburbia, and literally right next to a busy main road. Here in suburban little England, it’s typical for houses to have small areas of land surrounding the house – sometimes both at the front and the back, if you’re lucky – that we term gardens. If you’re very lucky, or rich, a garden can be over 20 m (60 ft) long, but more average is something 6 m (20 ft) by 10 m (33 ft) or smaller, and that’s what we have. In the photos that follows, remember that we’re seeing a tiny, enclosed area: not the edge of a large area of farmland or forest. Oh, to live in such a place. I hate watching TV shows set in the USA or continental Europe because it seems to be portrayed as normal in those places that people live on vast plots of land, surrounded by countryside. The UK is not like that for average people.

**Caption:** it is standard practise here in the UK to completely strip areas around houses of all vegetation, cut down all trees, prevent the growth of grasses and so-called weeds, and to replace all of it with paving or concrete. I have gone against this trend and am keeping this area wild. Some people see it as a total mess, others as a haven for wildlife. I think it’s both. Image: Darren Naish.

Enough preamble, let’s get to the animal photos. I’ve deliberately kept them the same size and format as I find it entertaining to see the respective animals appear and disappear.

Carnivorans. The area where I live might be bereft of animals like mustelids or wildcats, but we do, famously, have a conspicuous number of urban and suburban Red fox Vulpes vulpes and we know from the noise they make and from occasional sightings that individuals often visit our garden. The first week of camera-trapping reveals that at least one Red fox is a regular visitor to our garden. As you can see, the animal (I’m assuming for now that it’s the same one individual) often visits the area of the pond, and seemingly drinks from it. Most of the photos were taken at night but there are a few from daylight. Here are a few of the better shots…

               **And then there are pet cats.** In a small, overcrowded, overbuilt island nation with a small amount of declining wildlife, it should be obvious that free-roaming pet cats are an ever-present issue. It’s a very emotive one, to be sure. I feel – from decades of experience and observation – that pet cats are a massive problem for the health and persistence of wild animals, and I’ve gradually come to be annoyingly militant on this issue. I don’t think that cats should be out and roaming free *at all*, and I think that the damage they do to our handful of native reptiles and amphibians, to small mammals and birds, and even to arthropods (they’re good at killing butterflies, dragonflies and even crickets and grasshoppers), is self-evident and that letting your pet cat roam free outside is just not consistent with the claim that you care about wildlife (yes, I’m a cat owner, and yes my cats are indoor animals). I find that more and more people are coming round to this point of view and I no longer receive the total incredulity I used to when pushing this perspective.

**Caption:** we own two hilariously derpy cats, Mochi and Mocha, both looking quite grumpy on the left (Mocha is the one with the white ear spot). These are indoor cats who live happy, healthy lives in domestic surrounds. They’re placed on regular occasion inside a catio that basically resembles an aviary. We need to normalize the idea that pet cats should not be allowed to roam free outdoors. Images: Toni Naish; Darren Naish.

While I’m on this subject, pet dogs are a total disaster for beleaguered wildlife as well. It is extraordinarily depressing to visit places with the words ‘national park’ or ‘nature reserve’ in their name and see that people view them as dog playgrounds. Remember that dogs don’t necessarily need to kill or harass wildlife to cause problems; we know that the insecticides in dog shampoos and the de-worming agents expressed in their dung are ecologically damaging when dogs interact with the environment. But I digress.

Sure enough, a specific cat is revealed to be a regular visitor to the garden. It hangs around at the pond for extended lengths of time. It’s a black animal and hence a few of the photos are exciting as the first ones I got to see were mere glimpses of dark, curled tail and part of a hindlimb. Alas, that fluorescent collar is a bit of a giveaway. Look at the timestamps and note that the cat is standing in the same spot (lower two images) for over 20 minutes. This suggests that it’s stalking something.

      **Birds**. We get a reasonable number of small passerines in the garden – Dunnock *Prunella modularis*, House sparrow *Passer domesticus*, Blue tit *Cyanistes caeruleus*, Wren *Troglodytes troglodytes* and European robin *Erithacus rubecula*, predominantly – and it would be nice if individuals of at least some of those species got caught on camera. But for week 1, only robin was obliging, since a lone individual visited the shallow end of the pond, either to forage (they will eat aquatic arthropods, tadpoles and small fishes) or to collect mud or moss for nest-building. Obviously, the camera – despite being HD – isn’t quite up to capturing small passerines in detail.

**Caption:** hopefully you can see the robin at centre left. This is a proper robin, of course, not a pretender thrush or anything weird like that. Images: Darren Naish.

Common wood pigeons Columba palumbus are ever-present here and appear to be doing fine in suburbia, and individuals were often caught by the camera, often around the edge of the pond and sometimes wading right out to the edge of the shallow area.

      Also among larger birds, we have – in the past – had Eurasian magpies *Pica pica* nest in our garden (I wrote an article about this back in 2013), and it looks like this is happening again here in 2025, which is good. I’m a huge fan of corvids and don’t take them for granted.

**Caption:** what are likely the same pair of magpies have nested here on a few occasions, though I only know of one time (in 2013) in which they successfully fledged chicks. At left is one of the parents in 2013, and at right a montage of the two 2013 chicks. Of some relevance to the tone of my article here is that the several of the trees that these birds formerly used as vantage and resting places are now long gone because people in the suburbs sure do hate trees. I aim to start a ‘spite tree’ movement in response to this. Images: Darren Naish.

Magpies were captured on camera a few times in the first week, often at the pond’s edge. A very odd photo shows an individual flying at the top of frame with nesting material in its bill, and this bird also looks odd because it was lacking its long tail feathers when the photo was taken.

         Carrion crows *Corvus corone* are sometimes seen in the area, and in 2017 one discovered the magpie’s nest and set about harassing these birds until they abandoned their efforts and chose to nest elsewhere. However, I didn’t know until seeing the photos below that at least one crow is currently in the habit of visiting the garden. That’s good news. In the images here, the camera captured the crow standing on the path and taking off too.

      **Smaller animals.** A few cryptic small animals were captured by the camera too, but some imagination is required to work out what they are. You can see from the images (and the zoomed-in sequence) shown below that a small, dark animal with at least one large, reflective eye (I assume it had two) was photographed at the edge of the pond’s shallow end. If you look at the time stamps, you can see that it was in approximately the same location for about two hours.

      I’m pretty sure that this animal is a frog, by which I mean the only frog (and only amphibian) here: *Rana temporaria*, the Common frog. They do, of course, leave the pond and move about the garden when foraging or looking for a new area in which to take refuge.

Finally…. numerous humans were caught by the camera but I’ve opted not to showcase those pictures here. Except for those showing this frequently present, sometimes nocturnal individual…

   And thus, we come to an end of my first ever week of camera-trapping. I’ve got additional results since and will no doubt share results here again, but already I’ve essentially camera-trapped every species we can get in the garden. Really: that’s it. Oh, we have had rats and mice here before but none are in the area right now. This was a trial run to learn how the camera works and what it can do. The challenge next is to install it somewhere a bit more rural. If only such places existed nearby…

For previous articles on various of the animals mentioned here, see…

My local magpie family: four weeks of observation, 265 photos, and how good are the results?, June 2013
A Battle Among Blue Tits, February 2018
Birdwatching in Suburban China, May 2019
Tell Me Something Interesting About Dunnocks, July 2019
Surprising Diet of the House Sparrow, August 2022

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for Tet Zoo and the more productive I can be on those long-overdue book projects. Thanks!

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Yes… DINOCON!

If you’re at all aware of what I do on social media, you’ll know that I have – for some weeks now – been talking about DinoCon, our new, dinosaur-themed annual convention, a descendant (of sorts) of TetZooCon. DinoCon involves collaboration between myself and the PalaeoGames team, in addition to the University of Exeter’s Camborne School of Mines.

DinoCon 2025 happens at the University of Exeter in south-west England on the weekend of August 16th and 17th. And seeing as that’s only a little more than three months away, now is a good time to once more share some DinoCon news. Many announcements have been made recently.

   First of all, we’ve recently announced a good number of speakers and presenters, albeit not all (the full timetable is due to be announced soon). Talks will cover such issues as sauropod skin and palaeobiology, tyrannosaur biomechanics, David James Armsby’s dinosaur-themed animation projects, life a a fossil preparator, a behind-the-scenes view of dinosaurs in movies, the remarkable fossils of the Dorset coast, the science of camouflage and what it might mean for dinosaurs, *and more*.

We have a palaeoart discussion panel, a quiz (an evening event with prizes), an art exhibition, and a charity auction. We’ve now finished accepting applications for vendors and can confirm that we have a truly world-class collection of stallholders. We’re talking figures, models, books and book signings, merch, and tons more. If you’ve been successful in winning a table but haven’t yet responded or paid for this place, please do so asap.

**Caption:** a montage of promotional art created specially by Natalia Jagielska!

If you haven’t bought a ticket, now is the time. And be sure to sort your accommodation: our venue of choice is the university’s Holland Halls. We look forward to seeing you in Exeter in mid-August! This is going to be the UK’s biggest and best dinosaur-themed meeting by far and we can’t wait for you to join us.

For additional DinoCon news, be sure to follow us on social media (we’re @DinoConUK on Twitter/X and Instagram, and @dinoconuk.bsky.social at BlueSky), and for other articles on DinoCon, see…

Announcing DinoCon … Tickets Now on Sale!, February 2025
The UK’s First DinoCon – Britain’s Largest Dinosaur and Palaeontology-themed Convention, April 2025

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Let’s look at armadillos some more. Or, let’s look at more armadillos. I mean, let’s look more at armadillos. Whatever: armadillos! More.

**Caption:** a Pichi montage, showing wild individuals in (left) Torres del Paine National Park, southern Chile and (right) Chubut Province, Argentina. Note the overall covering of dark hairs and the long and gently curved claws on the forelimbs. The snout on the individual at left looks unusually short, I think because of foreshortening. Images: Wouter Potters, CC BY-SA 4.0 (**original here**); Mikelzubi, CC BY 3.0 (**original here**).

A relatively obscure yet apparently quite abundant armadillo, the Pichi, Dwarf armadillo or Pygmy armadillo is the sole species recognised within the genus Zaedyus, Z. pichiy. It’s native to southern Argentina and adjacent parts of eastern Chile where it’s associated with arid grassland. The southern part of its range – it occurs in areas bordering the Strait of Magellan – can be close to freezing in the (austral) winter, and a famous aspect of its biology is that it undergoes hibernation during times of environmental stress, it being the only extant xenarthran known to do this. Even when not hibernating, individuals can enter “prolonged torpor bouts lasting more than 24 hours” (Superina & Jahn 2013, p. 280), sometimes repeatedly.

Pichis can be easily distinguished from armadillos of other species thanks to their marginal scutes, which have triangular, pointed apices, this giving the carapace serrated borders. A pale longitudinal stripe along the dorsal midline is also supposedly diagnostic (Superina & Abba 2014), though I’m confused as I don’t see it in the photos here. The species is, however, similar enough to Euphractus (the six-banded armadillos) that some authors have considered the Pichi part of this genus, though this is not considered correct today.

**Caption:** museum taxiderm specimen of a Pichi at Museo Civico di Storia Naturale Giacomo Doria in Genoa, Italy. Image: Daderot, public domain (**original here**).

A niche debate on armadillo taxonomy. Current phylogeny and taxonomy has it that the Pichi is part of a clade that includes not just Euphractus but naked-tailed armadillos (Cabassous), three-banded armadillos (Tolypeutes) and others. Until recently, the convention was to include all extant armadillos within the family Dasypodidae. However, a deep split between the two main armadillo lineages – it appears to have occurred around 42 million years ago, within the mid Eocene (Gibb et al. 2016) – has led to the idea that long-nosed armadillos should be separated at the ‘family’ level from the others. Those ‘others’ are substantially more diverse, since euphractine armadillos, fairy armadillos, and tolypeutine armadillos (giant, three-banded and naked-tailed armadillos) all belong here (Gibb et al. 2016). The Pichi is a euphractine.

If we opt to recognise this split taxonomically, the correct name for the long-nosed armadillo lineage is obviously Dasypodidae. The apparently most correct name for the other armadillo clade is Chlamyphoridae, based on a name published by Charles Lucien Bonaparte in 1850 for Chlamyphorus, the Pink fairy armadillo. Gibb et al. (2016) wrote that Bonaparte used the name ‘Chlamyphorinae’ but he actually published ‘Chlamydophorina’ based on his understanding that Chlamyphorus should be written Chlamydophorus (McKenna & Bell 1997, p. 82), a view established in 1830 by German zoologist Johann Georg Wagler. Chlamyphorus had been published just five years prior by physician and naturalist Richard Harlan and was intended to mean ‘cloak carrier’. However, because the Greek root of the ‘chlamys-’, ‘cloak’ part of the word is actually ‘chlamyd-’, Wagler thought that Harlan’s 1825 Chlamyphorus should really be written Chlamydophorus*, hence Bonaparte’s Chlamydophorina.

Remember: phylogeny and taxonomy are not the same thing. However, there has always been a general preference to get taxonomy to match phylogeny.

Caption: the two extant fairy armadillos are currently placed in separate genera. This is the Pink fairy armadillo Chlamyphorus truncatus, the smallest living armadillo. Good photos are hard to get; this is a taxiderm specimen at the Naturmuseum Senckenberg, Germany. Image: Daderot, public domain (original here).

What to do? Shouldn’t we be following the ‘incorrect’, now out-of-favour formulation of the name Chlamyphorus (namely Chlamydophorus) in devising higher names, since that’s what Bonaparte did? Well, no. Article 35.4 of the ICZN basically says that family-level names shouldn’t be based on spellings of genus names now considered ‘incorrect’. One more thing on this: given that Bonaparte used ‘Chlamydophorina’ based on ‘Chlamydophorus’, should he really be credited with Chlamyphoridae, as he is in Gibb et al. (2016)? Views differ on this and a popular convention is to wave this away by saying that we’re going with (in this case) Bonaparte’s concept of the name, even though the formulation isn’t right according to modern use, and even though the ‘rank’ we’re now assigning the name isn’t the same as that originally preferred.

One of the funnest findings of molecular phylogenetics on armadillos is that glyptodonts are part of Chlamyphoridae (Delsuc et al. 2016, Mitchell et al. 2016): they’re deeply nested within armadillos, not an armadillo sister-group as we used to think. So, yes – it is now properly correct to refer to glyptodonts as ‘giant armadillos’ (whereas this wasn’t considered technically correct beforehand). I’ll come back to that issue in time, since oh boy do I intend to revisit glyptodonts, a group I haven’t really written about since 2008.

**Caption:** cingulate (armadillo) phylogeny as recovered by **Delsuc *et al*. (2016)** on the basis of mitochondrial DNA extracted from a carapace fragment of the giant Pleistocene-Holocene glyptodont *Doedicurus*. Glyptodonts are nested within armadillos, and within chlamyphorids. This result has been obtained in more than one study and is now backed by anatomical data too. Image: **Delsuc *et al*. (2016)**.

But back to the Pichi… I have to mention that the Pichi’s generic name Zaedyus – published by famed Argentine naturalist and palaeontologist Florentino Ameghino in 1889 – apparently means ‘very pleasant armadillo’ (Superina & Abba 2014).

Despite sometimes being called a ‘pygmy armadillo’, the Pichi isn’t that pygmy, head and body length reaching 35 cm (though between 22 and 31 cm is more typical) with an additional 13 cm or so for tail. It’s omnivorous, its diet including fungi, plants including mesquite pods, arthropods and small vertebrates of numerous sorts, and also carrion. If you recall the 2020 article from these here parts on armadillo carnivory and carrionophagy (err, yeah, let’s go with that), you might recall that the Pichi is one of several armadillos classified as an ‘omnivore-carnivore’ (Redford 1985), the species within the group being hard to group tidily in ordinarily dietary categories. It’s one of several mammals of arid habitats said not to drink water (Superina & Abba 2014).

**Caption:** another wild Pichi, specifically in Valdes Peninsula (a chunk of land that juts out into the Atlantic), north-east Chubut Province, Argentina. Valdes Peninsula is a nature reserve famous for the marine mammals that visit its shores. Image: Marianocecowski, CC BY-SA 3.0 (**original here**).

And that’s where we must end for now, with many thoughts on armadillos yet to be published.

For previous Tet Zoo articles on armadillos and othr xenarthrans, see…

Five things you didn’t know about armadillos, June 2007
What was that skull? (on glyptodonts), May 2008
I, Priodontes, the tatuasu, September 2008
Predation and Corpse-Eating in Armadillos, September 2010
The Fate of the Woolly Long-Nosed Armadillo of Peru, April 2024

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for Tet Zoo and the more productive I can be on those long-overdue book projects. Thanks!

Refs - -

McKenna, M. C. & Bell, S. K. 1997. Classification of Mammals: Above the Species Level. Columbia University Press, New York.

Mitchell, K. J., Scanferla, A., Soibelzon, E., Bonini, R., Ochoa, J. & Cooper, A. 2016. Ancient DNA from the extinct South American giant glyptodont Doedicurus sp. (Xenarthra: Glyptodontidae) reveals that glyptodonts evolved from Eocene armadillos. Molecular Ecology 25, 3499-3508.

Redford, K. H. 1985. Food habits of armadillos (Xenarthra: Dasypodidae). In Montgomery, G. G. (ed) The Evolution and Ecology of Armadillos, Sloths, and Vermilinguas. Smithsonian Institution, Washington, D.C., pp. 429-437.

Superina, M. & Abba, A.M. 2014. Zaedyus pichiy (Cingulata: Dasypodidae). Mammalian Species 46, 1-10.

Superina, M. & Jahn, G. 2013. Effect of low-quality diet on torpor frequency and depth in the pichi Zaedyus pichiy (Xenarthra, Dasypodidae), a South American armadillo. Journal of Thermal Biology 38, 280-285.

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One of my favourite mammal assemblages of all is Xenarthra, the ‘strange joint’ group that includes the remarkable and odd anteaters, sloths and armadillos…

**Caption:** a xenarthran montage. I’ve drawn a few xenarthrans over the years, but not as many as I should have by now. Here are a few illustrations I have to hand, but many more will be included in **my textbook** once I get to finishing it. Images: Darren Naish.

I’ve published a bit on those groups here, over the years (see the list of links below). I’ve just never published all that much. I aim to change that this year, and let’s start here, with assorted thoughts and observations on ARMADILLOS. Be sure to check the links for additional info.

Long-nosed armadillos. If you’ve heard of armadillos, the chances are high that you’re aware of the Nine-banded armadillo Dasypus novemcinctus, one of the best known, most intensely studied, and most geographically widespread of armadillo species. Until very recently, it was regarded as the only living long-nosed armadillo species that ranges into North America*. Incidentally, not all individuals have the eponymous nine bands: the actual number is variable across the range of the species. Some individuals have eight, others ten.

The text here initially stated – erroneously – that the Nine-banded armadillo is the only living armadillo species that occurs in North America, something that’s very much incorrect in view of the presence of naked-tailed armadillos in Mexico. The ‘Until very recently’ there refers to the 2025 splitting of the D. novemcinctus complex, something I will come back to in a future article.

Caption: the long-nosed armadillos include very familiar species – like the Nine-banded armadillo shown at left – as well as very obscure ones that are still poorly known and mostly unphotographed in living condition. Hence we have species like this one (on the right: the holotype of the Yungas lesser long-nosed armadillo D. mazzai), mostly only known from taxiderm specimens like this. More on D. mazzai in another article! Images: Mwcolgan8, public domain (original here); Feijó et al. (2018), CC BY 4.0.

Relatively little known is that D. novemcinctus is only one of eight or nine extant species in the genus Dasypus, collectively termed the long-nosed armadillos. Excepting D. novemcinctus, all are limited to South America. While their taxonomy has been considered mostly settled over recent decades (a real contrast to the 19th century), a few late 20th century and 21st century revisions have resulted in a bit of confusion with respect to the more poorly known taxa (Vizcaíno 1995, Feijó & Cordeiro-Estrela 2014, 2016, Feijó et al. 2018).

One thing to note is that some of these species are – in real contrast to the Nine-banded armadillo – of conservation concern and officially classified as ‘vulnerable’ or ‘near threatened’. That goes for the Southern long-nosed armadillo D. hybridus and the Hairy long-nosed armadillo D. pilosus, though a case has been made by the IUCN (Superina & Abba 2014) that the latter should better be treated as ‘Data Deficient’ (this is one of the worst categories to be in, as it creates the impression that the animal is ok). The Southern long-nosed armadillo, by the way, might not be a distinct species, since one recent revision found it to be a subspecies of the Seven-banded armadillo D. septemcinctus (Feijó et al. 2018).

**Caption:** yes, this is a real animal of the modern age, not a CG reconstruction of a Mesozoic mammal, not a rhinogradentian (**reference**). There are very few photos of live specimens of the Hairy or Woolly long-snouted armadillo, and this is the one that gets used the most. It was taken by André Baertschi and this version was taken from **the relevant IUCN page**. Other images of the species **can be seen at iNaturalist**. Image: © André Baertschi / wildtropix.com.

The Hairy or Woolly long-nosed armadillo. D. pilosus is radically odd. Endemic to the montane cloud forests of the Peruvian Andes (specifically to the departments of San Martín, La Libertad, Pasco, Huánuco, Junín and Amazonas), it’s very poorly known: just six specimens were known to science between its description in 1856 and recent decades, and even today few individuals are available in collections (Castro et al. 2015). The original specimen was described by Leopold Fitzinger based on a taxiderm specimen purchased in London by the animal dealer and merchant Ludwig Parreyss in 1833 (Feijó et al. 2018). It had no locality data beyond merely ‘Peru’ and Fitzinger was so perplexed by it that he wondered whether it might be an intermediate between armadillos and anteaters (Feijó et al. 2018).

**Caption:** the *Dasypus pilosus* holotype, the mounted taxiderm specimen NMW ST 222 at the Naturhistorisches Museum, Vienna, Austria, as featured in **Feijó *et al*.’s (2018)** excellent and very thorough taxonomic review of long-nosed armadillos. It’s labelled with the name *Praopus hirsutus*, this being given to the species by Hermann Burmeister in 1862 since he was unaware of Fitzinger’s earlier publication. Check **Feijó *et al*. (2018)** for more data on this specimen and the taxonomic history of *D. pilosus* in general. Image: **Feijó *et al*. (2018)**, CC BY 4.0.

Even today, almost nothing is known about the ecology, behaviour or biology of this species. Some authors have speculated that it might be a specialized insectivore that eats ants and/or termites (in which case it’s a myrmecophage or termitophage) (Castro et al. 2015), the unusual anatomy of its skull, teeth and jaw joint indicating that it’s even more specialized for this lifestyle than other Dasypus species.

However, its existence at altitude (2600-3400m above sea level; Castro et al. 2015) casts doubt on this suggestion, since surveys demonstrate that insects are comparatively rare in such places. Feijó et al. (2018) noted that D. pilosus has a number of similarities (including an especially elongate snout and mandible and reduced teeth) with Asian montane rodents that eat soft-bodied invertebrates, in particular worms. Could, then, D. pilosus be a (mostly) worm-eating armadillo? It’s a really interesting idea and more data is needed.

**Caption:** very nice image of Hairy or Woolly long-nosed armadillo in life, by Jorge González and included in Castro *et al*. (2015). It is not well known – even among people who know mammals – that there are armadillos that look like this. Image: Jorge González, Castro *et al*. (2015).

And it really is hairy, or woolly; being properly covered in a pelt, this giving it a body which superficially recalls that of a shaggy rodent or giant shrew. If you know about armadillos, you’ll know that there are species – most notably the hairy armadillos or peludos (Chaetophractus) – where a reasonable amount of hair is present across the unarmoured underside, and also here and there across the carapace. However, the Hairy or Woolly long-nosed armadillo is a wholly different thing. Scutes are only visible on the top of its head and snout, ears, hands, feet and slender tail. The hairs grow through pores on the scutes (Castro et al. 2015), it being one of several armadillos which show that you can be totally armoured and sport a pelage at the same time.

The case for Cryptophractus. D. pilosus is, in fact, so odd that a person who knows armadillos might doubt, purely intuitively, whether it should be included within Dasypus. It’s not just different because of the hairiness but also because it has a notably longer, more slender skull, especially small teeth, a reduced mandible with a more limited range of motion than other long-nosed armadillos, a higher number of movable bands than other long-nosed armadillos (as many as 11), and unusual osteoderms that lack sulci and have a unique pattern of pores (Castro et al. 2015). Accordingly, a long-standing idea is that it should be given its own ‘subgenus’ within Dasypus, namely Cryptophractus.

**Caption:** the Hairy or Woolly long-nosed armadillo is a special armadillo. As shown in these images – all from Castro *et al*. (2015) – the hairs emerge from numerous pores in the scutes, which appear to match the numerous foramina (bony openings) in the underlying osteoderms. Osteoderms = bony structures embedded in the skin. Scutes = keratinous structures overlying osteoderms (albeit not always!). There’s a long-standing discussion on whether foramina in bony structures are at all relevant to the presence or otherwise of hairs. This is a case where they apparently are. Scale bars = 50 mm. Images: Castro *et al*. (2015).

Inspired by the distinct nature of this species (and by the lack of detailed study), Mariela Castro and colleagues published an analysis in 2015, their conclusion being that it is indeed ‘distinct enough’ from other long-nosed armadillos to warrant exclusion from Dasypus (Castro et al. 2015), this being consistent with their phylogenetic analysis. This, if right, has implications for a number of fossil armadillos also named as species of Dasypus, since their position outside the Cryptophractus + Dasypus clade requires that they need distinct ‘genus-level’ names.

**Caption:** Castro *et al*. (2015) found this phylogenetic arrangement for dasypodids and other armadillos via their analysis of 58 anatomical characters. *Stegosimpsonia* is a fossil animal from the Eocene while *Peltephilus* and *Stegotherium* are Miocene; *Propraopus* and *Anadasypus* are dasypodids from the Pleistocene-Holocene and Miocene, respectively. The topology here has the Hairy or Woolly long-nosed armadillo outside a clade that contains all other *Dasypus* species, this being deemed consistent with the idea that the species might be distinct enough to be recognised as the distinct genus *Cryptophractus*. Does molecular data support this view? Read on. Image: Castro *et al*. (2015).

The case against Cryptophractus. However, 2015 is a little while ago now. How has this proposal been received? Hautier et al. (2017) looked at shape variation across long-nosed armadillo skulls using morphometrics, and did find pilosus to be an outlier in several of their analyses. They mostly found, however, that it was not so extreme that it should be excluded from Dasypus; rather, it was ‘extreme’ in terms of the sort of shape variation they possess.

**Caption:** **Hautier *et al*. (2017)** used principle components analysis to analyse the degree of shape variation present in the long-nosed armadillo species. In the plots shown here – analysing the shapes of mandibles – the Hairy or Woolly long-nosed armadillo is an outlier, being *mostly* outside the main cluster occupied by most other *Dasypus* species (it’s the red circles at lower left in A and B, and lower left in the left part of the main cluster in B). It’s not the only species that was found to be an outlier though. Image: **Hautier *et al*. (2017)**.

What seems to be the death knell for the ‘distinct Cryptophractus’ hypothesis came from the theatre of molecular systematics. Gibb et al. (2016) found a totally contradictory phylogenetic arrangement to Castro et al. (2015), since they recovered pilosus within the middle of the long-nosed armadillo radiation (not as its outgroup), and as a young lineage (not much older than Late Pliocene) whose “molecular divergence does not seem to match its morphological distinctiveness”. They were very much aware of Castro et al.’s study in saying this, since they took time to comment on it, noting that its anatomy-based conclusions were (according to their results) less reliable than their molecular ones.

Long-nosed armadillos, the book! One of the first things I checked when receiving Richard Webb and Jeff Blincow’s excellent A Field Guide to the Larger Mammals of South America (Webb & Blincow 2024) was the long-nosed armadillo section. Turns out that they give these animals excellent coverage, with full pages given to all species, four of which are depicted via paintings since no good photos exist! And there in the middle of this section is the Hairy long-nosed armadillo, smack-bang within Dasypus, and also featured in a location that doesn’t – contra Castro et al. (2015) – make it phylogenetically distinct relative to the others. I guess they’d seen Gibb et al. (2016) and followed it.

**Caption:** Richard Webb and Jeff Blincow’s *A Field Guide to the Larger Mammals of South America* is an extremely impressive book and I strongly recommend it. It features whole pages on most species, even tremendously obscure and recently named ones, and that includes all the obscure long-nosed armadillos. But good photos of some of those don’t exist, so they had to resort to the inclusion of artwork. **Get the book here.** Images: Darren Naish.

For all its obscurity, Dasypus pilosus is not – believe it or not – as obscure as other members of the long-nosed armadillo group. I was planning to talk about those (or, some of them, anyway) in this article… but it’ll have to wait. In fact, I had hoped in this article to talk about armadillo diversity in general. Alas, things expanded and time, as ever, is against me. I will be coming back to this fascinating group in time.

ADDENDUM: after I finished this article and prepared to get it published, I had a look to see if other bloggers have covered the species before. And turns out that Darin Croft – at his excellent blog The Rafting Monkey – published an article on Dasypus pilosus back in 2017. It essentially presages everything I’ve covered here, showing that great minds think alike, I guess. Check out that article for another take on this species.

For previous Tet Zoo articles on armadillos and othr xenarthrans, see…

Five things you didn’t know about armadillos, June 2007
What was that skull? (on glyptodonts), May 2008
I, Priodontes, the tatuasu, September 2008
Predation and Corpse-Eating in Armadillos, September 2010

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for Tet Zoo and the more productive I can be on those long-overdue book projects. Thanks!

Refs - -

Castro, M. C., Ciancio, M. R., Pacheco, V., Salas-Gismondi, R. M., Bostelmann, J. E. & Carlini, A. A. 2015. Reassessment of the hairy long-nosed armadillo “Dasypus” pilosus (Xenarthra, Dasypodidae) and revalidation of the genus Cryptophractus Fitzinger, 1856. Zootaxa 3947, 30-48.

Feijó, A. & Cordeiro-Estrela, P. 2014. The correct name of the endemic Dasypus (Cingulata: Dasypodidae) from northwestern Argentina. Zootaxa 3887, 88-94.

Feijó, A. & Cordeiro-Estrela, P. 2016. Taxonomic revision of the Dasypus kappleri complex, with revalidations of Dasypus pastasae (Thomas, 1901) and Dasypus beniensis Lönnberg, 1942 (Cingulata, Dasypodidae). Zootaxa 4170, 271-297.

Feijó, A., Patterson, B. D. & Cordeiro-Estrela, P. 2018. Taxonomic revision of the long-nosed armadillos, Genus Dasypus Linnaeus, 1758 (Mammalia, Cingulata). PLoS ONE 13(4): e0195084.

Hautier, L., Billet, G., de Thoisy, B. & Delsuc, F. 2017. Beyond the carapace: skull shape variation and morphological systematics of long-nosed armadillos (genus Dasypus). PeerJ 5: e3650.

Superina, M. & Abba, A.M. 2014. Dasypus pilosus. The IUCN Red List of Threatened Species 2014: e.T6291A47441122. Accessed on 30 April 2025.

Vizcaíno, S. F. 1995. Identificación específica de las "mulitas", género Dasypus L. (Mammalia Dasypodidae), del noroeste argentino. Mastozoologia Neotropical 2, 5-13.

Webb, R. & Blincow, J. 2024. A Field Guide to the Larger Mammals of South America. Princeton University Press, Oxford.

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It’s 2025, not 2024. But, as per last year, I still aim to rescue and republish the squamate-themed articles originally published at Tet Zoo ver 2 and 3, and today mostly ruined, paywalled, or removed by their hosters….

**Caption:** my location in western Europe means that Lacertidae is the lizard group whose species I’ve encountered most frequently, and here’s a lacertid montage. Clockwise from upper left: the African eremiadin *Gastropholis*, the European lacertin *Podarcis*, the Asian lacertin *Takydromus*, and the European lacertin *Zootoca*. Images: Darren naish; *Takydromus* (lower right) by TANAKA Juuyoh, CC 2.0 (**original here**).

Here, I’ve rescued a 2014 article that forms part of a series on lacertids, this being the Old World lizard clade that includes Eurasian wall lizards, green lizards and fringe-toed lizards. This article is devoted to racerunners – the Eremias species – and the original version is here at wayback machine. Let’s go…

Eremias lizards aren’t always called racerunners: ‘desert lacertas’ is in use too, and the name fringe-toed lizard is also used for some species. This makes sense given that some species specialised for life on loose sand have fringes or combs of subtriangular scales along their fingers and toes, but it’s already in use for another group of lacertids, namely the Acanthodactylus species.

**Caption:** Rapid fringe-toed lizard or Central Asian racerunner *Eremias velox*, a species mostly associated with Central Asia but occurring from Iran in the west to northern China in the east. This one was photographed in the Kyzylorda region of Kazakhstan. Image: Yuriy75, CC BY-SA 3.0, original here.

In general, racerunners are lizards of dry, sparsely vegetated semi-deserts, deserts and steppes, though members of some species live in sandy places in marshes, on coasts and in river deltas. Some tend to frequent rocky plains and the sides of wadis while others are psammophiles that inhabit sand dunes and burrow into sand when taking refuge. Arnold (2004) said that adaptation to aeolian habitats happened independently on two occasions (and see Mayer et al. 2007).

Subtle specialisation for different microhabitats is definitely present in the group: there are species that prefer stabilised sand where light vegetation is growing (like Anderson’s racerunner E. andersoni), others that occur on hard, sometimes pebbly, soil covered in vegetation (like the Black-ocellated racerunner E. nigrocellata), and yet others that frequent loose sand on dunes (like the Pointed-snouted racerunner E. acutirostris). Species that inhabit mountainsides and other rocky places (like Strauch’s racerunner E. strauchi) will happily climb steep slopes (Anderson 1999). Exactly how these microhabitat specialisations correlate with anatomical variation – with scale form, digit length, claw curvature and so on – is a question of great interest since a good understanding of any correlations here can help us make predictions about lizards where we have anatomical data but no (or less) data on microhabitat preference, and vice versa. Work of this sort has been done on several lizard groups but I’m not sure that it’s been done on racerunners.

A few papers have, however, looked at how body shape, limb proportions and so on correlate with life history in racerunners. Li et al. (2011) showed how body size and limb proportions in the Multi-ocellated racerunner E. multiocellata varied between populations according to local climate, with the ones from a warmer location being faster runners than those from a cooler location.

**Caption:** a particularly handsome and robust Gobi racerunner *E. przewalskii*, photographed in the Gobi Desert. There is some mild uncertainty about the status of this species since Gui *et al*. (2011) found it to be nested within the Multi-ocellated racerunner *E. multiocellata*. This could mean that *E. multiocellata* is a species complex, or that *E. przewalskii* doesn’t warrant recognition as a species. Image (c) Conrad Savy, CC BY-NC-ND (**original here**).

Introducing eremiadins. Where do racerunners fit within the lacertid tree? They’re part of a group that I’ve never had much reason to cover before: namely, Eremiadini or Eremiinae or Eremiainae or Eremiadinae, generally called eremiines. This is a pretty big group containing perhaps 15 genera and over 80 species, so I’m only going to talk here about one genus, namely Eremias*.

Arnold et al. (2007) argued that formulations including ‘-iadin’ are technically correct. If the group is regarded as a ‘subfamily’, it should therefore be Eremiadinae, and if it’s regarded as a ‘tribe’ within a ‘subfamily’, it should be Eremiadini. In view of this, the right vernacular terms should be ‘eremiadines’ and ‘eremiadins’.

Caption: my first ‘encounter’ with Eremias; namely, in the pages of Laňka & Vít's 1986 Amphibians and Reptiles, a popular book on European herpetology with really nice illustrations. These are by Libuše and Jaromír Knotek.

As someone who spent formative years reading fieldguides to European reptiles, I know one racerunner – the Steppe-runner E. arguta – as an obscure and exotic lacertid that only occurs in far eastern Europe and is even missing from some books on the European lizard fauna altogether. Sadly, it has disappeared or virtually disappeared from some regions where it used to occur and has been regarded as locally extinct in parts of Romania and Ukraine. I’ve looked for it in Romania but failed to find it. Many lacertid species seem to have declined in recent decades, predominantly as a consequence of agricultural development (Gherghel et al. 2007) and habitat degradation caused by over-grazing and heavy vehicle traffic.

**Caption:** a very charismatic Steppe-runner, the *Eremias* lizard most familiar to Europeans due to its presence in Romania and Turkey. Its range beyond this is huge, extending across to Central Asia all the way to Kyrgyzstan. Around six subspecies have been recognised. Image: Benny Trapp, CC BY-SA 3.0 (**original here**).

Where? How many? And how do you tell them apart? Racerunner species occur from southeastern Europe, across the Middle East, and all the way east to Pakistan, northern China and the Korean Peninsula. Given that we usually think of them as mostly based in Central Asia, the presence of at least one species literally on the shores of the Pacific is a bit surprising. The lizard concerned – the Mongolian racerunner E. argus – inhabits coastal dune systems on Yoobu-do Island and elsewhere on coastal South Korea, right on the edge of the Yellow Sea (Jin et al. 2013, Song et al. 2013).

**Caption:** racerunners as geopolitical icons. Steppe-runners *E. arguta* on stamps produced by Kyrgyzstan and Moldova. Unfortunately, I only have these images at low resolution, sorry. Images in public domain.

A reasonable number of racerunner species are recognised (something like 44) and a few new ones have been named since 2000; eight have been named since the first outing of this article in 2014. 21st century taxa include E. montana (originally written as E. montanus but later corrected by other authors), named in 2001, E. cholistanica, named in 2006, E. karivensis, named in 2007, and E. papenfussi, named in 2011; all are from Iran except for E. cholistanica, which is from Pakistan. The post-2014 taxa are E. dzungarica Orlova et al., 2017 of Mongolia and Kazakhstan, E. kakari Masroor et al., 2020 of Pakistan and Afghanistan, and E. isfahanica Rastegar-Pouyani et al., 2016, E. fahimii Mozaffari & Saberi-Pirooz, 2020, E. killasaifullahi Masroor et al., 2022, E. rafiqi Masroor et al., 2022, E. graphica Orlova et al., 2023 and E. pseudofasciata Orlova et al., 2023, all from Iran. Another supposedly new Iranian species – ‘E. novo’, named in 2006 (mentioned by Mozaffari & Parham 2007) – was never meant to be a new species, but a shorthand way of referring to a new population now thought to belong to E. montana.

As expected for any group of lizards, species predominantly differ from one another in the configuration, arrangement and number of their head shields and other scales, their pigmentation and patterns, and in the form of their subdigital scales. Head shape is reasonably variable within the group, some having far shorter, broader snouts than others. Some of the shallow-snouted ones sometimes don’t look all that different from Acanthodactylus lacertids. Hold that thought...

**Caption:** racerunner distribution shown on a handy colour-coded map, from Guo *et al*. (2011). This range raises questions: why no presence in India, south-east Asia, or Afro-Arabia? Is the group too geologically young to have moved into those regions, or was it in some of them in the past but later made extinct? And do they have the potential to move north as climate change continues across the northern parts of Asia?

Szczerbak (1974) decided to recognise the diversity within the racerunners by classifying Eremias species into five subgenera: Eremias, Ommateremias, Rhabderemias, Pareremias, and Scapteira. These were later raised to generic level by some authors. Oh, and Ommateremias is a junior synonym of yet another name – Aspidorhinus – according to some authors (Guo et al. 2011). As is so often the case with subgenera, the classification of species into these groups was done predominantly on phenetic grounds (that is, on general overall similarity) and distinct anatomical or molecular characters that distinguish them weren’t identified. Having said that, studies of hemipenial anatomy are more or less in agreement with the recognition of these groups (Arnold 1989). An rRNA analysis found Pareremias and Eremias to be clades whereas Rhabderemias, Ommateremias/Aspidorhinus and Scapteira were not monophyletic (Guo et al. 2011). The viviparous racerunners formed a clade within Pareremias*. What? Viviparous species?

Be careful with the diagrams in that paper, since some of the taxonomic names are spelt incorrectly.

Caption: phylogeny for Eremias generated by Guo et al. (2011). Some of Szczerbak's ‘subgenera’ corresponded to clades but others did not.

Life history and the viviparity thing. At least some racerunner species are reportedly short-lived, with average life expectancies of 2.5 years being given for the Rapid fringe-toed lizard (Pouyani 2009). This strikes me as surprisingly short-lived for a lizard, though I’m not wholly sure why I think this given that good longevity data on lizards is infrequently encountered in the literature. Longevity for the Mongolian racerunner is approximately 10 years (Kim et al. 2010), which is greater than that of other small lacertids.

Given the short lifespan of the Rapid fringe-toed lizard, I’m also surprised to find that it apparently only produces 2-5 eggs per year: you’d expect greater fecundity in such a small, short-lived animal. Juvenile racerunners often differ substantially from adults in colouration. This is a fairly common thing for lacertids and might be an evolutionary response to the different predation pressures that affect these animals at different points in their lives. Minor sexual dimorphism is present, with males having proportionally larger heads than females (Li et al. 2006) and, in some species, more prominent spots and stripes (Moravec 1994). However, the heads of males are apparently no broader than those of females (though they are longer) in Mongolian racerunners or Ordos racerunners E. brenchleyi, perhaps because head width is constrained in those species by its role in sand-burrowing, nor are males larger or longer overall (Kim et al. 2010).

**Caption:** Gobi racerunner *E. przewalskii*, a species first described from the Alashan Desert of China but also known from Mongolia, Kyrgyzstan and Russia. General Nikolai/Nikolay Mikhailovitch/Mikhaylovich Przewalski (also written Przhevalsky or Prjevalsky) was a Russian explorer and geographer. He died of typhus in 1888, aged just 49. Image: Alastair Rae, CC BY-SA 2.0 (**original here**).

I mentioned viviparity. Three Eremias ‘species’ are viviparous: E. buechneri, E. przewalskii and E. multiocellata, and I put the word ‘species’ in quote marks there because E. multiocellata is suspected to be a species complex, perhaps housing as many as 8 lineages that should probably be raised to ‘species’ level (Guo et al. 2010, 2011). Viviparity in racerunners might have arisen more than once but one recent study did find the viviparous species to form a clade within Pareremias (Guo et al. 2011). Why might viviparity have evolved in these lizards in the first place? Viviparity in squamates is usually thought to be linked to specialisation for life in cool climates. Guo et al*. (2011) used a molecular clock technique to come up with possible divergence times for the different racerunner clades and suggested that the viviparous clade might have originated at about the same time as the Tibetan Plateau expanded and affected climates in east Asia and elsewhere.

Przewalski is – so I’ve been told – pronounced something like ‘shu-val-ski’. Any further pronunciation tips appreciated.

Caption: Striped racerunner E. lineolata, a species named from Iran but which occurs across Central Asia as far north-east as southern Kazakhstan. Image: Yuriy75, CC BY-SA 3.0, original here.

How are racerunners related to other lacertids? The basic structure of the lacertid tree – we currently think – involves an early split between Gallotiinae and a large clade that contains all the other taxa, generally (but not universally) called Lacertinae. Within the latter clade, the long-tailed Takydromus grass lizards might (Fu 1998, 2000, Harris et al. 1998, Pavlicev & Mayer 2009) be outside a clade that includes (1) Lacerta and its mostly Eurasian kin (but see Mayer & Pavlicev 2007), and (2) the mostly African clade that contains Acanthodactylus and the racerunners. Arnold (1989) was first to establish this basic framework and it’s since been generally supported by subsequent studies. That second, ‘mostly African’ clade is Eremiadini or Eremiinae or Eremiainae or Eremiadinae, ugh... as noted above, Eremiadini and Eremiadinae are apparently the versions that are technically correct (Arnold et al. 2007).

**Caption:** sternum, sternal ribs, interclavicle and clavicles of *Acanthodactylus* (from Arnold 1989). Note the heart-shaped sternal fontanelle (other lacertids have round fontanelles). I wonder why these lizards have sternal fontanelles in the first place? Image: Arnold (1989).

Arnold (1991) supported a hypothesis of relationships where Eremias is part of what he termed the “advanced Saharo-Eurasian lacertids”: the clade that also includes Acanthodactylus, Mesalina and Ophisops. These lizards share an unusual internal stiffening system within the hemipenis termed an armature and hence have also been termed the ‘armatured clade’ on occasion (Harris et al. 1998). Pedioplanis was suggested to be the sister-taxon of the armatured clade, and both it and the members of the armatured clade share a number of derived anatomical characters (like a sternal fontanelle that’s often heart-shaped, and a reduced quadratojugal process on the jugal). Pedioplanis is from southern Africa and all the outgroups to this Pedioplanis + armatured clade (Meroles, Aporosaura, Ichnotropis, Heliobolus, Latastia, Philochortus and Nucras) are African too (Arnold 1991). We thus seem to have an ‘out of Africa’ model in this eremiadin group, lineages within the clade migrating out of Africa and into Eurasia once or several times.

Or do we? Fu (1998) found racerunners and the mostly Eurasian Ophisops to form a clade that was the sister-group to the rest of the ‘Saharo-Eurasian’ clade. Given that other lacertid lineages are Eurasian, and thus that this ‘Saharo-Eurasian’ clade must have dispersed into Africa from Eurasia at some point, this position would suggest that racerunners (and Ophisops) are primitively Eurasian, not recent invaders from Africa.

A molecular analysis that combined data from several genes also supported this model, though this time Eremias was closer to Adolfus than it was to Ophisops, Meroles, Pedioplanis, Latastia or Heliobolus, and Acanthodactylus and Mesalina were a long way away (albeit still in a ‘Saharo-Eurasian’ clade) (Fu 2000).

**Caption:** a substantially simplified lacertid phylogeny, showing the approximate structure pieced together in assorted studies. *Gastropholis* by Darren Naish, *Acanthodactylus* by Richard Hing, *Eremias* by Yuriy75 (CC BY-SA 3.0; original **here**), *Taky**dromus* by Acapella (CC BY-SA 3.0; original **here**), *Lacerta* by Darren Naish, *Gallotia* by Petermann (CC BY-SA 3.0; original **here**), *Psammodromus* by Wolfgang Wüster. Image CC BY-SA. This image was previously used in my **2020 article on another eremiadin, *Gastropholis*.**

That same study also includes a very different cladogram (based on data from the cytochrome oxydase I gene [COI] alone) where racerunners are shown as being part of a Eurasian clade that otherwise only includes Timon, Algyroides, Podarcis and Lacerta (sensu lato) (Fu 2000). I suppose the aim in Fu’s study was to see if single genes recovered the same relationships as did a combined analysis. Anyway, other molecular studies have found Eremias to be surrounded in the tree by African taxa (Harris et al. 1998, Mayer & Pavlicev 2007, Greenbaum et al. 2011, Kapli et al. 2011), so the ‘out of Africa’ model looks more robust than the 'primitively Eurasian' one. A lot could be said about the timing and precise details of lacertid biogeography: I have to ignore that whole area for now.

Arnold’s lament. Incidentally.... yes, lacertid diversity and phylogeny is confusing and ideas about the way the many constituent taxa might be related have varied substantially. One of the reasons that things are so difficult to resolve might be that the different lineages really did diversify very rapidly (Pavlicev & Mayer 2009). Among the most important studies prior to recent years are those of George A. Boulenger who published his monumental two-volume Monograph of the Lacertidae in 1920 and 1921. Arnold (1989) said the following: “This was Boulenger’s last task in a long and highly productive career. Even before the second volume of the monograph left the press, he quit the field, retired from the British Museum and returned to his native Belgium where he devoted his last years to studying roses. Anyone who has spent much time wandering in the confusing labyrinths of lacertid systematics will tend to sympathise” (p. 210).

Nick Arnold has been cited quite a few times in this article. If you’re familiar with his name, or his work, be sure to check out this 2024 Tet Zoo article on mite pockets.

**Caption:** where to go to find out about Eurasian lacertids? Here are the books that got me started: Arnold *et al*. (1992), Laňka & Vít (1986), and Anderson (1999). Note the very non-Eurasian frog and snake shown on the cover of Laňka & Vít!

I said earlier that I would only be talking about Eremias in this article and, despite having written over 3000 words on them so far, there’s an awful lot more that could be said. But as a wise and responsible Tet Zoo reader, you’ve surely been thinking “but what about all those other eremiadin genera that you’ve mentioned and not discussed at length?”. Yes, there are lots more eremiines eremiadines eremiadins to talk about. One day, one day...

Tet Zoo now features some fairly reasonable coverage of squamate diversity... but there is still so much to do. For previous Tet Zoo articles on lacertids (linking here to wayback machine versions to avoid issues with the hosting sites, SciAm in particular), see…

The New Forest Reptile Centre, August 2012
It’s high time you were told about Psammodromus, July 2013
Tale of the Takydromus, September 2013
The Mostly Arboreal African Gastropholis Lizards, February 2020

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for Tet Zoo and the more productive I can be on those long-overdue book projects. Thanks!

Refs - -

Anderson, S. C. 1999. The Lizards of Iran. Society for the Study of Amphibians and Reptiles, Saint Louis.

Arnold, E. N. 1989. Towards a phylogeny and biogeography of the Lacertidae: relationships within an Old-World family of lizards derived from morphology. Bulletin of British Museum of Natural History (Zoology) 55, 209-257.

Arnold, E. N. 1991. Relationships of the South African lizards assigned to Aporosaura, Meroles and Pedioplanis (Reptilia: Lacertidae). Journal of Natural History 25, 783-807.

Arnold, E. N. 2004. Overview of morphological evolution and radiation in the Lacertidae. In Pérez-Mellado, V., Riera, N. & Perera, A. (eds) The Biology of Lacertid Lizards. Evolutionary and Ecological Perspectives. Institut Menorquí d’Estudis. Recerca 8, 11-36.

Arnold, E. N., Arribas, O. & Carranza, S. 2007. Systematics of the palaearctic and oriental lizard tribe Lacertini (Squamata: Lacertidae: Lacertinae), with descriptions of eight new genera. Zootaxa 1430, 1-86.

Arnold, E. N., Burton, J. A. & Ovenden, D. W. 1992. Reptiles and Amphibians of Britain and Europe. Collins, London.

Fu, J. 1998. Toward the phylogeny of the family Lacertidae: implications from mitochondrial DNA 12S and 16S gene sequences (Reptilia: Squamata). Molecular Phylogenetics and Evolution 9, 118-130.

Fu, J. 2000. Toward the phylogeny of the family Lacertidae – why 4708 base pairs of mtDNA sequences cannot draw the picture. Biological Journal of Linnean Society 71, 203-217.

Gherghel, I., Strugaru, A. & Glavan, T. 2007. Eremias arguta deserti (Reptilia: Lacertidae) is not extinct from Romanian Moldavia. North-Western Journal of Zoology 3, 115-120.

Greenbaum, E., Villanueva, C. O., Kusamba, C., Aristote, M. M. & Branch, W. R. 2011. A molecular phylogeny of Equatorial African Lacertidae, with the description of a new genus and species from eastern Democratic Republic of the Congo. Zoological Journal of the Linnean Society 163, 913-942.

Guo, X.-G., Chen, D.-L., Wan, H.-F. & Wang, Y.-Z., 2010. Review of systematics of the racerunner lizard (Lacertidae: Eremias). Sichuan Journal of Zoology 29, 665-672.

Guo, X.-G., Dai, X., Chen, D., Papenfuss, T. J., Ananjeva, N. B., Melnikov, D. A. & Wang, Y. 2011. Phylogeny and divergence times of some racerunner lizards (Lacertidae: Eremias) inferred from mitochondrial 16S rRNA gene segments. Molecular Phylogenetics and Evolution 61, 400-412.

Harris, D. J., Arnold, E. N. & Thomas, R. H. 1998. Relationships of lacertid lizards (Reptilia: Lacertidae) estimated from mitochondrial DNA sequences and morphology. Proceedings of the Royal Society of London B 265, 1939-1948.

Jin, S.-D., Han, S.-w., Shin, H. C., Paik, I.-H., Paek, W.-K., Lee, H. & Kim, I.-K. 2013. Phylogeographical analysis of Eremias argus in Yoobu-do Island and Sohwang Sand Dune, Korea. Journal of Asia-Pacific Biodiversity 6, 455-458.

Kapli, P., Poulakakis, N., Lymberakis, P. & Mylonas, M. 2011. A re-analysis of the molecular phylogeny of Lacertidae with currently available data. Basic and Applied Herpetology 25, 97-104.

Kim, J.-K., Song, J.-Y., Lee, J.-H. & Park, D. 2010. Physical characteristics and age structure of Mongolian racerunner (Eremias argus; Lacertidae; Reptilia). Journal of Ecology and Field Biology 33, 325-331.

Laňka, V. & Vít, Z. 1986. Amphibians and Reptiles. Hamlyn, Twickenham.

Li, H., Ji, X., Qu, Y., Gao, J. & Zhang, L. 2006. Sexual dimorphism and female reproduction in the multi-ocellated racerunner (Eremias multiocellata) (Lacertidae). Acta Zoologica Sinica 52, 250-255.

Li, H., Qu, Y.-F., Ding, G.-H. & Ji, X. 2011. Life-history variation with respect to experienced thermal environments in the lizard, Eremias multiocellata (Lacertidae). Zoological Science 28, 332-338.

Mayer, W. & Pavlicev, M. 2007. The phylogeny of the family Lacertidae (Reptilia) based on nuclear DNA sequences: convergent adaptations to arid habitats within the subfamily Eremiainae. Molecular Phylogenetics and Evolution 44, 1155-1163.

Moravec, J. 1994. A new lizard from Iran, Eremias (Eremias) lalezharica sp. n. (Reptilia: Lacertilia: Lacertidae). Bonner Zoologische Beiträge 45, 61-66.

Mozaffari, O. & Parham, J. F. 2007. A new species of racerunner lizard (Lacertidae: Eremias) from Iran. Proceedings of the California Academy of Sciences 58, 569-574.

Pavlicev, M. & Mayer, W. 2009. Fast radiation of the subfamily Lacertinae (Reptilia: Lacertidae): history or methodical artefact? Molecular Phylogenetics and Evolution 52, 727-734.

Pouyani, E. R. 2009. The phylogeny of the Eremias velox complex of the Iranian Plateau and Central Asia (Reptilia, Lacertidae): molecular evidence from ISSR-PCR fingerprints. Iranian Journal of Animal Biosystematics 5, 33-46.

Szczerbak, N. N. 1974. Yaschurki Palearcktiki. [The Palearctic Desert Lizards] Axadeimiya Nauk Ukrainskoi SSR Institut Zoologii, Kiev.

Song, J.-Y., Chang, M.-H. & Koo, K.-S. 2013. Estimating the size of a Mongolian racerunner Eremias argus (Squamata: Lacertidae) population at Baramarae Beach, Taeanhaean National Park. Korean Journal of Herpetology 5, 9-13.

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Having recently republished an old, classic article on Speculative Zoology – that based around a 2014 interview with After Man creator Dougal Dixon – it seemed appropriate to republish another Tet Zoo classic...

I’m referring here to my 2015 article on Marc Boulay and Sébastien Steyer’s 2015 book *Demain, les Animaux du Futur*, originally outed at Tet Zoo ver 3, but now republished here, in revamped and updated form…

As will be obvious if you read the previous Tet Zoo article – or if you’re familiar with Dougal Dixon’s several books on the subject – speculative zoology is very much a ‘visually driven’ field, one where the look of the animals is important. And it’s in that vein that Boulay and Steyer’s Demain, les Animaux du Futur excels. First things first: this book is in French, and an English-language version does not exist, at least not to my knowledge. Back in 2015, efforts were made to get one produced but ultimately fell through. Whatever, Spec Zoo fans should obtain the volume for its illustrations alone. It has been the subject of an enormous amount of interest in France and I've seen numerous relevant interviews and articles in magazines, newspapers, and on TV.

**Caption:** once again, we have to give credit to *After Man* (Dixon 1981) as the foundational work behind this newer one. At right, what appears to be a new, 2025 Japanese edition of *Demain, les Animaux du Futur* (**from here**).

By total coincidence, I’ve only just discovered that a Japanese edition exists. I initially thought that this was brand new for 2025 but now know that it was published in 2017. You’ll recall from the Dougal Dixon article that Japan has a huge audience of Spec Zoo followers, meaning that publishers and broadcasters take it more seriously than they do elsewhere. Anyway… here’s hoping that an English language version does get the green light at some point. The book’s title – Demain, les Animaux du Futur – doesn’t translate especially well into English. Literally, it’s Tomorrow, the Animals of the Future, but I suspect it should be imagined as something more like ‘The Animals of Tomorrow’.

Co-authored and illustrated by artist Marc Boulay and palaeontologist Sébastien Steyer’s Demain, les Animaux du Futur is beautiful and lavishly illustrated in colour throughout, featuring spectacular renditions and diagrams of its hypothetical future creatures. The setting is the Early Neozoic of 10 million years in the future, a time in which humans are long-extinct, the 6th Great Extinction has been and gone, and a variety of new species inhabit the oceans and landmasses. The authors explain that they chose this time frame because it’s distant enough to allow sufficient novelty, is not so distant that truly different organisms might have evolved, and also represents a section of time mostly unexplored by previous authors (Boulay & Steyer 2015). On that note, it is implied that the work might exist in the same universe as certain other Spec Zoo projects.

**Caption:** life-sized models of two 'Dixonian epoch' animals on show at the at the Institut Royal des Sciences Naturelles de Belgique (IRSNB), Brussels, and photographed in 2010. At top, the giant penguin *Neopygoscelis* and, below it, the flightless pelagic petrel *Propellonectes*. *Neopygoscelis* here is shown as grey, albeit countershaded, but the version in *Demain* has a more penguin-like livery. I published a Tet Zoo article on these models (and others in the same exhibit) can no longer find an intact version. Images: Darren Naish.

The theme explored here is, I hardly need say, similar to that of Dougal Dixon’s After Man (Dixon 1981; Dixon 2021) and The Future Is Wild (Dixon & Adams 2004). Early sections of the book discuss this history (credit too is given to the Russell and Séguin dinosauroid; for more on that see this Tet Zoo article from 2021), and the authors explain how they were inspired by these works in crafting their own vision of the future. Incidentally, the genesis of their project is, I presume, reflected in the Institut Royal des Sciences Naturelles de Belgique’s installation on the ‘Dixonian epoch’, since that includes certain creatures that feature in the book (Boulay & Steyer 2015).

**Caption:***Demain* does a good job of setting the version of Spec Zoo it follows within its correct historical perspective. As you can see from the pages shown here, there’s fair coverage of Dougal Dixon and *After Man* and of Russell and Séguin’s dinosauroid. Image: Boulay & Steyer (2015).

In contrast to a few other books that focus on speculations about the lives of animals (I’m thinking here of the Walking With books in particular), the text doesn’t revolve around ‘day in the life’ style storytelling, which is good news as far as I’m concerned. Instead, the text is mostly devoted to describing the world we’re immersed in, and of the biology, ecology and evolutionary history of the organisms we meet (Boulay & Steyer 2015). Profile sections introduce the relevant species and describe facts about their biology, ecology and anatomy. They have technical scientific names, some of which include references to the Alien films, Blade Runner and Daft Punk. H. G. Wells, Gert van Dijk (of Furaha), palaeoartist Zdeněk Burian and others are commemorated too.

**Caption:** night-vision scene from Boulay & Steyer (2015) showing flightless birds interacting on the Euroafrican savannah, *Necropteryx* at left, *Tyrannornis* at right. Image: Boulay & Steyer (2015).

Full-colour images feature on virtually every page of Demain, les Animaux du Futur. There are also diagrams illustrating evolutionary sequences and numerous details of anatomy. The CG, photo-real artwork is spectacular and many of the complaints so often levelled at CG depictions of animals do not apply. The animals fit into their environments, lack crass efforts at motion-blurring, and are well colour-matched and correct in terms of ‘heat’ and shadow relative to the surroundings. Many of the images are memorable and extremely clever. One of my favourites is a night-vision scene showing an altercation between the carrion-eating, flightless raptor Necropteryx and the giant parrot Tyrannornis.

The animals of this world are a fascinating lot and I like a great many of them, finding them mostly plausible and anatomically sensible. The illustrations make them look very ‘real world’: some of the birds have frayed, gnarly keratin on their crests and bills, ruffled plumage and other details. These show that the artist has been paying a great deal of attention to what animals really look like.

**Caption:** a pair of *Neopygoscelis*, a large, deep-diving, pliosauromorph penguin. Soaring over the ocean above we see a giant bat, on which more below. Image: Boulay & Steyer (2015).

Creatures of the Neozoic oceans. There are in fact so many animals designed for the book that there are substantially more than I can write about or mention. We start off with the marine fauna. As can be predicted given the probable state of near-future conditions, over-fishing, pollution, acidification and warming have resulted in a collapse of the modern marine fauna. Numerous animal groups have disappeared, including several important plankton groups and many marine vertebrate lineages. A different ecosystem and set of species have arisen, the majority of biological wealth being benthic and well below the warm waters of the surface (Boulay & Steyer 2015).

Tiny catfish with bioluminescent lures are super-abundant and occupy the role taken in our time by such species as krill. Diverse squid – some of which are superficially fish-like, others of which are gigantic (Rhombosepia is 10 metres long) – occupy middle- and top-tier roles occupied in modern times by bony fish, sharks and cetaceans. Even tetrapod devotees have to grudgingly admit that cephalopods have enormous potential, especially in a world with massively reduced vertebrate diversity.

**Caption:** modern plankton are under all kinds of pressures, and the suggestion has been made several times that future oceans might be extremely different, ecologically, as a consequence. *Demain* proposes that the upper waters of the ocean are highly depauperate as a consequence, but that a radiation of new, miniature catfishes inhabit the depths. Image: Boulay & Steyer (2015).

Several flightless seabirds are present. The large, deep-diving penguin Neopygoscelis has inevitable similarities with the big pelagic penguins of After Man (Dixon 1981, Dixon 2021) and again hints at the idea that penguins might have the evolutionary potential to give rise to forms that resemble cetaceans or short-necked plesiosaurs. Penguins have been a mainstay of Spec Zoo projects even since After Man. The fact, however, that penguins are not doing well in a rapidly warming world – virtually all species are rapidly declining, even relatively abundant ones like the Adélie Pygoscelis adeliae (the global population is over 10 million, but entire colonies are disappearing as climatic warming is making their breeding islands unsuitable) – means that we might be sceptical of these ideas. I’m not the first to say this when discussing Spec Zoo scenarios.

**Caption:** news about penguin decline, about the loss or shrinking of breeding colonies, constantly makes the headlines. Most of the findable articles are about the big *Aptenodytes* penguins (the King and Emperor) and there’s less publicity on pygoscelids (the small Adélie and kin). However, they’re in trouble too, though the global situation is complex enough (colonies disappear *here*, but new ones appear *there*) that it can be hard to form a consensus.

Demain also features large, superficially hesperornithine-like flightless petrels (Propellonectes), though I find it ironic that they descend from one of the most terrestrial of petrel lineages.

Perhaps the most surprising creature of the Neozoic oceans is Benthogyrinus – a wing-headed, super-sized, suspension-feeding beast, 40 metres long (Boulay & Steyer 2015). Assorted arthropods and cephalopods lived on its surface. It might best be described as a gargantuan, armour-plated tadpole-whale. And that's an apt description, as it’s a paedomorphic, sea-dwelling frog. A descendant of Xenopus. In order to imagine such a beast, we have to envision frogs undergoing major developmental, physiological and morphogenic shifts. Contra popular wisdom, certain modern frogs do have some tolerance of salinity and can even swim in the sea; nevertheless, to account for the other changes we would need massive changes in anuran biology. But… hey, this is what speculative zoology is all about and I, for one, welcome our new gargantuan marine tadpole monster overlords.

**Caption:** why anurans have never evolved paedomorphosis whereas salamanders often has is an oft-asked question. The answer lies somewhere in the fact that sexual maturation in anurans is tightly linked to metamorphosis: they literally cannot reproduce while in larval form. *Demain* imagines that a solution to this has been found through evolution, and thus we have tadpole monsters. Image: Boulay & Steyer (2015).

Flightless birds: ‘new dinosaurs’. Moving now to the terrestrial realm, the large animal fauna is dominated by flightless birds, including modified ducks, waders, raptors, parrots, hummingbirds and crows. Many have converged on a thick-legged, shaggy-plumaged, ratite-like morph, meaning that terrestrial habitats are full of creatures that look something like giant kiwis and cassowaries. These are among my favourite animals of all, so I especially liked these designs.

Would gigantic descendants of geese, raptors and so on all really end up looking ratite-like? Well, it’s an important point that birds might be somewhat constrained in potential with respect to overall shape and form... can they do anything truly innovative? I don't want to give everything away, but I will say that the book includes quadrupedal birds (Boulay & Steyer 2015).

**Caption:** at left, ontogeny of the giant, herbivorous, quadrupedal goose *Giraffornis vandijki*. It grows quickly (which is in keeping with its goose ancestry), reaching 6 m in height for males, and is named in honour of Dr Gert van Dijk. The authors note that it’s somewhat reminiscent of sauropods. Image: Boulay & Steyer (2015).

Hadrornis and Giraffornis are giant, long-necked anseriforms. Necropteryx is a large, flightless descendant of the Lammergeier or Bearded vulture Gypaetus barbatus. It absolutely looks the part, with its long, pseudo-toothed bill (it's the creature depicted on the upper part of the cover). Tringapterus is a vaguely cassowary-like, sexually dimorphic flightless wader with a dagger-like bill. Tyrannornis rex – the T. rex of the piece – is a gargantuan pseudo-toothed parrot (and one of several species in this genus).

Neoviraptor is a short-faced flightless corvid that specialises on eating eggs. On that last point, a speculative animal can be specialised for whatever its designers wish, but did they base this idea on the supposed diet of Cretaceous oviraptorid theropods? The possibility that oviraptorids ate eggs still exists, but it looks more likely that they were predominantly herbivorous, or omnivores that consumed a large amount of plant material.

**Caption:** head-shot of male *Tyrannornis rex* (females are less gaudy and without the casque), a giant flightless parrot. I quite like this design but find the colour scheme a bit too reminiscent of certain macaws. The again, maybe this is ok for a forest-dwelling predator that interacts with species that have excellent colour vision. Image: Boulay & Steyer (2015).

We thus see a radiation of ‘new dinosaurs’, and it’s quite easy to see parallels – or, specifically, cases of evolutionary convergence – between these birds and the theropods of the Mesozoic. This is not ignored by the authors (Boulay & Steyer 2015). It again raises a very interesting possibility: as long are birds are around, it’s almost as if the world could once again become the domain of dinosaurs of a sort mostly associated with the Mesozoic. Not sauropod- or tyrannosaur-mimics, but of fuzzy, beaked theropods much like those that first appeared more than 100 million years ago.

I must mention the fact that Demain includes some smaller birds as well, including the eusocial, burrowing mole-bird Talpidornis, a highly modified weaver that builds massively complex burrow systems. How does it compare to other speculative burrowing birds? It does not look much like the spink of The Future is Wild but does look quite like a bird drawn whimsically by Gabriel Ugueto!

**Caption:** at left, the mole-like bird *Talpidornis sechani* from *Demain*. These Eurafrican birds have a caste system, with a large, long-lived queen, formidably armed soldiers, and smaller workers, explorers and nurses or nannies. At right, inspired by a 2019 discussion, Gabriel Ugueto illustrated a hypothetical ‘mole-bird’, and it’s interesting that his version is actually quite *Talpidornis*-like. Images: Boulay & Steyer (2015); Gabriel Ugueto.

A world of future bats. Mammals are still present in the Early Neozoic but are only represented by bats. The authors emphasise how this enormous, diverse and globally distributed group (remember: accounting for 20% or more of all mammal species today) could have an incredible future, assuming, that is, that they’re not too severely afflicted by anthropogenic changes. On that note, some studies have looked at the possible fate of bat diversity in tropical ecosystems in view of environmental and climate change. The news isn’t good, since diversity is expected to plummet and communities at the species level will homogenize (e.g., Meyer et al. 2016, Alroy 2017, Gonçalves et al. 2021).

**Caption:** the enormous, highly specialized sky-bat *Gigapterus troposherus* – a descendant of free-tail bats (molossids) – cruising at altitude. These animals look even more specialized for soaring than the most specialized of soaring pterosaurs and birds. Image: Boulay & Steyer (2015).

Then again, maybe it can be argued that the species integral to the evolutionary scenarios explored here will persist and thus have potential to do what they do in the book: at least some of the bats here descent from the highly abundant free-tailed bat Tardarida (Boulay & Steyer 2015). Some very, very interesting things have happened. An entirely new group of giant sky-bats have evolved and have developed a suite of innovations that allow them to fly, live and hunt at altitude.

Some of the species we’re introduced to are boldly patterned, mega-winged giants with wingspans of up to 15 metres. Giant lungs, an air-sac system and a pouch used to contain the young have evolved (Boulay & Steyer 2015). The bats also have a novel wing form. Many have incredibly high-aspect wings, and additional membranes and patagial lobes are present in some species. The hindlimbs are reduced to near-absence in the Velocipterus species, some of which look like long-winged, mammalian butterflies.

Whether bats might ever evolve pneumaticity, giant size and an ability to give birth on the wing are all questions asked at one time or another by those interested in bat evolution. They haven’t happened yet, but is there the potential for such features to evolve? These speculations are certainly thought-provoking.

**Caption:** the big, flightless, quadrupedal bat *Nesferapoda*, from Boulay & Steyer (2015). This image was borrowed from Gert van Dijk’s **review of the book from Furahan Biology & Allied Matters**. *Demain* includes several additional images of this animal. Image: Boulay & Steyer (2015).

Flightless bats. Yes, there are flightless bats here too. Fans of speculative zoology will immediately be thinking of Dixon’s Night stalker. Nesferapoda of the Neozoic is not especially similar. It’s a direct descendant of Desmodus and is a thick-limbed quadruped. As I said above, an interesting thing about the book is that the authors don’t discuss their creatures within an in-universe context only (as is the style for works like this); rather, they make proper reference to the ideas that have gone before, and they note how speculative terrestrial bats – including the Night stalker and the Future Predator of Primeval – have appeared before in the works of others.

Flightless bat are – much like those giant penguins – ‘classic’ beasts of the genre and, as the authors note, a certain number of constraints require that speculative bats can only really end up heading in a few given evolutionary directions (Boulay & Steyer 2015). On bats in general, I’m thrilled that this book explores so many new ideas on this group, I think for the first time in print.

Arthropods, echinoderms, plants. There’s more. A large Scolopendra centipede that has evolved a gliding ability is one of the most terrifying animals in the book. There are also large marine echinoderms – the predatory ‘terminator urchin’ Neocidaris schwarzenheggeri (erm, typo: that specific name should be schwarzeneggeri) – and crustaceans that ride aboard the giant Benthogyrinus.

And then there are plants. It’s all too tempting when creating a future world to put your future animals into an essentially modern flora. But plants are just as dynamic and changeable as animals – if not more so – and I respect the fact that Boulay & Steyer (2015) went to the trouble of thinking about, and designing, these crucial components of their world.

**Caption:** Marc Boulay in what I assume is his office space, with excellent promotional t-shirt. And what is that model?… I would love to know! Image: Ouest-France / Philippe Renault.

The big picture. There’s much more to the Neozoic world than outlined in this review. A section at the end discusses tectonics, climate and other big-picture stuff, while the book ends by looking at the creative process that led to the creation of the Neozoic world.

Echoing sentiments made previously here at Tet Zoo, it should be said that speculative zoology is not mere frivolous fun but inspires us to think about the shape of evolution, about the impact we’re having on the world and its fauna, and about the possibilities and limitations that exist as regards the organisms around us today. Remember that Dougal Dixon’s remit when designing After Man (Dixon 1981; Dixon 2021) was to discuss factual processes via speculative creatures (Thomann 2024). Overfishing, overhunting, pollution, climate change, human population expansion and a myriad other things may paint a view of the future somewhat bereft of hopefulness, but a future exists nonetheless and even a future with a depauperate fauna is still a future with a fauna.

Marc Boulay and Sébastien Steyer have succeeded in creating a fantastic and compelling view of a future that is not provincial or small in scope. It is a grand, vibrant, diverse vision of imaginary life, and it looks amazing. I’m sure that their book will inspire a new generation of fans and thinkers who will now become turned-on to the world of Dixonian fiction and discover the exciting works that already populate the genre.

**Caption:** Boulay’s scene showing part of ‘The infinite mangrove’ (though flipped horizontally relative to the one in the book). Note the bats and speculative vegetation. The CG render is really nice. Image: Boulay & Steyer (2015).

If you’re a Spec Zoo fan, you really must get hold of this book. And – again – here’s hoping that an English language version can be published, in which case this book will be bought to the attention of a new and large audience.

Marc Boulay and Sébastian Steyer. 2015. Demain, les Animaux du Futur. Softback, index, pp. 157. €23 $58 £15.87. Éditions Belin, Paris. ISBN 978-2-7011-5886-0. *Here at amazon.Here at amazon.co.uk.*

For previous Tet Zoo articles on speculative zoology, see...

Oh no, not another giant predatory flightless bat from the future, March 2007
Come back Lank, (nearly) all is forgiven, September 2008
Giant flightless bats from the future, November 2012
Of After Man, The New Dinosaurs and Greenworld: an interview with Dougal Dixon, April 2014
The LonCon3 Speculative Biology event, August 2014
Speculative Zoology, a Discussion, July 2018
The Dougal Dixon After Man Event of September 2018, September 2018
Dougal Dixon’s After Man, the Initial Pitch Document, June 2020
Speculative Zoology and the World of After Man; an Interview With Dougal Dixon, April 2025

Refs - -

Alroy, J. 2017. Effects of habitat disturbance on tropical forest biodiversity. Proceedings of the National Academy of Sciences 114, 6056-4516061.

Boulay, M. & Steyer, S. 2015. Demain, les Animaux du Futur. Éditions Belin, Paris.

Dixon, D. 1981. After Man: A Zoology of the Future. Granada, London.

Dixon, D. 2021. After Man: A Zoology of the Future (40th Anniversary edition). Breakdown Press, London.

Dixon, D. & Adams, J. 2004. The Future Is Wild: A Natural History of the Future. Dorling Kindersley, London.

Gonçalves, F., Sales, L. P., Galetti, M. & Pires, M. M. 2021. Combined impacts of climate and land use change and the future restructuring of Neotropical bat biodiversity. Perspectives in Ecology and Conservation 19, 454-463.

Meyer, C. F. J., Struebig, M. J. & Willig, M. R. 2016. Responses of tropical bats to habitat fragmentation, logging, and deforestation. In Voigt, C. C. & Kingston, T. (eds) Bats in the Anthropocene: Conservation of Bats in a Changing World. Springer, Cambridge, pp. 63-103.

Thomann, V. 2024. A “speculation built on fact”: on Dougal Dixon’s zoology of the future. In Castle, N. & Champion, G. (eds) Animals and Science Fiction. Springer Nature, pp. 345-362.

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Regular readers of Tet Zoo – and that group includes, by definition, people whose interests are similar to mine, hello! – will be aware of my affinity for what we now term Speculative Zoology, or Spec Zoo. That’s a topic that’s been covered here (and in previous incarnations of the blog) quite a few times, as you can see from the links below. And within that field, one person above all others has been foundational and worthy of repeated visitation.

**Caption:** in a scene from the future world of *After Man*, giant predatory rats harry a rabbuck. If you know the look of the animals in the final book, note how different these versions are… Dougal Dixon’s original concepts were, in cases, not exactly like the final animals ultimately illustrated by other artists. Image: (c) Dougal Dixon, used with permission.

I refer, of course, to British writer, author, visionary, artist and designer Dougal Dixon, generally and correctly regarded as the ‘father’ of modern speculative zoology… even though he denies this accolade in his own writings (Dixon 2021). Exactly where speculative zoology (and its more inclusive parent, speculative biology) fits in terms of genre has been relatively under-analysed but the fact that it involves an intersection of natural history writing, palaeontological knowledge, evolutionary theorising, imagination and artistry, and might also require the trappings of science fiction (Naish 2014, Thomann 2024) requires that relatively few have indulged in it.

**Caption:** a selection of Dougal Dixon Spec Zoo books in the Tet Zoo Towers library. The manga *The New Dinosaurs* and *The Future is Wild* volumes visible at left are especially hard to get here in the UK. Image: Darren Naish.

Dougal has been the go-to person on speculative zoology ever since the 1981 publication of his famous, beautifully illustrated and extraordinarily successful book After Man (Dixon 1981). Therein, we explore the biogeographical realms of our planet 50 million years in the future: an Earth where the continents have moved, many modern animal groups have disappeared, where large flightless bats inhabit islands, and where rabbits, rats, corvids and mongooses have given rise to a new assemblage of megafauna that also includes novel pigs, antelopes and marsupials. Giant swimming rodents, flightless auks and cetacean-like descendants of penguins inhabit the oceans (Dixon 1981).

**Caption:** there’s a lot of *After Man* fan-art out there, and I’ve created a bit of it myself over the years. This image (created for a *Fortean Times* article on speculative zoology) shows a Giantala (spotted animal at left), Zarander (tusked and trunked, lower left), a giant raboon (the maned biped), Gigantelope (far right) and (in the tree) a Slobber. I misunderstood the relative sizes of certain of these animals when creating this image and they’re not to scale. Image: Darren Naish, colouring by Rebecca Groom.

Published in multiple languages and countries worldwide, After Man was – and in ways still is – nothing short of a sensation, resulting in more than one dedicated TV show, a travelling exhibition featuring life-sized animatronics, and a worldwide following of fans and aficionados. After Man didn’t merely showcase Dougal’s ideas: the animals – all of them – originated via his own copious drawings and paintings, a fact not as well known as it should be given that the final art featured throughout the book is by others, among them Diz Wallis and Philip Hood.

After Man needed a follow-up and this was provided later in the decade by The New Dinosaurs (Dixon 1988), a book that told the story of a parallel Earth where the end-Cretaceous extinction event never occurred. Again, Dougal designed (and drew) the initial versions of the creatures, while the final art is by Amanda Barlow, Andy Farmer, Philip Hood, Denys Ovenden and several others. If you know British natural history books of the relevant era, some of these names will be familiar. A third volume – Man After Man (Dixon 1990) – appeared soon after, this one being a fantastic look at a speculative future for the descendants of humanity.

**Caption:** *After Man* has had several different covers over the years, and here’s what the 2021 40th Anniversary version looks like. **Buy it here from Breakdown Press.** *After Man* might be back in print… what about 1988’s *The New Dinosaurs*? Well…

Moving to the 21st century, Dougal was also involved in the TV series The Future Is Wild and coauthored the accompanying book (Dixon & Adams 2004). And while the After trilogy might be… well, a trilogy… was this ever the intention? Evidently not, because a project connected to After Man – Greenworld – came to fruition in the 2000s, though has (so far) only seen publication in Japan (Dixon 2010). More on that matter below (and see the relevant section on Dougal’s website here).

Dougal was born in Scotland but lives in Dorset in southern England, and thus in close proximity to me. We’ve met on numerous occasions (in recent years Dougal has been a regular attendee of TetZooCon), and I’ve rarely failed to pester him with questions about After Man and his various other projects. Back in August 2013, I opted to take advantage of this situation, and Dougal very kindly agreed to be interviewed exclusively for Tet Zoo. As you’ll see below, we covered the back-story to After Man, discussed the inspirations behind certain of his creatures and the way the accompanying artwork came to be, and also spoke about projects that were, at the time, not well known in Europe or the Americas. That interview was published in 2014 (the original can be seen here at wayback machine)… a lot has happened since.

**Caption:** I’ve met Dougal Dixon many times now and he’s spoken and presented at my TetZooCon events a few times. At left, here’s (left to right) Gert van Dijk (of **Furahan Biology and Allied Matters**), Darren Naish, Dougal Dixon. At right, Dougal and myself in London for the 2018 *After Man* event.

After Man in the 21st century. After Man saw republication via Breakdown Press in 2018 (Dixon 2018) and here’s where it became clear just how much ‘backstory’ material Dougal still owned. At a launch event hosted at London’s Conway Hall in the September of 2018, Dougal brought with him the initial pitch document, a huge number of his own initial sketches, a number of 3D models, and much else besides. I was blown away by this and wrote about it here at Tetrapod Zoology; I then cheekily borrowed the aforementioned pitch document and wrote about that too. It was obvious that there was massive public and fan interest in this ‘backstory’ material… would we ever see any of it in print?

Well, 2021 was the 40th anniversary of After Man’s initial publication, so here was the opportunity to republish After Man again. The result (again, from Breakdown Press) was a special, expanded version of the book, with a new Introduction and a substantial ’40 Years of After Man’ section (Dixon 2021). This includes copious ‘backstory’ matter, much in the way of previously unpublished art and imagery, and reproduced press material that publicised the book back in the 80s. The 2014 publication of my interview with Dougal and 2018 article on the initial pitch document played, I think I can say, some minor role in the release of this new version but, whatever, it’s a real treat for fans of Dougal’s work and you must be sure to get a copy for yourself even if you own the 1981 original.

**Caption:** more *After Man* fan-art, this time a very nice montage showing some of the book’s larger animals correctly scaled. It’s easy to forget how large some of them are meant to be. Image: Dragonthunders, from **here** (several other montage images showing scaled Dixonian creatures feature on the same page).

Greenworld still needs an English publisher. At the time of writing, rumours are afoot that new editions of certain others of Dougal’s Spec Zoo works will soon see publication as well. A modified version of Man After Man just saw print in Japanese. Dougal’s two-volume work on an alien world impacted by its human visitors – Greenworld – still needs publication in English and only parts of its complex story have been released outside of Japan so far. Inevitably, the tale of Greenworld – one where a lush, teeming biological paradise is filled with exotic alien organisms of numerous sort – recalls James Cameron’s Pandora of the Avatar franchise (first outed in December 2009), but that’s a coincidence and perhaps shows how alluring we find this vision to be. Greenworld and Avatar both function, of course, as parables of our impact on the natural world, an issue more pressing today than ever.

**Caption:** front cover of the 2024 Japanese edition of *Man After Man*. It is much modified relative to the original. Image: Darren Naish.

Anyway, the primary justification for the existence of the article you’re reading now is that I – perpetually frustrated that my old ScienceBlogs and Sci Am material is now unavailable, vandalized or even paywalled – recently decided to revamp and republish the 2014 interview text discussed above. There are a few statements of my own, made in the 2014 version, that I disliked or which are now wrong, so they’ve been modified or now have addenda. Without further ado...

Darren: First of all, do you realise how important and inspirational, how loved and cherished, the ‘After’ trilogy has been?

Dougal: Certainly. I get emails from established, professional people in the world of biology and palaeontology who say that my work inspired them 30 years ago. Some of these people were amateurs or beginners when they read the books.

**Caption:** over the years, numerous magazine articles have rehashed and summarised the main story points of *After Man*. The 1995 special issue of *Focus* include this feature on *After Man* (Anon. 1995), and it was surprising by showing illustrations novel to me at the time, like the predator rat bs rabbuck image.

  **Caption:** the 1995 *Focus* article also features various images based on those of *After Man* (like the family tree) or taken from it. That Vortex at right was novel and I haven’t seen it elsewhere. No author is credited for this piece so I have to cite it as ‘Anon. 1995’.

Darren: Do you follow the speculative zoology movement – are you aware of such things as SpecWorld (the Speculative Dinosaur Project) or the future creatures in Primeval?

Dougal: I have of course seen Primeval.

Darren: I always thought the Future Predator was inspired by your flightless bats (especially by the Night stalker of After Man).

Dougal: Oh yes, I’m sure that’s not a coincidence. I’ve been told that Tim Haines has a copy of After Man on his bookshelf.

Darren: Right. I asked him quite a few times about that (I worked at Impossible Pictures for a while a few years back) and he always cleverly evaded me. [UPDATE: while the Future Predator is canonically a bat today, the initial concept art and communications from the Impossible Pictures team show that it wasn’t initially any such thing. Instead, it started life as a vaguer, more reptilian predator. It’s not accurate to claim that it originated as an After Man homage or copy]. Moving on... can you remember (this is going back a long way!) what the original inspiration was for After Man?

**Caption:** the different editions of *After Man* now depict more than one version of the Night stalker. At left, the 1981 version; at right, the improved vision from Dixon (2018, 2021). It’s true that views differ on how ‘plausible’ it might be that bats would ever, or *could* ever, evolve in this direction, but I put it that the Night stalker remains an iconic vision in discussions about speculative bat evolution. Images: Diz Wallis, from Dixon (1981); Dougal Dixon, from Dixon (2018, 2021).

  **Caption:** photo – from the dustkjacket of *After Man* (Dixon 1981) – showing Dougal with his model of a Night stalker. Image: Duncan McNicol.

Dougal: As a child I was always struck by The Time Machine, by H. G. Wells, in particular by the part at the end where he goes to the far future and everything has changed: there are the giant crabs and so on. I was always artistic as a kid. I remember at the age of 8 or 10 years old, drawing comic strips, and one of them was actually my own retelling of The Time Machine. I had fun creating my own future creatures that had evolved from those of the modern day, featured there in the background. [UPDATE: for a more detailed retelling of these events, with relevant illustrations, see Dixon (2021).]

Darren: [incredulously] You came up with that stuff as a kid?

**Caption:** the giant, flightless, fully aquatic penguins of *After Man*, namely the mysticete-like Vortex *Balenornis vivipera*, and dolphin-like Porpin *Stenavis piscivora*. Image: Diz Wallis, from Dixon (1981).

Dougal: Well, yeah, not that there was any scientific point to it: it was just the background to the story. The next stage really – I was a teenager, so we’re in the 1960s now – would be when I was influenced by the conservationist movement that became important at the time. I remember in particular the ‘Save the tiger’ campaign. I was watching a TV programme about tiger conservation with my Dad, he turned to me and said “Why save the tiger? The tiger will become extinct. Everything becomes extinct, other things evolve”. And I thought – that’s a very unhelpful attitude, but it basically started me on a learning curve about evolution, and I essentially realised that he was right: everything does become extinct, other organisms develop to take their place. And I began to think to myself – just as idle musings – if things do become extinct, what evolves to take their place? Move on now to the late 1970s, I met a friend of mine I hadn’t seen from a long time and he was wearing a ‘Save the Whale’ badge. Well, that sparked off the whole idea again. I thought to myself: suppose the whale does become extinct, what might evolve to take its place? That, of course, is where the whole idea of the giant, whale-like penguins of After Man came from.

A couple of weeks later, I thought… I could use this. I could devise a popular-level book on evolution, but a book that did something quite different. Other popular-level books used evolution to tell the story of the past, but nobody had taken the whole process – the observable trends – and projected them into the future. I thought: I could do this. At this time, I was working in publishing – I was involved in encyclopedias – and I knew how to present a crazy idea to a publisher with a likelihood of being taken seriously. So I decided to create illustrated spreads, dummy text and so on… I took the idea to publishers the next time I was in London, and the rest is history.

**Caption**: one of several pages from the original BOOK PLAN for *After Man*, featured **here at Tet Zoo** but also featured in **Dixon (2021)**. Images: (c) Dougal Dixon.

Darren: Obviously the concept of After Man was a great idea that you’d had for a long time, but they (= the publishers) got it straight away? They understand the concept of the book you were aiming to create?

Dougal: Yes, I was very excited about the project, and there were lots of ideas about how it could be tied in to broader efforts concerning publicity. I was told by the publisher that a science-based radio programme on Radio 4 would be interested in covering the story behind the book, and that it would be Barry Cox reviewing it [Prof. C. B. Cox, then of King’s College London]. Well, when the review happened, let’s just say that it was a total demolition job. Cox hated it. And I thought… well, that’s it, I gave it my best shot. However, as other reviews began to roll in, things changed – certainly, the review in New Scientist changed the very low opinion I had of myself. BBC Wildlife and the Smithsonian’s magazine, they also did much to enhance the book’s credibility, and I also had publicity tours both in the UK and the USA.

**Caption:** *After Man* is one of so many books that I heard about, and read about, long before I ever saw a copy. The October 1981 issue of *Wildlife* magazine (the ancestor of *BBC Wildlife*) included an article devoted to the book, basically a long-form interview between Pamela Todd and Dougal Dixon (Todd 1981), and my secondhand copy of that included my first look at Dixonian future creatures. From Dougal’s point of view, publicity like this would have been golden.

Darren: It must have been very exciting. So far as I know, this was your first ‘big thing’.

Dougal: Yes. It made me think… there’s a future in this. That is, in popular-level books that use fictitious examples of factual processes, there’s definitely room for a few more. And that’s why I came up with the idea for The New Dinosaurs. Again, I wanted to do the same sort of thing but, this time, I was aiming to create a popular-level book on zoogeography, using fictitious examples to show what the dinosaurs might perhaps be like if they hadn’t become extinct.

Darren: That raises a question I have about that book. Were you coming up with the speculative animals first, or were you led by the zoogeography angle?

Dougal: It would have been a bit of both, I think (hard to remember all the details after all this time, of course). On balance, it was the pushing of the zoogeography – a fairly unfamiliar aspect to so many people – that was the main impetus here. I often wonder, however, with these books whether the people that look at them simply see them as picture books of funny animals.

Darren: I think that’s definitely true, to a degree, for some people... but not for the ones that matter! Quite a few people have said that some of the ideas that feature in The New Dinosaurs presaged modern discoveries. You’ve got small, tree-climbing theropods, fuzzy integument on dinosaurs, striding, terrestrial pterosaurs…

**Caption:** two of my favourite animals from *The New Dinosaurs* (Dixon 1988), the Lank (at left) and Balaclav. A couple of decades ago, I never much liked the animals of *The New Dinosaurs*, my thinking being that they mostly fail as realistic extrapolations of Mesozoic animal evolution. Other people who work on Mesozoic animals have said likewise, most notably Greg Paul. But more recently I’ve slowly come around to the idea that they might be better imagined as animals that deliberately mirror those of our own timeline, just re-imagined as the members of ‘Mesozoic’ groups. In other words: if we *have* to have a world with pandas, mountain goats, monkeys and rorquals, but *have* to have those animals represented by ‘Mesozoic’ groups, what might we get? Images: Steve Holden; Philip Hood, both from Dixon (1988).

Dougal: I remember at the time thinking that – to make this work, I’m going to have to embrace the extreme left-wing of vertebrate palaeontology, so that’s when I started reading work by Bakker and people like that, even though I was very much a traditionalist (in terms of dinosaurs) when I needed to be.

Darren: So you were definitely inspired by the controversial ideas of the Dinosaur Revolution during this project?

Dougal: Oh yes, very much.

**Caption:** we’re going through a phase right now of better appreciating the incremental nature of how our ‘modern view’ of Mesozoic archosaurs was compiled (see my comments on the Dinosaur Renaissance in 2021’s *Dinopedia*). Nevertheless, Robert Bakker’s and Greg Paul’s sleek, dynamic dinosaurs and pterosaurs ushered in a new age of how these animals might be visualised. Illustrations like (at left) Bakker’s fast-moving *Barosaurus* pair from the 1960s and (at right) Paul’s svelte, narrow azhdarchid pterosaurs and tyrannosaurs influenced the animals of Dougal's *The New Dinosaurs*. Images: (c) Robert T. Bakker; (c) Gregory Paul.

Darren: There have been a couple of recent articles that have drawn attention to this. There’s an online article by Riley Black highlighting the fact that things considered pretty radical in The New Dinosaurs now appear on the ball, or at least pretty reasonable, in view of recent discoveries [that article is here].

Dougal: Well, I’ve got pterosaurs running around like giraffes!

Darren: Exactly, yes. So the animals of The New Dinosaurs weren’t coincidental inventions: they were extrapolations based on what Bakker and Paul had said. Having said that, there are creatures in The New Dinosaurs that don’t look modern in terms of their anatomy and/or behaviour. There’s a specialised scavenging tyrannosaur – the Gourmand – for example. That’s not a Bakkerian dinosaur, it’s a Beverly Halstead dinosaur: it’s a giant, sluggish, carrion-eating tyrannosaur.

**Caption:** the Gourmand *Ganeosaurus tardus*, a giant, scavenging tyrannosaur from *The New Dinosaurs*. Image: Steve Holden, from Dixon (1988).

Dougal: Remember that the whole of The New Dinosaurs was very much a cognitive exercise in the same vein as After Man. I was interested in patterns, and pushing the patterns to an extreme. So, in the lineage leading up to the Gourmand, the idea is that we’re seeing a pattern in which the forelimbs become smaller and smaller and smaller, eventually disappearing.

Darren: If you were to do The New Dinosaurs today, would you do it any different?

Dougal: Yes I would. If you’re asking for specific examples – it’s difficult to know where to start...

Darren: Absolutely. Obviously we have so many feathery theropods now they’ve even been described as mundane, plus now there are even filamentous structures on ornithischians. One of the things I like about the book is that it doesn’t just depict a ‘dinosaurs only’ world. There are mammals in there, there are big pterosaurs, some plesiosaurs. One of the things I think we’ve learnt about the Mesozoic that makes it more interesting is that it’s not just a ‘dinosaurs only theme park’: you’ve got these other groups, and since the book was published, since the 1990s, we’ve learnt that, for example, in Late Cretaceous South America we’ve got an enormous radiation of terrestrial or semi-terrestrial crocodyliforms living alongside theropods. Obviously crocs went on to have their own, rich Cenozoic history, but if you have more complex animal communities at the end of the Cretaceous, and then don’t have the end-Cretaceous event, you end up with a completely different Cenozoic and modern day biota, perhaps with these groups doing things that we don’t see in our timeline.

**Caption:** ugh, yet more Dougal Dixon fan-art made by myself. This fanciful montage shows various smaller and ‘mid-sized’ creatures from *The New Dinosaurs*, including Lank, Springe, Northclaw, Treepounce, Nauger, Cutlasstooth, Cribbum, Pangaloon and (in the water) Watergulp, Pouch, Plunger and Glub. The animals are not scaled at all correctly. Colouring by Ethan Kocak. Image: Darren Naish and Ethan Kocak.

Dougal: Indeed. Imagine then, that, with the formation of the South American landbridge, we’d see migration and diversification involving these animals as they moved north.

Darren: Greg Paul did a review of The New Dinosaurs. It’s kind of critical, but it does basically say at the end “Well done, it takes guts to do something like this and I respect that”. It gives him an excuse to invent his own ‘modern’ non-bird dinosaurs, including weird new horned dinosaurs, cursorial tyrannosaurids...

Dougal: Yes, I’ve seen that illustration.

**Caption:** Dougal has made several large-scale models of theropods and other animals. This photo, taken in August 2013, shows Dougal talking with palaeoartist Bob Nicholls; Dougal’s feathered oviraptorid model is at left. Image: Darren Naish.

Darren: Going back to the subject of pitching these books to publishers in the first place, this would mean that it was your artwork, your illustrations, that led to these projects being a success. What I’m getting at here is that... thinking of, say, the future animals of After Man, we know that you invented these creatures as extrapolations based on extant animals. And I’ve seen your illustrations, the models you’ve made; I know you’re a very talented artist. So why, in these books, don’t we see your artwork, your illustrations? Instead, we see depictions of these animals re-imagined by other artists (usually wildlife artists): people like Denys Ovenden, Philip Hood. Is the decision to use art by these people made by the publishers?

Dougal: It is a publisher decision, yes. At the beginning of the production of After Man, I did produce detailed illustrations for the artists to follow... and the artists would ignore them, and draw any old science fiction-based creature. So I really did have to bring them to heel. I ended up producing draft line illustrations that were so detailed, they just needed to ink them in to produce the final products. It’s a shame. In reality, my stuff should have been in there, yes.

**Caption:** a set of original Dougal Dixon illustrations, showing a foraging gigantelope herd, and detailed profile drawing. Numerous such sketches exist and were unpublished and mostly unseen when I first published this interview text in 2014. They and many others are now at **Dougal Dixon’s website**. Image: (c) Dougal Dixon, used with permission.

Darren: Where are all those illustrations of yours now? Do you still have them? Keeping in mind that the world of publishing is very different now thanks to self-publishing, ebooks and online pdfs and so on, I just wonder whether you’ve ever thought of getting them published?

Dougal: I do still have them. As for getting them published... I hadn’t thought of that, actually. [UPDATE: see Dixon (2021), since some large amount of this original art has now been published.]

Dougal: So, we’ve spoken now about After Man, which is about future evolution, and The New Dinosaurs, which is about zoogeography. Then there’s Man After Man – a project I was never keen to be involved in, the title of which was originally being kept for a project of my own. And that project again involved fictitious examples of factual processes. I thought: right, let’s have the current world collapsing through overpopulation, famine and so on, and the idea that mankind needs to escape destruction. What does mankind do? Invents time travel and moves 50 million years into the future and sets up civilization then. Then what we’ll have is that all the man-made catastrophes, all the ecological disasters... they happen all over again. So I’ve got this world already created in After Man, and I’m now going to destroy it... this was going to be Man After Man. But the name Man After Man was taken for that other disaster of a project.

**Caption:** proof that the *Greenworld* volumes exist, and that I own them (thanks to Dougal). As Japanese books, they operate ‘backwards’ relative to what English speakers are used to. Both volumes are substantial, containing way over 300 pages each. It would be great if they existed in English! Images: Darren Naish.

Darren: So, Man After Man would originally have been near-future humanity’s impact on the far-future creatures of After Man?

Dougal: Yes. What I did with that concept, quite a few years previously, is create an alien biota for an Earth-type planet in a far solar system, but based on the same biochemical processes that created life today. It was actually done as a design exercise to show to the local science fiction group. I had the whole ecosystem worked out. I thought: I’m going to use the Man After Man scenario on that. I’m going to have colony ships going to this planet. This became a book, called Greenworld.

The concept is that human colonists arrive in a pristine natural environment and immediately set about screwing it all up. The way it’s arranged is as a series of short stories that follow different generations of the same few families, thereby building up into a sort of dynastic epic, covering a thousand years of colonisation on this planet. The planet ends up as a smoking ruin, echoing the planet that was left behind at the start of the story. All of the illustrations are done by me, for a change (except for a couple of things that involved techniques that I’m unfamiliar with). The illustrations reflect the idea that the reader is basically eavesdropping on the lives of the characters of the book. We get to see excerpts from field guides, from herbals, and recipes, warning signs, bounty notices, advertisements, and all sorts of stuff like this – the idea being that, if you look at this combination of images, you build up an idea of what the biota is like.

**Caption:** pages from within both of the two *Greenworld* volumes, showing just a few of the many alien species encountered on the planet. Body plans very different from the ones we’re more familiar with are the norm, but the starting point was a pentameral pattern like that of echinoderms. There are no native humanoids, blue-skinned or otherwise. Images: (c) Dougal Dixon.

  **Caption:** the human colonists of *Greenworld* cause extinctions that occur in a staged, sequential pattern (among the earliest to go are large ‘pest’ species, like the Tallun shown here) and are described on a generational, long-term basis. What unfolds on the planet is of course a mirror of events that have occurred, and are still occurring, here on our homeworld. Images: (c) Dougal Dixon.

Darren: This is great. And has this material been collated? I mean, are there plans to get it published?

Dougal: It has been published, as a book, but only in Japan. My stuff is extremely popular in Japan. In fact my name has a hell of a lot more clout in America and Japan than it does here in the UK.

Darren: I do recall seeing some of the images from Greenworld on TV. I remember in particular the Strida creatures.

Dougal: That’s right – this was before I put it together, so parts of the project appeared here and there. There was the BBC programme Natural History of an Alien...

**Caption:** a *The Future is Wild* banner, **from the (still active!) website**, showing several star creatures of the series, including Carakiller, Gannetwhale, Megasquid and Toraton. Image: (c) *The Future is Wild.*

Darren: Yes. This featured the tripodal leaping creature and the giant conch-like animal. Let’s talk about The Future is Wild. The book is co-authored with Joanna Adams, is that right?

Dougal: Yes, Adams is a producer of television documentaries with a very good track record, who had always wanted to do a project based around the concept of evolution projected into the future. So, when this started crystallising about 15 years ago, she brought together a few consultants who might have ideas along these lines, like R. McNeill Alexander and Jack Cohen. And Jack said “Have you read After Man, by Dougal Dixon?”. So that’s where my name was mentioned, and hence I became part of the project. I was brought in as a consultant but, what with all these other consultants, my role evolved to become that of a designer, inventing the animals that had been suggested by this suite of consultants. I ended up designing all the animals to their briefs and passing along the designs to the animation studio. Since I’m an established writer, I also got to write the tie-in book. So I didn’t start any of this and was a mere hired hand in it.

**Caption:***The Future is Wild* features a far-future scenario that’s been entertained several times in speculative fiction…. Squid World! Here are two terasquids from the Northern Forest of 200 million years hence, the arboreal and intelligent Squibbon (at left) and the giant, graviportal Megasquid. Image: (c) *The Future is Wild.*

Darren: Ah. My imagining was always that they’d seen After Man, thought “this would make a great TV series”, but then found that – due to copyright or some other reason – they were unable to use the original creatures and hence had to invent new ones, similar but slightly different.

Dougal: Well, there is a constraint of that sort. By that time, DreamWorks SKG had bought the rights to After Man and optioned it, so we could do nothing with the original animals meaning, indeed, that a whole new suite of beasts had to be invented.

Darren: So we have giant flightless gannets instead of giant penguins.

Dougal: Exactly. They came up with giant penguins, but then the lawyers were --- no no, you can’t!

**Caption:** I think we can question the idea that gannets – which are plunge-divers, and wing-propelled when underwater – would give rise to long-jawed animals of such seal-like form, but giant, flightless, marine seabirds were needed for *The Future Is Wild*, so gannetwhales it was. Image: (c) *The Future is Wild.*

Darren: Did anything ever come of the DreamWorks project?

Dougal: No. DreamWorks finally abandoned it and it was then picked up by Paramount. It’s still getting optioned; in fact the option’s up for renewal this month, I need to find out what’s going on there. [UPDATE: according to Dixon (2021, p. 136), “No film ever resulted from this option so the scenario is now available again”.]

Darren: In the Earth of The Future is Wild, Amazonia is now a grassland, what’s currently Bangladesh is now flooded, forming the great Bengal Swamp...

Dougal: Yes, and this is completely a different tack from what I did in After Man. The geography had changed in After Man because of plate tectonics, but the various vegetation zones and biomes – I thought I’d keep them as close as possible to those of today, since the reader – being faced with all these new and terrible animals – will at least recognise the background.

Darren: There are four-winged birds in The Future is Wild called windrunners, specialised for flight over the Himalayan Plateau.

Dougal: They were created by Phil Currie, the consultant providing ornithological expertise on the project. I was amazed: a palaeontologist coming up with speculative evolution, with something as radical as four-winged birds. I thought: let’s go for it, this is going to be spectacular.

Darren: I always wondered if it was a coincidence that the windrunner was invented at about the same time as Microraptor was announced.

**Caption:** at left, the four-winged Great blue windrunner of *The Future is Wild*, a crane built for a lifestyle that involves migration across a vast, elevated plateau (formed when Australasia collided with Asia). At right, a *The Future is Wild* promo image, showing a flish of the far future. Flish are air-breathing actinopterygians (ray-finned fishes) capable of true flight and among the last surviving actinopterygians, most of which died out in an extinction event 100 million years hence. Images: (c) *The Future is Wild.*

Dougal: No, it [= the invention of the windrunner] was before that... unless Phil knew about Microraptor before that.

Darren: my thinking was that windrunners were so weird, so revelatory – why would anyone create a four-winged bird? – that a coincidence here would be remarkable, but I’ve since remembered that there was this early 20th century concept of the Tetrapteryx. In other words, there were ideas about four-winged fliers long prior to the discovery of Microraptor.

Dougal: I should finish here by saying that Jo Adams and team were all fired up to work on a big screen version of The Future is Wild with Warner Bros (I think) and had raised the funding for the television series The Future is Wild 2. However, on the big screen side, Avatar had raised the bar so high that Warner Bros (I think) pulled out. And on the small screen side, Discovery Channel announced that they were going to stop doing documentaries; instead, they were going over to dancing chefs and celebrity idiots and so the funding for that collapsed. Not a good year!

Darren: Any future ideas? What’s next? You’ve spoken about Greenworld.

Dougal: Yes, an English language edition of that needs to appear. [UPDATE: see Dougal’s website section here.]

**Caption:** we experience the flora and fauna of *Greenworld* through the literature, images and advertisements created by the people who live there. This poster – advertising an Artemis product – features a Strida... and I think a human as well. But it turns out that this image is not exactly representative of how things unfold on the planet, and a real Strida rider ends up more like the model shown at right (model by Dougal Dixon). Images: at left, Julius Csotonyi, from Dixon (2010); at right, (c) Dougal Dixon.

Darren: After Man has been repackaged a few times – there are several different designs with different covers. I saw one recently with a Night stalker on the cover. Oh – any thoughts on the 2013 movie After Earth? [UPDATE: After Earth is irrelevant and forgotten today but was new enough when I did the interview in 2014. Its inclusion of future animals, including a novel big cat, a giant, condor-like raptor and North American baboons, led to suggestions that it was inspired by After Man.]

Dougal: I haven’t even seen it, but I’m told by people who have seen it that it has no connection whatsoever with After Man. In any case, it more or less sank without a trace.

And that’s where we end things. I hope you enjoyed this updated version of a transcribed 2014 interview, and I hope that it provided interesting – and new – background to Dougal’s several projects, all of which have been a major inspiration to so many of us. This interview wouldn’t have happened without Dougal’s co-operation, and I sincerely thank him for his patience, kindness, time and assistance. Thanks also to the team at Breakdown Press for providing books.

For previous Tet Zoo articles on Dixoniana and other aspects of speculative zoology, see…

Oh no, not another giant predatory flightless bat from the future, March 2007
Come back Lank, (nearly) all is forgiven, September 2008
Giant flightless bats from the future, November 2012
Of After Man, The New Dinosaurs and Greenworld: an interview with Dougal Dixon, April 2014
The LonCon3 Speculative Biology event, August 2014
Speculative Zoology, a Discussion, July 2018
The Dougal Dixon After Man Event of September 2018, September 2018
Dougal Dixon’s After Man, the Initial Pitch Document, June 2020

Refs - -

Anon. 1995. Life on earth – 50 million AD. Focus January 1995, 56-60.

Dixon, D. 1981. After Man: A Zoology of the Future. Granada, London.

Dixon, D. 1988. The New Dinosaurs: An Alternative Evolution. Salem House, Topsfield (MA).

Dixon, D. 1990. Man After Man: An Anthropology of the Future. Blandford, London.

Dixon, D. 2010. Greenworld (two volumes). Diamond, Tokyo.

Dixon, D. 2018. After Man: A Zoology of the Future. Breakdown Press, London.

Dixon, D. 2021. After Man: A Zoology of the Future (40th Anniversary edition). Breakdown Press, London.

Dixon, D. & Adams, J. 2004. The Future Is Wild: A Natural History of the Future. Dorling Kindersley, London.

Naish, D. 2014. Speculative zoology. Fortean Times 316, 52-53.

Thomann, V. 2024. A “speculation built on fact”: on Dougal Dixon’s zoology of the future. In Castle, N. & Champion, G. (eds) Animals and Science Fiction. Springer Nature, pp. 345-362.

Todd, P. 1981. After man. Wildlife 23 (10), 16-18.

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I haven’t written much on pinnipeds (seal, sea lions and walruses) here at ver 4. Here’s a recycling of material originally published (here) at ver 2 way back in 2007, enjoy! I haven’t updated it…

**Caption:** captive Southern sea lion (at the Cornish Seal Sanctuary in Gweek, photographed in 2012). Note the massively built muzzle, the very dark pelt, and the stained, almost blackish hue to the teeth. This is common in sea lion teeth and I wonder what causes it. Various terrestrial mammals develop blackish teeth due to staining, but this is more of a herbivore thing than a carnivore one. What might cause this? Image: Darren Naish.

The Southern sea lion is a large pinniped that inhabits virtually the whole of the Pacific coast of South America, occurring as far west as the Juan Fernandez Islands. It also occurs on the south-eastern Atlantic coast as far north as Uruguay and southern Brazil, occurring also around the Falkland Islands. A dead one was found on the Galapagos in 1973 (Bonner 1994), and I think this is the most northerly record for the species. Adult males can reach 2.3 m and over 300 kg; females 1.8 m and c. 140 kg. The species is variable in colour, ranging from black to dark grey and reddish brown, with adult males being particularly dark. Females can be dull yellow on the head and neck. Males have an incredibly massive head and neck and a distinctive broad and up-turned snout.

**Caption:** a wild group of Southern sea lions at Peninsula Valdes, Argentina, showing adult males and females and numerous pups. Image: Reinhard Jahn Mannheim, CC BY-SA 2.0 (**original here**).

Taxonomy. Two scientific names are in use for the Southern sea lion: Otaria byronia (originally Phoca byronia de Blainville, 1820) and Otaria flavescens (originally Phoca flavescens Shaw, 1800). Because O. flavescens is older, many authors have used it in preference to O. byronia. However, Shaw’s “yellow seal” is not definitely a Southern sea lion, despite Rodriguez & Bastida’s (1993) argument that it was, as other otariids can sometimes have unusually pale coats.

In contrast, the type specimen for de Blainville’s Phoca byronia was undoubtedly a Southern sea lion as he referred to key cranial characters unique to this species. For this reason at least some authors have used O. byronia (e.g., King 1983, Berta & Sumich 1999, Brunner 2004, Brunner et al. 2004). I agree with them and it’s the technical name I prefer to use. For reasons that aren’t quite clear to me, however, O. flavescens currently appears more popular among specialist researchers.

**Caption:** a drawing of a male *Otaria* I created for the in-prep textbook (**cough cough**). Otariids are well able to hold the body high up off the ground when walking. The massive bulk of the head and habitually elevated head pose (where the snout is tilted upwards) are characteristic of this species. Image: Darren Naish.

A gnarly skull, a massive mandible. Adult male Southern sea lions have by far the most robust skull of any otariid, with a particularly prominent sagittal crest, a proportionally wide, short and robust rostrum, and an incredibly deep mandible. Large, shelf-like supraorbital processes are prominent, the palatal morphology is unique, with a palate that reaches all the way back to the level of the glenoid fossae. In the lower jaw, the masseteric fossa (the big concave area on the side of the jaw’s posterior section) of Otaria is immense, occupying about half the length of the dentary in some individuals.

Superficially, some of these details are similar to those of walruses and it’s interesting that, in her study of cranial variation among otariids, Brunner (2004) found Otaria to group separately from other otariids: a result that contrasts with previous classifications of this taxon alongside the New Zealand sea lion Phocarctos hookeri and the Australian sea lion Neophoca cinerea within a monophyletic Otariinae (Berta & Sumich 1999). There’s an awful lot that could be said about otariid phylogeny. Most molecular studies actually find Otaria to be close to Arctocephalus (the southern fur seals)… I’ll leave it at that for now.

**Caption:** an old and not good photo I took in the collections of the Natural History Museum, London, back in the early 2000s (when Stig Walsh and I were working on fossil pinnipeds). The insane gnarliness of this specimen (and others like it) never ceases to impress me. Look at the depth of that lower jaw! Image: Darren Naish.

With a skull morphology like this, it figures that male Southern sea lions are not limited to a diet of fish, squid and crustaceans (although these prey items do form the bulk of their diet). They will also catch and eat South American fur seals Arctocephalus australis, and not just the pups, but adult females too (Gentry & Johnson 1981, King 1983).

Penguins as prey. They also kill and eat penguins, including rockhoppers, gentoos and Magellanic penguins (Boswell 1972). Pinniped predation on penguins has now been reported in several species, and in some cases particular individuals develop a penguin-killing habit and can then have a significant impact on a colony. There is one case for example where, between 1997 and 1999, a Southern elephant seal Mirounga leonina single-handedly killed and ate 88 Magellanic penguins in one Argentinean colony (Clark & Boersma 2006). Predation on Yellow-eyed penguins Megadyptes antipodes by (apparently) a single female individual of Hooker’s sea lion Phocarctos hookeri on New Zealand is also posing a significant threat to the viability of the Otaga Peninsula Yellow-eyed penguin population (Lalas et al. 2007).

**Caption:** portraits of captive Southern sea lions, again at the at the Cornish Seal Sanctuary in Gweek. Features to note include the length of the longest whiskers, the short and scroll-like pinnae (external ears), and the slightly goofy, bulging eyeballs. Images: Darren Naish.

Sea lions vs fur seals. Intraspecific abduction, harassment and killing of pups is fairly widespread in pinnipeds and endemic in some species. In an Alaskan colony of Northern fur seals Callorhinus ursinus, Kiyota & Okamura (2005) described how each pup in the colony was harassed, attacked or abducted an average of 3.8 times (by adult Northern fur seals) during the breeding season. Interspecific aggression directed towards other pinnipeds is rarer, in part because species tend not to form colonies in the same place.

But we now know that subadult male Southern sea lions grab, shake and bite the pups of South American fur seals – often with fatal results – and not for food, but apparently as a form of misplaced aggression: these individuals have yet been able to win mating battles with older males (Cassini 1998). In cases, the fur seal pups are shaken and thrown about for hours and, in about 40% of the cases recorded by Cassini (1998), the pups died as a result. Mothers attempted to rescue their pups in about a third of the attacks, but in only one case out of 31 was a mother successful.

**Caption:** I suppose you don’t often get to see images of swimming sea lions – Southern sea lions in particular – from under the water, but the enclosure at the Cornish Seal Sanctuary allows views like this. I like this image, since it has a mysterious aura that gives the animal a slightly amorphous, frightening aura. Image: Darren Naish.

  **Caption:** pinnipeds of many sorts routinely swim upside down, and *Otaria* evidently does this too. Maybe they do this because they can, maybe because it’s fun, or maybe it’s because it allows them to better see what’s happening on the seafloor (or, in this case, the floor of the pool). Image: Darren Naish.

Female sea lions generally don’t get up to the same sort of nasty behaviour as males, but they still exhibit a fair amount of aggression towards each other, with biting and open-mouth threat displays being common in crowded colonies. Mostly this is due to defence of their own pups, and of their own little patch of the colony.

Aggression in females was studied by Esteban & Cassini (2007) who found that females were more aggressive to each other when the environment around the breeding colony limited their access to tide pools. Sea lions use tide pools in order to help control their temperature, so when these are in short supply, females are more likely to act territorially. If crowding and lack of access to a resource (tide pools) results in increased aggression, you might wonder why female sea lions (and other pinnipeds) bother to form colonies at all. Colony formation is a much investigated subject, with recent studies arguing that the main benefit that females derive from it is that their grouping together cuts down on the amount of harassment they’d otherwise get from males. And at the risk of spinning off at a tangent, I’ll stop there.

**Caption:** a male Southern seal lion with ‘his’ group of females, photographed at Chubut, Argentina. Image: Nestor Galina, CC BY 2.0 (**original here**).

Pinniped vs human. All in all, big otariids like Otaria are hefty and quite scary predators. One of my favourite mammals is the Leopard seal Hydrurga leptonyx, a species well known for its ability to kill other pinnipeds (including juvenile Otaria), large penguins, and even humans. Sea lions haven’t yet been reported to kill a person, but Californian sea lions Zalophus californianus in particular have bitten plenty, with the number of reported attacks increasing (Berkeley marina suffered a spate of attacks in 2006). In April 2007 a 13-year-old girl was pulled off her surfboard and apparently came close to death.

And that’s where we’ll end. Pinnipeds have been covered a few times at Tet Zoo before and I certainly plan to cover them more in the future. But it doesn’t look like there’s any material at ver 4. Here are links to other Tet Zoo articles on pinnipeds.. Essentially all of them have been ruined thanks to the failures of ScienceBlogs and Sci Am, so I’ve had to dig around for intact versions at wayback machine…

The most inconvenient seal, December 2006
The Long-necked seal, described 1751, September 2008
England ‘does a Montauk’ (mostly on the Grey seal), January 2009
Harbour seal kills and eats duck, March 2009
Statistics, seals and sea monsters in the technical literature, March 2009
The most inconvenient seal, June 2010
Pinnipeds Descended from One Ancestral Line, Not Two (was: Seals, the Early Years), June 2014
Incredible Elephant Seals, Part 1, June 2017
Incredible Elephant Seals, Part 2, June 2017

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for TetZoo and the more productive I can be on those long-overdue book projects. Thanks!

Refs – -

Berta, A. & Sumich, J. L. 1999. Marine Mammals: Evolutionary Biology. Academic Press, San Diego.

Bonner, N. 1994. Seals and Sea Lions of the World. Blandford, London.

Boswell, J. 1972. The South American sea lion Otaria byronia as a predator on penguins. Bulletin of the British Ornithologists Club 92, 129-132.

Brunner, S. 2004. Fur seals and sea lions (Otariidae): identification of species and taxonomic review. Systematics and Biodiversity 1, 339-439.

Brunner, S., Bryden, M. M. & Shaughnessy, P. D. 2004. Cranial ontogeny of otariid seals. Systematics and Biodiversity 2, 83-110.

Cassini, M. H. 1998. Inter-specific infanticide in South American otariids. Behaviour 135, 1005-1012.

Clark, J. A. & Boersma, P. D. 2006. Southern elephant seal, Mirounga leonina, kills Magellanic penguins, Spheniscus magellanicus, on land. Marine Mammal Science 22, 222-225.

Esteban, F.-J. & Cassini, M. H. 2007. Intra-sexual female agonistic behaviour of the South American sea lion (Otaria flavescens) in two colonies with different breeding substrates. Acta Ethologica 10, 23-28.

Gentry, R. L. & Johnson, J. H. 1981. Predation by sea lion on northern fur seal neonates. Mammalia 45, 423-430.

King, J. E. 1983. Seals of the World. British Museum (Natural History), London.

Kiyota, M. & Okamura, H. 2005. Harassment, abduction, and mortality of pups by nonterritorial male Northern fur seals. Journal of Mammalogy 86, 1227-1236.

Lalas, C., Ratz, H., McEwan, K. & McConkey, S. D. 2007. Predation by New Zealand sea lions (Phocarctos hookeri) as a threat to the viability of yellow-eyed penguins (Megadyptes antipodes) at Otago Peninsula, New Zealand. Biological Conservation 135, 235-246.

Rodríguez, D. H. & Bastida, R. O. 1993. The southern sea lion, Otaria byronia or Otaria flavescens? Marine Mammal Science 9, 372-381.

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Sea monsters (or sea serpents; same thing) have been covered here a fair amount over the years. And thanks to a brand-new book, it’s time to cover them once more…

Among those few truly noteworthy tomes on sea monsters or sea serpents, it remains that Bernard Heuvelmans’s 1968 *In the Wake of the Sea-Serpents* will forever be the great standard work. Such is its size, scope and impact. Notable others include Rupert Gould’s 1930 *The Case for the Sea Serpent*, Michael Bright’s 1988 *There Are Giants in the Sea* and Robert France’s *Entangled* of 2014. To this short list we can add Adrian Shine’s brand-new (2024) *A Natural History of Sea Serpents*, for noteworthy it most certainly is, and – I might add – for all the right reasons.

**Caption:** there aren’t really that many books on sea monsters that warrant repeat consultation, but the majority that exist are included in this photo. The new book that forms the focus of the review you’re reading now is in the middle. Image: Darren Naish.

I will say immediately that Shine’s volume should be prominently displayed at the front of any collection of these books, not tucked away or kept out of view. I say that because A Natural History of Sea Serpents is not just beautifully designed, compellingly argued, and well written, but also because it might be the most important book on the subject so far. I hold that view not just because of the case studies it reviews and reframes, but also because it provides a masterclass in how we should approach the entire subject going forward.

Adrian Shine is widely recognised as the preeminent expert on the Loch Ness monster, and if you’ve ever seen a TV documentary on Loch Ness or read any book on the topic it’s to be assumed that you know full well who Shine is, and certainly what he looks like. Of interest is that his expertise on the subject of Loch Ness has not so much been conveyed via his command of relevant facts and figures (to be sure, he knows them), but more via his skill in working out how we might test the performance of eyewitnesses claiming to see monsters in the loch.

**Caption:** Adrian Shine in the field. At left, a scene from the Scholarly Research of the Anomalous meeting, held at the Counting House in Edinburgh in February 2015. Loch Ness researcher Dick Raynor is at left, journalist and Forteana expert David Clarke at right. At right, Adrian (with Darren Naish on the left) at the Loch Ness Centre, Drumnadrochit; a photo from September 2016.

‘Doing science’ is as much about knowing how to test hypotheses, in addition to working out which hypotheses we should be testing in the first place, and what we have in Shine’s collected works is a decades-long exposition in how evidence pertaining to the Loch Ness phenomenon might and should be assessed (Shine 1984, 1993, 2006, Shine et al. 1988, 1993a, b, Paxton et al. 2016, 2025).

As ever, we here collide with the eternal issue of whether cryptozoology is, is not, can be, or cannot be a science, and then of whether someone ‘doing science’ is or is not a cryptozoologist. I am not aware of Shine leaning hard either way on that matter, but I am fully in agreement with his contention that investigations of monster sightings should (in general) begin with the assumption that witnesses saw something, it’s ‘just’ that that something was misinterpreted, misunderstood, misremembered or even misdescribed or misdrawn. Sometimes there are cultural reasons behind relevant events – this is one of the primary contentions of my 2017 Hunting Monsters (Naish 2017) – but also of utmost importance is that looking at animals is hard.

An evening at Regent’s Park in 2011. Proof enough that Shine and I are on the same page is that the book begins with his recounting of a 2011 meeting of the Zoological Society of London, titled Cryptozoology: Science or Pseudoscience?, in which one of the three speakers – a Dr Darren Naish – wrapped things up by arguing that “you can ‘do science’ with cryptozoological data, and I hope that we [have] succeeded in showing that at least some people interested in mystery animal reports are trying to look critically and objectively at the data” (Shine 2024, p. 1).

**Caption:** the ZSL meeting of July 2011 was quite the event and a list of people known for their association with cryptozoology were in attendance, Adrian included. At right, a 2011 version of this blog’s author with Adrian Shine. Images: Darren Naish; John Conway.

That meeting was phenomenally well attended, so much so that the society had to remove a partition wall at the back in order to allow guests to fill up a second room. And it’s good that Shine remembers it in positive terms; I, personally, was not far off from being a bright-eyed naïve literalist at that point, by then entering my post-Heuvelmans phase but still labouring under the assumption that there were real aquatic cryptids to find (Naish 2000, 2001).

**Caption:** more images from the 2011 ZSL meeting. At left, the ‘welcome to the meeting’ talk being delivered by Henry Gee, whose connection to the subject comes from his prominently publicised comments on the possible late survival of *Homo floresiensis*. At right, the cover slide for my talk, with me lurking in the darkness at the side. Several write-ups of this meeting exist, the most memorable being a *Guardian* piece by Carole Jahme. Images: Darren Naish; John Conway.

A Natural History of Sea Serpents describes Shine’s own personal journey through the subject of sea monster investigation, both as he recalls the history of sea monster sightings and gives us a tour of the various discussions and arguments provided by Rupert Gould in The Case for the Sea Serpent, a book that Shine was fortunate enough to encounter at a young age (Shine 2024, p. 3).

Across eight chapters (and there’s a prologue and epilogue to boot), Shine discusses multi-humped monsters of the sort associated with New England, multi-finned accounts, those sightings that involve slender appendages, horse-headed and dragon-like accounts of the sort associated with British Columbia, the Daedalus event of 1848, ‘necky’ sightings, the Valhalla creature of 1905, and a potential new explanation for some New England sea monster accounts.

**Caption:** thanks to images like these, authors – and the public – have tended to think that we have an incredibly firm view of what objects like the Daedalus sea monster of 1848 really looked like. The two shaded illustrations are from the Illustrated London News; the plainer one is from Gould’s *The Case for the Sea Serpent*. This is a pivotal case (in 2019, Charles Paxton and I argued that it was partly responsible for changing the public’s view of what sea monsters were and are like; Paxton & Naish 2019) and there’s a vast amount to say about it. Shine has an interesting take on it, one that makes me wish that I’d stuck with the hypothesis I endorsed in 2001 (Naish 2001).

The well informed natural historian. In assessing Shine’s interpretations, three generalisations – three valuable take-homes – stand out.

Firstly, consideration of an account requires that the author has a level of experience and knowledge of what known animals look like when observed in the field. On that point, authors have not been sufficiently catholic in their consideration of possible monster identities. This is not a novel argument. I remember being struck (and concerned) during the early 2000s when one of Shine’s colleagues and collaborators – Charles Paxton – made the exact same point: Charles’s argument was that those researchers interested in sea monster sightings have not considered a sufficient number of potential explanations when evaluating accounts. I was “concerned” because my experience at the time was highly deficient. Sure, I ‘know’ and have known for decades what basking sharks, cetaceans, molid sunfishes and so on look like in photos and art, but do I really ‘know’ what they look like in the field? Today, my experience of seeing big animals at sea is extensive, but there’s still a list of animals that I haven’t seen at all, and some of those that I have seen I’ve only seen once or twice, always in a familiar and expected part of their range, and (so far) while they adopt familiar swimming poses and perform typical behaviours.

**Caption:** a montage of big marine animals I’ve seen within recent years, all around the shores of the UK or in the north-east Atlantic. There are no mystery or unknown species here (even though some of these specific images don’t reveal the key traits that might demonstrate their identity); we’re seeing an assortment of seals, beaked whales and rorquals. Images: Darren Naish.

It's not the case that Shine’s (or Paxton’s) argument posits anything like out-of-the-box thinking, since it’s truer to describe it as… I don’t know, simply ‘well-informed’, involving a more thorough understanding of what big marine animals look like and what they do. This is the very anathema of the “it must have been an oarfish” logic that still looms large (Shine terms this the ‘universal’ approach, since it involves the author naively assuming that a single explanation has universal application across multiple accounts). Part of the reason that this ‘well-informed’ approach is so hard to employ is that our knowledge is still so deficient. That said, a key point Shine makes is that this was certainly more so in the past, especially in the late 19th and early 20th centuries given the impact of industrial whaling and the nature of maritime travel (Shine 2024, p. 211).

Take the famous Valhalla incident of 1905, in which the ornithologists Michael Nicoll and Edmund Maede-Waldo observed a large, purportedly long-necked animal with a squared-off dorsal fin off the coast of Brazil. I didn’t want to steal Shine’s thunder on this case, but the primacy of his argument is that those previously interpreting the account (this includes myself) have essentially been unable to appreciate key elements that appear to show what the animal most likely was. And, yes, this means that I’m giving up on Cameron McCormick’s otariid suggestion (Naish 2017, p. 50).

**Caption:** just as with the Daedalus object shown above, artistic depictions of the Valhalla creature are not necessarily especially realistic renditions of whatever was seen. This whole montage is a slide from a talk I used to give.

A point worth making in connection with the ‘well-informed’ approach is that a number of potential explanations exist for various accounts, at least some of which will always remains remain on the table, and none of which – in the absence of time travel – can ever be falsified. I suppose we have to have faith in the idea that self-correction will occur over time; we’ll never really know which proposed identity for a given sighting is the ‘most correct’, but the point is that some have higher explanatory power than others.

**Caption:** one of many sea monster sightings that might be explainable, Shine proposes, is the HMY Osborne account of June 1877, made off the coast of northern Sicily. As shown in the montage here (from Shine 2024), a row of fins were seen initially, then an immense creature with two “large flappers”. It has been tempting to interpret the latter as a big turtle but – as shown in the montage here – it could well have been a humpback. The fins (which were of irregular height) were most likely of different animals, presumably dolphins. Image: Shine (2024).

We don’t all have the chance to all see the same thing. The second major take-home is that what people have reported – that is, what they think they’ve seen, which you’ll note is not the same as what they have seen – depends very much on the circumstances of observation, and this is something that isn’t static, but has changed over the course of history.

Among the most familiar and vexing of sea monster accounts are those concerning long lines of repeating humps, most famously associated with the coastal seas off New England. Robert France argued that these can be explained as sightings of large animals – big tuna, perhaps – entangled in netting, the ‘humps’ being rows of cork or glass floats (France 2019). Shine’s (2024) interpretation of these observations is also tied to human use of the sea, but this time to the new steamships that were plying the seas at the same time, and creating new kinds of wakes that people simply hadn’t seen before (pp. 32-36). These wakes can persist on the surface for at least 20 minutes, and their individual ‘humps’ look solid and inky black. The case is surprisingly good and emerged from Shine’s experience at Loch Ness (Shine 2024, p. 36).

**Caption:** it isn’t true that multi-humped aquatic monster sightings only come from places where people used nets lined with multiple rounded floats, since these illustrations all depict observations reported from Loch Ness. This montage is included in Shine (2024) but the individual drawings come originally from Rupert Gould’s 1934 *The Loch Ness Monster and Others* (I own a 1976 reprint, but am still in quest of a first edition).

On original sources. Finally, the third take-home is one always worth repeating, and it’s one that’s become paramount to me in my time as a researcher on mystery animals: finding and considering the original source is paramount. A great many classic monster accounts are most widely known thanks to retellings in popular books and articles written – it’s important to note – by authors generally doing their best to make the account as sensational as possible.

Shine (2024) doesn’t discuss the U-28 account of July 1915 (in which an immense ‘marine saurian’ was blasted skyward by the explosion of the torpedoed British steamer the Iberian) but it’s one of the best examples of this sort of thing. Thanks to archival research, we can say with confidence today that the event never happened, and that authors like Heuvelmans (1968), who assumed that it did, relied on those aforementioned popular retellings (in this case, Richard Hennig’s 1957 book Les Grandes Enigmes de l’Univers).

**Caption:** the alleged U-28 event of 1915 is one of the most famous sea monster accounts. It did not happen (if you want the full details, I wrote a TetZoocryptomegathread **which you can find here**). The reason for the montage here is to emphasize that Heuvelmans (1968) relied on popular retellings – where the animal was depicted as a sort of flippered marine reptile (as shown in the two images at right) – and not on the original discussions of the event. In the first illustration, shown here at left, the animal looks very much as if it was based on a stuffed baby crocodile… which I think it was.

Time and time again, Shine shows via his checking of original sources that the version of a sea monster account we usually regard as canonical is – yikes – quite a bit flawed. One of the biggest problems here is the popularity of sea monster accounts, since journalists cannot, of course, help but commission artists to depict what the witnesses saw. The impact of this precise phenomenon and what it means for our imagining of unusual, rare events has been addressed several times by Fortean researcher and historian Mike Dash; see his 2010 Our Artist Pictures What the Witness Saw. The Daedalus event of 1848, the Pauline sighting of 1877 and many others – yes, even the Valhalla incident – have all been affected by misreporting and misrepresentation.

I like to think that I’m familiar with most of these misreportings, or embellishings, but one in particular was new to me: the classic illustration of the 1893 SS Umfuli observation from the South Atlantic. It shows a long-necked creature with a lumpy back and massive body. But… nope! Shine (2024, pp. 134-135) shows that this depiction is simply not realistic based on what was described in the original report. In fact, it now seems that the creature can be explained by a known species.

**Caption:** one of the best long-necked, or plesiosaur-like, sea monster illustrations that’s based on an eyewitness account is this take on the SS Umfuli encounter of 1893. But that long, elevated neck with a distinct head at the end is simply not an accurate depiction of what was described at all. As Shine (2024, pp. 134-135) explains, it was actually said to be shaped like a conger eel, and that ‘neck’ was described as a long upper jaw that was seen to open and close.

On that note: as much as I’d like to say more about the specific identities that Shine proposes, I’m going to apply uncharacteristic self-restraint. The book is simply full of nuggets that propose basking sharks, killer whales, beaked whales, grey whales, rorquals, humpbacks, leatherback sea turtles and so on as the true identities behind many of our most cherished sea monster accounts. Time and again, I found myself thinking of Thomas Huxley’s immortal line “How extremely stupid not to have thought of that”.

A Natural History of Sea Serpents is extremely well illustrated and does a great job of depicting the interpretations and specific eyewitness events referred to in the text. We owe special credit to Shine’s collaborator – his wife Maralyn – for the excellent work here. Many of the photos are in colour and the book simply looks great, with a very pleasing picture to text ratio (I am not ashamed to say that I like pictures, and I know I’m not alone). In particular, I really appreciate the images that show – sometimes via superimposition of the animal such that you can see the whole of it through the water – how a proposed identity might explain a given account. Inspired, I’d like to suggest that we should get into the habit of producing illustrations of this sort as standard when proposing identifications.

**Caption:** this montage illustrates the fact that some long objects depicted in illustrations (note those words) as representing the necks and heads of long-necked animals actually sound – if you read the written version of events – more like whales engaging in breaching or head-slapping. The case illustrated here was reported by people aboard the City of Baltimore when in the Gulf of Aden (between Yemen and Somalia) in 1879. The digitally composited humpback and minke photos at bottom show what the observations in question might actually have involved.

Shine writes well, and the book is immaculately edited. The index is thorough. On mistakes, the putting of ‘family-level’ taxonomic names in italics is a no-no that should have been caught, and a printing error affects the look of the text on one page (p. 184) of my copy.

Concluding thoughts. If it’s not already abundantly clear, I really like this book. A few years ago, cryptozoologist and author Karl Shuker noted that we’re in a golden age of cryptozoological books. I’m still wavering on that view. Sure, there are a lot of books out there on the subject now, but a great many are poor and shoddy and don’t contribute anything of value. What at least some monster advocates and promoters dislike is that the books that do have value are the sceptical ones. This perception of ‘value’ isn’t an idiosyncratic preference (I may be what many term ‘a sceptic’, but in fact I dislike knee-jerk or faux scepticism); instead, that ‘value’ more reflects the fact that the sceptical works involve the deep investigation and analysis that proper research should.

Shine’s new book has this in spades. This is a brilliant discussion of classic cases, many of which now have a revised interpretation, and it essentially covers the entire gamut of sea monster diversity. I should add that Shine (2024) appropriately cites and discusses the work of his predecessors and contemporaries in extremely fair fashion – a rare thing in the cryptozoological literature – and he also knows who and what should be ignored for reasons of bias or sophistry (his text on pp. 211-212 makes veiled reference to a specific pro-monster author I ignore where possible). Given the pre-eminence of Shine’s expertise as a well-informed natural historian, as discussed above, A Natural History of Sea Serpents has – unusually for a cryptozoology-themed book – proper relevance to people interested in known marine species, among them big sharks, cetaceans familiar or otherwise, and sea turtles. It should be read by people interested in marine wildlife in general, not just by the niche cryptozoological crowd.

**Caption:** ***A Natural History of Sea Serpents*** has an excellent picture to text ratio, and these pages are a good demonstration of that. Very interesting photos, reproduced in colour and in good resolution.

I will be coming back to this book again and again, am destined to cite it in probably every article I publish on sea monsters for the remainder of my tenure, and I congratulate the author, his illustrator (Maralyn Shine), and the publishers in a job so well done.

Shine, A. 2024. A Natural History of Sea Serpents. Whittles Publishing, Caithness. ISBN 978-1-84995-588-1, softback, illustrated, index, pp. 233. *Here from the publisher. £18.99.*

For previous Tet Zoo articles on sea monsters and related topics, see…

Books on the Loch Ness Monster 1: Ronald Binns’s The Loch Ness Mystery Reloaded, March 2019
Books on the Loch Ness Monster 2: Gareth Williams’s A Monstrous Commotion, March 2019
Usborne’s All About Monsters, April 2019
Sea Monster Sightings and the ‘Plesiosaur Effect’, April 2019
Books on the Loch Ness Monster 3: The Man Who Filmed Nessie: Tim Dinsdale and the Enigma of Loch Ness, August 2019
A Review of Robert L. France’s Disentangled: Ethnozoology and Environmental Explanation of the Gloucester Sea Serpent, November 2019
The 1972 Loch Ness Monster Flipper Photos, August 2020
Monsters of the Deep, a Ground-Breaking Exhibition at the National Maritime Museum, Cornwall, October 2020
The Lake Dakataua ‘Migo’ Lake Monster Footage of 1994, February 2021
What Was the Montauk Monster? A Look Back to 2008, October 2021
Santa Cruz’s Duck-Billed Elephant Monster, Definitively Identified, November 2021
Nessie Point and Counterpoint; Who Owns the ‘Facts’ on the Loch Ness Monster?, November 2024
A Cryptozoologist’s Bibliography: Matt Bille’s Of Books and Beasts, February 2024

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

Refs - -

France, R. L. 2019. Disentangled: Ethnozoology and Environmental Explanation of the Gloucester Sea Serpent. Wageningen Academic Publishers, Wageningen, The Netherlands.

Heuvelmans, B. 1968. In the Wake of the Sea-Serpents. Hill and Wang, New York.

Naish, D. 2000. Where be monsters? Fortean Times 132, 40-44.

Naish, D. 2001. Sea serpents, seals and coelacanths: an attempt at a holistic approach to the identity of large aquatic cryptids. In Simmons, I. & Quin, M. (eds) Fortean Studies Volume 7. John Brown Publishing (London), pp. 75-94.

Naish, D. 2017. Hunting Monsters. Arcturus, London.

Paxton, C. G. M. & Naish, D. 2019. Did nineteenth century marine vertebrate fossil discoveries influence sea serpent reports? Earth Sciences History 38, 16-27.

Paxton, C. G. M., & Shine, A. J. 2016. Consistency in eyewitness reports of anomalies: examples from accounts of aquatic “monsters”. Journal Of Scientific Exploration 30, 16-26.

Paxton, C. G. M., Shine, A. J. & Popov, V. M. 2025. Identifying biases and the relevant statistical population: the case of the Loch Ness monster. Journal of Statistics and Data Science Education 1-8, https://doi.org/10.1080/26939169.2025.2455195.

Shine, A. 1984. A very strange fish? In Brookesmith, P. (ed) Creatures From Elsewhere. Macdonald & Co, London, pp. 66-70.

Shine, A. J. 1993. Postscript: surgeon or sturgeon? Scottish Naturalist 105, 271-282.

Shine, A. J. 2006. Loch Ness. Loch Ness Project, Drumnadrochit.

Shine, A. 2024. A Natural History of Sea Serpents. Whittles Publishing, Caithness.

Shine, A. J. & Martin, D. S. 1988. Loch Ness habitats observed by sonar and underwater television. Scottish Naturalist 105, 111-199.

Shine, A. J., Martin, D. S. & Marjoram, R. S. 1993a. Spatial distribution and diurnal migration of the pelagic fish and zooplankton in Loch Ness. Scottish Naturalist 105, 195-235.

Shine, A. J., Minshull, R. J. & Shine, M. M. 1993b. Historical background and introduction to the recent work of the Loch Ness and Morar Project. Scottish Naturalist 105, 7-22.

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If you were following the hominin-themed discoveries of that long-ago era known as the 1990s, you may well remember the hot news, published in Science in 1994 and 1996, on Homo erectus…

As announced in those papers (Swisher et al. 1994, 1996), these archaic, Asian hominins seemingly had an extremely long tenure in the continent. They were there from a surprisingly early point in hominin evolutionary history and persisted there until extremely recently. That latter fact means that they overlapped in time with modern humans like us and also – far to the west in Iberia – with the last of the Neanderthals. This was (and still is) a big deal because it was one of several discoveries which proved that hominin evolutionary history didn’t follow a ‘one species at a time’ model but instead involved a bush-like pattern of lineages where several species were contemporary.

The main story told in the book I want to discuss today – Garniss Curtis, Carl Swisher and Roger Lewin’s 2000 *Java Man: How Two Geologists Changed the Course of Human Evolution** – concerns efforts to pin down the ages of some famous Javan hominid fossils via the application of argon-40/argon-39 dating. These specimens are, most prominently, the Mojokerto child skull, discovered in 1936, and the Sangiran cranium, found the following year. These were conventionally assumed to be somewhere round about 700,000 years ago, and thus from close to the end of the Middle Pleistocene (or Chibanian), but the radiometric dating was shocking: the Mojokerto child proved to be about 1.81 million years old, and the Sangiran specimen 1.66 million years old. That’s close to the end of the Early Pleistocene (or Galesian).

I’m working from the 2001 UK edition, so am citing it as ‘2001’ throughout.

Caption: Ngandong on the map; it’s in the northern part of East Java, the nearest city being Surabaya to the east. The term ‘Ngandong’ today is synonymous with the archaeological site that yields the H. erectus specimens discussed here. Stegodonts, extinct cattle, rhinos, tapirs and a large tiger (Panthera tigris soloensis) are also known from the site. Image: Google maps.

A separate set of fossils – the 12 partial skulls of Ngandong in Java that came to characterise a form of hominin known as Solo man (after the area’s Solo River) – form the focus of the book’s final chapters. The Solo hominin has been regarded as a distinct species (Homo soloensis), as a form of Neanderthal, as part of Homo erectus, and as part of Homo sapiens. Most workers today do regard it as part of Homo erectus. Curtis et al. (2001, pp. 223-229) regarded the fossils as being between 53,000 and 27,000 years old and thus from near the end of the Late Pleistocene, this being the radically surprising result they published in 1996 (Swisher et al. 1996). More recent dating work has pushed the age of the specimens back in time to somewhere around 100,000 years ago (Indriati et al. 2011), which is still Late Pleistocene but puts them tens of thousands of years prior to the time when H. sapiens moved through Asia, or migrated to Australasia.

**Caption:** a montage of various of the Solo *H. erectus* fossils, showing skullcaps in various stages of completeness. Those shown here mostly come from adult individuals. By extrapolating complete skull size, we can see that these were large animals, with skulls generally larger than those of *H. sapiens*. How so many crania came to be preserved in the same area has been the subject of some speculation: was this a product of taphonomy or transport by water, or had these remains been deliberately collected or assembled before discovery? Images: from Weidenreich’s 1951 monograph on Solo man, in public domain.

This book is effectively the backstory to the team’s research on these fossils, and it does a good job of telling the tale engagingly and in a way that connects it with the various debates occurring within palaeoanthropology at the time. The title is a bit superlative and also misleading given that I honestly doubt that the two geologists referred to in the title – Garniss Curtis and Carl Swisher – really did all that much in terms of contributing to the human gene-pool. But you can see what the title is getting at.

One of the reasons that we like reading about palaeoanthropology – part of the reason for it being the perennial subject of a massive number of first-person narratives about work on specific fossils and the controversies surrounding them – is the politics; the internecine turf wars, fall-outs and squabbles; the human drama. I didn’t pull Java Man off the shelf in order to win titillation from my reading of such skirmishes, but the book does deliver on this front, and it’s interesting stuff.

**Caption:** at left, the Mojokerto child skull photographed at Sangiran Museum, Sragen, Indonesia. At right, Dutch palaeontologist G. H. Ralph von Koenigswald with the skull, other specimens nearby. von Koenigswald is discussed throughout the book and is an important figure in the discovery, history and interpretation of these fossils. Images: Midori, CC BY 3.0 (**original here**); Tropenmuseum / National Museum of World Culture collection, CC BY-SA 3.0 (**original here**).

The Mojokerto child incident. The book stats with a cold open whereby Swisher and Curtis are examining the fabled skull of the Mojokerto child while at Gadjah Mada University in Java in the September of 1992. They want to pin down its age using geochronology, but that can only be done if a sample of the adhering matrix is collected. So Swisher opts to use a knife to scrape away a small amount of material. That might not sound like a big deal, but in the world of fossils and other rare objects it's what’s known as destructive analysis. You can’t just do destructive analysis on a whim: you need special permission, something that might involve the consultation of any number of high-ups, and – sometimes – controlled conditions and written agreement. Clear from the text is that the fossil’s caretaker – Indonesian archaeologist Teuku Jacob – was unprepared for and surprised by the application of a tool, and he takes Swisher aside and aims to politely but obliquely explain why his actions were absolutely not acceptable (Curtis et al. 2001, p. 14).

**Caption:** at left, Garniss Curtis, the geologist who’s essentially the primary character of the book. Curtis died in December 2012; this photo shows him in Wyoming in 2001. At right, a c 1960 photo showing Curtis with (at left) Jack Evernden and a mass spectrometer. Images: Gilbert WG, CC BY 2.5 (**original here**); J. Hampel / UC Berkeley (taken from **this obituary at UC Berkeley News**).

There’s the implication from the book’s early chapters that this incident resulted in all sorts of fallout, and I had high hopes for an exciting tale of cross words, banishment from Indonesia for the book’s authors, and even a court case or two. But… no, we don’t really return to this saga and I felt a bit cheated. The book’s implication on Swisher’s actions is that the ends justify the means, since the data that Swisher and colleagues obtained did pin down the Mojokerto child’s age, and ultimately were important in improving our knowledge of Homo erectus’s evolutionary history on the Asian continent. However, Swisher’s actions were wrong and disrespectful and the book certainly doesn’t convey this. It would be a pretty dangerous precedent if people examining fossils – no matter how credentialed and senior those people are – thought it within their rights to remove fragments, no matter how small, for their own pet projects.

On Louis Leakey and Don Johanson. A minor part of the background story to Curtis’s innovations in geochronology concern his disagreements with Louis Leakey. Working with Jack Evernden in 1961, Curtis used potassium/argon dating to establish that Leakey’s famous ‘nutcracker man’ Zinjanthropus (now Paranthropus) boisei was a surprising 1.75 million years old, much older than Leakey thought. However, “Louis repeatedly refused to believe dates that Garniss and Evernden produced for other fossils, when the dates did not jibe with what Louis wanted or believed” (Curtis et al. 2001, p. 19) and they ended up falling out.

**Caption:** replica of the famous 1959 ‘Zinj’ skull at the Natural History Museum, London. It is today classified as *Paranthropus* *boisei* and surely deserves the massive amount of publicity achieved at the time of its publication. The taxonomic history of *Australopithecus*, *Zinjanthropus* and *Paranthropus* is complex. Image: Emőke Dénes, CC BY-SA 4.0 (**original here**).

Another famous palaeoanthropologist – Don Johanson; Curtis et al. (2001, p. 106) quote Tim White’s description of him as “a nail-polish salesman in Yves St. Laurent pants and Gucci sneakers” – is also initially a colleague but later becomes an enemy. Early in the book, Curtis and colleagues have as their academic base the Berkeley Geochronology Center (BGC), which since 1985 was allied with the Institute of Human Origins (IHO). But this was not a happy marriage. The BGC view was that the IHO – and in particular its president, Johanson – were “spending far too much time on public relations activities – such as television appearances and writing popular books – to the detriment of fund-raising for the institution as a whole” (Curtis et al. 2001, p. 110). In turn, the BGC was accused by the IHO as being a bunch of “fucking nerds” (p. 110). There’s then more discussion of belittlement, jealousy, personality clashes, mischaracterisations of the BGC’s work and accomplishments, and descriptions of outbursts (from Johanson) that involved shouting and the throwing of objects.

Things in the book come to a head when, in April 1994, members of the BGC meet up with philanthropist Phyllis Wattis for lunch at the Berkeley restaurant Chez Panisse, the aim being to discuss their new Javan discoveries. Johanson, apparently by coincidence, unexpectedly turned up at the same restaurant and gave some hard stares on arriving and leaving. From there, things went rapidly downhill. It turned out that Johanson had been speaking to Wattis with funding in mind, and his interpretation – and that of other people on the IHO board – was that the BGC team were fishing for cash. The Chez Panisse incident, as it became known, then precipitated a series of events that resulted in the IHO tearing itself apart and ejecting the BGC from its premises, locking the BGC out of their lab, and ultimately in a June 1994 court case. My treatment here is a highly condensed summary of a long-running and complex series of events.

**Caption:** I became curious about the appearance of Chez Panisse and discovered that it has its own Wikipedia page. Apparently, it’s famous in the farm-to-table movement. Images: Calton, CC BY-SA 3.0 (**original here**); stu\_spivack, CC BY-SA 2.0 (**original here**).

Multiregionalism and regional continuity. A good section of the book – maybe a third of it – isn’t about the specific specimens that Curtis, Swisher and co worked on, nor about the politics that surrounded the study of these remains, but instead covers ‘big picture’ issues in Asian hominin evolution, and in particular the conflict over the multiregional hypothesis (this is where modern humans are posited as having evolved from a network of interbreeding pre-sapiens populations distributed across the modern range of our species). I admit to feeling some degree of frustration on seeing how much of the book was devoted to this topic. I get this a lot (I mean: frustration while reading books due to the apparent familiarity of the content) and it’s definitely a me problem, not a problem with the book.

**Caption:** a cast of Sangiran 17 on show at Beijing Museum of Natural History, one of the best Javan *H. erectus* fossils and a key specimen used in linking the Javan hominins to those of Zhoukoudian in China. Image: Bjoertvedt, CC BY-SA 4.0 (**original here**).

The bottomline is that the substantial geological age discovered for Javan erectus fossils like the Mojokerto and Sangiran specimens was and is important to the multiregional hypothesis, essentially because it’s at odds with the regional continuity required if erectus fossils in eastern Asia are posited as having evolved into the sapiens of the same area. What of the younger age that Curtis et al. (2001) thought correct for the Solo specimens? They argue that the anatomically archaic nature of these fossils is fully at odds with the idea that these are part of a population that could be considered continuous with H. sapiens. When they were writing, they thought that the Solo specimens were contemporaneous with Asian H. sapiens populations and thus needed to emphasize the anatomical disparity (Curtis et al. 2001, pp. 229-230). The anatomical argument remains valid, of course, but today we’re back to thinking of the Solo people as rather older than Asian (and Australasian) H. sapiens populations in any case (Rizal et al. 2020).

Towards the latter part of the book, Curtis et al. (2001, pp. 185-200) describe how proponents of multiregionalism (Milford Wolpoff in particular) responded to their new data. Primarily, this response was inconsistent and contradictory with previous multiregional arguments; in addition, Curtis et al. (2001) argue throughout that their work is consistent with the Out of Africa, single origin hypothesis for H. sapiens, and at odds with multiregionalism.

**Caption:** palaeoanthropologists are famously resistant to the use of computer-assisted phylogenetics… nevertheless, there are at least few published cladograms within the field. This tree (from **Ni *et al*. (2021)**, on the Harbin hominin lineage) shows how *H. erectus* is outside the clade that includes *heidelbergensis*, Neanderthals and moderns. What’s also notable here is that the lineages conventionally included within *H. erectus* form a paraphyletic group. We can agree that that group is a species if we want (and most researchers do want to maintain that view), but it means that some erectines, if you will, are phylogenetically closer to the *heidelbergensis* + Neanderthal + modern clade than are others. Image: **Ni *et al*. (2021)**.

Wrap-up. If you’re interested in the history of studies on Homo erectus in Asia, in the politics and personalities involved in hominin research, and in debates about hominin evolution, taxonomy and phylogeny, I recommend this book and really enjoyed reading it. It is, however, concerned almost entirely with specific discussions and discoveries pertaining to a particular set of Javan fossils, so shouldn’t be imagined as covering the whole history of Homo erectus in Java, in east Asia more broadly, and certainly not in Eurasia in entirety. How does Curtis et al. (2001) complement other popular books devoted to Homo erectus? I’ll come back to that at some point in the near future.

**Caption:** there are a massive number of hominin-themed books. Here are just a few. Image: Darren Naish.

For previous Tet Zoo articles on hominins and other hominids, see…

Zihlman’s ‘pygmy chimpanzee hypothesis’, October 2012
Why Humans Are Important to Studies of Primate Diversity, July 2014
The Cautious Climber Hypothesis, March 2019
Piltdown Man and the Dualist Contention, March 2023

Huge thanks to those who help support this blog at patreon. You can see behind-the-scenes and in-prep stuff relevant to projects for as little as $1 per month.

Refs - -

Curtis, G., Swisher, C. & Lewin, R. 2001. Java Man: How Two Geologists Changed the History of Human Evolution. Little, Brown and Company, London.

Indriati, E., Swisher, C. C., Lepre, C., Quinn, R. L., Suriyanto, R. A., Hascaryo, A. T., Grün, R., Feibel, C. S., Pobiner, B. L., Aubert, M., Lees, W. & Antón, S. C. 2011. The age of the 20 meter Solo River Terrace, Java, Indonesia and the survival of Homo erectus in Asia. PLOS One 6, e21562.

Ni, X., Ji, Q., Wu, W., Shao, Q., Ji, Y., Zhang, C., Liang, L., Ge, J., Guo, Z., Li, J., Li, Q., Grün, R. & Stringer, C. 2021. Massive cranium from Harbin in northeastern China establishes a new Middle Pleistocene human lineage. Innovation (Camb) 2 (3), 100130.

Rizal, Y., Westaway, K. E., Zaim, Y., van den Bergh, G. D., Bettis, E. A., Morwood, M. J., Huffman, O. F., Grün, R. Joannes-Boyau, R., Bailey, R. M., Sidarto, Westaway, M. C., Kurniawan, I., Moore, M. W., Storey, M., Aziz, F., Suminto, Zhao, J.-x., Aswan, Sipola, M. E., Larick, R., Zonneveld, J.-P., Scott, R., Putt, S. & Ciochon, R. L. 2020. Last appearance of Homo erectus at Ngandong, Java, 117,000–108,000 years ago. Nature 577, 381-385.

Swisher, C. C., Curtis, G. H., Jacob, T., Getty, A. G., Suprijo, A. & Widiasmoro. 1994. Age of the earliest known hominids in Java, Indonesia. Science 263, 1118-1121.

Swisher, C. C., Rink, W. J., Antón, S. C., Schwarcz, H. P., Curtis, G. H. & Widiasmoro, A. S. 1996. Latest Homo erectus of Java: potential contemporaneity with Homo sapiens in Southeast Asia. Science 274, 1870-1874.

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It’s true… DinoCon 2025 is go!

**Caption:** our website banner, featuring art by Natalia Jagielska.

Regular readers of Tetrapod Zoology will know – should know, I think – about the annual Tetrapod Zoology Convention, aka TetZooCon. Following an 11-year run – during which it underwent steady expansion and improvement – it had to come to an end, and 2024’s TetZooCon, the biggest and busiest of them all, was the last. We went out with a bang, as you can see from the article here.

**Caption:** scenes from the last ever TetZooCon, held at King’s College, central London, in September 2024. It ran smoothly and was a pretty epic success. Images: Georgia Witton-Maclean; Neil Phillips, used with permission.

But that 2024 event was an end, and never intended to be the end. Running and hosting conventions has proved such a rewarding line of work that my aim has long been to expand things further. If only I had the know-how and right set of contacts. To cut a long(ish) story short, plans for the future have worked out. After teaming up with the people behind PalaeoGames – my colleagues Annie Barling, Nathan Barling, and Mike O’Sullivan – I’m proud to announce the existence of an entirely new venture: starting August 2025, we’re hosting the UK’s first large-scale dinosaur-themed convention…. DinoCon, this year occurring on the weekend of August 16th and 17th at the University of Exeter in England’s southwest. We’re collaborating with the university’s Camborne School of Mines in pulling things together. Tickets are already on sale.

At the time of writing, we haven’t announced a schedule nor explained exactly what’s happening, but that’s coming soon. Watch the website (and our activity on social media; here on Twitter, here on BSky, here on Facebook) for breaking news. Rest assured that an exciting itinerary involving dinosaur-themed talks, panel discussions, fun workshop events, exhibitions, quizzes and more is coming together. A substantial number of stalls, selling (mostly) dinosaur-themed merch, artwork, models, figures, books, and more will be in attendance, in greater number than at any previous UK event. DinoCon is family-friendly and there will be plenty of things to keep kids interested. There will also be plenty of content for adults as well.

Dedicated, dinosaur-themed events have, of course, been hosted in the UK before, but nothing like this. DinoCon really will be on the same scale as the large Comic Cons. I should mention that my previous collaborator in running TetZooCon – palaeoartist and industry mogul John Conway – is still involved and will be running one of our palaeoart workshops.

On some practicalities. For those who don’t know southern England, Exeter is some distance (about 317km, or about 200 miles) from London, but it’s close to Bristol (104km or 65 miles) and other cities in the south (the port city Plymouth is 72km/45 miles away; Southampton is 179km/111 miles away). It’s well connected by rail and motorway (the M5 connects it to Wales and the midlands) and has an airport (with direct connections to Amsterdam, Belfast, Dublin, Edinburgh and Glasgow).

**Caption:** for those of you who don’t know where Exeter is, this map should help. It’s in south Devon, over in the west. Nearby tourist and holiday destinations include Dartmoor National Park, Exmoor National Park, and Lyme Regis and other parts of the Jurassic Coast. Image: © google maps.

On accommodation, numerous hotels, Airbnbs and so on are within a ten-minute drive of the university campus, but our accommodation of choice is the university’s own Holland Halls, places for which can be booked here. Space is limited so don’t leave it to the last minute. If you’re interested in hosting a stall you need to apply for a place here on our website; be aware that spaces are filling up fast, and that April 7th is the deadline for submissions.

**Caption:** a view from one of the halls of residence… pretty nice! This photo was taken in October 2024. Image: Darren Naish.

Finally, we have every confidence that DinoCon 2025 will be a success, and that other DinoCons will occur in the future, perhaps elsewhere in the UK. While DinoCon is certainly dinosaur-focused and themed, we don’t necessarily intend for DinoCon to be a dinosaurs-only thing, and in time will likely expand to include content on natural history, zoology and biology more broadly. So that’s that for now… more announcements are coming. I hope you’re sufficiently interested to come along, if you can. I and the rest of the team look forward to seeing you in Exeter in August.

Some useful and relevant links…

DinoCon website
DinoCon ticket sales
DinoCon stallholder applications
Holland Halls accommodation

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It is once more that time of year when the lone amphibian species here – the Common frog Rana temporaria – gathers to breed in ponds and pools. How are things going this year?

**Caption:** a large scrum of frogs battle and compete in the pond’s shallowest part; a photo from 6th February. It is not unusual for male frogs to leave the pond and move about the edges, presumably in an effort to intercept females as they approach. Common frogs are highly recognisable as individuals due to their bars, spots and stripes, and sometimes injuries and deformities. The individual in the foreground has a damaged left eye. Image: Darren Naish.

Some necessary background. I’m in Southampton, way in the south of England, and hence in one of the warmest, mildest parts of the UK. Every time I talk about frogs spawning during January or February, I have surprised and even incredulous people in continental Europe and North America telling me that surely I’m kidding. No. It’s normal in this incredibly mild, maritime, high latitude but warm part of the world for amphibians to begin breeding in February, January and even in late December. For the current breeding season, the UK’s earliest reported spawn clutch dates to 21st December 2024 in the Isles of Scilly (just off the south-west coast of Cornwall); additional records from Cornwall were made throughout late December 2024, and Devon, a bit further east, had spawn by January 1st. When I was a kid back in the 1980s, Common frogs generally started breeding during the second week of February. As the climate has changed, their spawning has happened earlier and earlier, and at the moment we expect spawn to start appearing during the last week of January.

**Caption:** a photo from 4th February, the day on which the first spawn clutch appeared. Note that the frogs mass and compete in the shallowest part of the pond, where the water is less than 10 cm deep. Image: Darren Naish.

  **Caption:** a photo from 5th February, the day on which the fourth and fifth spawn clutches appeared. Note the enthusiastic individual at right grabbing an attractive stone. Image: Darren Naish.

This year, things have been a bit weird. Late January was cold, heavily rainy and extremely windy here, and while frogs did begin gathering in my main pond, there was no spawn until February 1st. A massive number of frogs – over 60, the highest count so far – was visible by Feb 6th, so I predicted a record-breaking amount of spawn (we had 31 clutches last year: the highest so far). Six clutches were present on Feb 6th, but the temperatures here then dropped several degrees (from approximately 9° C to around 3° C) and all frog activity ceased. The frogs themselves disappeared from view (presumably taking refuge in the deeper water), and no new spawn clutches appeared, resulting in a plateau on the graph that lasted for about a week. I became convinced that this cold spell had prematurely curtailed the whole breeding season, meaning that 2025 would turn out to be a poor year for overall productivity.

But… no. Temperatures have increased again (today, the daytime temperature here is 13° C) and frog activity is in full swing. They were highly active during the night, over 50 frogs were observed in the pond yesterday, and today we’re at 17 clutches. Will we exceed 2024’s 31? Let’s see.

**Caption:** I keep records of when spawn clutches appear, but this is the first time I’ve tabulated it properly. The long plateau here was a cause for concern but the massive spike at right shows that we’re now back on track. Image: Darren Naish.

The pond that’s the focus of attention here is newly renovated relative to its 2024 condition, and now has a massive shallow area, lined with stones and surrounded by sloping banks, specifically designed for frog breeding. One interesting aspect of behaviour I didn’t foresee is that Common frogs are really good at inadvertently moving stones around as they compete, fight and push around, and hence shallow parts of the pond lined with stones end up being stone-free, which is upsetting for the aesthetic I was aiming for.

**Caption:** frogs and spawn massed in the pond’s shallow end. I see Paley, a distinctive and unusually pale frog (yes, I’m trying to identify and keep track of individuals year on year). The patch of exposed pond liner at upper left is a consequence of frogs inadvertently moving stones across the pond floor while competing. Image: Darren Naish.

Finally: I’ve done everything I can to help frogs in the area immediately around my house, and it’s clear that it’s really paid off. This area is now quite frog-rich, and a recently constructed pond about 100 m to the east of us is also now serving as a frog breeding site. That’s a win in an area where people are otherwise working as hard as they can to destroy and remove all areas that might serve as refuge, feeding and breeding sites for frogs. I’ll talk more about ponds and how to create a good one later in the year. For live news on spawnwatch activity here in my part of the world, follow #spawnwatch on Twitter/X and BSky.

**Caption:** 17 separate clutches present as of 21st February. Several are obviously only a few hours old and still have the tight, dense look typical of new clutches. Note that Common frogs really prefer to spawn in the shallowest water possible, literally just a few cm deep. No frogs are visible here as the arrival of a delivery worker caused them to rush away and hide. Image: Darren Naish.

For previous articles here on frogs and ponds, see…

A Love Letter to the Common Frog, August 2020
Live Spawnwatch Action From Pond 2 at Tet Zoo Towers, February 2024

Tetrapod Zoology is dependent on funds raised at patreon. Please help support this blog if you consider it worthwhile and want to see me continue doing it, thanks!

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It’s time once more to do a zoo review…

… and this time I’m going to talk about another Japanese zoo: Tama Zoological Park (or Tama Zoo) in Hino, in the western part of Tokyo, Japan. I was lucky enough to visit it during the August of 2024 and the visit is still relatively fresh in my mind. Tama opened in 1958 and started out as a satellite branch of Ueno Zoo, its original function being to house larger animals that needed to be kept outside of the more urban setting of Ueno.

Like all the places I wanted to visit in Tokyo, Tama Zoo was easy to get to via public transport and is serviced by a subway station that’s immediately adjacent to the main entrance (which features tall arches, columns and elephant statues). The zoo is large and extensively vegetated, but a downside on access is that it’s hilly, with steep concrete slopes and steps in places. Roads are present throughout, and a small bus runs on a circuit around the zoo; bus stops are present along its route. The zoo is divided into three main sections: the African, Australian and Asian Zones, the first of which is closest to the entrance/exit. There’s also an Insectarium section which features a large insect house in addition to a large, paved area with giant metal insect statues.

**Caption:** the zoo’s entrance/exit features a bridge-like bar hanging above the gates and booths. A central supporting column is decorated with a large Asian elephant sculpture. At right is the entrance/exit seen from within the zoo looking out, the gift shop at right. Image: Darren Naish.

I should mention that my visit to Tama was unusual in that I just happened to travel there as a major tropical cyclone – Typhoon Shanshan – was travelling north-east across the region, the consequence being high winds and enormous quantities of torrential rain. It wasn’t so bad that things needed to shut down, at least not where I was, but it meant that the zoo was being visited by a substantially lower number of people than normal, plus that parts of the zoo were closed and that certain animals were off show.

The bad weather means that my photos were exceptionally bad, even for me, so I’ve generally had to really up the contrast to make things more visible.

**Caption:** Waterfowl Bridge, the adjacent area involving not just a large pond complex but a series of vegetated islands. Wild birds, including ducks and grebes, use the ponds as well as captive ones. Image: Darren Naish.

To business. Rather than turning right (to the north) and visiting the large African Zone, I opted to continue ahead and ultimately left (so, west and south-west) through the Asian Zone. And what did I encounter first? A large pond complex – the Waterfowl Bridge area – was home to wildfowl, including Cackling goose Branta hutchinsii, Snow goose Anser caerulescens, Greylag geese A. anser and Bar-headed goose A. indicus. Lesser white-fronted goose A. erythropus were in a separate enclosure nearby. I’m a massive wildfowl nerd, so this goose-themed feature was good, even if the majority of species were familiar.

**Caption:** two of the several goose species kept at the zoo. At left, Bar-headed goose, a Central Asian species that winters in India and elsewhere in southern Asia. It’s significantly different from other *Anser* species and has at times been regarded as worthy of its own genus, *Eulabeia*. Both images at right show Greylag goose, with the long, pink bill and pink legs and feet showing that this must be the eastern subspecies *A. a. rubirostris*. Images: Darren Naish.

  **Caption:** Cackling goose group with food plants visible on the ground. The several Cackling goose subspecies were generally considered conspecific with the Canada goose *B. canadensis* until the last few decades. Images: Darren Naish.

A family of wild Little grebes Tachybaptus ruficollis also lived in one of the pools, and signs drew attention to their presence: a nice feature, since zoos should bring attention to the wild animals that live in their grounds and benefit from the environments they provide. I got great views of the mother feeding a dragonfly to her chicks.

**Caption:** at left, this wall in the tapir house provides abundant info on the zoo’s tapirs and their history. At right, the large colour sign on the outside of the house. Images: Darren Naish.

TAPIRS. Nearby is the large and impressive Malayan tapir Tapirus indicus exhibit, which includes large indoor and outdoor sections and a large amount of tapir-themed art. Indoors, a clear partition on one side means close views of the tapirs on land, and a pool on the other side – also fronted by a clear partition – allows you to see the tapirs should they go swimming or wading. The outdoor section was very green, very heavily vegetated, and with an area of marshy ground that the tapirs presumably approve of.

**Caption:** the tapir pool as seen from inside the house. The tapirs didn’t do any swimming, let alone bottom-walking, during the time of my visit, alas. Images: Darren Naish.

  **Caption:** a glass (or transparent plastic) wall allows you to see the tapirs up close and unobstructed if they’re in the house at the same time as you are. Note the variation in pale colouration at the snout and how much white is present around the hooves. Images: Darren Naish.

  **Caption:** view inside the tapir house. Lots of tapir-themed art – I approve! Images: Darren Naish.

  **Caption:** the zoo’s two tapirs as seen from a viewing platform. I presume this was a male and female but was never able to check. The presumed male has a pale scar on the dorsal surface of his snout. Images: Darren Naish.

  **Caption:** the tapirs traversing the marshy, grassy outdoor area of their enclosure. This photo gives some idea of how green and leafy the zoo grounds are. Images: Darren Naish.

A few domestic animals – ducks, goats and bunnies – were nearby, but the big surprise in this part of the zoo is the famous Mole House, an installation devoted to the keeping of the endemic Japanese mole Mogera wogura. A few shrew species are there too. For shame, I wholly forgot about the existence of this feature, and the bad news is that I never visited it, nor was properly aware of it. I’ve been trying to figure out how this happened. Looking back at my photos, I see that I did get close to the house, but emergency hazard cones visible in one of my photos might mean that it and the adjacent area was closed to visitors at the time of my trip. I could also have simply missed it, because (a) I’m not very smart and (2) it’s tucked away and located on a road that I don’t recall exploring.

**Caption:** the building in the middle of this photo is the fabled Mole House. You might be able to see bright yellow hazard cones close to the bottom of the staircase. Images: Darren Naish.

Otters were nearby (not sure what species), as was the large Walk-in Aviary (alas, closed) housing Black-faced spoonbill Platalea minor and Red-crowned crane Grus japonensis. Waitminute: if the aviary was closed (presumably due to concerns over H5N1 bird flu), is this connected to my missing of the Mole House?

**Caption:** a spoonbill montage. African spoonbill at left (with a somewhat washed-out face: the facial skin is ordinarily redder than it is in this bird), Black-faced spoonbill at right. Images: Darren Naish.

Tahr, takin and more. Further to the west – but still within the Asian Zone – was a large area devoted to hoofstock, among them Himalayan tahr Hemitragus jemlahicus, Mouflon Ovis gmelini (identified as O. aries musimon on signage: see the comments for some discussion on the taxonomy) and domestic Water buffalo Bubalus bubalis.

A large Asian elephant Elephas maximus enclosure was here too. The zoo was home to three Asian elephants at the time of my visit, the male of which is kept separate from the two females. The large indoor sections are viewable from outside via windows and from inside via a raised observation deck. The outdoor section includes a bathing pool, and replica rock faces and other structures to give the elephants things to do.

**Caption:** the outdoor section of the Asian elephant enclosure (this from the female section). Image: Darren Naish.

  **Caption:** part of the indoor area for the female Asian elephants. The elephants remained indoors during the time of my visit, perhaps because of the weather. I didn’t get to see the male. Image: Darren Naish.

  **Caption:** viewing windows for the large indoor area for the female Asian elephants. This entire area is roofed, which was useful at the time of my visit in view of the rain and wind. Image: Darren Naish.

An area for Indian one-horned rhino Rhinoceros unicornis was nearby but… I didn’t see one, and that’s a shame because a baby was born there in 2024. Tahr are (from my western European perspective) unusual, unfamiliar bovids of decidedly Asian theme (even though extinct Hemitragus species occurred across Europe) and sit in the same part of my brain as a few other exotic bovids, like serow and takin.

**Caption:** Himalayan tahr are – arguably – less visually impressive and more goat-like than sometimes conveyed in art (I generally picture them as longer-coated, shaggier, weirder). Molecular studies show that the three tahr groups (the other two are Arabian tahr and Nilgiri tahr) do not form a clade and hence they’re no longer included in the same genus. Images: Darren Naish.

And members of both groups were present as well. The serow – quelle surprise – was a Japanese serow Capricornis crispus. As for the takins…. I’ve seen takins in captivity before (both at Edinburgh Zoo in Scotland and Marwell Wildlife in England), but they were the dark brown, nominate form. These were different: they were the very beautiful Golden takin Budorcas taxicolor bedfordi, recognised as the distinct species B. bedfordi by some authors (Castelló 2016). The enclosures (more than one was devoted to takin) were flanked by steep, rocky walls to make it look like the animals were living in mountainous surrounds, and a concrete ramp in one of the enclosures had built-in rocky steps that the takins had no trouble in ascending and descending. One enclosure included a sloping area of ground, and two takins were at the top of it. Certain of the takins were moderately noisy and kept making grunts and hoot-type sounds.

**Caption:** I was really happy to see these animals and took some substantial number of photos. Look at the elegant sweep of those horns, the ‘overbuilt’ look to the forelimbs, and the very unusual, elongate hooves. Image: Darren Naish.

  **Caption:** Golden takin with down-sloping neck, stood in energy-saving pose on a rock. An interesting idea inspired by the incredible coat of this animal is that the pelt of one was perhaps inspirational to, or connected to, the origin of the golden fleece legend of ancient Greece. I have no real idea how plausible this might be. Image: Darren Naish.

  **Caption:** takin in profile, showing the obvious difference in robusticity between the fore- and hindlimbs. While much of the pelt is ‘golden’ – though I suppose it’s more like blonde with reddish tints here and there – note that there are large grey areas too. This is very variable across individuals. Image: Darren Naish.

  **Caption:** takin facial detail. Takins have a very tall, deep nasal aperture, with the nasal bones arched and high up on the skull. The *bedfordi* of the name honours the Duke of Bedford, who funded the British expedition that resulted in the scientific discovery of this animal. Image: Darren Naish.

  **Caption:** part of the takin enclosure, giving some idea of the rockiness of it, and the deliberate use of stone steps to mimic the elevated, hilly terrain they inhabit in the wild. Image: Darren Naish.

  **Caption:** the deep space here looks like a moat of the sort often built into zoo enclosures in order to keep animals always from visitors (and vice versa). In this case, however, the sloping inner wall features a set of large rocks that function as steps, and it’s clear here that the takins are adept at using them. Image: Darren Naish.

An enclosure for another sure-footed bovid was nearby – namely, a chamois – but a sign stated that the exhibit was temporarily closed. Based on a photo next to this sign, I tried working out what sort of chamois they have… and couldn’t. But I’ve learnt thanks to a video uploaded to YouTube in 2019 that it was an Alpine chamois Rupicapra rupicapra. Chamois don’t occur further east than the Caucasus (excepting the feral ones on New Zealand), so they’re properly exotic for a Japanese zoo visitor. Some authors argue that as many as six extant chamois species should be recognised (Castelló 2016), and – even if you don’t accept that taxonomy – there are numerous supposed subspecies.

Surrounded by these hoofed mammals and elephants was a group of Great white pelicans Pelecanus onocrotalus, the pelican species most frequently kept in captivity. A feature worth commenting on is that the outdoor part of their enclosure was floored with a carpet of felt-like fake grass fabric that’s meant to look like a lawn… but totally doesn’t. I don’t need to tell you that having material like this outside is bad, since it sheds plastic microfibres into the environment and ultimately into ecosystems and the bodies of animals.

**Caption:** the Great white, Eastern white, Rosy or White pelican is a mostly African species but also occurs patchily across Asia as far east as Indonesia. An apparently increasing number also occur across Europe, mostly in the east. It’s strongly migratory and will likely substantially alter its range in step with climatic change over the next few decades. Image: Darren Naish.

Père David’s deer or Milu Elaphurus davidianus. Readers with good memories will recall the long article I published in 2011 on this animal and its evolutionary history and scientific discovery (it’s here, but ruined by Sci Am). Several details of Milu anatomy are familiar and get mentioned in most discussions of the species: the antlers look like they’re being worn back-to-front, the face is slightly skewed to the side, and the tail is long and superficially recalls that of a horse more than of a deer. But there are several things about Milu that I didn’t know about before, and you can see them here.

**Caption:** the lone male Milu at the zoo. I watched as he took some time rubbing his face and neck against the stone wall of a large feature in the middle of his enclosure. Note the blackish areas on his neck, side, and the sides of his limbs, the brush-like tail, and the series of white spots along the shoulder. Image: Darren Naish.

  **Caption:** strange features of the Milu head and neck on show. The antlers are atypical, there are hair whorls on the neck and side of the shoulder, and the face is unusual, and seemingly skewed or asymmetrical. Images: Darren Naish.

The most surprising of these (to me) is the presence of very prominent hair whorls on the side of the neck. Hair whorls are conventionally said to be an adaptation for water-shedding (that goes for the whorl typically present in human head hair). I don’t recognise these from the other deer species I know (I checked, they totally don’t have them), so I wonder if the Milu is unique here. Also of interest is the black patch and series of white spots along the top of the shoulders and midline of the back. The zoo did a good job of celebrating Milu weirdness via a life-sized metal statue, replica antlers mounted on a display board, and signage which drew attention to its ‘designed by committee’ nature.

**Caption:** a nice little installation on the unusual antlers of the Milu (they’re weird in that the brow tine – the first forward-projecting growth from the beam – is as prominent as, or more prominent than, the beam itself). I own a Milu antler myself. Image: Darren Naish.

  **Caption:** Milu statue. Metal installations in places like Tokyo can become very hot in the sun, so much so that warning signs are placed nearby. Images: Darren Naish.

  **Caption:** one of the things said several times about the Milu is that it looks like a mashup of deer- and horse-like features, and that’s the idea explored here on this sign. Image: Darren Naish.

Asian big cats, orangutans, wolves and horses. Several additional Asian mammals were nearby, including Red panda Ailurus fulgens, Siberian tiger Panthera tigris altaica, Masked palm civet Paguma larvata (not showing during my visit), Lar gibbon Hylobates lar, Bornean orangutan Pongo pygmaeus and Snow leopard Panthera uncia. One of the gibbons appeared to suffer from a condition that forced his lower jaw to project well ahead of the upper, giving him a slightly scary ‘tusked’ appearance.

**Caption:** I’ve read about the agility of red pandas but never really seen it in life, since the only things I’ve seen them do when climbing involves walking along branches or thick ropes. However, this individual demonstrated its ability to hang upside down from the roof of its enclosure. Here’s your regular reminder that the Red panda is the original panda (I said exactly the same thing **the last time I mentioned pandas**, and I’m sure I’ve said it on previous occasions as well). The Giant panda *Ailuropoda melanoleuca* is an imposter, made known to science some decades later. Images: Darren Naish.

  **Caption:** a male and female Lar gibbon shared this enclosure. This is the male: females are black. As you can see, he suffers from some kind of unusual jaw configuration. Image: Darren Naish.

  **Caption:** the Siberian tiger’s enclosure was heavily planted with bamboo, a good design feature that I’ve seen at a few other zoos. I’m used to tigers avoiding eye contact but this one purposefully stared at me while I was photographing it. Image: Darren Naish.

I didn’t see any Snow leopards (which is ok, I’ve seen them plenty enough times in zoos closer to home), but I heard one, specifically a female on heat, yowling (it sounded like ‘ra-oooow’, with the ‘oooo’ as in ‘coot’). I tried to record this with my phone but only succeeded once. Most times I started recording she went quiet for a few minutes… meaning that I would then have to stop recording, at which point she would of course start calling again.

The orangutans were active and doing interesting things. A juvenile was playing on ropes, a large, flanged male was ant-fishing (the enclosure includes a large, clear plastic box housing an ant colony), and a female was interacting at a window with a visiting woman who was showing the orang how to apply makeup. The orangutan watched with what looked like polite indifference and even yawned during the woman’s performance. Right at the far south-western corner of the zoo is an orangutan forest exhibit, with an ‘orangutan skywalk’ that allows the orangs to climb overhead. Unfortunately, I didn’t have time to get to that.

**Caption:** a Bornean orangutan watches a woman applying foundation. Signs inside this house reminded you to be aware of (human) pickpockets. Image: Darren Naish.

  **Caption:** outdoor component of one of the orangutan enclosures. The replica rock surfaces, again, are impressive, and the signs do a good job of telling you about the history of the specific animals kept at the zoo. Image: Darren Naish.

Wolves and horses. Wolf Canis lupus – labelled as ‘Common wolf’ – were at the heart of the Asian Zone. I only saw a single one, and it was lying on its belly inside its house during one of the rainstorms. The wolf enclosure is mostly viewable via windows in a concrete tunnel that has a bunker-like aesthetic. This was actually quite a useful place for me, as I was able to hide there during the most extreme part of the storm. Outside, various announcements echoed across the zoo while I was taking refuge, and I hoped with all my sincerity that they weren’t storm warnings telling people to evacuate and go home. Evidently, they were not, for I wasn’t the only visitor to emerge from shelter once the rain had slowed.

**Caption:** wolf resting indoors on a rainy, stormy day. I don’t know if the animal was locked inside the house or if it just chose to be there. Image: Darren Naish.

  **Caption:** it should be clear throughout this article that Tama Zoological Park is strong on signage, as is its sister zoo, Ueno. This panel – on the outside of the wolf enclosure – explains wolf body language and predatory behaviour in the wild. Image: Darren Naish.

  **Caption:** my refuge site during the height of Typhoon Shanshan. The outdoor part of the wolf enclosure is visible through the windows on the left. There were no other people in this tunnel during my time there, but there was a large katydid (which I photographed, and which I now see that I never uploaded to iNaturalist). Image: Darren Naish.

A group of Przewalski’s horse or Takhi Equus przewalksii were kept in a large, concrete-floored enclosure nearby. Signage included photos of all individuals with a bit of info on each one. The text was all in Japanese, so the only thing I know is that the horses were all mares. I got to see them standing about in the rain, their ear positions indicating that they weren’t exactly enjoying themselves. While these horses had the robust, deep faces and chunky bodies characteristic of this sort of horse (it’s another species I know well due to its presence at zoos local to me, in particular Marwell), at least some of them looked floppier and softer in the mane than seemed right. The zoo also keeps domestic horses, I assume of an interesting or rare breed, but I didn’t get to them.

**Caption:** Przewalski’s horse enclosure in the rain. As you can see, the horses are mostly using the shelter. It’s a shame that they aren’t kept in an enclosure with a grassy, steppe-like flooring. However, I know that there are husbandry and welfare advantages to keeping hoofstock on hard surfaces like this. Image: Darren Naish.

  **Caption:** closer view of horses in the rain. As noted in the text, these individuals look a bit less like ‘pure’ Przewalski’s than ones I’ve seen elsewhere. Przewalski’s horses are, of course, caballine horses close to domestic horses, and you should be able to see the chestnuts on the forelimbs of the individual at far right. Image: Darren Naish.

In my article on Ueno Zoo, I noted the existence there of an animal cenotaph, a memorial to the animals that have died during their time at the zoo. Tama has one of these too. After finishing with the wolves and horses, I trekked along a long, snaking road that led back east in the direction of the entrance. I shouldn’t have walked this – I could have better spent my time by exploring the African Zone over in the north – and I think you’re expected to traverse this road via bus.

Birds and more birds: ibises, cranes, pigeons and more. Anyway, near the end of that road is another aviary, the bulk of it devoted to ibises and spoonbills. One enclosure housed Common goldeneye Bucephala clangula, Bush stone curlew Burhinus grallarius, American white ibis Eudocimus albus, Black-headed ibis Threskiornis melanocephalus, Black-faced ibis Theristicus melanopsis and African spoonbill Platalea alba. Others were home to Eurasian eagle owl Bubo bubo, Straw-necked ibis T. spinicollis, Puna ibis Plegadis ridgwayi and Black-headed gull Chroicocephalus ridibundus, Black stork Ciconia nigra, and Red-headed wood pigeon Columba janthina nitens.

**Caption:** a montage of some of the larger birds kept at the zoo. Left to white: Great white pelican, Black stork, Puna ibis. The stork looks a bit affected by the day’s weather. Images: Darren Naish.

  **Caption:** a mixed Black-headed gull / Puna ibis group in an aviary. The former is a very common gull here in western Europe, so seeing it in captivity is novel. It occurs across Japan as well though. Gulls are, relatively speaking, rarely kept in captivity, an issue discussed once or twice here in the past. Image: Darren Naish.

  **Caption:** Straw-necked ibis, a species new to me. Juvenile at left, adult (with eponymous ‘straw’ on neck) at right. This is a partly migratory ibis, mostly associated with eastern Australia. Image: Darren Naish.

  **Caption:** Tama Zoo has several enclosures dedicated to the Red-headed wood pigeon, and (I think) breeds this endangered, geographically restricted animal. It’s endemic to the Bonin Islands and Iwo Jima and is one of three subspecies of the Black wood pigeon. Image: Darren Naish.

Other Asian birds housed in a different part of the Asian Zone were Siberian crane Leucogeranus leucogeranus (their signs of course featured the older name Grus leucogeranus), Edward’s pheasant Lophura edwardi, Pied imperial pigeon Ducula bicolor, White-belled green pigeon Treron sieboldii, Chinese bamboo partridge Bambusicola thoracicus and Himalyan monal Lophophorus impejanus. That is a pretty good collection, including quite a few superstars of the Asian avifauna. I was especially happy to see all those ibises in close proximity. You’ll recall that Ueno Zoo was also strong on ibises.

**Caption:** at left, White-belled green pigeon, one of the 30 or so species within the Afro-Asian *Treron* green pigeons. At right, Himalayan monal male and female, though with the bad lighting making it hard to see the iridescent, metallic colours of the male. Images: Darren Naish.

Here’s where I discovered that more Asian Zone animals were north of the main road, opposite to the Waterfowl Bridge I mentioned earlier. Among these was a lone Asian black bear Ursus thibetanus, though it lacked the thick amount of neck fur typically associated with this species. This made me wonder if it was a member of the Taiwanese subspecies U. t. formosanus, since it’s apparently unique in lacking this feature. But no, the signs had it labelled as the Japanese U. t. japonicus. The bear looked a bit shabby (admittedly, a good part of this could have been due to the day’s high rainfall), and again I’ll say that bears in captivity rarely look happy, especially in small enclosures.

**Caption:** Asian black bear walking (at reasonably brisk pace) about enclosure, showing good view of what happens to the hand and foot during locomotion. The white chest marking characteristic of the species is just visible in one of these photos. Images: Darren Naish.

  **Caption:** the better part of the bear’s enclosure, which does a reasonably good job of making the most of the sloping elevation. It is not, of course, big enough or with enough things to do to keep a bear happy. Image: Darren Naish.

A Japanese giant flying squirrel Petaurista leucogenys was also in this part of the zoo. The animal was visible, albeit only its tail and part of its rump, since it was asleep in a nestbox, only parts of its body sticking out through the opening. Nearby was a large enclosure devoted to a group of Japanese macaque Macaca fuscata, and thus similar in feel to the macaque enclosure at Ueno Zoo. The Ueno exhibit features a substantial replica mountain, but this one has concrete steps and platforms and wooden climbing structures.

**Caption:** at left, a flying squirrel sleeping in its nestbox. At right, Japanese macaques. Macaques as a group are highly variable in tail length. Images: Darren Naish.

  **Caption:** macaque enclosure at Tama Zoological Park. The main display area is elevated, with concrete slopes and a moat on all sides. Image: Darren Naish.

I failed to get to a number of displays at the northern edge of the Asian Zone, including those housing Raccoon dog Nyctereutes procynoides, Oriental stork Ciconia boyciana, various owls and raptors, and reindeer of some sort. As a consequence, I’m left wondering what sorts of owls they have but I have a strong suspicion that among the species they keep is Blakiston’s fish owl Ketupa blakistoni. Why do I think this? We’ll come back to that…

**Caption:** at left, entrance sign to the Australian Zone with adjacent image of the globe. At right, big sign on the front of the Koala House. The images there portray a good likeness of the koala and even correctly portray their vertical slit pupils. Images: Darren Naish.

To the Australian Zone. The Asian Zone mostly done, I moved north to the Australian Zone. As you approach, signs give indications that animals on show will include kangaroos, koalas, assorted small marsupials, and birds like kookaburras and emus. I walked straight to the north-west corner to get to the Koala House, not just for the Koalas Phascolarctos cinereus but for the promise of more obscure marsupial species. I never did get to the emus by the way.

**Caption:** vertically arranged trees and branches, with freshly provided eucalyptus, in the large central atrium of the Koala House. I see two koalas in this photo but others might be in shot as well. Image: Darren Naish.

Anyway, I was quite impressed: the house was imposing and spacious (remember that hardly any people were present at the time of my visit), with a large, well lit, open area for the koalas, as well as a series of glass-front ‘nocturnal’ rooms that house Sugar glider Petaurus breviceps and Woylie or Brush-tailed bettong Bettongia penicillata. I’d previously seen Woylie at Ueno Zoo, but the enclosures at Tama allowed for closer and clearer viewing.

**Caption:** inside the Koala House, darkened halls provide the viewing area for enclosures that house sugar gliders and other species. I liked the whole setup, but the species on show were familiar ones if you’ve seen marsupials in captivity elsewhere. Image: Darren Naish.

  **Caption:** sugar glider on nestbox, at left, and bettong at right. I’ve seen sugar gliders several times in captivity but only climbing about, not gliding. I wonder how frequently they do that in captivity: do they even need to glide at all when in a small enclosure? Image: Darren Naish.

A few other macropods – that’s kangaroos and wallabies – are housed nearby and outside the Koala House, including Red kangaroo Osphranter rufus, Yellow-footed rock wallaby Petrogale xanthopus and Parma wallaby Notamacropus parma…. understandably, the zoo is still using the older name Macropus parma for that last species and doesn’t account for the new taxonomy proposed by Celik et al. (2019). Common wallaroo Osphranter robustus were present but must have been indoors. Shame, as I’ve never seen a wallaroo of any sort.

**Caption:** the kangaroos and wallabies were mostly kept in glass-walled enclosures, and the rain of the day made it especially hard to photograph them. Here are the best photos I got of a Yellow-footed rock wallaby. Rock wallabies are close relatives of tree kangaroos and are excellent climbers, well able to ascend and descend strongly inclined rock faces and even sloping tree trunks. Images: Darren Naish.

The highlight, however, was the Tasmanian devil Sarcophilus harrisii exhibit…. which I suppose I have to count as a highlight even if the animals weren’t showing. As you can see from the photos, the outdoor area was quite large, viewing areas were substantial, signage was good, and there were features like raised platforms, ramps and large rocks for the devils to interact with. Perhaps because of the weather, both devils (they had two) were indoors and asleep during my visit, but this wasn’t all bad as a live camera feed allowed you to see them in their bedroom. I’m now confused as to whether I’ve ‘seen’ live Tasmanian devils or not… I mean, I saw the ones that were there, but only via a screen.

**Caption:** two Tasmanian devils, resting or sleeping in a box indoors, and viewable on what basically looks like a TV set contained itself within a wooden box. There aren’t many zoos outside of Australia that keep devils, but they are (or were!) at Copenhagen, Duisburg in Germany, Prague, Beauval in France, Columbus Zoo in Ohio, and elsewhere too. Images: Darren Naish.

  **Caption:** Tasmanian devil enclosure, showing a large quantity of glass, various structures inside, and a large amount of devil-themed signage. Image: Darren Naish.

Unfortunately, it was now close to closing time. It was too late for me to even gain entry to the massive African Zone (the entrance road had been part closed off with mobile fencing). This houses lion, cheetah, serval, giraffe, African elephant, chimpanzee and flamingo, at least. Missing the cheetahs Acinonyx jubatus was the greatest tragedy, as the zoo famously has four King cheetahs, variously born there 2011, 2012 and 2013 (though I don’t know that all four are there today). I opted to rush quickly around the Insectarium, a large area with a butterfly-shaped tropical house as well as an Insectarium Center. Big metal locust statues were out in the paved area between these buildings.

**Caption:** the Insectarium Center, locust statues (and obligatory warning cones) nearby. Image: Darren Naish.

Amphibians! I rushed around the tropical house and didn’t give it appropriate time. It was very impressive though, with trees reaching up to the transparent ceiling and abundant shrubs and other plants flanking an oval walkway that descended toward a paved central area at ground level. Large butterflies of several sorts were free flying. A few terrariums around the edges housed amphibians, namely Schlegel’s green treefrog Zhangixalus schlegelii and Forest green treefrog Z. arboreus (both are rhacophorids endemic to Japan), Mountain brown frog Rana ornativentris, Eastern Japanese common toad Bufo japonicus formosus, Tokyo salamander Hynobius tokyoensis and Japanese fire-bellied newt Cynops pyrrhogaster.

**Caption:** amphibians on show inside the Insectarium. The salamander at left is, sadly, not a hynobiid but a fire-bellied newt. The frog at right is a brown frog (the group of ranid frogs most familiar to me, here in the herpetofaunally-depauperate UK), specifically a Mountain brown frog. Images: Darren Naish.

Only the ranid frog and a single newt were visible during my short visit, and it’s ironic that these are the animals most ‘familiar’ to a European visitor (Cynops is kept widely in captivity). The hynobiid salamander would have been the biggest score, since hynobiids are the stuff of legend and I don’t think I’ve ever seen a live one. Well, at least I can say that I‘ve been close to one, even if I didn’t see it.

Finally, the shop. Yet again, Tama’s shop, located right next to the large entrance feature, succeeded in doing what the zoo shops here in the UK so often fail at: namely, by providing a massive amount of material that (1) might appeal to a zoo nerd, and (2) is specifically relevant to the animals kept at the zoo! An excellent range of models and figures were on sale, as well as origami animal kits, cuddly Tasmanian devils (I bought one for my wife), and more. Impressive. I’m kicking myself for not buying more than I did.

**Caption:** I am so impressed by these origami animals… but I didn’t buy them, since I absolutely lack the patience or skill to construct such things. You will note that nearly all the models shown here represent species kept at the zoo, the notable exception being the Jaguar. Also, that tiger looks like a white one, and the zoo doesn’t keep those. Remember what I said earlier about Blakiston’s fish owl? The bird at upper right represents that species, so I’m guessing that it, likely, is kept at the zoo and that I missed it. Image: Darren Naish.

Final wrap-up. And that was that. As should be obvious from this review, Tama Zoological Park is huge and there’s a massive amount to see. I lost a fair chunk of my time there due to the weather, but – even so – I would have needed a few more hours had I aimed to see everything. Maybe I spent too much time with the wildfowl and tapirs…

It should also be obvious that Tama is well landscaped, with massive quantities of green and some impressive water features, including a nice waterfall. The shuttle bus was a good feature and would definitely save on walking, but using it would mean missing at least some of the enclosures. If you do traverse the zoo on foot, as I did, there’s a lot of walking to do, at least some of which involves slopes and stepped areas. As noted earlier, there are sections of the zoo that are definitely not ok if you’re of restricted mobility or in a chair.

**Caption:** view across part of the Asian Zone, looking east toward the long, winding road and in the direction of the entrance/exit, and with the Milu enclosure in the middle of frame (you can see the animal standing close to stonework in the centre of the enclosure, as it’s also doing in the other photos used earlier in the article). There’s extensive greenery and a good number of trees here. Image: Darren Naish.

  **Caption:** another view of greenery within the zoo, this time flanking the road within the Asian Zone that heads west and is near the tiger and orangutans. You can see the pelicans at lower right. Image: Darren Naish.

  **Caption:** steps leading to the area where the wolves, horses and cenotaph are. I want to make it clear that you don’t *have* to use the steps to get to this area though: the same area can be reached via a road that doesn’t have steps along its route, but the route is much longer. Image: Darren Naish.

The zoo’s signage was pretty good, the enclosures looked good and environmentally appropriate, and a good number of installations – statues and the like – assisted the educational mandate. I was impressed, really enjoyed my visit, and was pleased with the range of animals on show. As ever, we finish this review with my highly idiosyncratic scoring system…

Selection of species: 8 out of 10
Zoo nerd highlights: tapir enclosure, mole house (even though I missed it!), Golden takin, chamois (if there!), Tasmanian devil, good ibis selection, hynobiid salamander
Quality of signage: 8 out of 10
Value for money: 9 out of 10
Overall worthiness: 10 out of 10

A housekeeping thing: for no particular reason, I’ve never used justified text here at Tet Zoo. But I’ve now decided to start using it. What do you think? Should I stick with it, or should I go back to non-justified text?

For previous articles in my zoo reviews series, see…

Tet Zoo Reviews Zoos: Edinburgh Zoo, August 2016
Tet Zoo Reviews Zoos: Colchester Zoo, October 2017
Tet Zoo Reviews Zoos: the Isle of Wight Zoo, February 2020
Tet Zoo Reviews Zoos: ZSL Whipsnade Zoo, June 2021
Tet Zoo Reviews Zoos: Bristol Zoo, July 2021
Tet Zoo Reviews Zoos: Tripoli Zoo in Libya, January 2023
Tet Zoo Reviews Zoos: New Forest Wildlife Park, April 2024
Tet Zoo Reviews Zoos: Tokyo’s Ueno Zoo, September 2024

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Castelló, J. R. 2016. Bovids of the World: Antelopes, Gazelles, Cattle, Goats, Sheep, and Relatives. Princeton University Press, Princeton and Oxford.

Celik, M., Cascini, M., Haouchar, D., Van Der Burg, C., Dodt, W., Evans, A. R., Prentis, P., Bunce, M., Fruciano, C. & Phillips, M. 2019. A molecular and morphometric assessment of the systematics of the Macropus complex clarifies the tempo and mode of kangaroo evolution. Zoological Journal of the Linnean Society 186, 793-812.

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Let’s look at a remarkable cryptozoology-themed book, published in 2023…

**Caption:** front cover of Frost (2023). The striking colour cover is by Daniel Falconer and features a reconstruction of the face inspired by a night-time photo featured within the book.

Fatfoot: Encounters With A Dooligahl extensively recounts researcher Neil Frost’s thoughts and experiences pertaining to the supposed humanoid cryptids – yowies – of Australia’s south-east, the eponymous Fatfoot being a nickname that Frost has given to one of the individuals that, he thinks, live in his study area (Frost 2023). Frost writes with erudition and displays a level of science literacy and understanding of natural history often lacking in cryptozoological writings. If you’re well read on yowie speculation and lore (Joyner 1977, 1990, 2003, 2009, Healy & Cropper 1994, 2006, Gilroy 2001, Cropper 1996, Smith 1996, Gilroy & Gilroy 2006, Naish 2017), Frost’s research and conclusions might be familiar, for they’re summarised in one of the few books devoted to this subject: Tony Healy and Paul Cropper’s2006 The Yowie: In Search of Australia's Bigfoot (Healy & Cropper 2006). The key word there is summarised, for this new work sure ain’t no summary.

**Caption:** there aren’t many books that cover the yowie phenomenon, but here are a few. Some books on the subject are much sought after, most notably Graham Joyner’s 1977 *The Hairy Man of South Eastern Australia*. Image: Darren Naish.

Indeed, the text you’re reading now is not a long, proper review of the sort the book deserves, partly because I haven’t finished reading it. I figure that the least I can do is bring attention to its existence (buy it here). Let me say right off the bat that – for those interested in the relevant subjects (Australian cryptozoology, unusual hypotheses pertaining to novel megafauna, and investigations of cryptids traditionally regarded as mystery hominids) – Fatfoot is both a must-read and a unique, very special must-have.

**Caption:** a few yowie encounters – I’m talking here of the phenomenon in general, not the specific section associated with Neil Frost – have resulted in illustrations that supposedly depict the creature that was observed. As you can see, they appear to show hominids. Here are two such drawings featured in Healy & Cropper (2006). That on the left depicts the creature seen by Katrina Tucker in Acacia Hill, Northern Territory, in August 1997; the one on the right was drawn by Melba Cullen in 2001, albeit from recollection of an event that occurred in 1930!

Two other points about this book deserve initial comment. One is that this is a monster of a book, 715 pages long and 5 cm thick. The second point is that one early chapter, around 40 pages long, is a diary-like ‘log’ that discusses adventures on a day-to-day basis. I confess to finding this one chapter such a slog that it proved a total obstacle to my reading. It should have been substantially condensed or even stripped out.

**Caption:***Fatfoot* is a fat book. Remarkably, I’ve just discovered that my copy has increased in size while I’ve owned it, either because my use of the book has roughed up the pages a little, or because of humidity. It *was* 48 mm thick but is now 50 mm. Image: Darren Naish.

The primary hypothesis that forms the core of this book is as follows: yowies are real and have been encountered by Frost and his associates in the forested landscape of the Blue Mountains over a span of more than five decades. Frost suggests use of the indigenous term Dooligahl for them. The animals are large, heavy, forest-dwelling, bipedal mammals with highly sensitive night vision, flexible and dextrous forelimbs, and clawed hands (Frost 2023). They are predatory and kill lambs, ducks and other animals, use tools, are highly adept when it comes to the scaling of cliffs and mountainsides, and make loud roars commensurate with a massive thoracic cavity.

These attributes are based on a high number of observations and encounters described within this book, the oldest of which date to 1966. They involve the creatures visiting campsites at night, walking around houses and peering into windows, and a concerted effort by Frost, his collaborators and local police to photograph, document and identify them. Indigenous elders were consulted and provided information consistent with what Frost and other experiencers believe true of the creatures.

**Caption:** at left, Neil Frost (as of a few decades ago) holding a case of a partial track believed to be that of a yowie. At right, a Blue Mountains scene showing the sort of landscape associated with the events discussed here. Both images are taken from **here at The Fortean**.

As I’ve said before of Neil Frost’s thoughts on yowies – this was in my 2010 review of Healy & Cropper (2006) – it’s all very strange stuff. A dismissive, sceptical approach would be to argue that all relevant accounts involve wild-living people, and perhaps misidentified wild pigs, owls, possums or other known animals. Those latter suggestions would have to be invoked to explain the sightings of red-eyed animals seen among trees, some of which have resulted in photos that are included within the book (Frost 2023) (frustratingly, none of these images are findable online). A critical stance, and this includes my own, would also have it that Frost and his friends and colleagues are over-reaching in their interpretation of experiences, falsely amalgamating diverse events because they have ‘yowies on the brain’. As ever, I hope I’m wrong.

Frost’s contention is that these creatures do not only exist, but are, further, not hominids as most authors have supposed, but only superficially hominid-like. They are, in fact, gigantic macropods – kangaroos – of a new sort. The idea that yowies might be marsupials isn’t novel (Joyner 2003, 2009) – those authors who’ve taken the phenomenon seriously within a zoological paradigm have occasionally suggested that yowies might be diprotodontoids of some sort (Greenwell 1994) – but the idea that they might be humanoid kangaroos is new to the literature so far as I know. Needless to say, a bold and daring claim. I will resist the urge to mention Tank Girl. Oops, too late.

**Caption:** the concept of the marsupial yowie is not novel to this book, and in fact a few artists who specialise on speculative animals and cryptozoology have imagined the yowie as a sthenurine kangaroo before. This infographic depicting the yowie as a big striding kangaroo is by Fabio Alejandro.

The final chapters go as far as comparing the identikit image of the Dooligahl that Frost has built up to various kinds of kangaroos living and fossil, and if you’re aware of research positing that the large sthenurine kangaroos were bipedal walkers or striders, rather than hoppers… well, yes, Frost is aware of this too.

**Caption:** Frost’s giant macropods are striders and runners, and don’t use the paired-limb, ricochetal gait of most kangaroos. But here’s your reminder that this isn’t as hard a rule as we generally think: tree kangaroos and the extinct sthenurines moved their hindlimbs alternately, and even specialised ricochetal species like this Red kangaroo *Osphranter rufus* move the limbs in conventional fashion during swimming. This photo is from Wilson (1974).

The book’s cover states that “three marsupial enigmas” are covered, for Dooligahl isn’t the only cryptid that Frost discusses. He proposes that two similar animals, related to the Dooligahl but taxonomically distinct, exist as well: the Junjudee and the Quinkan.

While I have more to say, that’s where I’ll end for now. As I’ve said in the past, I’m genuinely nonplussed about the relevant accounts. Like most of us, I struggle to accept the idea that a massive, scientifically unrecognised mammal – let alone three such species – await discovery in modern Australia. Furthermore, the data compiled by Frost is anecdotal and suggestive, but never convincing (Healy & Cropper 2006, Frost 2023). But I remain open to the idea that there’s a valid and perplexing zoological mystery here, and I’m impressed with Frost’s efforts to document and catalogue everything that’s happened.

**Caption:** another artistic image that depicts a yowie as a marsupial, though in this case as a diprotodontoid rather than a kangaroo. I won’t deny that the concept of the marsupial yowie is great fun and has involved some great creature-building. Image: AThrillosopher (taken from **here**).

In short, the weirdness and complexity of this case, the undeniable creepiness of the accounts, and the novel creature-building inherent to Frost ‘marsupial hominoid’ hypothesis make this a fascinating work, despite its flaws. Specialist researchers should definitely get hold of it, and those interested in building a cryptozoological library will want it too. It is certainly a unique work.

Fatfoot: Encounters With A Dooligahl is well edited, organised and designed, illustrated throughout with maps, photos and drawings, includes footnotes, and has a good index. Its weight and size requires that it’s not cheap, and I do expect this, unfortunately, to limit sales.

One final point: a second yowie-themed volume appeared in 2023, namely Tony Healy and Paul Cropper’s The Yowie File: Encounters With Australian Ape-Men (Healy & Cropper 2023). I’ll write about that one too, in due time.

Frost, N. 2023. Fatfoot: Encounters With A Dooligahl. Coachwhip Publications. ISBN 978-1-61646-575-9, softback, illustrations, pp. 714. *Here from the publishers. £43.95 / $44.95 ($29.95 for black and white version available only in the US).*

For other Tetrapod Zoology articles on this subject and related ones, see…

What to make of the Yowie?, January 2010
Bigfoot’s Genitals: What Do We Know?, August 2018
eDNA, Footprints and the Biological Bigfoot: Comments on an Interview With Jeff Meldrum, April 2021
Santa Cruz’s Duck-Billed Elephant Monster, Definitively Identified, November 2021
Legend of the Black Dog, August 2022
A Cryptozoologist’s Bibliography: Matt Bille’s Of Books and Beasts, February 2024

This blog benefits from your support. Thank you to those who help via patreon!

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Cropper, P. 1996. Two yowie reports. The Cryptozoology Review 1 (2), 25-28.

Frost, N. 2023. Fatfoot: Encounters With A Dooligahl. Coachwhip Publications.

Gilroy, R. 2001. Giants From the Dreamtime: the Yowie in Myth and Reality. URU Publications, Katoomba.

Gilroy, R. & Gilroy, H. 2006. Out of the Dreamtime: the Search for Australasia's Unknown Animals. URU Publications, Katoomba.

Greenwell, J. R. 1994. The whatsit of Oz. BBC Wildlife 12 (2), 53.

Healy, T. & Cropper, P. 1994. Out of the Shadows: Mystery Animals of Australia. Ironbark, Chippendale, Australia.

Healy, T. & Cropper, P. 2006. The Yowie: In Search of Australia's Bigfoot. Strange Nation, Sydney.

Healy, T. & Cropper, P. 2023. The Yowie File: Encounters With Australian Ape-Men. Fortean Publishing, Sydney.

Joyner, G. 1977. The Hairy Man of South Eastern Australia. Graham Joyner, Canberra.

Joyner, G. C. 1990. Scientific discovery and the place of the yahoo in Australian zoological history. Cryptozoology 9, 41-51.

Joyner, G. 2003. Scientific reaction to evidence for the yahoo or 'Australian ape', 1882-1912. Journal of the Royal Australian Historical Society 89, 162-178.

Joyner, G. 2009. Monster, Myth or Lost Marsupial? The Search for the Australian Gorilla in the Jungles of History, Science and Language. Hayes UK & Thomas, Canberra.

Naish, D. 2017. Hunting Monsters. Arcturus Books, London.

Smith, M. 1996. Bunyips & Bigfoots: In Search of Australia’s Mystery Animals. Millennium Books, Alexandria, New South Wales.

Wilson, G. R. 1974. How kangaroos swim. Search 5, 11-12.

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You’re kidding… another year has passed? Yes, it’s late January, meaning that Tetrapod Zoology, the world’s best and most famous zoology-themed blog, has reached another birthday. As ever, I here take a very long-form look back at 2024….

**Caption:** my 2024, summarized.

There will come a time – very soon – when Tetrapod Zoology the blog is 20 years old. Two entire decades of blogging. But that day isn’t here yet. Today, Tet Zoo is 19, still a teenager. As is tradition for January (for Tet Zoo started life on January 21st 2006), we once more embark on my exceedingly long, rambling, tediously introspective review of the previous year’s events. I suppose these articles are mostly written for me, mostly as I strive to ever keep track of things, and make sense of them. I also like to see things I’ve learned, experienced and recorded shared, otherwise I’m just keeping stuff to myself.

**Caption**: Flame the dragon is alive and well and had a happy and healthy 2024 that involved going outside during the warmer parts of the year and foraging among the greenery. Flame is now over 11 years old, and members of her species only rarely make it to 15 (the world record is 18). Fingers crossed. Image: Darren Naish.

2024 in summary. What was the last year of operation like, from the Tet Zoo perspective? It was a packed year, with multiple things happening, a good amount of paying work, and numerous adventures and trips relevant to my interests. I went to Norway, Japan and Germany, attended five conferences, participated in an Atlantic whale survey, co-hosted a momentously successful TetZooCon and attendant tour, gave nine invited talks (in Hull, Bristol, Dorchester, Lyme Regis, Chichester, Southampton, Mettmann in Germany, Glasgow, and one online), and published three academic papers, several popular articles, a Chinese edition of Dinosaurs: How They Lived and Evolved, and both Chinese and Spanish translations of my book Dinopedia.

**Caption:** hey, have I ever mentioned the Natural History Museum book I co-authored with Prof Paul Barrett, titled *Dinosaurs: How They Lived and Evolved*? Despite being out since 2016, it has so far only been published in a handful of languages. This Chinese edition appeared in 2024. Image: Darren Naish.

I visited a good number of local woodland areas, wildlife parks and zoos, and did what I could in terms of observing wildlife local and foreign (images relevant to those trips are more or less randomly included in the article here). Some significant things were set up for 2025 and beyond. A lot was achieved here at the blog. As ever, there were significant failings in the completion of various works minor and major.

**Caption:** I feel very lucky to live in close proximity to the New Forest, one of the largest remaining unenclosed areas of medieval-style woodland, heathland and pasture, and I get to go there on numerous times throughout the year. This photo is from February 2024. Image: Darren Naish.

A huge number of things happened throughout the year that I can’t talk about. On that note, I’m extremely fortunate to still be working at BBC Studios. This, at a time when the TV industry is going through one of its darkest and most trying episodes, when massive numbers of people have been laid off or have found it impossible to remain in work.

Amphibians and a trip to Hull. February started with my attendance at the Herpetofauna Worker’s Meeting in Fareham, not far from where I live. Herpetology-themed meetings are always fun, and deeply relevant to me what with my involvement in pond construction and renovation, and local conservation. A few comments on the meeting in question were included in the February article published here, and I won’t ever be able to think about my attendance there without linking it to my buying of Jean Raffaëlli’s enormous, and enormously expensive, book Salamanders and Newts of the World. I’ve never opted to spend this much money on a book, and I don’t think I will again.

**Caption:** so, buying a copy of Jean Raffaëlli’s *Salamanders and Newts of the World* is quite the commitment. You need shelf space to house it (and your shelves must be able to take some substantial amount of weight), and you need some arm strength to lift it. The selection of books shown alongside it at left is pretty random – that shelf is an unsorted mess. Images: Darren Naish; Toni Naish.

It was also spawnwatch season, wherein I report the sexual shenanigans of the Common frogs Rana temporaria that inhabit the area around our house. I could say a lot about that, but of course I already have in this article from February 2024. It was a record year in terms of numbers of frogs and clutches of spawn, and I keep my fingers crossed that the situation will continue to improve in 2025 and beyond. And with ecologists Phil Budd and John Poland, I visited various ponds and wetland areas across the north and east of Southampton to monitor the amphibian situation, for concerns are that the invasive Alpine newt Ichthyosaura alpestris is here and expanding its range across the region. We didn’t find any, but let’s see what things are like in 2025.

**Caption:** Palmate newts *Lissotriton helveticus* at the degraded Hum Hole ponds in Southampton. The animal on the left had climbed to the top of a vertical wall that separates two of the tiered ponds, but on reaching the top it let go and fell back down (note the gammarid crustacean in shot too). Animals handled under licence. Image: Darren Naish.

  **Caption:** in February 2024, I visited the abandoned swimming pool and its nearby ponds at Southampton’s Outdoor Sports Centre. We found abundant Smooth newts *Lissotriton vulgaris* but not the Alpines we were looking for. Also present were a low number of Common toad *Bufo bufo*; calling male at left. Images: Darren Naish.

In which I’m outed as a disclosure agent for the Reptilians. And now for something entirely different. As regular readers will know, I dabble on occasion with cryptozoology, both academically and in terms of things at the more popular end. A few people heavily invested in monster belief – and who just happen to possess a good number of attendant conspiratorial ideas – tend to have opinions about those of us who view the subject through a sceptical, critical lens. I’m pretty familiar with this sort of thing. But what happened in February 2024 was kinda new.

Over on Twitter/X, the anonymous proprietor of the Beasts of Belial account (who I’m aware of due to previous exchanges) explained their complex, three-step reasoning behind the following modest proposal: Darren Naish is, most likely, “a disclosure agent in disguise, masquerading as either a skeptic or a disinformer”, who is actually “providing us clues about cryptids” while belonging to “a co-ordinated network of pro-disclosure people (probably all reptilian) who are using the disinformation conceit as a means to disclose things”. ‘Reptilian’, in this case, refers to the humanoid ‘reptilian aliens’ of conspiracy culture, not the group of animals that includes lizards and turtles.

**Caption:** I think that the tweets speak for themselves. There were many of them, but I’ve only bothered sharing the initial salvo here.

Sooo…. yup, outed at last. Someone told me that getting ‘conspiratorialised’ like this is quite the event, and that I should print the tweets out and get them framed on the office wall. I took screengrabs but the tweets themselves have all been deleted, and indeed the Beasts of Belial account no longer exists at all. Boo-hoo.

**Caption:** 2024 was a good year in terms of acquiring new animal figures and models. At left, a random assortment of recently obtained pieces, among them the giant AAA *Smilodon*, an unusual bigfoot, the Chap Mei orangutan, some Starlux mammals, and the various Safari Ltd crurotarsans from the toob set. At right, the very impressive *Prehistoric Planet* *Hatzegopteryx* from **Xotic Sculpts**. Yes, it has a technical issue concerning its feet. Images: Darren Naish.

  **Caption**: more models and figures new to the collection for 2024. At left, the pike, trout and ceramic bowhead from **TheYoungerEarth**, the *Homo diluvii testis* and *Corythoraptor* made by **Splendid Editions**, and a **Jed Taylor** *Deinonychus* acquired at TetZooCon 2024. At right… 2024 was the year in which Haolonggood dinosaur models really arrived on the scene, and here are some, with their megatooth shark figure and a giant salamander. Haolonggood figures are available via **Everything Dinosaur**. Image: Darren Naish.

Back to the world of sanity… I travelled to the University of Hull to deliver a lunchtime seminar on dinosaurs and what we currently think about their behaviour, a talk originally designed to accompany the release of my 2021 book Dinopedia. That gave me an excuse to visit The Deep, a giant multi-level aquarium that houses a good number of species and some great exhibits and displays. Thanks to Rob Knell for his help with this visit.

**Caption**: The Deep in Hull, as seen from Millennium Bridge looking south toward the Humber Estuary. It’s one of the largest aquariums in the UK, opened in 2002, and is a centre for research and conservation. Image: Darren Naish.

  **Caption**: I was impressed with many things about The Deep, and one of them is the giant rock wall that extends parallel to the walkway near the entrance. As you can see, it’s decorated with replica (but very real-looking), life-sized, fossils of spectacular marine animals from throughout history. Those shown here are (at left) the stem-whale *Dorudon*, (at upper right) the placodont *Placodus*, and (at lower right) the Cretaceous sea turtle *Archelon*. Images: Darren Naish.

My friend and colleague Paul Stewart – who I’ve mentioned here and there in connection with his roles in Prehistoric Planet and The Velvet Claw – visited in late February for a bunch of reasons, one being that we could photograph all editions of the seminal Bau und Leben der Rhinogradentia, that’s The Snouters: Form and Life of the Rhinogrades by Professor Dr Harald Stümpke in the same one place. Has such a thing ever been done before? I highly doubt it. The range of All Yesterdays figures made by Sam St Leger of Splendid Editions, made with my co-operation as consultant, went on sale at Etsy.

**Caption**: have more than three editions of Stümpke’s classic volume been photographed together before? I don’t know, but I doubt it. At left, the German original. In the middle, the 1967 Doubleday English first edition. At right, the 1981 University of Chicago softback. Other versions of the book do of course exist but I doubt I’ll ever get to own them. Image: Darren Naish.

  **Caption:** batch 1 of the *All Yesterdays* figures from Splendid Editions include Sleepy Stan the *T. rex*, the Swan, and the therizinosaur. Buy them (and others in the range) **here at Etsy**. Image: Darren Naish.

Wildscreen in Bristol and Ancient Sea Reptiles in Dorchester. During March, I visited Norway – for reasons I can’t discuss – and the Boneheads crew (that’s YouTubers Ben, Hamzah and Eddy) dropped in at Tet Zoo Towers. An on-stage discussion themed around the Apple TV / BBC Studios Prehistoric Planet series happened in Bristol at the natural history film and TV event Wildscreen, involving myself, palaeobotanist and palaeoclimatologist Bob Spicer, TV producer Simon Bell, and pterosaur expert Liz Martin-Silverstone. Tori Herridge led the whole thing. We did a good job of discussing the process behind the making of Prehistoric Planet – the event was titled Prehistoric Planet: Where Science Drives the Story – and Tori very graciously took time to explain the global significance and impact of Tetrapod Zoology. Yes, really – she did!

**Caption**: the Wildscreen 2024 *Prehistoric Planet* event. Tori Herridge (at far left) talks to (left to right) Simon Bell, Darren Naish, Bob Spicer and Liz Martin-Silverstone. People in the TV world generally don’t see the value of continual scientific input, and in fact often don’t want it, so *Prehistoric Planet* was unusual in having scientists as full-time members of staff. Image: © Wildscreen.

The day after, and it was time to visit Dorset County Museum in Dorchester to talk, yet again, about Mesozoic marine reptiles in connection with my book Ancient Sea Reptiles. I’ve lost track of the number of times I’ve given that talk. I substantially under-estimated the number of copies of the book I might need and many people who wanted to buy one went home without. Events relating to Ancient Sea Reptiles continued throughout 2024 and I’m pleased with the reception the book has had. In fact, the publishers (Natural History Museum, London) sold the last of their stock during the July of the year, which can only mean…. a third edition at some point, right? Stay tuned.

**Caption**: at left, a promotional image for my talk at Dorset Museum and Art Gallery, Dorchester, knocked up by my friend Mark North (who has proved himself quite the enabler when it comes to those accursed animal toys and figures). The artwork featuring the pliosaurid is by Konstantin Gerasimov. At right, the spectacular skull of the Weymouth Bay pliosaurid, holotype of *Pliosaurus kevani*, on show in Dorchester. Image: Darren Naish.

On plastic pollution, again. I assisted in another beach clean during March, again at the tidal stretch of the River Itchen known as Chessel Bay, famous for the epic quantity of plastic pollution that afflicts it. The situation, at least based on my experience, is worse than ever, with great swathes of the beach being composed of plastic fragments: discarded domestic junk as well as tiny polystyrene fragments, many of which come from the marina industry (where they’re used in pontoons).

What I think is happening is that several waterside companies and industries – involving shipping and distribution, leisure boating and watercraft mooring, and plastic manufacture – are woefully unregulated, and have been able to be lax on pollution at every turn. We understand this now (an organisation called Friends of Chessel Bay, here on Twitter/X, has led investigation and cleanup operations) but it does feel like too little, too late. We will keep working though.

**Caption**: if you don’t think that plastic pollution is a problem… well, maybe ignorance is bliss I guess. Small fragments of packaging of all sorts, plastic sticks used in toiletry products, plastic nurdles… Images: Darren Naish.

  **Caption**: this is the situation we’re in. Millions – literally – of polystyrene fragments, with known origins. You can pick up this stuff for the whole of your lifetime and not make much difference to an affected beach. And here’s a reminder that plastics are not inert in the environment. They’re chemically active and destructive to living things. Image: Darren Naish.

Moving now to something entirely different, at the end of March I published another of my TetZoocryptomegathreads, this one devoted to Tim Dinsdale’s Loch Ness Monster footage of April 1960. The text was shared at Twitter/X and my Patreon, but not at the blog so far. As I’ve said before, the plan is to compile all this text for a book. I’m working on that, and progress was made during the year.

**Caption:** it seems almost unbelievable today, but there were years during the 1960s and 70s when a select number of sane and intelligent people actually believed that the Dinsdale Loch Ness film of 1960 (a still from which is shown at left, diagrammatic interpretations of which are shown at right) really showed a large, scientifically unrecognised animal. These images, and numerous others, featured in my megathread on Dinsdale… **which is here on Twitter/X**.

During April I visited the New Forest Wildlife Centre – hence this article – as well as Jamie Jordan’s Fossils Galore museum and shop in March, Cambridgeshire. A number of very interesting marine reptile, dinosaur and Pleistocene mammal specimens are in Jamie’s care, and it would be good to see them studied and written up. One of the many things going on in the background at this point was the arranging of 2024’s TetZooCon and the accompanying tour. By late April, we essentially knew that we were going to be at Bush House, central London, once again, and we’d also booked the majority of our speakers and presenters. We knew also that this was to be the last TetZooCon of them all, for change was on the horizon once again.

**Caption**: the **Caspar *et al*. (2024)** paper on dinosaur intelligence proved popular with journalists. Some of the coverage was good, some was ok, but some was shallow and weak. “Scientists change their minds … yet again’” for example, is a terrible title that completely misses the point, since it’s written to imply that scientists all share the same interpretation of a given body of data. Here are screengrabs of just two of the many articles that covered the story.

Late April saw the publication of the year’s first technical paper, ‘How smart was T. rex? Testing claims of exceptional cognition in dinosaurs and the application of neuron count estimates in palaeontological research’ by Kai Caspar and a team of colleagues (Caspar et al. 2024). A version of the paper – a preprint – had already been online for a while at this point, so those properly invested in this subject were already aware of the article’s existence and primary contention. Nevertheless, publication makes a work more formal and definitive than it was before, and the study received substantial coverage and discussion. A particularly good article by Rachael Funnell can be found at IFLS: T. Rex Was A “Smart Giant Crocodile”, Not A Massive Brainy Baboon.

**Caption:** the extremely picturesque saltmarsh and salt dune complex of East Head, West Wittering, West Sussex, as of April 2024. This is yet another place in the UK where dogs are allowed and even encouraged by their owners to run about, despite the signs, ropes and fencing designed to encourage people to keep their dogs out of these areas. Nothing works. The only solution is to ban people and dogs from such areas, but no-one wants to take draconian action of that sort. Image: Darren Naish.

  **Caption:** another relatively local green area within easy reach of where I live: River Hamble Country Park, April 2024. A large pond in this park gets good quantities of frogspawn, but all of it – I’m not exaggerating, *all of it* – disappeared, and presumably died, during the early months of the year. Image: Darren Naish.

  **Caption:** I visited Blenheim Palace in April 2024 (specifically for the Icons of British Fashion exhibition). This is the toy wooden elephant that belonged to Winston Churchill during his childhood. I like it very much. Image: Darren Naish.

May was nuts in terms of work, with background events involving the Brian Ford paper, my speaking at the second of Verity Burke’s The Unnatural History Museum conferences, and prep for both the Lyme Regis Fossil Festival and TetZooCon. The Unnatural History Museum conference occurred online on May 22nd and was devoted to cryptozoology in museums. My talk – Conveying 'Sea Monster Science' in a Museum Setting – was about the story behind Monsters of the Deep, my National Maritime Museum exhibition. I believe that a recording will eventually be released online; until then, the script is available at the Tet Zoo patreon for those interested.

**Caption:** Steve Etches talks Kimmeridge Clay fossils at the *New Scientist* Jurassic Coast Weekender event in Exeter, 2024. In this particular moment, Steve is discussing the spectacular new pliosaurid discovered by Philip Jacobs in early 2022, then excavated by Steve and colleagues from its cliff-side location, and which was featured in the documentary *Attenborough and the Giant Sea Monster* on New Years Day 2024. In my capacity as a consultant working at BBC Studios, I helped devise the events featured in that film, though I’m not adequately credited. Image: Darren Naish.

Do not talk about the state of the British countryside. Later in the month, I headed west for the New Scientist Jurassic Coast Weekender event in Exeter where – yet again – I spoke about Mesozoic marine reptiles. While there I also, after years of knowing her online, met geologist Anjana Khatwa; my friend and colleague Liz Martin-Silverstone was a speaker at the event too. While travelling back home on the train, I made a point of filming a section of the Dorset countryside that I’ve long noted as being completely devoid of wildlife. I stand by that contention. If you visit the same area on foot… it’s dead. No birds, no insects, just agricultural monoculture with a small amount of denuded coppice. Quite a few sections of the British farmed landscape are like this. To make a point, I created a short video recording and uploaded it to social media. And… oh boy.

**Caption:** there are a number of short movies that show green, European landscapes… and which have a famously triggering effect (I’m talking about the K-fee jump scare commercials). Well, now there’s another triggering movie of a green landscape. Here’s a still, taken from a train as it thundered through the English countryside. Because this image is a little boring, I hid three small bigfoots in the background. Images: Darren Naish.

A few conservation advocates (some with a very large social media footprint) share the fairly negative view of the state of the countryside outlined above, and use what opportunity they can to point it out. And thus it was that I won a major signal boost. But another contingent of people hate any suggestion that things might be off in the British agricultural landscape. Rural places are, they contend, just peachy: natural forest is healthy and expanding, wildlife is thriving, and farmers and gamekeepers are just and wise stewards of the land, fully cognizant of the way things work and how they’re meant to be. A massive pile-on happened as a few hundred angry men descended to tell me what a complete idiot I am, the whole thing extending for the better part of a day (May 20th 2024, for those interested).

It was an interesting experience, for sure, but – for what it’s worth – a notable thing is that the very vocal, very visually obvious complainers were far fewer in number than those who ‘liked’ (in the social media sense) or retweeted the point being made. Despite the anger, the state of the UK countryside is dire. We’re hammering what’s left of our wildlife into the dirt and things – on average – are getting worse.

**Caption:** it wouldn’t be an annual review without me sharing at least something on gulls. This is a Herring gull *Larus argentatus* pair that accompanied me while I was having lunch next to the Avon in Bristol one day. What’s interesting about one of these birds (the presumed male on the left) is that its iris was speckled with black, not just pale yellow as is more common. This is a known intraspecifically variable feature of this species, speckled irides being rarer than plain ones. Image: Darren Naish.

The Chinese translation of Dinopedia appeared in May (a Chinese edition of Dinosaurs: How They Lived and Evolved appeared in print at the start of the year), slow progress on the Cryptids of Bernard Heuvelmans project continued (as it did throughout the year), Plumpmot the frog was photographed tackling and consuming an earthworm, and I discovered my long-lost Safari Ltd Great crested newt Triturus cristatus while undergoing loft clearance. What a win. Tetrapod Zoology articles published at around this time included those on the taxonomic vandalism of Ray Hoser, dibamids, atractaspidids – the burrowing asps or stiletto snakes – and the lamprophiine snake Bothrolycus. Those articles all focus on squamates, since I’d promised myself to rescue and republish what squamate-themed articles I could from the archives of ver 2 and 3.

**Caption:** views of the Chinese edition of *Dinopedia*. It’s a condensed, reduced version of the book relative to the original, but I hope that it still carries enough content to be considered worthwhile by its readers. The red-brown hue of the illustrations also marks it as different from the English edition. Images: Darren Naish.

The 2024 Lyme Regis Fossil Festival. I’ve said in previous birthday articles that the Lyme Regis Fossil Festival – once a sad and forlorn event that looked to be circling the drain – has very much found its feet in recent years and is now one of my highlights of the year. 2024’s happened over the first weekend of June. I gave my marine reptiles talk again, attended a book signing (both Ancient Sea Reptiles and Dinosaurs: How They Lived and Evolved were on sale), and went to a special showing of Tony Pinto’s 2024 documentary movie Why Dinosaurs?

**Caption:** it’s very appropriate that my involvement in the Lyme Regis Fossil Festival generally revolves around Mesozoic marine reptiles. At left, a stack of books to sign. At right, title slide from my talk, featuring images from the Apple TV / BBC Studios series *Prehistoric Planet*. Images: Darren Naish.

  **Caption:** it’s apt when attending a *Why Dinosaurs?* screening to do the red carpet thing and have your photo taken in front of the official backdrop. Here’s a group of us at the entrance to the Marine Theatre, Lyme Regis. Left to right: Stella Ludwig, Nizar Ibrahim, Hamzah Imran and Darren Naish.

I also arrived early enough on some days to go birdwatching along the beautiful Lyme Regis coast. Massive thanks to my buddy Kieran Satchell for help with accommodation, and to Natalia Jagielska and other staff at Lyme Regis Museum for organisational help.

**Caption:** I’ve now shared scenic photos of Lyme Regis’s East Cliff Beach on numerous occasions, but here it is at high tide on a sunny morning in June. The water is clear and blue, and birds including Common whitethroat *Curruca communis* sing from the cliff-top greenery. Image: Darren Naish.

When Mary met Flip. My talk at the Fossil Festival was followed by one given by another of my colleagues – Luke Muscutt – who works on plesiosaur locomotion (Muscutt et al. 2017). As you’ll know if you’ve seen the 2024 BBC documentary Attenborough and the Giant Sea Monster (which I had a hand in developing), or if you attended TetZooCon 2023, Luke has manufactured an accurately scaled robot plesiosaur. It’s called Flip and has been used in various tests to see how live plesiosaurs might have performed as swimmers.

I’m pleased to say that Flip was in attendance at the 2024 Fossil Festival. Late on Sunday, Luke took Flip to meet the Mary Anning statue, a great photo opportunity. 2024 was the 200th anniversary of the publication of Mary Anning's Plesiosaurus, so this was doubly appropriate.

**Caption:** Mary Anning holds Flip the robot plesiosaur during the 2024 Lyme Regis Fossil Festival, with marine reptile workers (left to right) Richard Forrest, Judyth Sassoon, and Luke Muscutt. Down at the bottom is little Darren Naish, who’s evidently shorter in some photos than in others. Image: Hel Naish.

Numerous little froglets were leaving pond 2 at this time, and I photographed what I could. We (John Conway and I) finally got the advertising for TetZooCon 2024 off the ground, and tickets immediately started selling and selling well. New articles at Tetrapod Zoology included my tribute to the late Richard Ellis, and more on squamates (the small-eyed snake Micropechis, the Abronia alligator lizards, and grayiid water snakes). The new North American ceratopsian Lokiceratops saw print in June (June 20th, actually), and I got to write a news piece about it for BBC Wildlife. I also got to cover the very exciting Namibian stem-tetrapod Gaiasia of the Permian for BBC Wildlife in early July.

The philosophy of All Yesterdays. Deeply relevant to my own interests and to the history of this blog was a late June talk by Adrian Currie for Chris Manias’s Popularising Palaeontology workshop. Titled ‘Palaeoart as Science?’, it focused on the philosophy of All Yesterdays, a little book described by some as “potentially the most influential book on palaeoart ever written”.

Conflict exists between the need of the palaeoartist to depict ancient organisms as holistic objects (and to thus avoid speculation) versus their epistemic responsibility, so… what to do? Well, that was the whole point of the talk. I sure wanted to attend the whole thing, but it clashed with a meeting at work so I only caught part of it.

**Caption:** frogs arrive in great numbers in pond 2 during late January and early February, but adults are obvious – as here – throughout the summer. This photo from June 2024 shows at least eight frogs at the water surface. Part of the reason that frogs do this is to ambush insects that come to the pond to drink or (in the case of damselflies and such) lay their eggs. Common pondweed *Lemna minor* is visible in this photo but has since been successfully eliminated from the pond. Images: Darren Naish.

Watching cetaceans and seabirds. In early July, I travelled to Plymouth to board the Pont-Aven for my annual bout of whale surveying – organized and led by ORCA – across the Cornish Sea and the Bay of Biscay. Again, it was spectacularly successful. Of cetaceans, we saw Common Delphinus delphis, Striped Stenella coeruleoalba and Bottlenose Tursiops truncatus dolphin, Harbour porpoise Phocoena phocoena, Cuvier's beaked whale Ziphius cavirostris, Northern minke Balaenoptera acutorostrata, and an incredible 17 Fin whales B. physalus. The Cuvier’s were the highlight for me – this is about the fourth time I’ve seen beaked whales in the wild – and included a substantially scarred male. A Bottlenose dolphin calf (seen adjacent to adults off the Spanish coast) and some Fin whale juveniles seen with their mothers were special moments too.

**Caption:** sea-watching on trips that revolve around marine mammals involves a lot of what you might expect… peering for long periods at large stretches of water. You see animals of many sorts, but the skies in this part of the world are amazing too. I don’t take for granted the complex, dynamic cloudscapes we get here on the Atlantic fringes. Climatic change means that the atmosphere in general is becoming cloudier. Image: Darren Naish.

  **Caption:** I remain absolutely awed by the fact that I’ve now observed live, wild ziphiid whales on a number of occasions. This is a male *Ziphius* with a very white head, observed in the Bay of Biscay. The extensive scarring across his back is very distinctive and this individual could likely be identified from some of these markings if witnessed again. Images: Darren Naish.

  **Caption:** not the best photo in the world, but this *probably* shows a lone Harbour or Common porpoise witnessed in the afternoon, on the approach to Spain. Its brief, secretive reveal was typical for its species, but that very rounded, blunt-tipped apex to the dorsal fin is a bit unusual. Image: Darren Naish.

After photographing some dolphins, I lifted my binoculars off my neck, not realising that the strap for my Canon E380 was tangled up with the binocular strap. Down the camera went, the lens hood smashing hard on the deck. The hood wasn’t broken, and the camera and its lens functioned fine for the rest of the trip, but the thread on the lens was broken and… goodbye lens # 3.

**Caption:** at left, a guillemot showing characteristic disturbance on the water surface. At right, part of a Manx shearwater flock. Images: Darren Naish.

Seabirds are seen on these trips as well as cetaceans, our list for 2024 including Northern gannet Morus bassanus and both Manx Puffinus puffinus and Cory's Calonectris borealis shearwater. I love these trips and will keep going on them as long as I can. ORCA is a marine conservation charity and you can find out about their trips here. Oh, thanks to train delays, some of us almost missed departure, and – thanks to additional train delays – some of us then suffered additional difficulties in getting back home. I use trains here in the UK a lot, and oh my god are they a joke.

**Caption:** I’ve said before that I aim to photograph every Common slow-worm *Anguis fragilis* I encounter, and I guess that goes for deceased ones too. This dead adult female was discovered under a log near Minstead in the New Forest in September 2024, cause of death unknown… but there are what *look like* two holes caused by canine teeth on the left side of its head. A burying beetle was in attendance. In life, slow-worm teeth are all but invisible, since they’re concealed by the lips and gums. But you’ll note that a specimen doesn’t need to be skeletonized for dehydration and decomposition to make the teeth visible in both the upper and lower jaws. Images: Darren Naish.

  **Caption:** much of the coastline of southern England is pretty spectacular if conditions are right. But maybe this can be said about coastlines everywhere. This is the coast of Barton on Sea in west Hampshire, right opposite The Needles on the Isle of Wight, looking east during July 2024. What might you see in the water? Well….

  **Caption:** from high up on the shore at Barton on Sea, I saw the rectangular head of a good-sized animal in the water, and knew instantly what it was. A person and dog at the water’s edge were watching it too, and it didn’t seem worried about getting away in the hurry. So I aimed to get closer…

  **Caption:** as suspected, it was a Grey seal *Halichoerus grypus*, and I got photos of it both as a long and amorphous dark body in the water, and as a co-operative animal that pointed its snout sky-ward and also peered at people and other animals on the land. Grey seals are amazing animals and I’ve had good, close views of them on several occasions now. Images: Darren Naish.

A trip to Tokyo. During early August I published a long-in-development article on dogman at Tetrapod Zoology. Predictably enough, it proved popular and also resulted in my appearance as a guest on ep 46 (season 3) of MonsterTalk (which you can listen to here). We also did a podcats episode of our own on the same subject… the only podcast we got round to recording and releasing during the whole of the year! The big event of the month was a family trip to Tokyo, my first ever visit to Japan and one that could only happen due to a special, all-inclusive discount deal we were lucky enough to find.

I did enough things in Tokyo to write an entire essay (don’t worry, I won’t). Highlights relevant to the TetZooniverse include trips to Ueno Zoo (go here for a review article), Tama Zoological Park (which I have yet to write about), and Tokyo's National Museum of Nature and Science. As with so much in Tokyo, the museum is overwhelming in terms of content and quality. There are exhibits on modern natural history, the diversity of life as a whole, palaeoanthropology, dinosaurs, fossil mammals and ... oh my, the shop. I am forever disappointed by museum shops in the UK, but Japan sure knows how to do them right.

**Caption**: the National Museum of Nature and Science in Tokyo is incredible, and stuffed full of so much fantastic stuff that it’s hard to get round it all in a day. Here’s part of the dinosaur hall, showing numerous fantastic display pieces, some of which are familiar if you know the technical literature on these animals. Image: Darren Naish.

  **Caption**: a montage of just three great items on show at the National Museum of Nature and Science in Tokyo. Left to right: the Miocene desmostylian *Paleoparadoxia tabatai* (I took many photos!), the elasmosaurid *Futabasaurus suzukii* (ditto), and Hachi, the famous faithful dog. Hachi is one of the museum’s most popular exhibits, and the room housing him was one of the busiest there during my visit. Images: Darren Naish.

On that note, I should mention in passing that I succeeded in obtaining a fair number of animal figures while there, essentially all of which are unavailable here in Europe. I wish I’d obtained more, but I couldn’t go completely nuts on the spending. We visited quite a few Godzilla-relevant locations in Tokyo. I didn’t see much wildlife, excepting arthropods.

**Caption:** I’ve said it before and I’m sure I’ll say it again… spending time in green spaces is important, possibly vital, to human wellbeing, and we should see their preservation from a selfish perspective as much as an environmentally conscious one. Here is part of the beech-dominated section of Telegraph Woods that I walk to on regular occasion. This place and others proved vital to me during the covid pandemic. Image: Darren Naish.

SVPCA in Southampton. Almost immediately on returning, I had to get things sorted for 2024’s SVPCA (Symposium on Vertebrate Palaeontology and Comparative Anatomy), this year held at the University of Southampton’s School of Biological Sciences, where I have visiting researcher status. It was a good meeting with much of interest: Chris Barker gave a talk on work he, Lucy Handford, Neil Gostling, myself and others have done on a theropod dinosaur tooth assemblage from the Wealden of East Sussex, and more on that in a minute.

**Caption:** oh wow, *Ancient Sea Reptiles* mentioned and even promoted in someone else’s talk! Ok, the speaker happens to be a good friend and co-author of mine – it’s plesiosaur locomotion expert Luke Muscutt again – but that still counts as a win. Images: Darren Naish.

I also gave a talk myself – an invited talk – and in fact it was the very last talk of the entire conference. It was devoted to my experience of dealing with independent researcher and author Brian Ford and his 'all dinosaurs were aquatic' rhetoric. Near the end of the talk, I included a slide on ‘other contrarians I’ve written about’, and among those is Professor Alan Feduccia, the ‘birds are not dinosaurs’ (BAND) guy (see the 2023 article Alan Feduccia’s Romancing the Birds and Dinosaurs: Forays in Postmodern Paleontology for more). Mike Howgate – who also pushes the naïve, erroneous BAND narrative – was present in the audience (in fact, he’d earlier given a talk about the supposed absence of the furcula across non-bird Theropoda) and responded during the Q&A to my inclusion of Feduccia as a 'contrarian'. It was, apparently, a disgrace that I dare accuse Feduccia of being a contrarian, one that I should be ashamed of. We argued back and forth, and it certainly wasn’t a constructive end to what had otherwise been a positive event.

**Caption:** Darren Naish talks about contrarianism and pseudoscience. Brian Ford and his aquatic dinosaurs might be irrelevant nonsense in one little corner of the intellectual world (remember that his book only sold a few thousand copies, at best), but… is it part of the same ecosystem as pseudoarchaeology and other brands of crankery? Images: Darren Naish.

I’m angry with myself, since I didn't do a sufficiently good job of explaining WHY Feduccia is a contrarian, and indeed why his approach is anti-scientific. As I tried to make clear in my review of Feduccia’s most recent book… yes, the specifics of the argument are faulty, but perhaps more important is the illogical style of argument that's employed: the use of whataboutism, naive falsification and personal incredulity as if they're good arguing tactics. I also wish that I'd thought more beforehand on what it is that makes someone a contrarian, or a crank, since I gave a poor answer when asked about that too and wish I'd done better. I've been thinking a lot about the matter of crankery since that exchange and am preparing an article on it.

The last TetZooCon, the first TetZooTour. SVPCA out of the way, and I gave another dinosaur-themed talk, this time on dinosaur behaviour, at The Novium in Chichester. It was a tie-in for their soon-to-close exhibition on dinosaur ontogeny, amusingly titled Dinosaurs: Hungry Hatchlings!

**Caption:** there might be little… perhaps no… real fossil material on show, but I really like this style of display, here at The Novium, Chichester, and photographed here in September 2024. The combination of life-sized reconstructions, artwork, and an assortment of fossils with data panels, arranged together behind glass, has an appeal… though I wouldn’t arrange things exactly as per this specific montage. Image: Darren Naish.

  **Caption:** more from The Novium, showing (upper left) *Protoceratops* and (right) *Psittacosaurus* models, with a *Tyrannosaurus* montage at lower left. This psittacosaur model is supposed (I think) to represent *P. gobiensis*, so don’t go assuming that it ‘must’ be shown with tail filaments. Not sure about all the armour though… very hypothetical. Images: Darren Naish.

But the big event of September was of course… the last TetZooCon, held at Bush House, London. This event involved a massive amount of work and organisation, and the good news is that it went extremely well. A long article here reports what happened. A live, on-stage recording of the podcast was made but still hasn’t been released because John is a slacker. TetZooCon out of the way, I joined a coach-load of others as we set off on the first ever TetZooTour, and of course I don’t need to talk about that either since a long article already exists on it too. It was also a massive success and I look forward to running similar ventures in the future (albeit not in 2025).

I owe huge thanks to everyone who helped make TetZooCon and the TetZooTour what they were: John Conway, Chris Manias, Steve White, all the speakers, stall-holders and assistants, Georgia Witton-Maclean, Mike and Sue at Everything Dinosaur, driver Marc Bacon, Hel, Toni, and everyone who attended and participated.

**Caption:** I have about a million photos of things that happened during the TetZooTour and still haven’t had a chance to share them (though they *were* shared with the tour attendees). Here are two taken on our Lyme Regis fossil hunting trip, led by Natalia Jagielska of Lyme Regis Museum. Images: Darren Naish.

  **Caption:** if you know anything about TetZooCon, you know that we had a vast amount of artwork, merch and such on sale… things are going to continue, and expand, with 2025’s DinoCon. Here are a few relevant items I obtained at TetZooCon 2024: work by Jay Balamurugan (dodo), Matt Dempsey (dinosaur skeleton and *T. rex*), and Natalia Jagielska (Jurassic stickers). Image: Darren Naish.

While one of those ‘background’ things that occurred during the year was the disbanding of TetZooCon, another was the creation of DinoCon, a new venture fronted by a new team, and overall more palaeo-themed than TetZooCon. DinoCon 2025 is being held at the University of Exeter in England’s south-west on the weekend of August 16th and 17th. It’s going to be epic and we look forward to seeing you in Exeter later this year. Be sure to check out our website here.

**The Cobb stegosaur donation.** On several occasions during 2024, Professor Matthew Cobb of the University of Manchester indicated that he might one day donate his *entire lot of accrued model and toy stegosaurs* to the Tet Zoo Towers collection. That fateful day arrived in early October. Several large boxes, packed full of stegosaurs large and small, diverse in appearance and material composition, were successfully received. A few of the more delicate models were, unfortunately, broken. Anyway… if only I had the space to have then all out on display. One day, one day.

**Caption:** part of – yes, part of – the Cobb stegosaur donation. You’ll perhaps recognise some familiar toys and models, but note as well that there are stegosaur-themed objects of diverse sort here. Image: Darren Naish.

Cryptozoology and Neanderthals in Germany. Also in October… another dream trip, this time to the Neanderthal Museum at Mettmann in North Rhine-Westphalia, Germany, where I was an invited speaker at the German Cryptozoology Symposium 2024. My hosts – Tobias Möser and Markus Bühler – very kindly took me to Cologne Zoological Garden before the meeting began; we were also joined by my long-standing friend Kristina Henschke. It was great, and a long article on the zoo is still due to appear here. For now, all you get is a few pictures…

**Caption:** a scene from the 2024 German Cryptozoology Symposium. Markus Bühler is a long-time reader of Tetrapod Zoology (he has often published comments here as well) and it was great to finally meet him. Here, Markus is discussing discoveries pertaining to European *Gallotia* lizards. His talk also covered surprising finds made in the world of lake-dwelling trout. Image: Darren Naish.

  **Caption:** Cologne Zoo houses a phenomenal collection of birds. At left, Magellanic or Fuegian steamer duck *Tachyeres pteneres*. At right, Blue, Paradise or Stanley crane *Anthropoides paradisea*. Images: Darren Naish.

  **Caption:** at left, Black stork *Ciconia nigra*, one of several Eurasian species housed in a really interesting open-air structure combining vintage stonework with large trees. At right, White-browed, Burchell’s or Lark-heeled coucal *Centropus superciliosus*. Images: Darren Naish.

  **Caption:** more from Cologne Zoo. From left to right: Green peacock *Pavo muticus*, a life-sized photo of a Siberian or Amur tiger *Panthera tigris altaica* with this blog’s author, Blue-throated macaw *Ara glaucogularis*. Images: Darren Naish.

  **Caption:** some carnivorans of Cologne Zoo. A vocal California sea lion *Zalophus californianus* at left, a resting Spectacled bear *Tremarctos ornatus* at right. Images: Darren Naish.

As for the meeting itself, I gave the Mesozoic marine reptiles talk again. Yeah, it’s mostly on the evolution, ecology and natural history of Mesozoic animals but it does of course have relevance to cryptozoology as well given all those hypotheses about the survival of mosasaurs, plesiosaurs and so on beyond the Cretaceous and into the present. The meeting done, and I was very much looking forward to visiting the Neanderthal Museum itself, mostly because of the famous and excellent hominin sculptures by the Kennis brothers. They didn’t disappoint, and the museum itself is… interesting, albeit not as Neanderthal-themed as I was hoping. It’s more arranged around ‘that which makes us human’, which – I submit – is substantially less interesting.

**Caption:** woodland scenes in the Neander Valley, basically at the spot where Neanderthal people fossils were first discovered. Well, actually… the first Neanderthal found in the scientific era comes from Gibraltar where it was discovered in 1848, a bit before the first German find of 1856. A large metal arrow and a stone carving of a brain have both been added to the site. I dislike them both and think that they distract from its aesthetic. Images: Darren Naish.

  **Caption:** some (not all) of the Kennis brothers hominin models on show at the Neanderthal Museum, Mettmann. Left to right: *Homo ergaster*, *H. sapiens*, adult male Neanderthal (Mr. N!), Neanderthal girl. There are more, including other *H. sapiens* individuals and an australopithecine. Images: Darren Naish.

After the meeting, a group of us trekked to the Neanderthal discovery site in the Neander Valley before visiting the nearby Heck cattle and horses. I’m not sure what I was expecting, but the Neander Valley is very much a conventional, picturesque, northern European deciduous woodland, the specific rocky outcrop that yielded the Neanderthal fossils no longer existing. The cattle and horses can both be viewed at close range. The whole thing was a real thrill that I was hugely grateful to experience: thanks Tobias, Markus and everyone else involved.

**Caption:** Heck cattle that live between Mettmann and Düsseldorf, North Rhine-Westphalia, Germany. They’re good looking cattle, but I don’t know how closely they really resemble Aurochs. They’re friendly and very approachable for one thing, and they’re quite a bit smaller than the originals. The two males here are sparring, not fighting. Images: Darren Naish.

  **Caption:** Heck horse close to Mettmann, North Rhine-Westphalia, Germany, where it’s part of a small group that live in semi-captive conditions. They’re supposed to resemble Tarpan but lack the erect manes of that extinct horse. At least some do have a low number of leg stripes though. Image: Darren Naish.

My copies of the Spanish edition of Dinopedia arrived in October. Spanish-speaking countries – including Argentina, Mexico, Chile and Spain itself – have been extremely important in the building of our views on Mesozoic dinosaurs, so it’s fitting and significant that the book now exists in this language. A long article on my recollections of working with Sir David Attenborough was published at the patreon (sorry, I can’t share it publicly. At least, not yet).

**Caption:** ¡Mi libro *Dinopedia* ya existe en español! Espero que los lectores de España, México, Chile y otros países de habla hispana lo disfruten. Muchos de estos países tienen una importancia enorme en términos de lo que nos han enseñado sobre los dinosaurios. [My book *Dinopedia* now exists in Spanish! I hope that readers in Spain, Mexico, Chile and other Spanish-speaking countries enjoy it. Many of these countries have massive importance in terms of what they’ve taught us about dinosaurs.] Images: Darren Naish.

Goodbye Zar. November had a glum start. For more than a decade now, we've had pet guinea-pigs, and for about seven years we've kept Zar, a Teddy (a breed marked for their short, dense, 'rex-type' coat). During September and October, Zar went rapidly downhill, with one issue occurring after the other in different parts of the body. I've learnt a lot about guinea-pigs, most of this thanks to Zar in particular, and I'm sad that he is no more. While digging his grave, I discovered a crumpled shirt in the bottom of the hole I’d made. I extracted it and opened it up. Within was the complete skeleton of a snake, most likely a Corn snake Pantherophis guttatus. What are the odds.

**Caption:** a Zar montage. The two photos at right were taken in 2019 and 2020, and it’s weird now to see how young he looked. I was pretty attached to this little animal, RIP. Images: Darren Naish.

November saw publication of my second technical paper of the year, that on the Ford ‘all (non-bird) dinosaurs were aquatic’ nonsense mentioned above (Naish 2024). An accompanying Tet Zoo article is here. Ford asked for a pdf, and then responded "Interesting - all the more recent evidence substantiates my view. All I need is time to revisit it. But I shall". My plan at one point was to put out a press release on this article (which appeared in Historical Biology), since one angle the paper takes is to compare Ford and his tactics with Trumpian 'post-truthism'. I couldn't make time for this, wasn't ready when the paper saw release, and am frankly too depressed about Trump that using this connection as a light-hearted but informative way of bigging up the research isn't something I want to do.

**Caption:** a new-ish feature at Marwell Wildlife, a great zoo close to me, is the Thriving Through Nature exhibit, which opened in early 2023. It includes (at left) this very impressive rock-dominated aridland display, which features hyraxes, several lizards, and pupfishes. At right, typhlonectid caecilians are on show in the zoo’s tropical house and can be observed swimming and foraging under the water. Images: Darren Naish.

  **Caption:** another view from Marwell Wildlife. Scimitar oryx or Scimitar-horned oryx *Oryx dammah* have been kept at Marwell since its inception in the 1970s and served as the zoo’s official emblem for some time. Marwell is also good on zebras; Imperial or Grevy’s zebra *Equus grevyi* are visible here (there’s a Tet Zoo article on that species **here**). Image: Darren Naish.

November’s real big science news concerned the publication of the Siberian Homotherium cub (yeah… cub, not kitten), a huge deal if you’re interested in Pleistocene animals, extinct cats, or both. I didn’t cover it at Tetrapod Zoology but did write about it for BBC Wildlife. I attended another local herpetology meeting, this time the joint meeting of the British Herpetological Society (BHS) and Amphibian and Reptile Conservation (ARC) at the Bournemouth Natural Science Society, and my efforts to rescue more squamate-themed articles from the archives continued with contributions on leiosaurids and chameleons.

**Caption:** at left, the Bournemouth Natural Science Society (BNSS) building, a location now regularly associated with meetings and conferences relevant to my interests. I have no idea whether this was true prior to the mid-2000s! After the recent BHS-ARC conference, a group of us went on an, erm, urban fieldtrip about Bournemouth, and here’s a captive musk turtle we encountered. Musk turtles have been covered at Tet Zoo in the past, but the article concerned has been destroyed by *Scientific American* and I can’t find it anywhere. Images: Darren Naish.

This was also the time of the year in which Steve White and I were mostly done with the compiling of Mesozoic Art II, the successor to our successful and much lauded Mesozoic Art of 2022. At the time of writing (January 2025), the final cover for MAII has been released and the final proofs have just recently been checked. We’re due to publish in September 2025, unfortunately too late for a launch event at DinoCon. Mesozoic Art II is already available for pre-order here.

**Caption:** there was never an aim to have these books serve as ‘portfolios’ of where we’re at with respect to modern palaeoart, but there’s no avoiding that this is the role they serve, at least in part. We aim to boost and showcase the work and art of world-class contributors, worldwide, and it’s been very rewarding to see the excitement with which these books are received.

  **Caption:** another New Forest scene, this time from October 2024. Taken from somewhere in the vicinity of Beaulieu Heath. Good luck spelling ‘Beaulieu’ if you’re dyslexic. Image: Darren Naish.

Pond adventures. A pond renovation job occupied my weekends throughout November and part of December too. Over recent years, the relatively young-ish pond 2 on my property has become used by a gradually increasing number of frogs. Which is great. But the shallow part of the pond that the frogs use for spawning is too small for the number of frogs and amount of spawn now expected. In addition, the pond contains an old carpet buried beneath it that I really wanted to remove and dispose of properly. And thus changes were made.

**Caption:** the pond I’m mostly talking about in the text here is still too much of a mess for me to want to share photos. But this additional, smaller pond (in Sheila’s garden) is good and ready to go. I want to dig more ponds! If you’re local to me, contact me and I’ll come dig and design one for you. Images: Darren Naish.

The pond was emptied (my aim being to cause minimum disturbance to wildlife, hence the end-of-year timing), the hole containing it was very much modified, and it was reassembled. Water was put back in, and rain happened too. But the water level didn’t go up, it went down, and then it went down some more. There was evidently a hole somewhere in the liner. I got another one and had to repeat part of the process. I went to install the liner aaaand realized immediately that it was too small. I’d ordered one that was the wrong size. So I got another one. Things worked out eventually, and right now pond 2 is complete and fully ready for the frogs that will start calling, competing and fighting during the latter parts of January. The small liner obtained by mistake was used to create another pond on another property, and that’s in good shape and ready to be used by wildlife as well.

**Caption:** birds of many sorts were observed during 2024. Here are Ruddy turnstones *Arenaria interpres* in winter plumage – we just call them turnstones in the UK, seeing as we only have the one species – at the edge of the land in Milford on Sea, southern Hampshire, in November. Image: Darren Naish.

Wealden theropods, Scotland, and hello 2025. December saw the publication of the year’s third of my technical papers, this one being another contribution to our understanding of theropod diversity in the English Wealden Supergroup. As discussed in the Tet Zoo article here, it concerns our analysis of isolated theropod dinosaur teeth discovered in the Hastings Group from the older part of the Wealden (Barker et al. 2024). It received a surprising amount of coverage in the popular press.

In my capacity as a designer and curator of museum exhibits, I travelled by train to Glasgow, specifically the University of Strathclyde, for a one-day workshop event on museum displays organized by Will Tattersdill and Jordan Kistler. There are many things I’d like to see brought to life for show in a museum space – after all, the primary subjects you read about here at Tet Zoo all involve physical things and imagery – so fingers crossed for the future.

**Caption:** I went to Glasgow and talked sea monsters, and that explains the unusual maned animal on the name badge here. At right, Glasgow was cold and misty during my stay. Images: Darren Naish.

As 2024 drew to a close and 2025 kicked into gear, I found myself massively occupied with book editing, DinoCon organisation, and the setting up of media projects that will develop throughout 2025 and beyond. Here in the first month of the year I’m massively overloaded and somewhat despairing about my chances to write regularly for the blog. Almost everything I do professionally arose because of work and time put in here at Tet Zoo, and I have no plans to cut back or reduce my blogging output. Yes, generating novel content for the blog comes at some personal cost, but I want the material I write to be out there, online, and accessible.

**Caption:** one of the last things I accomplished in the year (as in, between Jan 21st 2024 and Jan 21st 2025) was the very overdue review of Creative Beast’s Beasts of the Mesozoic (BOTM) giant *T. rex* figure. I’ve released it on social media but not yet uploaded it to YouTube. Images: Darren Naish.

Subject coverage across the year. Soooo…. let’s play the good ol’ game of phylogenetic representation. I’ve given up on any chance of things ever being ‘fairly’ balanced round here, and I guess I should resign myself to the fact that each year is, basically, a thing of chaos and compromise. In blogging, as in life I guess. Anyway, here’s a list of the year’s articles arranged by subject…

Miscellaneous

Tet Zoo Reviews Zoos: New Forest Wildlife Park, April 2024
In Memory of Richard Ellis, Influential Artist and Author, June 2024
Announcing TetZooCon 2024, the 11th Tetrapod Zoology Convention, June 2024
Tet Zoo Reviews Zoos: Tokyo’s Ueno Zoo, September 2024
The Last TetZooCon, October 2024
The Inaugural 2024 TetZooTour of Southern England, October 2024

Amphibians

Live Spawnwatch Action From Pond 2 at Tet Zoo Towers, February 2024

Turtles

Megalochelys, Truly a Giant Tortoise, January 2024

Squamates

Do Lizards Really Have ‘Mite Pockets’?, March 2024
Meeting Lake Zacapu’s Garter Snake, March 2024
Ray Hoser, Number 1 Taxonomic Vandal, May 2024
The Mysterious Dibamids, May 2024
Side-Stabbing Stiletto Snakes, Redux for 2024, May 2024
The Rehabilitation of Günther’s Black Cameroonian Snake, May 2024
Ikaheka and Other ‘Palatine Draggers’, Cryptozoic Elapid Snakes of Melanesia, June 2024
Arboreal Alligator Lizards of Mesoamerica... and Beyond!, July 2024
Grayiids: Africa Has Water Snakes Too, July 2024
Smaug, Ouroborus, Flat Lizards and More, August 2024
Leiosaurid Lizards: South America, Land of Iguanians, November 2024
What's With All These New Chameleon Names? Chameleons, Part 1, November 2024

Pterosaurs

Azhdarchid Progress, a Personal View, October 2024

Non-bird dinosaurs

The Third Edition of Naish and Barrett's Dinosaurs: How They Lived and Evolved, February 2024
T. rex the Tool Maker -- Testing Controversial Claims About Dinosaur Smarts, April 2024
Post-Truthism, Brian J Ford’s Aquatic Dinosaurs, and the Fate of ‘Too Big To Walk’, November 2024
A New Theropod Dinosaur Assemblage from the Older Part of the English Wealden, December 2024
My Weird 2000 Paper on Tree-Climbing Dinosaurs, December 2024
Heptasteornis, My Beloved; Alvarezsaurids in Europe, the Backstory, January 2025
An American Tyrant in London, Collector's Edition, January 2025

Cryptozoology

A Cryptozoologist’s Bibliography: Matt Bille’s Of Books and Beasts, March 2024
Werewolves in America; the Tale of Dogman, August 2024
Nessie Point and Counterpoint; Who Owns the ‘Facts’ on the Loch Ness Monster?, November 2024

And here’s that data portrayed as a graph…

It's obvious from the graph that 2024 was unusual in terms of coverage here. I’m surprised to see that I never found time to write about mammals (or indeed synapsids of any sort) or birds: that seems very weird. I’m also really weirded out by the total absence of fossil marine reptiles here, given the importance they obviously had what with the many events that revolved around my *Ancient Sea Reptiles* book.

The ‘equal’ coverage received by amphibians, turtles and pterosaurs is vaguely encouraging but for the fact that we’re talking about lone articles. Non-bird dinosaurs were well served, both because I had a reasonably healthy year in terms of technical publications (Barker et al. 2024, Caspar et al. 2024, Naish 2024) but also because of a few articles relating to articles of the past. A few cryptozoology-themed articles kept that subject alive at the blog as well. But of course – as deliberately planned – 2024 was Squamate Year at Tet Zoo, in which I aimed to rescue and republish as many articles on that group as I could (for those who don’t know, my older articles at ver 2 and ver 3 have been ruined by their now defunct hosters, and I feel compelled to rescue all of this otherwise lost material). I didn’t get through half as many articles as I wanted to, so I’d like to make 2025 a Squamate Year as well. I’d be interested to know what you, dear reader, feel about this… is that enough squamates for now, or are you good for me to continue?

**Caption:** here’s Teddy the West Highland terrier in two of his guises. He came with me on numerous walks throughout the course of the year, often accompanying these adventures with observations on urban planning and the socio-political landscape. A leg operation during the latter part of 2024 put a temporary end to this, however. Images: Darren Naish.

And that’s where I must end. I found 2024 an unbelievably full-on year with too many things happening for me to really keep track of. That’s not because I deliberately seek out all of these overlapping tasks and events: it’s an inevitable consequence of what I’m involved in at this point of my life and career. 2025 is looking to be the same, with tons of stuff happening. It all feels very chaotic.

**Caption:** I just know that you want to hear more about the fabled rediscovery of the Safari Ltd Great crested newt model I mentioned earlier in the article. Well, here are assorted images, including (lower left) a screengrab from the ‘discovery’ video I shared on social media and (at right) a montage created for Instagram. At upper left is a screengrab from eBay, giving you some idea of what some people think that this figure is worth. I might have made a few rash amphibian-themed purchases in my time… but even I think this is ridiculous.

As ever, some major slow-burn giant projects weren’t finished during the year, which frustrates me now just as much as it has before. Nevertheless, things are looking good for 2025, so stay tuned for some very exciting announcements. Huge thanks as ever to everyone who reads the blog, to everyone who leaves comments (except for the haters and weirdos), and especially to those who provide support at patreon. And here’s to another year of operation!

For previous TetZoo articles on birthdays and other landmarks, see…

Happy first birthday Tetrapod Zoology (part I), January 2007
Happy first birthday Tetrapod Zoology (part II), January 2007
Happy second birthday Tetrapod Zoology (part I), January 2008
Tetrapods of 2007 (happy birthday Tet Zoo part II), January 2008
Happy THIRD birthday Tet Zoo, January 2009
Tet Zoo = 4 years old today, January 2010
2009, a year of Tet Zooery, January 2010
Four years of Tet Zoo: to infinity... and beyond!, April 2010
It is with some dismay that I announce Tet Zoo's first hemi-decade, January 2011
Tet Zoo 5th birthday extravaganza, part II, January 2011
Happy Birthday Tetrapod Zoology: SIX YEARS of blogging, January 2012
Happy 6th Birthday, Tetrapod Zoology (part II), January 2012
Tetrapod Zoology enters its 8th year of operation, January 2013 (I cannot find any intact versions of this article, thanks SciAm)
Today marks NINE YEARS of Tetrapod Zoology, January 2015
Tetrapod Zoology 10th-Birthday Extravaganza, Part 1: 2015 in Review, January 2016
Tetrapod Zoology 10th Birthday Extravaganza, Part II: the Rest of 2015 Reviewed, January 2016
Tetrapod Zoology 10th-Birthday Extravaganza, Part 3: Tet Zoo's Tetrapod Treatment in 2015, January 2016
Today Is Tet Zoo's 11th Birthday, January 2017
The 12th Year of Tet Zoo, January 2018
The Tet Zoo 12th-Birthday Event, Part 2, January 2018
The Much Belated Final Part of the Tetrapod Zoology 12th Birthday Event, December 2018
Tetrapod Zoology Is A Teenager Now, January 2019
Tetrapod Zoology's 14th Year of Operation, 2019 in Review, January 2020
On Tetrapod Zoology’s 15th Birthday, the Year in Review, January 2021
Happy 16th Birthday, Tetrapod Zoology, February 2022
Four Years of Tetrapod Zoology ver 4, July 2022
The 17th Year of Tetrapod Zoology: 2022 in Review, January 2023
Five Years of Tetrapod Zoology ver 4, July 2023
Tetrapod Zoology Reaches 18 Years of Age, January 2024

Refs - -

Barker, C. T., Handford, L., Naish, D., Wills, S., Hendrickx, C., Hadland, P., Brockhurst, D. & Gostling, N. J. 2024. Theropod dinosaur diversity of the lower English Wealden: analysis of a tooth-based fauna from the Wadhurst Clay Formation (Lower Cretaceous: Valanginian) via phylogenetic, discriminant and machine learning methods. Papers in Palaeontology 2024, e1604.

Muscutt, L. E., Dyke, G., Weymouth, G. D., Naish, D., Palmer, C. & Ganapathisubramani, B. 2017. The four-flipper swimming method of plesiosaurs enabled efficient and effective locomotion. Proceedings of the Royal Society B 284, 20170951.

Naish, D. 2024. The response to and rejection of Brian Ford’s Too Big to Walk, a 21st century effort to reinstate the aquatic dinosaur hypothesis. Historical Biology 10.1080/08912963.2024.2421268

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I’m overloaded with work and deadlines, and unable to complete anything new. So please enjoy this classic Tet Zoo article from the archives, originally published at ver 2 back in 2008 (an archived version is here)…

**Caption:** *Dynamosaurus imperiosus* wasn’t named for great material, namely an incomplete mandible with teeth, some cervical and dorsal vertebrae, and parts of the pelvis and hindlimb. This was originally accessioned at the AMNH in New York as AMNH 5866, but was relabelled BMNH R7994 when it joined the collections of the NHM London in 1960. The mandible has, on occasion, been on display: here it is in December 2016 (without flash above, with flash below). Images: Darren Naish.

When Tyrannosaurus rex was named by Henry Osborn in 1905, it was described alongside a second gigantic theropod, the armour-plated, Ceratosaurus-like Dynamosaurus imperiosus, collected in 1900 near the Cheyenne River, Wyoming. Unlike T. rex, Dynamosaurus possessed irregularly shaped bony plates on its back and sides. These later turned out to be from an ankylosaurid (probably Ankylosaurus), and in fact small craters on the scutes look like T. rex tooth marks, suggesting that the scutes were part of the tyrannosaur’s stomach contents (Carpenter 2004, Breithaupt et al. 2008a, b). Osborn later realised that Dynamosaurus was synonymous with T. rex*, but continued to think that the animal possessed armour plates (Osborn 1917).

Ankylosaurid armour may well have helped protect these animals from tyrannosaurid predators, but the awesome bite strengths reconstructed for T. rex suggest that the biggest tyrants might still have been able to do serious, fatal damage to an ankylosaurid. Scott Hartman once depicted this in a reconstruction titled ‘Armor? What armor?’.

Caption: there aren’t that many colour photos of the NHM T. rex mount, but this one has been reproduced in several popular books. Note that the specimen is a half panel-mount, the skull, much of the ribcage and various of the other elements being reproductions (made of fibreglass, I think). The whole thing adheres to an environmental painting. The gallery held several of these and I don’t know anything about their history, not have I seen good, complete versions of them. Thanks to Jim Robins for sourcing this picture in the first place. Image: NHM archives.

In 1960 the Dynamosaurus specimen was (together with parts of three other T. rex specimens) sold to the then British Museum (Natural History), and here it was mounted in the museum’s old dinosaur gallery in a rather ‘modern’ pose: that is, with its body and tail near-horizontal and its tail well up off the ground. Those who’ve commented on this have usually noted that the museum’s Barney Newman deliberately chose to depict the animal in this way. Indeed, Newman (1970) explained how correct articulation of the vertebrae meant that the animal simply had to be arranged like this.

**Caption:** Erwin S. Christman’s illustration of the vertebral column and pelvis of *T. rex*, as understood at the time (from Osborn 1917). This depicts AMNH 5027, the *T. rex* specimen discovered by Barnum Brown in 1908, later described by Henry Osborn, and today on show in the AMNH Hall of Saurischian Dinosaurs. It was remounted in a horizontal pose in 1996.

Newman was not, actually, the first person to depict a tyrant dinosaur in a horizontal-bodied posture. Erwin S. Christman illustrated the T. rex back and tail in horizontal pose for Osborn’s 1917 paper on theropods (Osborn 1917), and one might conclude from this that Christman (and perhaps Osborn too) actually advocated a wholly ‘modern’ posture for tyrant dinosaurs. In, however, an earlier paper, Osborn (1913) had drawn attention to Christman’s on-going work on reconstructing the life postures of theropods, and it seems from words used here that Christman imagined theropods to walk with their backs held diagonally, their tails sloping down to the ground. Various dinosaur reconstructions supervised by Osborn during the 1920s are depicted this way too, so I think that Osborn regarded bipedal dinosaurs as ‘diagonal-bodied’, not ‘horizontal-bodied’.

Newman (1970) noted that Ned Colbert had depicted other large theropods (Allosaurus and Gorgosaurus) in horizontal postures in his 1965 book The Age of Reptiles, and I think that what Newman was referring to here are Margaret Colbert’s life reconstructions of these animals. They indeed show horizontal-bodied big theropods in a book of 1965.

**Caption:** Margaret Colbert’s 1965 reconstruction of a Late Cretaceous ‘Belly River’ scene in western Canada, showing the tyrannosaurid *Gorgosaurus* at left. There might be amphibious lambeosaurines and tail-dragging ankylosaurids here, but the theropod is fairly modern in appearance. Presumably the Colberts (both Ned and Margaret) imagined theropods to walk or run with raised tails and mostly horizontal bodies, and evidence from published statements implies that many workers thought this through the 20th century. Image: Margaret Colbert, from Colbert (1965).

So, was Newman’s tyrannosaur mount as far-sighted and innovative as might appear? It soon entered the ‘mainstream’ literature. Halstead (1975), for example, illustrated T. rex in the postures depicted by Newman, also repeating Newman’s idea that tyrannosaurs might have used their short didactyl forelimbs as props when raising themselves from a resting posture. But Newman’s ideas weren’t entirely modern. He thought that the tail must have swung far to the side with each step, and that these dinosaurs must have walked with an “ungainly waddling” rather “than the formerly postulated majestic striding” (p. 123). Modern work has not supported this idea, which never seemed reasonable anyway.

**Caption:** Newman (1970) illustrated *T. rex* lying on the ground, and then showed how it might have used its forelimbs as props and an upward throw of its head to raise itself up and stand. This image is the fourth in the sequence. The animal isn’t fully horizontal, what with a diagonally sloping tail. The overly flexed (curled) look of the left foot is probably outside the range of what was possible, the forelimb is disarticulated at the shoulder joint, and the neck is likely too erect relative to what was normal. Versions of this diagram appeared in several popular dinosaur books of the 1970s and 80s.

What interests me in particular is Alan Charig’s claim that the BM(NH) T. rex “was mounted with its body in a far too horizontal position: this was done because it would otherwise have been too tall for the Gallery. Newman, who made the mount, has attempted to rationalise this (1970) by stating that the posture was much more bird-like than is suggested by earlier mounts” (Charig 1972, p. 137). Charig and Newman were at loggerheads, and this exchange on the posture of the BM(NH) T. rex “did nothing to improve relations between the two” (Moody & Naish 2010). But, personal spats aside, this must mean that Charig regarded Newman’s re-posed T. rex to be too horizontal, that Charig wanted it to be more upright. Maybe not bolt upright like a standing kangaroo, but more diagonally posed, at least.

Whatever the true explanation behind the posture of the London T. rex, I remain upset that I never got to see it. And I’ve decided that I want this article to serve as a repository for images of that now lost mount, so I’ll be adding additional pictures of it here, as I find them. Thanks if you know of any that aren’t already shared here.

**Caption:** another image of the NHM’s old *T. rex* mount, but a poor, black and white one. I’ve lost track of its source! Note the small chain-link fence at the bottom of the photo, this showing that it was taken at a different time from the other black and white picture here.

  **Caption:** at left, colour image from Michael Benton’s 1984 *Pocket Dinosaurs* (though this is the 1987 edition), showing the *T. rex* in the background. This photo confirms that the half skull used in the panel mount is wholly separate from the complete skull replica housed in the case (I knew that already, because the half skull is kept in the collections while the complete skull is on show in the galleries). At right, a black and white photo from Michael Tweedie’s 1977 *The World of the Dinosaur*. The accompanying caption states that “The skeleton is displayed in the animal’s most probable standing posture against a background of Upper Cretaceous scenery”, and thus includes obvious reference to Newman (1970).

  **Caption:** another colour version, but this time composited such that the skeleton is stepping out of the original and overlapping the adjacent sections of the book. This is from Michael Benton’s 1993 *Dinosaurs: Living Monsters of the Past*, a valuable compendium of unusual and rarely reproduced pictures. It includes another view of the gallery, this time the *Diplodocus* side.

  **Caption:** the first episode of the 1975 BBC series *Fabulous Animals* (for so long regarded as lost media, but now online at YouTube) includes a segment on dinosaurs, and here we see David Attenborough in the fossil reptile gallery, alongside the *T. rex*.

  **Caption:** the credits to the 1982 Longman Group children's documentary *Dinosaurs: Fun, Fact & Fantasy* include this view of the mount in side view. Thanks to Brian Choo for reminding me about this.

For previous Tet Zoo articles on tyrannosauroids and related issues, see…

Naish and Barrett's Dinosaurs: How They Lived and Evolved, November 2016
Ornithoscelida Rises: A New Family Tree for Dinosaurs, March 2017
The Second Edition of Naish and Barrett’s Dinosaurs: How They Lived and Evolved, November 2018
Dr Angela Milner and the Discovery of Baryonyx, August 2021
Two New Spinosaurid Dinosaurs from the English Cretaceous, September 2021
The Eotyrannus Monograph Is Finally Published, July 2022
The Third Edition of Naish and Barrett's Dinosaurs: How They Lived and Evolved, February 2024

Huge thanks as ever to everyone who supports me and what I do at patreon. That funding allows me to make time for this blog, and for the technical research I publish on the side. Click here to join the collective and see unpublished work as it comes together.

Refs - -

Breithaupt, B. H., Southwell, E. H. & Matthews, N. A. 2008a. The ‘powerful imperial lizard’ Dynamosaurus imperiosus: the world’s first Tyrannosaurus rex comes to London. In Moody, D., Buffetaut, E., Martill, D. & Naish, D. (eds) Dinosaurs and Other Extinct Saurians: A Historical Perspective. The Geological Society of London (London), pp. 22-23.

Breithaupt, B. H., Southwell, E. H. & Matthews, N. A. 2008b. Wyoming’s Dynamosaurus imperiosus and other discoveries of Tyrannosaurus rex in the Rocky Mountain West. In Larson, P. & Carpenter, K. (eds). Tyrannosaurus rex: the Tyrant King. Indiana University Press (Bloomington and Indianapolis), pp. 57-61.

Carpenter, K. 2004. Redescription of Ankylosaurus magniventris Brown 1908 (Ankylosauridae) from the Upper Cretaceous of the Western Interior of North America. Canadian Journal of Earth Sciences 41, 961-986.

Charig, A. J. 1972. The evolution of the archosaur pelvis and hind-limb: an explanation in functional terms. In Joysey, K. A. & Kemp, T. S. (eds) Studies in Vertebrate Evolution. Oliver & Boyd (Edinburgh), pp. 121-155.

Colbert, E. H. 1965. The Age of Reptiles. Weidenfeld and Nicolson, London.

Halstead, L. B. 1975. The Evolution and Ecology of the Dinosaurs. Peter Lowe, London.

Moody, R. T. J. & Naish, D. 2010. Alan Jack Charig (1927-1997): an overview of his academic accomplishments and role in the world of fossil reptile research. In Moody, R. T. J., Buffetaut, E., Naish, D. & Martill, D. M. (eds) Dinosaurs and Other Extinct Saurians: A Historical Perspective. Geological Society, London, Special Publications 343, pp. 89-109.

Newman, B. H. 1970. Stance and gait in the flesh-eating dinosaur Tyrannosaurus. Biological Journal of the Linnean Society 2, 119-123.

Osborn, H. F. 1913. Tyrannosaurus, restoration and model of the skeleton. Bulletin of the American Museum of Natural History 32, 91-92.

Osborn, H. F. 1917. Skeletal adaptations of Ornitholestes, Struthiomimus, Tyrannosaurus. Bulletin of the American Museum of Natural History 32, 133-150.

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Among the most fascinating of Mesozoic theropods are the alvarezsaurids, a mostly small-bodied group of maniraptoran coelurosaurs characterized by modified, ‘pick-like’ forelimbs, a lightly built, shallow lower jaw, tiny, simple teeth and elongate, slender hindlimbs…

**Caption:** when alive, at least some alvarezsaurids (specifically those in the same clade as *Mononykus* from Mongolia) might have reminded us of long-legged birds like bustards or plovers, albeit equipped with shortened forelimbs and an owl-like facial disk. At left is the *Mononykus* design I and others devised for *Prehistoric Planet*; at right is the reconstruction included in **my 2021 book *Dinopedia***. Images: © Apple TV / BBC Studios; Darren Naish.

The story of how we came to discover alvarezsaurids and work out what they are is complex and I don’t want to cover it here – see the section in Dinopedia (Naish 2021) for a summary – but suffice to say that we currently regard them as an early-diverging group within Maniraptora that are apparently specialized for a diet of social insects. Mononykus of Mongolia, named in 1993, was the first member of the group to become scientifically well known, and much of the early discussion of what these animals were like and where they might fit, in phylogenetic terms, revolved around Mononykus* specifically.

It was originally published in April 1993 as Mononychus but this proved preoccupied by a weevil. A replacement spelling for the dinosaur appeared in May 1993.

Caption: it’s old hat now and no longer the source of discussion, but the phylogenetic position and unusual anatomy of Mononykus – the first alvarezsaurid to be well understood – was hot news in the 1990s, and a great many news pieces and toings and froings in the technical literature were devoted to where it might fit in the dinosaur family tree. Included here are the original Nature paper of 1993 as well as articles from Natural History, New Scientist and The Auk.

The possession within alvarezsaurids of highly sensitive hearing and a highly reduced hand that (in some taxa) is monodactyl have kept them in the news over recent years. At the time of writing, a new study on the evolution of body size within the group has also been deemed newsworthy, since it shows how modification in body size over the course of alvarezsaurid history was linked with key events that affected other aspects of their anatomy (Meso et al. 2024).

‘Seven Town’ bird backstory. In this article I want to talk about one alvarezsaurid – or possible alvarezsaurid, anyway – and that’s the poorly known Heptasteornis andrewsi Harrison & Walker, 1975 from the Maastrichtian of Sânpetru in the Haţeg Basin, Romania. The name means ‘Andrews’s Seven Town bird’, the ‘Seven Towns’ being a reference to “the area of origin” (Harrison & Walker 1975) (these authors used the old, Hungarian name for Sânpetru, namely Szentpéterfalva). The name applies specifically to A4359, a partial hindlimb element that involves the distal end of the tibia and the adjacent, mostly fused astragalus, one of the proximal tarsal bones. Because we’re talking about a compound bone that involves both the tibia and tarsus, it gets termed the tibiotarsus. A4359 has a width of about 3 cm and a thickness at its distal end of nearly 2 cm.

**Caption:** to my knowledge, the precise location from which Nopcsa collected his maniraptoran specimens remains unknown, but it was somewhere in the vicinity of Sânpetru (shown here), a village in Hunedoara County [UPDATE: I previously featured the wrong Sânpetru here, since there’s another, further to the east and in Brașov County, Transylvania]. The Retezat Mountains loom nearby. Image: Țetcu Mircea Rareș, CC BY-SA 3.0 (**original here**).

My aim here is to talk in particular about the backstory to a paper that Gareth Dyke and I published on this animal 20 years ago (Naish & Dyke 2004). Somewhat surprisingly, I’ve never done this before. So, yes, this is another introspective article.

Heptasteornis is one of several maniraptorans known from the Romanian Maastrichtian, nearly all of which are enigmatic and known from frustratingly incomplete remains. The exception is Balaur bondoc Csiki et al., 2010, known from a partial articulated skeleton as well as a partial forelimb from a second individual. Balaur is found in some studies to be a velociraptorine dromaeosaurid (e.g., Brusatte et al. 2013, Turner et al. 2021), and in others to be part of Avialae, the bird clade (Cau et al. 2015). I think that the latter is more likely correct, but – whichever hypothesis we prefer – this is an interesting case of different data sets finding different positions for the same taxon. Either way, Balaur was wholly unknown when the research discussed here was being performed.

**Caption:** this reconstruction, by Jaime Headden and created for **Cau *et al*. (2015)**, depicts *Balaur* as it might look as a jeholornithiform-grade bird, rather than a dromaeosaurid. The differences are only obvious to a specialist but involve the form of the tail, shape of the snout and configuration of the teeth. As yet, no part of *Balaur*’s skull has been recovered. Image: Jaime Headden / **Cau *et al*. (2015)**.

While discussing Late Cretaceous Romanian theropods during the early 2000s, Gareth and I realized that the ascending process of the astragalus in Heptasteornis was similar to that of Asian alvarezsaurids. In particular, it had an unusual wavy or notched median margin, a feature not seen in other theropods. Gareth had recently contributed to a paper that described a new specimen of the Mongolian alvarezsaurid Shuvuuia deserti (Suzuki et al. 2002), and his familiarity with the tibiotarsus of that animal (and other indisputable alvarezsaurids) is what led to the idea of Hepasteornis’s potential alvarezsaurid nature.

A4359 was not at all new at this time. It had, in fact, been discovered by famed nobleman and palaeontologist Baron Franz Nopcsa during the early 1900s and – together with several other, similar bones – was gifted by him to the British Museum. Here, they were studied and described by Charles W. Andrews (1913) who thought that they represented a new sort of cormorant-like bird that he named Elopteryx nopcsai. Andrews actually attached this name to a partial femur that Nopcsa had found. A4359, Andrews thought, probably belonged to the same species because both shared a “close similarity of the sculpturing of the surface of the bone” (p. 196) – remember that detail – even though he regarded the femur as being from a swimmer and the A4359 tibiotarsus as from an “ambulatory” bird.

**Caption:** at left, a montage of the *Elopteryx* specimens described by Andrews (1913). In the upper row, we see the proximal end of a left femur (specimen A1234), and in the lower row A4359, the fragment of tibiotarsus later named *Heptasteornis andrewsi*. At right, Jenny Halstead’s reconstruction from 1989 of *Elopteryx* as a “cormorant-like” seabird. Images: Andrews (1913); Jenny Halstead.

For much of the 20th century, authors were generally happy to regard Elopteryx as some sort of Cretaceous proto-cormorant, and a few books feature reconstructions of such an animal. However, things began to become a little more chaotic during the 1970s.

In fact, it might be fair to say that these specimens are, today, notorious in that they’ve been allocated various positions in the theropod family tree over the last six decades, and the whimsical title I chose for our paper (Naish & Dyke 2004: ‘Heptasteornis was no ornithomimid, troodontid, dromaeosaurid or owl: the first alvarezsaurid (Dinosauria: Theropoda) from Europe’) is a deliberate reference to this. A cynical response is to mock this repeated reclassification and imply that the seemingly eternal reinterpretation of dinosaur remains that occurs in some corners of the literature is akin to tasseography or palmistry (Feduccia 2020). That’s naïve as well as unfair, since what it really reflects is that theropods of several groups – including birds – are highly similar in the anatomy of their tibiotarsi, and it can be difficult to spot the small details specific to the groups in question. Reclassification and reinterpretation of the sort discussed here also reflects the self-correction and scepticism that’s inherent to the scientific process.

The age of giant Cretaceous owls. Perhaps the best-known aspect of the Heptasteornis saga is that this taxon was identified by Harrison & Walker (1975) as an owl, as was Bradycneme draculae, another Romanian Nopcsa taxon recognised by the same authors, also based on the distal end of a tibiotarsus. Both were united by Harrison & Walker (1975) within Bradycnemidae, a group of supposed archaic owls. This explains why owls have been depicted as denizens of Late Cretaceous wildlife scenes on a handful of occasions.

**Caption:** a scene depicting Late Cretaceous life as imagined during the mid-1970s, by Italian artist Giovanni Caselli. It’s true that early members of modern groups of squamates, insects, plants and so on were present during the Late Cretaceous, but the once popular idea that archaic kin of rails, gulls, waders and so on were present at this time no longer appears correct. And... what’s that bird at far left? Image: Giovanni Caselli, from Halstead (1975).

This was always a fascinating idea, especially given that the owls concerned would have been gigantic. You’ll recall me saying that the A4359 tibiotarsus is about 3 cm wide. For comparison, in the biggest living owls the tibiotarsus is about 1 cm wide, so a conservative interpretation of Heptasteornis as an owl is that it was about 2 m tall in vertical standing pose. This was also a problematic idea given how other data on neornithine phylogeny shows that diversification within Neoaves (the clade that owls and their kin belong to) was predominantly a post-Cretaceous event.

**Caption:** yup, that’s an owl at left in the Caselli scene (from Halstead 1975) shown above. The owl doesn’t look that big; it looks something like an *Athene* owl (among the most familiar of which is termed the Little owl!). My photo isn’t blurry: this part of the painting in the book is slightly out of focus. It’s of some interest that Caselli’s painting was published so close in time to the publication of Harrison & Walker (1975). This makes me think that Caselli, presumably via Halstead, knew of that research before it was out, perhaps because Halstead was on the editorial board of the journal where it was published (his own research appeared there – that is, in the journal *Palaeontology* – several times). At right, Greg Irons’s 1978 illustration of a gigantic *Bradycneme* (accidentally mis-spelt *Brachycneme*), from the 1980 book *The Last of the Dinosaurs*. After discussing the small ankylosaur *Struthiosaurus*, the text states “He must have fled from the colossal owl *Brachycneme* [sic], standing five to six feet from beak to claw! The closely related *Heptasteornis* was nearly as large”.

Indeed, those authors who commented on the Cretaceous owl proposal – beginning with influential palaeornithologist Pierce Brodkorb in 1978 – disagreed with it, most usually saying that the specimens in question were indeterminate but surely non-avian (see Naish & Dyke 2004). Storrs Olson, never afraid to hold back when he sensed blood in the water, described Harrison and Walker’s owl idea as “Certainly one of the most egregious errors ever made in avian paleontology” (Olson 1985, p. 129).

An ode to Harrison and Walker. It’s useful at this point to have some awareness of Colin J. O. Harrison and Cyril Walker’s work on bird fossils and their approach to identification, since their paper on the Romanian ‘Cretaceous owls’ is no outlier.

**Caption:** at left, screengrab of that *egregious* Harrison & Walker (1975) paper. At right… there’s a Tet Zoo ver 2 article on Cyril Walker, published on his death in 2009, but an intact version is no longer findable. **It’s here.**

From the mid-1970s onwards, Harrison and Walker embarked on a career of identifying fragmentary fossil bird fragments (mostly from the European Paleogene) on the basis of their similarity to the bones of modern species, which they had ready access to thanks to the massive, world-class collections of the Natural History Museum today archived at Tring. Harrison and Walker were literally noting genus-level similarities for isolated toe bones, partial shoulder blades and mere fragments of skull bones. They got a lot wrong – this making it easy to criticise them – but they also got a lot right, and an argument can be made that a vast number of bird fossils were only reported in the published literature because Harrison and Walker regarded them as worthy of identification. I never did meet Colin, but I knew Cyril and always found him to be extremely generous with time and information, and also appropriately fair when faced with criticism. He’s also one of the few senior academics who – back when I was a new PhD student – asked me for my opinion during a conversation, something I wasn’t used to and which I’ve never forgotten.

**Caption:** this figure – from Harrison & Walker (1975) – gives an idea of how Harrison and Walker made the identifications they did. They noted superficial similarities between the fossil fragments and the bones of extant birds (initially coming up with these identifications thanks to their prior knowledge of anatomical characters), and then used detailed similarities and differences to work out how close the fossils were to their supposed extant kin. Image (a) here shows an extant Barred owl, this representing one of three groups deemed by these authors especially similar to the Romanian fossils. The others were (d) true falcons and (e) caracaras. Image: Harrison & Walker (1975).

Beyond the owls. In a ‘post-bradycnemid owl’ world – where the Romanian specimens were universally regarded as being from non-bird theropods but of uncertain sort – authors suggested various solutions to their identity.

In 1988, Greg Paul argued that these specimens were sufficiently similar to the tibiotarsi of troodontids to be included in that group, and perhaps even regarded as representing a European species of Troodon (Paul 1988). A few subsequent authors followed this troodontid identification. Le Loeuff et al. (1992), however, thought that the remains were better referred to Dromaeosauridae, the Velociraptor family. Csiski & Grigorescu (1998) then argued that the tibiotarsi weren’t from maniraptorans at all. They thought that A4359 possessed a “cranial transverse groove on [the] astragalar condyles”, a structure lacking in maniraptorans. But that’s an error, since what they interpreted as a transverse groove is in fact a break, and it’s in the wrong place for the groove in question anyway (Naish & Dyke 2004).

**Caption:** identifications that have been published for *Heptasteornis* (my drawing of the holotype at upper left). 1. Alvarezsauridae. 2. Ornithomimidae. 3. Troodontidae. 4. Dromaeosauridae. Images: Darren Naish.

One final comment on Hepasteornis appeared in the literature prior to Naish & Dyke (2004), this time in a poorly known article (Martin 1997) published by palaeornithologist Larry Martin. Martin, who died in 2013, was a bit of a wildcard in his contribution to palaeontology and a lot could be written about him. He was fully committed to the ‘birds are not dinosaurs’ (BAND) movement, and in that context said and published things that were patently false and also inherently dishonest. But he was also brilliant, with an exceptional eye for detail and a skill in interpreting and identifying vertebrate fossils of all sorts. He was also a champion talker and highly entertaining speaker.

Anyway, Martin was very much against the idea – popular among some researchers for a time during the 1990s – that alvarezsaurids might be highly modified birds (this is another complex aspect of the alvarezsaurid story, and one I have to avoid here). He instead thought that they were ornithomimids (the group popularly known as ‘ostrich dinosaurs’ or ‘ostrich mimics’), and in a 1997 article titled ‘The difference between dinosaurs and birds as applied to Mononykus’, went through all the skeletal features of Mononykus in order to show that it was more like an ornithomimid than it was like a bird (Martin 1997). On getting to the tibiotarsus, he noted (citations removed for ease of reading)…

“It is fair to point out that many workers recognized the nonavian status of Mononykus. This is undoubtedly the result of its being well described and based on excellent material. More scrappy remains of a very similar animal Elopteryx from the Cretaceous of Transylvania was also described as a bird, and some of its referred material erroneously as a Cretaceous owl” (Martin 1997, p. 341).

**Caption:** Martin’s 1997 figure that compares the skeletal elements of *Mononykus* with those of ornithomimids. He wasn’t the last person to compare alvarezsaurids with ornithomimids and in fact other workers since have regarded the groups as closely related; this is less popular, however, than the hypothesis that alvarezsaurids are maniraptorans. (20) shows one of the Romanian tibiotarsal fragments and (22) the *Elopteryx* femur. Image: Martin (1997).

This wasn’t, actually, the first time that Martin had referred to the Romanian specimens. Back in 1983, and hence prior to the early 1990s recognition of alvarezsaurids, he discussed them in a section of an article titled ‘Fossils that are not birds’ and wrote “they appear to be struthiomimids in view of the extreme enlargement of the astragalus and reduction of the calcaneum (Martin 1983, p. 301). For ‘struthiomimids’, read ornithomimids.

What this means is that Larry Martin, not Naish and Dyke, wins credit as the first person to draw comparison between Heptasteornis and alvarezsaurids. However, I don’t recall being influenced by this when writing the paper.

On publishing fragmentary remains. When it comes to the publication of Naish & Dyke (2004), we were spurred to publish not just by the influence of the Suzuki et al. (2002) paper mentioned above, but also by Hutchinson & Chiappe’s (1998) then-recent suggestion that remains from the Hell Creek Formation of Montana also represented an alvarezsaurid. The material they used to make their claim wasn’t of the very best sort, involving a right pubis and part of an attached ischium.

**Caption:** initial evidence pointing to the presence of alvarezaurids in the Maastrichtian of North America was poor, but today enough is known that we consider their presence sufficiently supported by specimens. *Trierarchuncus prairiensis*, published in 2020, was named for ungual phalanges and a few other limb fragments, and other Maastrichtian remains from western North America presumably belong to it as well. Image: Badlands Dinosaur Museum / Boban Filipovic.

There’s a phenomenon here worth talking about. Once authors start proposing identifications based on fossil material of a certain, shall we say… lesser quality, others are inclined to follow suit. I do NOT mean to imply that this is a bad thing, since fragmentary and isolated fossil remains should be published, often being important and useful, and hypotheses of identity often alert others to the possible presence of the relevant animal group in a respective area or sedimentary unit.

Another issue worth noting here is that making identifications on the basis of fragmentary remains is a substantially risky endeavour. Speaking from experience, I know how easy it is to become convinced that a single skeletal element can look like that of a group you know, only to be proved completely wrong down the line. One random example: ichthyosaur expert Chris McGowan kept identifying ichthyosaur coracoids from the Late Cretaceous of North America. But, as he admitted himself (McGowan 1991), he had ‘ichthyosaur bias’, and other workers showed that his specimens were plesiosaurian pelvic bones, not ichthyosaurian coracoids at all (Baird 1984). Seen from that point of view, our alvarezsaurid identification is risky and – while I’m fine to see it advocated for now, in the absence of better data – I won’t be surprised if it proves wrong in future. Ditto for other supposed alvarezsaurid fragments from Romania.

A paper is (eventually) published. Whatever the caveats, by early 2003 we were ready to attempt publication. We initially submitted a manuscript (with Cyril Walker on the authorship) to Journal of Paleontology but got rejected for the usual ‘we have too many submissions’ reason. We then tried Proceedings of the Geologists’ Association (by now without Cyril; he didn’t feel that he’d contributed enough to warrant authorship), who rejected it for being ‘too specialized’ for their readership. In August 2003, it was submitted to our third and final venue: Neues Jahrbuch, a good venue for short papers on mildly controversial specimens, and it was out and published by November 2004. At least, that’s when my reprints arrived.

Incidentally, one version of the manuscript included a throw-away speculation that the specimen’s relatively large size relative to other alvarezsaurid tibiotarsi might be related to the evolution of this animal in an island setting. Gareth made me remove that bit, wisely I’m sure. So I was interested to see Meso et al. (2024) recently calculate that Heptasteornis would have a mass of 6.1 kg and a length of 1.8 m, which is smaller than the sizes estimated for ancestral members of the alvarezsaurid lineage but larger than a good number of the taxa close to Mononykus (Meso et al. 2024).

**Caption:** Greg Paul kindly provided a skeletal reconstruction of a mononykin alvarezsaurid for use in **Naish & Dyke (2004)** (Mononykini is a clade within Parvicursorinae, the alvarezsaurid clade that includes the majority of taxa and not the early-diverging ones). It’s mostly based on the Djadochta Formation taxon *Shuuvia deserti*, specimens of which were originally assumed to be additional *Mononykus* individuals. Image: © Greg Paul.

Post-2004, I have to say that the paper has mostly been mentioned and cited fairly and appropriately in relevant literature (Kessler et al. 2005, Csiki et al. 2010, Brusatte et al. 2013, Csiki-Sava et al. 2015, Mayr et al. 2020, Stoicescu et al. 2024, Meso et al. 2024). A common complaint among publishing scientists is that there exist all too many cases whereby authors have failed to credit the work of their colleagues, or are so insufficiently familiar with the literature that they’ve missed relevant stuff. Gareth and I did kick up a bit of a fuss when the paper wasn’t mentioned in Xu et al.’s (2011a) article on alvarezsaurid evolution and distribution (Dyke & Naish 2011), and Xu et al. (2011b) put up a spirited defence to this challenge.

Our proposed identification has also had the impact we hoped it would among Romanian specialists, who post-2004 started to look at theropod remains from their country with a potential alvarezsaurid identification in mind. Kessler et al. (2005) looked anew at a partial Haţeg Basin femur – the ‘Scoabă specimen’ – and found it more similar to the corresponding region of the alvarezsaurid skeleton than anything else, and suggested that this was an additional Romanian alvarezsaurid specimen.

**Caption:** at left, the distal part of a right femur known from the La Scoabă locality at Sânpetru, and described by Kessler *et al*. (2005). At right, the proximal end of a right femur from Nălaț-Vad (lower row) compared with Andrews’s original illustrations of the partial *Elopteryx* femur from Sânpetru (here flipped for comparison), reported by Stoicescu *et al*. (2024).

What about the other Romanian maniraptorans? Because the Heptasteornis distal tibiotarsus is only one of several Late Cretaceous theropod fragments from the same region of Romania – all of which tend to get discussed alongside one another – it didn’t seem right to ignore those others in our project. The original partial Elopteryx femur? It lacks alvarezsaurid characters and the presence of a trochanteric crest (a tall ridge on the bone’s proximal edge that overhangs the shaft on its anterior face) and posterior trochanter (a mound-like muscle attachment site on the bone’s outside edge) led us to prefer either a troodontid or early-diverging pygostylian identity (Naish & Dyke 2004).

What about the Bradycneme tibiotarsus? It doesn’t possess a distinctively curvy ascending astragular process like Heptasteornis, but is at least shaped like a maniraptoran tibiotarsus, so we identified it as Maniraptora indet. And a second partial tibiotarsus referred by Harrison & Walker (1975) to Heptasteornis (A1528) is also not sufficiently well preserved to allow an identification beyond Maniraptora to be resolved.

**Caption:** the maniraptoran specimens discussed in this article share a very unusual sculptured, textured or wrinkled bone surface. It’s definitely a real anatomical thing and not a product of preparation or, apparently, taphonomy, and I understand that it’s associated with an unusual histology. At left is the posterior face of the *Bradycneme* holotype, at right the left foot and tibiotarsus of *Balaur* (from Csiki *et al*. 2011). Images: © Natural History Museum, London; Csiki *et al*. (2010).

The elopterygid hypothesis. I said above that the alvarezsaurid hypothesis proposed by Naish & Dyke (2004) was risky, based as it is on scant data. On that note, if these remains aren’t those of alvarezsaurids, what might they be from? You see, there’s another, very different hypothesis out there. I’m calling it the ‘elopterygid hypothesis’.

Here’s where we come back to that intriguing detail of the Romanian maniraptoran fossils mentioned earlier in this article: the “sculpturing of the surface of the bone” that Charles Andrews mentioned in 1913. The weird thing is that this sculpturing is present in the Elopterx femora, the Heptasteornis and Bradycneme tibiotarsi, the Scoabă specimen and Balaur too.

This similarity is hard to explain away unless these animals are all close relatives, and part of a group that’s distinct from other maniraptorans. If this is so, suggestions that Heptasteornis might be an alvarezsaurid, that Balaur is a dromaeosaurid, or that some Elopteryx remains might be from troodontids would be in error. Instead, we’d be looking here at a new clade, perhaps deserving of the name Elopterygidae (a moniker first suggested, albeit as a dromaeosaurid clade, and as the ‘subfamily’ Elopteryginae, by Le Loeuff et al. (1992). UPDATE: that’s not correct, since Elopterygidae was used by Kálmán Lambrecht in 1933 for the supposed pelecaniform bird family that includes Elopteryx). Thus, ‘elopterygid hypothesis’.

**Caption:** a much simplified maniraptoran cladogram, showing some possible positions for an ‘elopterygid’ lineage. If these animals are within Paraves, they might be outside the Troodontidae + Avialae clade, or maybe they’re closer to birds (and thus perhaps *within* Avialae). Image: Darren Naish.

Or… would this hypothetical clade not be termed Elopterygidae, but Gargantuaviidae? A partial pelvis from Romania, reported by Mayr et al. (2020), looks consistent in size and anatomy with the Elopteryx femora, has similarities with the pelvis of Balaur, and has features suggestive of an affinity with Gargantuavis, an emu-sized maniraptoran from the Maastrichtian of France and Campanian of Spain, initially interpreted as a flightless bird belonging to Ornithuromorpha, the clade that includes crown-birds but excludes enantiornithines. A few authors have now mooted this (Mayr et al. 2020, Stoicescu et al. 2024).

If this proves correct, it might be that Balaur and Elopteryx are synonymous. Heptasteornis would presumably remain distinct, since it looks different from the more strongly fused distal tibiotarsus of Balaur (then again, is this difference a product of variation or ontogeny?). Again, it’s inconsistent with all of those specific hypotheses proposed for the identification of Balaur (a supposed dromaeosaurid), Heptasteornis (a supposed alvarezsaurid), and Gargantuavis (a supposed ornithuromorph).

**Caption:** the *Balaur bondoc* holotype EME (Transylvanian Museum Society, Dept. of Natural Sciences, Cluj-Napoca, Romania) PV.313, on display in 2011. It represents an animal that would have been about 2 m long when complete, though some ambiguity about this length exists as a bird identity for the animal would suggest a slightly shorter tail than a dromaeosaurid one. Image: Darren Naish.

A lament we often made on fieldwork while in Romania was that the additional fragmentary fossils forever being added to the canon were just making the situation ever more complex and confusing. Ultimately, we can’t resolve any of this without substantially better fossils. Will good Heptasteornis remains turn up and vindicate an alvarezsaurid identity? Or might we be shocked when it turns out to be an ‘elopterygid’ with Balaur*-like features? As ever, we live in hope, cursed by a cruel and fickle fossil record…

Shortly after publishing our paper, Gareth and I were both told that alvarezsaurid material had been discovered in the Upper Cretaceous rocks of France and were soon to see print. Twenty years later though and nothing further there has transpired.

Caption: I’m one of many researchers who’s visited the Balaur discovery site (Sebeş Glod, Alba County, Romania) in the hopes of finding more material of this intriguing taxon. It, and other fossils from the site, are found literally on the riverbed as well as in the banks. The bones are white, and hence stand out from the red, silty mudstone that forms the outcrop. Brandon Jardine is standing at right, a photo from March 2013. Images: Darren Naish.

The mistakes and the regret. Finally, published works are very often strewn with errors. I blame a number of reasons for the errors I’ve introduced into the publishing literature, but the primary one is that virtually every bit of science I’ve ever published – or tried to get published – has been done in a rush, in moments snatched between the actual jobs that burn up my time. It’s a terrible excuse, but a non-trivial one as far as I’m concerned.

Among the things I don’t like about Naish & Dyke (2004) is that we used an awful photo of A4359 (it’s Fig. 2A in the paper). We did actually arrange to get proper photos of the specimen taken, but a mix-up meant that we ended up with high-quality images of the Bradycneme holotype, not the Heptasteornis one. I still own them and they haven’t been published.

**Caption:** at left, the *Bradycneme draculae* holotype in anterior view, a photo provided by © The Natural History Museum, London, and unpublished to date. The specimen figures in **Naish & Dyke (2004)** should look like this. But they don’t: instead we have the image shown here at upper right, which shows the *Heptasteornis andrewsi* holotype. It’s not great. At lower right is my diagrammatic interpretration of the same specimen. Images: © The Natural History Museum, London; **Naish & Dyke (2004)**.

And a major embarrassment is that at the last minute – literally on the day of submitting the final version of the manuscript to the publisher – I opted to change the spelling of nopcsai to the incorrect ‘nopscai’, purely because I’m not capable of remembering the correct spelling of Nopcsa (how goddam Anglocentric of me, I know). That incorrect version is the one that made it into print.

And there we have my thoughts on Naish & Dyke (2004). I really didn’t intend to write so much, but… here we are.

 For previous Tetrapod Zoology articles connected to issues covered here, see…

Cyril Walker, May 2009
The Romanian Dinosaur Balaur Seems to Be a Flightless Bird, June 2015
The “Birds Are Not Dinosaurs” Movement, November 2017
My New Book: Dinopedia from Princeton University Press, December 2021
Alan Feduccia’s Romancing the Birds and Dinosaurs: Forays in Postmodern Paleontology, October 2023

Refs - -

Andrews, C. W. 1913. On some bird remains from the Upper Cretaceous of Transylvania. Geological Magazine 10, 193-196.

Baird, D. 1984. No ichthyosaurs in the Upper Cretaceous of New Jersey … or Saskatchewan. The Mosasaur 2, 129-133.

Brusatte, S. L., Vremir, M., Csiki-Sava, Z., Turner, A. H., Watanabe, A, Erickson, G. M. & Norell, M. A. 2013. The osteology of Balaur bondoc, an island-dwelling dromaeosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Romania. Bulletin of the American Museum of Natural History 374, 1-100.

Cau, A., Brougham, T. & Naish, D. 2015. The phylogenetic affinities of the bizarre Late Cretaceous Romanian theropod Balaur bondoc (Dinosauria, Maniraptora): dromaeosaurid or flightless bird? PeerJ 3: e1032.

Csiki, Z. & Grigorescu, D. 1998. Small theropods from the Late Cretaceous of the Hateg Basin (western Romania) – an unexpected diversity at the top of the food chain. Oryctos 1, 87-104.

Csiki, Z., Vremir, M., Brusatte, S. L., Norell, M. A. 2010. An aberrant island-dwelling theropod dinosaur from the Late Cretaceous of Romania. Proceedings of the National Academy of Sciences of the United States of America 107, 15357-15361.

Csiki-Sava, Z., Buffetaut, E., Ősi, A., Pereda-Suberbiola, X., Brusatte, S. L. 2015. Island life in the Cretaceous - faunal composition, biogeography, evolution, and extinction of land-living vertebrates on the Late Cretaceous European archipelago. ZooKeys 469, 1-161.

Dyke, G. J. & Naish, D. 2011. What about European alvarezsauroids? Proceedings of the National Academy of Sciences, USA 108, E14.

Feduccia, F. 2020. Romancing the Birds and Dinosaurs: Forays in Postmodern Paleontology. BrownWalker Press, Irvine, Boca Raton.

Halstead, L. B. 1975. The Evolution and Ecology of the Dinosaurs. Peter Lowe / Eurobook Ltd, Frome, Somerset.

Harrison, C. J. O. & Walker, C. A. 1975. The Bradycnemidae, a new family of owls from the Upper Cretaceous of Romania. Palaeontology 18, 563-570.

Hutchinson, J. R. & Chiappe, L. M. 1998. The first known alvarezsaurid (Theropoda: Aves) from North America. Journal of Vertebrate Paleontology 18, 447-450.

Kessler, E., Grigorescu, D. & Csiki, Z. 2005. Elopteryx revisited – a new bird-like specimen from the Maastrichtian of the Haţeg Basin (Romania). Acta Palaeontologica Romaniae 5, 249-258.

Le Loeuff, J., Buffetaut, E., Mechin, P. & Mechin-Salessy, A. 1992. The first record of dromaeosaurid dinosaurs (Saurischia, Theropoda) in the Maastrichtian of southern Europe: palaeobiogeographical implications. Bulletin de la Societe géologique de France 163, 337-343.

Martin, L. D. 1983. The origin and early radiation of birds. In Brush, A. & Clark, G. A. (eds) Perspectives in Ornithology, Essays Presented for the Centennial of the American Ornithologists’ Union. Cambridge University Press, Cambridge, pp. 291-338.

Martin, L. D. 1997. The difference between dinosaurs and birds as applied to Mononykus. In Wolberg, D. L., Stump, E. & Rosenberg, G. D. (eds) Dinofest International: Proceedings of a Symposium Sponsored by Arizona State University. Academy of Natural Sciences (Philadelphia), pp. 337-343.

Mayr, G., Codrea, V., Solomon, A., Bordeianu, M. & Smith, T. 2020. A well-preserved pelvis from the Maastrichtian of Romania suggests that the enigmatic Gargantuavis is neither an ornithurine bird nor an insular endemic. Cretaceous Research 106, 104271

McGowan, C. 1991. Dinosaurs, Spitfires, & Sea Dragons. Harvard University Press, Cambridge, Mass. & London.

Meso, J. G., Pol, D., Chiappe. L., Qin, Z., Díaz-Martínez, I., Gianechini, F., Apesteguía, S., Makovicky, P. J. & Pittman, M. 2024. Body size and evolutionary rate analyses reveal complex evolutionary history of Alvarezsauria. Cladistics doi doi.org/10.1111/cla.12600

Naish, D. 2021. Dinopedia: A Brief Compendium of Dinosaur Lore. Princeton University Press, Princeton NJ.

Naish, D. & Dyke, G. J. 2004. Heptasteornis was no ornithomimid, troodontid, dromaeosaurid or owl: the first alvarezsaurid (Dinosauria: Theropoda) from Europe. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 2004, 385-401.

Olson, S. L. 1985. The fossil record of birds. In Farner, D. S., King, J. R. & Parkes, K. C. (eds) Avian Biology, Volume III. Academic Press, New York, pp. 79-238.

Paul, G. S. 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.

Stoicescu, V., Codrea, V. A., Bordeianu, M. & Solomon, A. A. 2024. Elopteryx at Nălaț-Vad: new theropod material described from the Hațeg Basin (Romania). North-Western Journal of Zoology 20 73-80.

Suzuki, S., Chiappe, L. M., Dyke, G. J., Watabe, M., Barsbold, R. & Tsogtbaatar, K. 2002. A new specimen of Shuvuuia deserti Chiappe et al., 1998 from the Mongolian Late Cretaceous with a discussion of the relationships of alvarezsaurids to other theropod dinosaurs. Natural History Museum of Los Angeles County, Contributions in Science 494, 1-18.

Turner, A. H., Montanari, S. & Norell, M. A. 2021. A new dromaeosaurid from the Late Cretaceous Khulsan Locality of Mongolia. American Museum Novitates 3965, 1-48.

Xu, X., Sullivan, C., Pittman, M., Choiniere, J. N., Hone, D., Upchurch, P., Tan, Q., Xiao, D., Tan. L. & Han, F. 2011a. A monodactyl nonavian dinosaur and the complex evolution of the alvarezsauroid hand. Proceedings of the National Academy of Sciences, USA 108, 2338-2342.

Xu, X., Sullivan, C., Pittman, M., Choiniere, J. N., Hone, D., Upchurch, P., Tan, Q., Xiao, D., Tan. L. & Han, F. 2011b. Reply to Dyke and Naish: European alvarezsauroids do not change the picture. Proceedings of the National Academy of Sciences, USA 108, E148.

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Every now and again – speaking here as someone who’s published some number of articles, books and technical papers – I find it worthwhile to look back at the completed works of the past. For whatever reason, I’m doing that a fair amount right now, and today I want to talk about a peculiar short paper of mine that appeared in 2000…

**Caption:** juvenile *Deinonychus* in the trees! Image: Darren Naish.

Titled ‘Theropod dinosaurs in the trees: a historical review of arboreal habits amongst nonavian theropods’ and featured in the German journal Archaeopteryx, the paper concerned provides a brief overview of published comments on the subject before evaluating the concept of the tree-climbing (non-bird) theropod (Naish 2000a). I tried to make clear in the paper that while evidence for arboreality in non-bird theropods lacked support, I found it likely that scansoriality was plausible (Naish 2000a). Arboreality refers specifically to living in the arboreal environment and scansoriality to the ability to climb in trees; the two are not synonymous.

**Caption:** a physical copy of the paper – **Naish (2000a)** – that’s the focus of the article here. Unusually, I own both the specific issue of the journal *and* reprints of the paper itself. That’s a fluke and I can’t recall now how and where I got hold of the whole issue. Incidentally, the same issue also features John Videler’s paper on the idea that *Archaeopteryx* might have been capable of running over the surface of water, basilisk-style. Images: Darren Naish.

And – rightly or not – one of my foundational assumptions was that some degree of scansoriality should be expected or assumed for non-bird theropods given that avian origins likely involved use of trees. We’ll come back to that.

In fact, the article basically opens by noting that tree-climbing (non-bird) theropods are “currently popular in the non- and semi-technical palaeontological literature” (Naish 2000a, p. 35), my idea being that Greg Paul’s tree-climbing Ornitholestes from Predatory Dinosaurs of the World (Paul 1988) was a seminal image of the time. That’s arguable, given that many researchers interested in dinosaur palaeobiology neither pay attention to what people like Greg Paul were and are saying, nor remember or receive influence from artistic reconstructions. In my defence, I was seeing things more from the perspective of a social historian interested in palaeoart than that of a palaeobiologist: my understanding that Paul, Robert Bakker and the palaeoartists inspired by their writings and reconstructions had a paradigmatic impact on views of Mesozoic life (Naish 2021) is still partly at odds with that of many palaeontologists, especially here in the UK.

**Caption:** Greg Paul’s illustration of a climbing *Ornitholestes*, from *Predatory Dinosaurs of the World* (Paul 1988). The style of printing used in that book means that the illustrations do not photograph well: the original drawing is grey, not just black and white as it looks here. A 2024 update of this illustration, included in the third edition of Paul’s *The Princeton Field Guide to Dinosaurs*, has a modified head and lacks a nasal horn. Image: (c) Greg Paul.

Also on an introspective note, I very much regret the way in which I framed the view of 1980s/90s dinosaur palaeobiology discussed above: I said that we partly owed the concept of tree-climbing dinosaurs “to the highly artistic members of the ‘dancing dinosaur’ school” (Naish 2000a, p. 35). The term ‘dancing dinosaurs’ had been used by Bakker himself but the implication that there was a ‘dancing dinosaur school’ looks derogatory, and I shouldn’t have used it.

Anyway, the idea that certain smaller dinosaurs might have been capable of tree-climbing certainly goes back much further than the 1980s, and was mentioned here and there – mostly in connection with the Wealden ornithischian Hypsilophodon – during the 1910s, 20s and 30s. During archival research on the theropods of the English Wealden, I was interested to find that the Reverend William Fox – finder of several of the Isle of Wight Wealden Supergroup theropods I was working on – mentioned this idea in his brief articles on the enigmatic theropod Calamospondylus oweni, published in 1866 (Fox 1866a, b). This makes him the first person, so far as we know, to suggest tree-climbing in non-bird dinosaurs, though it should be said that he proposed this idea on quite spurious grounds, namely the perceived lightness and pneumaticity of the bones (Fox 1866a, b).

Fox’s idea was commented on favourably by William Swinton in 1936, one of the few researchers to keep dinosaur studies alive in the UK during the quiet, post-Victorian period (Martill et al. 2001). Swinton published several articles on Wealden dinosaurs during the 1930s and 40s and is also credited with writing the first ever dinosaur textbook, namely his 1934 The Dinosaurs: A Short History of a Great Group of Extinct Reptiles.

**Caption:** illustrations from **Naish (2000a)** that are relevant to the issue of scansoriality in non-bird theropods. Carnivorans like wolverines are not specialist climbers, but the flexibility of their limbs, sharpness of their claws and so on mean that they can climb if need be (this drawing is based on a photo). Theropods and carnivorans are fundamentally different, and what goes for one is not necessarily applicable to the other. Nevertheless, this is not a wholly invalid analogy. At right, a figure devoted to toe pad specialisations in trunk-climbing passerines, the idea being that we might test for trunk-climbing should we find preserved toe pads on non-bird dinosaurs (we now have, and the relevant specialisations are not present). Images: Darren Naish.

Hypsilophodon and Galton. Skip on a few decades to the 1970s, and tree-climbing in non-bird dinosaurs was mentioned again, albeit in a very different context. Peter Galton’s redescription of Hypsilophodon led to his re-evaluation of the scansorial hypothesis promoted by Othenio Abel, Swinton and those beforehand (Galton 1969, 1971a, b, 1974). Hypsilophodon did not, it turns out, have the opposable hallux, prehensile hands and feet and especially flexible forelimbs that earlier authors said it did, and its anatomy was instead that of a more cursorial animal committed to fully terrestrial life. Hypsilophodon is not, of course, a theropod, but it’s not irrelevant here because reconsideration of its alleged climbing abilities meant that Galton (1971a, b) did credit the idea that scansoriality remained a possibility for certain smaller non-bird dinosaurs, at least.

**Caption:** during the first half of the 20th century, *Hypsilophodon* was interpreted as scansorial or arboreal, and it’s *possible* that this led workers of the time to look especially favourably at the idea that other dinosaurs, like small theropods were tree-climbers too. For all that, only a few reconstructions of the time show these animals among the branches. Galton’s papers of the 1970s ultimately showed that *Hypsilophodon* lacked climbing adaptations and was fully terrestrial, as depicted in the images at left (from Galton 1974; the illustration is by Bakker). At right, we see (E) Galton’s reconstruction of the *Hypsilophodon* foot versus (F) Othenio Abel’s ‘grasping’ reconstruction of the foot from 1912. Images: Galton (1971, 1974).

Ostrom and beyond. Also on theropods, John Ostrom argued in his work on Archaeopteryx and bird origins that both archaic birds (like Archaeopteryx) and near-bird theropods (like Deinonychus) lacked structures that might suggest any adaptation for climbing (Ostrom 1974, 1979); these animals were “ground-dwelling cursorial bipeds with no obvious scansorial or arboreal adaptations” (Padian & Chiappe 1998, p. 19).

And that’s the ‘mainstream’ view that’s remained prevalent today. It’s notable that other suggestions from the late 20th century pertaining to scansoriality in non-bird theropods – there’s Anatoly Rozhdestvensky’s 1970 suggestion of climbing in ostrich dinosaurs and Sankar Chatterjee’s and Svend Palm’s 1990s proposals of trunk-clinging in Deinonychus-like maniraptorans (Naish 2000a, b) – have come from ‘outside’ voices not influential to the mainstream.

**Caption:** in his 1997 book *The Rise of Birds* (and in technical papers too), Sankar Chatterjee argued that “The elongated forelimbs and swivel wrist joint in proavian dromaeosaurs can best be interpreted as features evolved for the climbing of vertical tree trunks” (Chatterjee 1997, p. 166) (excuse the teleological framing); the diagrams at left were featured in the context of that claim. Also in 1997, Svend Palm’s*The Origin of Flapping Flight in Birds* argued that ‘tree-mounting’ behaviour evolved in dinosaurs, and it was in this setting that bird flight evolved (Palm 1997). Hypothetical pre-birds like those shown at right were featured in this work. Images: Chatterjee (1997), Palm (1997).

Several studies of claw curvature and digit proportions in non-bird theropods have been published since my 2000 article appeared, and as a generalisation they conclude that none of these animals were regular climbers (e.g., Birn-Jeffery et al. 2012)… with the possible exception of the very odd, very small scansoriopterygids (Dececchi et al. 2016). By quirk of fate, the first member of this group to become known to science was announced and discussed at conferences at about the time that a follow-up article to Naish (2000a) went to press (Naish 2000b).

**Caption:** the partial skeleton (part at left; counterpart at right) of the tiny scansoriopterygid *Scansoriopteryx heilmanni* Czerkas & Yuan, 2002, today generally considered synonymous with *Epidendrosaurus ninchengensis* Zhang *et al*., 2002. This animal is *tiny*, with a total length estimated at about 16 cm. It’s generally agreed that scansoriopterygids were climbers, and probably arboreal. Images: Czerkas & Yuan (2002).

Why Archaeopteryx anyway? My decision to publish this article in the pages of Archaeopteryx also deserves comment, since I know today that this journal is considered obscure, off the radar for most researchers, and hard to get as well.

Fact is, I just didn’t appreciate that at the time. One of my many failings as a young academic is that I didn’t have a clue on what I was doing, and I certainly didn’t follow any sensible course of action when it came to where or how I published things. And I wasn’t part of a research group where we were told or encouraged to aim for the top, or to ‘standard’ journals in the field… things were far more, shall we say, provincial. I chose to submit an article to Archaeopteryx because it seemed like a venue where articles on matters relevant to bird origins might be welcome. Indeed, a few articles devoted to the possibility of climbing in Mesozoic theropods had recently been published there (Chiappe 1997, Yalden 1997).

And that brings things to a close. Naish (2000a) is here online; it’s a weird little paper but not an especially bad one, and it’s been cited a few times where appropriate. It’s been missed a few times too, but that’s understandable in view of my comments above.

For previous Tetrapod Zoology articles on climbing dinosaurs, bird origins, hypotheses of bird ancestry and other connected issues, see…

A quick history of tree-climbing dinosaurs, July 2008
Epidexipteryx: bizarre little strap-feathered maniraptoran, October 2008
Did Velociraptor and Archaeopteryx climb trees? Claws and climbing in birds and other dinosaurs, December 2012
Yi qi Is Neat But Might Not Have Been the Black Screaming Dino-Dragon of Death, May 2015
The Second Edition of Naish and Barrett’s Dinosaurs: How They Lived and Evolved, November 2018
Heilmann, Thompson, Beebe, Tetrapteryx and the Proavian, January 2019
Alan Feduccia’s Romancing the Birds and Dinosaurs: Forays in Postmodern Paleontology, October 2023

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Birn-Jeffery, A. V., Miller, C. E., Naish, D., Rayfield, E. J., Hone, D. W. E. 2012. Pedal claw curvature in birds, lizards and Mesozoic dinosaurs – complicated categories and compensating for mass-specific and phylogenetic control. PLoS ONE 7 (12): e50555.

Chatterjee, S. 1996. The Rise of Birds. The Johns Hopkins University Press, Baltimore and London.

Chiappe, L. M. 1997. Climbing Archaeopteryx? A response to Yalden. Archaeopteryx 15, 109-110.

Czerkas, S. A. & Yuan, C. 2002. An arboreal maniraptoran from northeast China. In Czerkas, S. J. (ed) Feathered Dinosaurs and the Origin of Flight. The Dinosaur Museum (Blanding, Utah), pp. 63-95.

Dececchi, T. A., Larsson, H. C. E. & Habib, M. B. 2016. The wings before the bird: an evaluation of flapping-based locomotory hypotheses in bird antecedents. PeerJ 4: e2159.

Fox, W. 1866a. Another new Wealden reptile. Athenaeum 2014, 740.

Fox, W. 1866b. Another new Wealden reptile. Geological Magazine 3, 383.

Galton, P. M. 1969. The pelvic musculature of the dinosaur Hypsilophodon (Reptilia: Ornithischia). Postilla 131, 1-64

Galton, P. M. 1971a. Hypsilophodon, the cursorial non-arboreal dinosaur. Nature 231, 159-161.

Galton, P. M. 1971b. The mode of life of Hypsilophodon, the supposedly arboreal ornithopod dinosaur. Lethaia 4, 453-465.

Galton, P. M. 1974. The ornithischian dinosaur Hypsilophodon from the Wealden of the Isle of Wight. Bulletin of the British Museum (Natural History) 25, 1-152.

Martill, D. M., Naish, D. & Hutt, S. 2001. Introduction. In Martill, D. M. & Naish, D. (eds) Dinosaurs of the Isle of Wight. The Palaeontological Association (London), pp. 11-24.

Naish, D. 2000a. Theropod dinosaurs in the trees: a historical review of arboreal habits amongst nonavian theropods. Archaeopteryx 18, 35-41.

Naish, D. 2000b. 130 years of tree-climbing dinosaurs: Archaeopteryx, ‘arbrosaurs’ and the origin of avian flight. The Quarterly Journal of the Dinosaur Society 4 (1), 20-23.

Naish, D. 2021. Dinopedia: A Brief Compendium of Dinosaur Lore. Princeton University Press, Princeton NJ.

Ostrom, J. H. 1974. Archaeopteryx and the origin of flight. Quarterly Review of Biology 49, 27-47.

Ostrom, J. H. 1979. Bird flight: how did it begin? American Scientist 67, 46-56.

Padian, K. & Chiappe, L. M. 1998. The origin and early evolution of birds. Biological Reviews 73, 1-42.

Palm, S. 1997. The Origin of Flapping Flight in Birds. Svend Palm, Ballerop.

Yalden, D. W. 1997. Climbing Archaeopteryx. Archaeopteryx 15, 107-108.

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Late in the afternoon of Friday November 8th, my paper ‘The response to and rejection of Brian Ford’s Too Big to Walk, a 21st century effort to reinstate the aquatic dinosaur hypothesis’ saw digital publication in Historical Biology (Naish 2024)…

I wasn’t aware that it was about to see print (I’ve been too occupied), and hence: no press release, no lengthy announcement of any sort. This brief article is all you’re getting.

**Caption:** at left, cover of *Too Big To Walk* (2018 edition). At right, a montage of dinosaurs (from **Naish 2024**) to show how form and function reveals these to be predominantly terrestrial animals: in making this claim, I do not dispute that dinosaurs of all sorts might have waded, wallowed, or swam, nor that some, like certain (g) spinosaurids, were potential amphibious specialists. Images: Darren Naish.

Regular readers will know that I’ve written several times in the past about Brian Ford’s claims about dinosaurs, and I can appreciate how some might feel that I’ve devoted too much time to the topic already. I’m hoping that this will be the last time. My new paper approaches Ford’s aquatic dinosaur rhetoric from a different angle. Non-bird dinosaurs – as a whole* – were not aquatic, as Ford has argued (Ford 2012a, b, 2018, 2019), and it’s obvious that the ‘support’ he offered for the aquatic hypothesis is unsatisfactory, to put it mildly. I would be wasting your time and mine in going through the key points again (Naish 2012, 2024).

The caveat – as I have gone to trouble to point out – is that some species or groups may have been aquatic to some degree. Ford (2012a, 2018, 2019) has used the dishonest tactic of claiming that hypotheses pertaining to one or a few species (like Spinosaurus aegyptiacus or spinosaurids in general) can be applied to non-bird dinosaurs as a whole.

But what’s obvious in reading Too Big To Walk is how Ford built his case via techniques very much recalling the ‘post-truthism’ associated with certain political movements. And, ugh, what a bad time it is to be talking about that. Anyway, obvious from the book is the dismissal and belittling of experts, a strong and overly prevalent narrative claiming that palaeontologists are too blinkered to see the ‘truth’ of Ford’s wisdom, and abundant errors of fact and interpretation (Naish 2024). The book is also full of padding and (at 516 pages) overly long. For an author who’s devoted his career to the promotion and popularisation of science, this is pretty sorry stuff.

**Captions:** I own, and have extensively consulted, all three English-language editions of *Too Big To Walk*. The second edition (Ford 2019) is the small, paperback one with the bold typeface. It contains minor corrections and updates relative to the first (Ford 2018). *Too Big To Walk* is 516 pages long but doesn’t introduce the concept of aquatic dinosaurs until p. 282. Images: Darren Naish.

The reception of Too Big To Walk. The good news is that the failings of Too Big To Walk have been widely recognised, and that the book – published, in two editions(!), by famous and respected publisher William Collins – has, variously, been criticized, mocked and ignored, and scores poorly on aggregate sites (Naish 2024).

I have no doubt that I write from a provincial, Anglocentric bias, but what this means is that the substantial coverage that journalists gave to Ford and his views was thoroughly misleading in terms of the book’s wider reception. People mostly saw it as the baseless grandstanding it was. Here I’ll mention in passing that a trusted insider source has obtained sales data for the book on my behalf, and wow are they interesting. It didn’t do especially well, certainly not enough to justify a second edition.

**Caption:** a montage of reviews and amazon reviewer scores pertaining to *Too Big To Walk*. It’s not hyperbole to say that the book has not been well received, and has not received high scores overall. The Inquisitive Biologist review, screengrabbed at left, **can be read here**.

An interesting question is whether the poor reception of Too Big To Walk was driven by, or connected to, the pushback that came from palaeontologists, science writers and bloggers. Did our complaining help alert people to the pointlessness and time-wasting inherent to the existence of Too Big To Walk, or did they discover this themselves? As ever, my argument is that pushing back against bad science, anti-science, pseudoscience and crankery is important: not everyone is privileged enough to find out for themselves that a given view or publication belongs to those categories. Indeed, pushing back is more important than ever, given the way things are going, and I have no time for the view that scientists should keep their heads down and ignore crank and anti-scientific views when they see them expressed.

These topics and others are explored in the new paper (a few typos were introduced at a late stage in the publication process; I’m seeing if they can be corrected), and thanks to everyone who’s expressed interest or asked for a copy. The review history of this paper is interesting: it won the most positive reviews I’ve ever had for any submitted work.

As for Brian J. Ford…. My effort throughout writing this paper was to fairly recognise Brian Ford’s very active role in the popularisation of science across his career, and to give fair credit to his contributions. But why he thought it was a good idea to declare all experts wrong on the matter of what dinosaurs were like when alive, and to simply press forward with Too Big To Walk is a question that only he is able to answer.

**Caption:** I’ve published two whole articles on the Brian Ford ‘aquatic dinosaurs’ thing now. I think we can agree that that’s enough. **Naish (2012) is here**.

For previous articles on Ford’s aquatic dinosaurs and related issues, see…

Brian J. Ford's Aquatic Dinosaurs, 2014 Edition, February 2015 (I cannot find an intact version of this article)
Tetrapod Zoology Is A Teenager Now, January 2019 (reviews Naish v Ford Conway Hall event)

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Ford, B. J. 2012a. A prehistoric revolution. Laboratory News April 2012, 24–26.

Ford, B. J. 2012b. Aquatic dinosaurs under the lens. The Microscope 60, 123–131.

Ford, B. J. 2018. Too Big To Walk: The New Science of Dinosaurs. London: William Collins.

Ford, B. J. 2019. Too Big To Walk: The New Science of Dinosaurs. London: William Collins.

Naish, D. 2012. Palaeontology bites back… Laboratory News May 2012, 31–32.

Naish, D. 2024. The response to and rejection of Brian Ford’s Too Big to Walk, a 21st century effort to reinstate the aquatic dinosaur hypothesis. Historical Biology 10.1080/08912963.2024.2421268

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Regular readers of this blog might be aware of my book Hunting Monsters: Cryptozoology and the Reality Behind the Myths, initially published as an ebook in 2016 (Naish 2016) and appearing in hardcopy in 2017 (Naish 2017)…

**Caption:** at left, cover of the **2016 ebook version** of *Hunting Monsters*, and – at right – the **2017 hardcopy version**. I really like both covers. Because the two works differ in minor ways and have different publication dates, I’m treating them here as separate publications.

Hunting Monsters discusses hypotheses and cases relevant to six main subject areas: (1) the history, function, role and future of cryptozoology; (2) sea monsters; (3) lake monsters; (4) bigfoot and such; (5) mokele-mbembe and such; and (6) Australian cryptids. It’s mostly an exploration of a cultural, ‘post-cryptid’ view of cryptozoology. This posits that sightings of and encounters with monsters – while often or sometimes representing genuine events of some sort – mostly stem from the creation and maintenance of cryptids as objects within the ‘cultural landscape’ inhabited by the experiencer. As I’ve always tried to make clear, I’m not in the least bit original in stating any of this, and the sociocultural theory I promote has its roots in the writings of several who went before me (e.g., Binns 1983, 2017, Meurger & Gagnon 1988, Loxton & Prothero 2013).

**Caption:** noted books on mystery animal/monster research that are relevant to, or promote, the ‘post-cryptid’, sociocultural view of cryptozoology also endorsed in *Hunting Monsters*. I read or consulted most of the books shown here prior to writing *Hunting Monsters*, but not Samantha Hurn’s *Anthropology and Cryptozoology: Exploring Encounters with Mysterious Creatures* (published 2017). I’m currently re-reading **Meurger & Gagnon (1988)** because I need better notes on it.

Eight years after publication, I can say that the response to Hunting Monsters has been mostly predictable: people with a sceptical or scientific approach to cryptozoology generally like it and have said positive things about it, and people who want cryptids to be accepted as real, and dislike the critical approach employed by sceptics, mostly dislike it and have said how it represents a close-minded, blinkered, elitist mindset of the sort typical among nay-sayers. The former set of people are better at recording their thoughts than the latter, and it’s partly for that reason that Hunting Monsters has had a net positive reception, currently scoring 4.2 out of 5 at amazon. It’s hardly perfect, of course, and I accept that a good number of valid criticisms have been directed at it. Those are topics I’ll address at another time. Incidentally, the hardcopy version of the book (Naish 2017) is now out of print, and indeed hard to get and generally only sold at high price. The ebook version (Naish 2016) continues to sell well though.

**Caption:** several Nessie models exist around the edges of Loch Ness. This green plesiosaur – presumably based specifically on an elasmosaurid (if you want to know about plesiosaur diversity and evolutionary history, **see my 2023 book *Ancient Sea Reptiles***) – is on show at the Clansman Hotel in Bracla, on the loch’s north shore, north of Drumnadrochit. The sign near the model says DO NOT RIDE.

Among those who’ve criticised the book is Loch Ness Monster advocate and independent researcher Roland Watson, author of several Nessie-themed books and the blog Loch Ness Monster. I’ve had reason to address Watson’s comments on previous occasions and am not a fan of his approach, most of which revolves around the idea that (1) Nessie’s existence should be considered more plausible than the opposite and (2) that Nessie sceptics can only be framed as biased, sloppy-thinking, pseudo-sceptical rejectionists.

Back in 2016, Watson published a review of Hunting Monsters at his blog, and at amazon. Much of what he said was contestable or just factually wrong, and as a consequence I felt compelled to write a long response, which I also shared on amazon on August 21st 2016. However, I’ve recently discovered that it’s no longer there. Why, I have no idea, but there’s no trace of it. That bothers me because it was my fair right of reply, and a worthwhile contribution that has been cited in the literature (Binns 2017, p. 203). So, what I’ve done is dig it out and republish it here. While some of the text below does require a bit of background knowledge, I think that it’s mostly easy to follow, even if you haven’t read Watson’s original review. Nevertheless, the picture captions should help add context when that’s otherwise absent. Ok, here we go…

**Caption:** here’s proof that my initial response to Watson’s review was published at amazon during August 2016. It’s a bit concerning that amazon then later removed it. Does the site routinely do that to comments written as responses to reviews? If so… yikes.

In a comment written in response to his review of my book Hunting Monsters, Roland Watson “[wishes] that these comments [pertaining to the Loch Ness Monster] … would actually address the issues I raise than rather than cast general aspersions which target nothing expect [sic] the person”. Fair enough, challenge accepted. Let’s look in detail at Watson’s various claims…

Claim number 1: Watson dislikes the contention that “the diversity of creatures described points more to human imagination than actual animals awaiting discovery” and argues that “Even though [witnesses] may have seen something large and alive, the finer the detail described, the greater the room for error”.

**Caption:** Nessie proponents like Watson want us to accept that Nessie sightings represent the same one animal species, whereas what we actually have is a disparate mess of basically any and everything you could imagine… a fact more consistent with the idea that people are interpreting observations of all kinds of things as ‘the monster’. The variety of reported monsters is reflected in efforts to imagine Nessie as real. Is it a friendly, small-headed plesiosaur, a shaggy, humped quadruped, a rough-skinned horror with an undulating appendage, or a twin-humped, snorkel-headed animal with great diamond-shaped flippers? In reality, it is all of these things, and also none of them. Images: Darren Naish.

Response: in promoting (via his blog and books) the view that the large number of Loch Ness Monster accounts might actually describe a genuine undiscovered large animal species, Watson is routinely guilty of both confirmation bias and a seeming inability to understand the principle of parsimony. In Hunting Monsters I stated repeatedly how the diversity of Loch Ness Monster accounts makes it likely that diverse animals and phenomena have been witnessed, reported, interpreted and misinterpreted within the context of both a folkloric belief and sociocultural expectation that monsters might be seen. This is a clearly stated hypothesis noted several times in the book. How does it stand up to the alternative: that people are reporting an unknown species but failing to describe it consistently? When we combine this with the impressive lack of compelling evidence of any sort, the literalist interpretation has to be considered a failure. To argue otherwise does have a wilful desperation and a tinge of romanticism, so good luck with it.

Claim number 2: Watson says that the proposal that the Spicer sighting was influenced by King Kong “is not a convincing theory”.

Response: well, it’s not a “theory” for starters. Nevertheless, it remains an interesting possibility worth noting (the exact wording used in Hunting Monsters is that “a case can also be made that the account was inspired by the release of King Kong” (Naish 2017, p. 86). In other words, this is put out there as an interesting possibility, and it remains so; not as a point that can be dismissed simply because it fails to accord with personal bias.

**Caption:** it’s now a well-known idea that the amphibious, aggressive brontosaur of 1933’s King Kong might have been inspirational to the Spicers, and somehow affected their recalling and retelling of their Loch Ness encounter. **Loxton & Prothero (2013)** are among several authors who’ve discussed this proposal, and it would be wrong – in a review that discusses the evolution of ideas on the Loch Ness Monster – to not mention it. I was kinder in *Hunting Monsters* than I am to this concept today though, since there’s an argument (made by monster researcher Charles Paxton) that we don’t know for sure that the Spicers saw the movie prior to their sighting.

Claim number 3: Watson says it’s a mistake to say that the Fordyce sighting is relevant to accounts of the early 1930s (to quote Watson: “that story was not made public until 1990 and had nothing to do with the mood "at the time"”).

Response: Fordyce’s sighting apparently occurred in 1932, and it is thus fully justifiable to say that it is relevant to things “afoot at the loch at the time”. Having said that, I have modified the choice of wording for the hardcopy version of the book, a disclaiming clause (“though note that this report did not come to light until 1990”; Naish 2017, p. 89) now added.

**Caption:** in July 1933, Mr and Mrs Spicer reportedly witnessed a large, unusual animal cross the road (General Wade’s Military Road, or the B852) adjacent to Loch Ness’s southern shore, between Dores and Inverfarigaig. It was said at times to have what looked like the head of a “lamb or small deer” neat its middle, a component that later morphed into a tail tip (in which case the body and tail were bent around to the side). Author Rupert Gould interviewed the Spicers, and eventually concluded that they’d seen a group of deer bounding quickly across the road. The interpretation of the event shown here comes from the third, 1976 edition of Tim Dinsdale’s *Loch Ness Monster*.

Claim number 4: Watson regards it as “an example of exaggerated narrative” to say of the Spicer sighting “Over the years, the description became increasingly sensational. It started out as 2–2.5 m in length but gradually increased to 9m”. Watson goes on to say that it “is completely wrong” to refer to this as a “monster [story that grew] with the telling”, and quotes Gould’s quoting of Spicer’s justification of this increase in length.

Response: Hunting Monsters states that “Over the years, the description became increasingly sensational. It started out as 2-2.5 m in length but gradually increased to 9 m” (Naish 2017, p. 85). The fact that Mr Spicer explained to Gould – within the space of 10 months of the initial sighting – how an initial leap in size occurred as a consequence of his direct measuring of the road is said by Watson to be an “omitted detail”. He then objects to my implication that the monster “gradually” increased in size with retelling. I concede that my idea of the monster “gradually” increasing in size was inspired by Ronald Binns (“Their estimate of the monster’s size also changed dramatically. From being between six and eight feet in length it grew and grew as Spicer retold his story until it was twenty-five, even thirty feet long”; Binns 1983, pp. 90-91), and also by the fact that there are sources where Mr Spicer stated the monster to be 30 ft long, and even 40 ft and 50 ft long. In short, it remains true that the Spicer’s monster did increase in length with the retelling of the story, and did so in step with an increasingly sensational interpretation of the account. Couple this with Gould’s conclusion that “I think the Spicers saw a huddle of deer crossing the road”, and it again seems clear that a pro-monster interpretation of the sighting appears to be the problematic one.

**Caption:** an artistically brilliant but very much hyper-sensationalized view of the Spicer sighting, now depicting their undulating, amorphous creature with a possible lamb-like head as a giant predatory plesiosaur. This is by the brilliant Gino D’Achille.

Claim number 5: Watson claims – contra Hunting Monsters – that the Hugh Gray and Peter O’Connor photos are not images of, respectively, a swan and an inverted kayak, and that he has ‘dismantled’ these proposals.

Response: I leave others to judge whether the proposals endorsed in my book (that the Hugh Gray photo depicts a swan and the O’Connor photo depicts an inverted Tyne Prefect kayak) are reasonable and likely and serve as superior alternative explanations to Watson’s use of these images as depictions of an unknown giant animal or animals. Further details on why the swan and kayak interpretations should be supported are included within the book (Naish 2016, 2017). The kayak hypothesis is discussed at length in Dick Raynor’s article ‘A Study of the Peter O'Connor photograph of the Loch Ness Monster’, available here.

**Caption:** the Peter O’Connor Nessie photo of 1960 (one version of which is shown at upper left) is of very dubious provenance. I agree with Dick Raynor that it likely shows a specific kayak (Dick’s photo of that boat is at lower left). At right are illustrations from my discussion of the photo and how its ‘anatomy’ matches that of the kayak. Images: © Peter O’Connor; **Dick Raynor**; Darren Naish.

Claim number 6: Watson opines, contra a suggestion in Hunting Monsters, that the Fordyce sighting does not resemble a pony (not a “donkey”, as per Watson) carrying a deer, and – says Watson – it would be “Better to say nothing and take a neutral position”.

Response: Watson might prefer it if the door were left as far open as possible as goes the proposed existence of large unknown animals at Loch Ness. Again he is, alas, all too keen to demonstrate his inability to adopt the principle of parsimony. The fact is, a pony carrying an antlered stag really does look suspiciously like the creature Fordyce reported. It might be difficult to find a photo that exactly matches what Fordyce described, but it is easy – based on the photos findable online – to imagine one, unless one is too biased, of course.

**Caption:** the Fordyce ‘camel’ sighting is one of the weirdest Loch Ness Monster sightings. So: what to do? Should we just ignore it as potentially fictional (in which case… why not do the same with most or even all other accounts? That hardly seems right), or should we aim to rationalise it? At left is the drawing of the creature as reimagined for Mike Dash’s *Fortean Times* article of 1991 (Dash 1991); at right is an 1873 painting by Richard Ansdell showing the technique whereby a pony is used to carry a deceased stag. Watson used a dirty trick in deliberately saying “donkey” (instead of pony) throughout his discussion of this idea. Images: Dash (1991); The Cheltenham Trust and Cheltenham Borough Council (**original here**).

Claim number 7: Watson argues that my “handling of the folklore of the Loch Ness Water Horse is unsatisfactory”, since I dismiss all pre-1933 accounts. Specifically, Watson notes that my dismissal of “Richard Franck's 17th century "floating island" at Loch Ness, as a man-made raft runs completely counter to what even Franck theorised about this strange object from 1658”.

Response: I have read Watson’s book on his interpretation of pre-1933 accounts and found nothing more than credulity and confirmation bias, as mentioned above. As noted in Hunting Monsters, my approach to the pre-1933 accounts is heavily inspired by Ulrich Magin’s 2001 article ‘Waves without wind and a floating island – historical accounts of the Loch Ness Monster’ (Fortean Studies 7, 95-115). I would encourage those curious to read Magin’s article and see whether it makes a more convincing set of arguments than Watson’s pro-monster claims and book.

**Caption:** views of Urquhart Castle as seen from the water. A few Nessie encounters have been reported from around the castle, some involving the creature supposedly crawling onto land. As you can see from these shots (taken in 2016), sufficient vegetation has grown up around the banks and the ruins to obscure them relative to how they looked in Nessie’s heyday (the 1930s, or the 60s and 70s, take your pick). Images: Darren Naish.

Claim number 8: Watson ends his review with a series of bizarre and childish suggestions that I might be being “groomed” (his term) as “the successor” to noted and respected sceptical investigators Dick Raynor and Adrian Shine. To quote: “Perhaps Darren is seen as "The One", but in my view, once Dick and Adrian get out their slippers and pipe, Loch Ness scepticism will go down the plug hole”.

Response: Firstly, it is an honour to be compared to these upstanding individuals who have done so much to carefully, thoroughly and diligently bring critical thinking and scientific analysis to the Loch Ness enigma. Secondly, there are weird and sinister overtones as goes the reference to ‘grooming’. Nevertheless, the opinion put forward here is nonsense: it implies that Raynor and/or Shine sought me out and wilfully modified my thoughts or intellectual approach for personal means. As is well documented in a paper trail that stretches back about 20 years (see my articles from the 1990s, published in cryptozoological periodical Animals & Men, my 2000 article ‘Where be monsters?’ in Fortean Times 132, my 2001 article ‘Sea serpents, seals and coelacanths: an attempt at a holistic approach to the identity of large aquatic cryptids’ in Fortean Studies 7, and other publications), I have followed a distinct individual pathway of cryptozoological investigation that has been evidence-led and sceptical for quite some time.

**Caption:** nobody cares enough to find out (why would they?), but an interested person could chart the changing of my cryptozoology views over time. I started out as substantially more credulous than I was in 2016/2017, or am today. The images here show publications of mine dating to 1995, 1996, 2000 and 2001.

For years now I have corresponded with Dick and Adrian and have found my views to match theirs. It is insulting and ridiculous to imply that there has been any effort on their part to influence or lead my own views and conclusions, and I put it that this proposal does the accuser no favours; in fact it indicates that he inhabits a worrying and idiosyncratic intellectual landscape. Thirdly and finally, there is no effort involving myself or anyone else to have me installed as “The One” when it comes to Loch Ness scepticism; as is obvious from everything we might glean about Loch Ness, the cryptozoological literature, and collective human knowledge in general, scepticism – if not rejection – of the Loch Ness Monster’s purported existence is the mainstream, widely accepted view for obvious reasons (these being the impressive lack of evidence and the unsatisfactory, clutching-at-straws nature of the arguments coming from LNM proponents). It is thus highly amusing to speak of “Loch Ness scepticism going down the plug hole”. Next you’ll be saying that the theory of evolution and the heliocentric model of the solar system are in danger too.

Again, I thank the reviewer for reading my book and for taking time to encourage others to read it too.

For previous TetZoo articles on the Loch Ness Monster, lake monsters, and cryptozoology more generally see…

The Loch Ness monster seen on land, October 2009
Dear Telegraph: no, I did not say that about the Loch Ness monster, July 2011
Photos of the Loch Ness Monster, revisited, July 2013
My New Book Hunting Monsters: Cryptozoology and the Reality Behind the Myths, February 2016
Books on the Loch Ness Monster 1: Ronald Binns’s The Loch Ness Mystery Reloaded, March 2019
Books on the Loch Ness Monster 2: Gareth Williams’s A Monstrous Commotion, March 2019
Usborne’s All About Monsters, April 2019
Sea Monster Sightings and the ‘Plesiosaur Effect’, April 2019
A Review of Robert L. France’s Disentangled: Ethnozoology and Environmental Explanation of the Gloucester Sea Serpent, November 2019
The 1972 Loch Ness Monster Flipper Photos, August 2020

If you enjoyed this article and want to assist me in what I do — and see behind-the-scenes stuff in development — please support me at patreon. Click anywhere on this text in bold.

Refs - -

Binns, R. 1983. The Loch Ness Mystery Solved. Open Books, London.

Binns, R. 2017. The Loch Ness Mystery Reloaded. Zoilus Press.

Dash, M. 1991. The camels are coming. Fortean Times 58, 52-53.

Loxton, D. & Prothero, D. R. 2013. Abominable Science! Columbia University Press, New York.

Meurger, M. & Gagnon, C. 1988. Lake Monster Traditions: A Cross-Cultural Analysis. Fortean Times. London.

Naish, D. 2016. Hunting Monsters: Cryptozoology and the Reality Behind the Myths. Arcturus, London.

Naish, D. 2017. Hunting Monsters: Cryptozoology and the Reality Behind the Myths. Arcturus, London.

Abominable Science!: Origins of the Yeti, Nessie, and Other Famous Cryptids

Lake monster traditions: A cross-cultural analysis

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I’m a big fan of the fact that views on a given topic develop piecemeal…

**Caption:** *Prehistoric Planet* promotes **Azhdarchid Supremacy**. © Apple TV.

… that short bursts of dialogue and argument – occurring over decades – are crucial in the construction of a case, and that only after years and years of slow, incremental addition can we look back and realise how far we’ve come. Big, ‘game changer’ discoveries are great, of course, but we need to appreciate, too, that paradigm shifts take time and the accruing of relatively small bits of data.

Over the last few decades, our knowledge of one of my favourite groups of animals – the often gigantic, long-skulled azhdarchid pterosaurs – has improved substantially, and I say this as someone lucky enough to have been involved in research on this group. Writing (as ever) from a biased personal perspective, I thought it would be interesting to chart ‘azhdarchid progress’ as I’ve seen it from my own research interests. The words here are mostly recycled from the summary texts published at my publications site, but I decided it would be useful to have them together. I owe thanks to the other researchers I’ve collaborated with, in particular Mark Witton and the late Mátyás Vremir.

**Caption:** this life reconstruction of a foraging *Quetzalcoatlus* group accompanied **Witton & Naish (2008)**. It’s a very dated image today and Mark probably won’t enjoy the fact that I’m sharing it (sorry Mark). But it’s significant in the story told here, since it was widely shared in news articles reporting our 2008 conclusions. It probably is, in fact, one of the most widely shared, most often reproduced, azhdarchid-themed images. Image: Mark Witton, from **Witton & Naish (2008)**.

The ‘terrestrial stalking’ paradigm. Azhdarchids have to be regarded as the most distinctive and unusual of pterosaur groups, and views on how they might have lived have varied considerably. In our opening salvo published in 2008, Mark Witton and I examined the skeletal proportions, cranial anatomy and sedimentological setting of azhdarchids and concluded that they were strongly adapted for quadrupedal walking in terrestrial settings (Witton & Naish 2008). They were likely striding predators of small and mid-sized prey, analogous to modern ground hornbills. We termed this the ‘terrestrial stalking’ model. Our proposal has been supported by additional studies (Witton & Naish 2015, Naish & Witton 2017). Some minor pushback came from other pterosaur specialists, on which read on…

**Caption:** this diagram, from **Witton & Naish (2008)**, shows how joint flexion in the fore- and hindlimbs of an idealized azhdarchid (this is the Chinese form *Zhejiangopterus*) allows these otherwise very leggy animals to reach the ground with their jaw tips. Image: Mark Witton, from **Witton & Naish (2008)**.

Sympatry and niche partitioning was probably normal in azhdarchids. In a project resulting from collaboration on the vertebrate palaeontology of the Romanian Late Cretaceous, I and colleagues described a new azhdarchid – Eurazhdarcho langendorfensis Vremir et al., 2013 – represented by cervical vertebrae and part of the wing skeleton. The specimen is represented by adult remains and is not a juvenile specimen of the much larger Romanian azhdarchid Hatzegopteryx (Vremir et al. 2013). The presence of both of these azhdarchids in the same geological unit (the Sebeş Formation of the Transylvanian Basin) is significant, since it indicates that niche partitioning was present in sympatric azhdarchids. We noted the presence of several locations worldwide where sympatric azhdarchids occur, these taxa differing in size and hence presumably in ecology and habits. Eurazhdarcho is one of many new taxa discovered by Mátyás Vremir (who died in 2020).

**Caption:** some geological units reveal evidence of two or even three sympatric azhdarchid species. Since this diagram was published, one of the Dinosaur Park Formation azhdarchids has been named *Cryodrakon boreas*, and the Javelina Formation has two additional azhdarchids (*Quetzalcoatlus lawsoni* and *Wellnhopterus brevirostris*). Diagram produced by Mark Witton and map used with kind permission of Ron Blakey, Colorado Plateau Geosystems, Inc; from **Vremir *et al*. (2013)**.

Some azharchids had ‘short’ necks. The idea that azhdarchid pterosaurs were anatomically homogenous – an assumption more or less inherent to Witton & Naish (2008) – is challenged by several poorly known taxa. These indicate that neck length and proportions, and body size overall, was variable in the group (an idea first mooted by Dave Unwin in his 2006 book The Pterosaurs From Deep Time). In this paper, we described LPB (FGGUB) R.2395, a Haţeg Basin cervical vertebra from a mid-sized azhdarchid, estimated to have a wingspan of 3-4 m (Vremir et al. 2015). Despite its small size (89 mm long), its smooth (non-pitted), polished external bone texture shows that it belonged to an adult. It’s probably a cervical IV.

**Caption:** the Romanian azhdarchid neck vertebra R.2395 is one of several specimens which helped reveal that this group includes relatively short-necked forms in addition to the more familiar, giraffe-necked kinds like *Quetzalcoatlus*. R.2395 represents an additional taxon that lived alongside the very different *Eurazhdarcho* and *Hatzegopteryx*, but it hasn’t yet been named. Another azhdarchid (*Albadraco*) has been named from the Maastrichtian of Romania (albeit from a different faunal assemblage) and doesn’t appear to represent the same taxon as R.2395. The life reconstruction shown here is, of course, speculative. Images: Mark Witton; **Vremir *et al*. (2015)**.

The most interesting thing about R.2395 is that it is relatively broad for its length, this indicating that the animal had a neck length c 30-40% shorter than that of other azhdarchids of similar size. It indicates the presence of an additional azhdarchid taxon in the Haţeg Basin fauna (Vremir et al. 2015). We also argued in this paper that efforts to sink all Haţeg Basin azhdarchids into a single taxon do not withstand scrutiny. Due to an editorial mishap, the final published version of this paper is not as richly illustrated as planned in earlier drafts.

More thorough support for ‘terrestrial stalking’. The view that azhdarchid pterosaurs were ‘terrestrial stalkers’, proposed by Witton & Naish (2008), was challenged in 2013 by Alexander Averianov. Averianov argued that the depositional settings of azhdarchid fossils were inconsistent with our proposal, that big theropods made our proposal problematic, and that azhdarchids were more likely ‘scoop-netters’. In our response, Mark Witton and I re-examined the environmental context in which azhdarchids have been discovered and showed that the evidence was consistent with them being animals of continental settings (Witton & Naish 2015). We also looked at the behaviour and ecology of those theropods contemporaneous with azhdarchids to see if they would really present the problem that Averianov argued they would, and examined his ‘scoop-netting’ idea in order to test its viability (Witton & Naish 2015).

**Caption:** the terrestrial stalking hypothesis is based on multiple lines of evidence. It does not (like so many other hypotheses about pterosaur behaviour and ecology) rely on cherry-picking one or two anatomical or palaeoenvironmental features. This image – from **Witton & Naish (2015)** – shows how several independent pieces of data all provide support for our hypothesis.

Some azhdarchids were giant, thick-necked predators. The view assumed in the writing of Witton & Naish (2008) and Witton & Naish (2015) – that azhdarchids were essentially alike in proportions and hence similar in lifestyle – was again shown to be incorrect via our 2017 analysis of cervical vertebrae belonging to the giant Late Cretaceous Romanian azhdarchid Hatzegopteryx. We showed that the neck of this animal was broad and thick relative to that of other giant azhdarchids (like Arambourgiania) and mechanically able to resist substantial loads (Naish & Witton 2017).

**Caption:** skeletal reconstructions of the giant azhdarchids (A) *Hatzegopteryx* and (C) *Arambourgiania* to scale, showing the markedly different body shapes of these animals. (B) shows how broad the back of the skull and neck was in *Hatzegopteryx*. Contrast this with D-E: the small *Quezalcoatlus* species, with its 4.6 m wingspan and long, slender neck. Scale bar = 1 m. Credit: **Naish & Witton 2017**.

This view is consistent with the absence of large predators (like theropods) from the same region and suggests that Hatzegopteryx was a predator of animals that could have weighed tens of kilos. This study is consistent with our previous proposal (Vremir et al. 2015) that some azhdarchids were relatively short-necked.

Finally…. what else? The research discussed here focuses on North American, Afro-Arabian and eastern European azhdarchids, but this is a group with a near-global distribution. In addition, the iconic North American taxon Quetzalcoatlus was (at least in terms of technically published data) poorly known throughout the entire time that the studies discussed above were being compiled and published, this hindering our understanding of azhdarchid anatomy and phylogeny. In late 2021, the long-awaited monograph on Quetzalcoatlus was finally published, meaning that this was a good time to round up recent discoveries from the azhdarchid world and look at then-current views on their diversity and phylogeny.

**Caption:** montage depicting some azhdarchid finds and publications of the 2010s and 20s. At left, the new reconstruction of *Quetzalcoatlus* published by Padian *et al.* (2021). There’s some controversy over the accuracy of this image, since an argument has been made that the hindlimbs have been made too long. At upper right, a montage from Novas *et al*. (2012) showing the mid-sized Argentinian *Aerotitan* (A and B, and reconstructed in E) compared to the Moroccan *Alanqa* (C and D; *Alanqa* is now regarded by some authors as being outside of Azhdarchidae, and within the separate group Alanqidae). At lower right, a speculative reconstruction of *Wellnhopterus*, depicting this animal as a thalassodromid, from Campos (2021). Scale bar = 10 cm.

Many basic questions about the biology, ecology, distribution and anatomy of these animals remain, and there’s tons of science left to do. I’m one of many people aiming to publish on them again, and with new data, new ideas, and new hypotheses. Those things will emerge in time. But the point of this article is that we have – I contend – turned a metaphorical corner in terms of what we think azhdarchids were like, and that, I put it, is significant.

For previous TetZoo articles on azhdarchids and other pterosaurs, see…

Tiny pterosaurs and pac-man frogs from hell, March 2008
Daisy’s Isle of Wight Dragon and why China has what Europe does not, March 2013
In Rio for the 2013 International Symposium on Pterosaurs, May 2013
Quetzalcoatlus: the evil, pin-headed, toothy nightmare monster that wants to eat your soul, August 2013
Were azhdarchid pterosaurs really terrestrial stalkers? The evidence says yes, yes they (probably) were, December 2013
Some Azhdarchid Pterosaurs Were Robust-Necked Top-Tier Predators, February 2017
Postcranial Palaeoneurology and the Lifestyles of Pterosaurs, August 2018
Did Dinosaurs and Pterosaurs 'Glow'? Extinct Archosaurs and the Capacity for Photoluminescent Visual Displays, March 2020
New Perspectives on Pterosaur Palaeobiology, the TetZoo Review, September 2020
Baby Pterosaurs Were Excellent Fliers and Occupied Different Niches From Their Parents, July 2021
The Quetz Monograph Lives and Other News on Azhdarchid Pterosaurs, December 2021

If you enjoyed this article and want to assist me in what I do — and see behind-the-scenes stuff in development — please support me at patreon. Click anywhere on this text in bold.

Refs - -

Campos, H. B. N. 2021. A new azhdarchoid pterosaur from the Late Cretaceous Javelina Formation of Texas. Biologia doi: 10.1007/s11756-021-00841-7.

Naish, D. & Witton, M. P. 2017. Neck biomechanics indicate that giant Transylvanian azhdarchid pterosaurs were short-necked arch predators. PeerJ 5:e2908.

Novas, F. E., Kundrát, M., Agnolín, F. L., Ezcurra, M. D., Ahlberg, P. E., Iasi, M. P., Arriagada, A. & Chafrat, P. 2012. A new large pterosaur from the Late Cretaceous of Patagonia. Journal of Vertebrate Paleontology 32, 1447-1452.

Padian, K., Cunningham, J. R., Langston, W. & Conway, J. 2021. Functional morphology of Quetzalcoatlus (Pterodactyloidea: Azhdarchoidea). Society of Vertebrate Paleontology Memoir 19. Journal of Vertebrate Paleontology 38 (supp 4: 1), 218-251.

Witton, M. P. & Naish, D. 2008. A reappraisal of azhdarchid pterosaur functional morphology and paleoecology. PLoS ONE 3 (5): e2271. doi:10.1371/journal.pone.0002271

Witton, M. P. & Naish, D. 2015. Azhdarchid pterosaurs: water-trawling pelican mimics or “terrestrial stalkers”? Acta Palaeontologica Polonica 60, 651-660.

Vremir, M., Kellner, A. W. A., Naish. D. & Dyke, G. J. 2013. A new azhdarchid pterosaur from the Late Cretaceous of the Transylvanian Basin, Romania: implications for azhdarchid diversity and distribution. PLoS ONE 8(1): e54268. doi:10.1371/journal.pone.0054268

Vremir, M., Witton, M., Naish, D., Dyke, G., Brusatte, S. L., Norell, M. & Totoianu, R. 2015. A medium-sized robust-necked azhdarchid pterosaur (Pterodactyloidea: Azhdarchidae) from the Maastrichtian of Pui (Haţeg Basin, Transylvania, Romania). American Museum Novitates 3827, 1-16.

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If you read the previous article – recapping the events of TetZooCon 2024 – you’ll know that things went well. But what was that cryptic mention toward the end of a tour?

**Caption:** basic ingredients for a TetZooTour… (1) a snazzy coach, (2) stops at hotels, (3) dinosaurs in museums (here, a cast of the *T. rex* specimen AMNH 5027), (4) perplexing taxidermy (here, a koalabat at the Horniman Museum). Images: Darren Naish.

Yes: for the first time ever, I decided to run an official TetZooTour of southern England, the aim being to take a number of guests to such attractions as museums, zoological parks and palaeontological sites. On the morning of Monday September 30th – yeah, the next morning after TetZooCon, what was I thinking? – I and around 25 other people boarded a beautiful, brand-spanking-new tour coach at our designated pick-up in central London, and off we went. I’ve worked as a tour guide in the past and am aware of how much information has to be compiled in advance. Working in co-operation with Marc Bacon of South East Coaches, a tour was devised that would involve three days of travel, two overnight stays at hotels, and stops at a number of TetZoo-relevant locations.

**Caption:** we board and prepare to leave London. Our pick-up and drop-off points were essentially right next to the TetZooCon venue (Bush House, Kings College), and you can’t get more convenient than that. Images: Darren Naish.

Dinosaur rEvolution at the Horniman. I’m pleased to say that everyone arrived on time, and off we went to the first of our stops: the Horniman Museum and Gardens in Forest Hill, London, specifically to see the exhibition Dinosaur rEvolution. This is one of those separately ticketed exhibitions that’s not part of the museum’s main galleries, and its time at the Horniman is due to come to an end within the next few weeks. It’s based around the work and art of palaeoartist Luis Rey and I’m very pleased to report that Luis was able to accompany us and give us what was effectively a guided tour. The roaring dinosaur sounds sure made it hard to speak to a group, but we managed.

**Caption:** the outside of the Horniman Museum, as seen from the South Circular Road looking north. The museum opened to the public in 1901 and is named for its founder, Frederick John Horniman (1835-1906), who collected numerous items as a consequence of lifetime ownership of the Horniman Tea Company. The history of British tea companies is something else and *in part* involves Britain’s exploitation of Victorian China and its opium production. Image: Darren Naish.

I hadn’t seen the exhibition before and was impressed. While broadly functioning as a showcase of Luis’s art (and with his famous ‘chicken’ Deinonychus – not my term! – at centre stage), it’s designed to show how two of the main dinosaurian lineages (Theropoda and Ornithischia) evolved and existed in parallel, only one persisting beyond a certain mass extinction you might have heard of.

**Caption:** a (roughly) life-sized model of the Early Cretaceous North American dromaeosaurid *Deinonychus*, in the red-wattled, vibrantly feathered guise invented by Luis and used in his books. Incidentally, the dappled lighting across the model isn’t a random effect: I might be wrong, but *I think* we’re seeing light being projected through an image of a thin section of sauropod bone. Image: Darren Naish.

  **Caption:** at left, Luis Rey talks therizinosaurs to our tour group while standing alongside a replica skeleton of the large, North American *Nothronychus mckinleyi*. The large hand claws, small teeth, long and slender neck and large overall size of this animal are among the key distinctive features of this theropod group. At right, a closer view of the skeleton with some of Luis’s artwork making an appearance at edge of frame. Images: Darren Naish.

The exhibition features a large number of cast or replica fossils, including of archaic birds and other maniraptorans, a posed and mounted Velociraptor, Avimimus, the therizinosaurs Falcarius and Nothronychus, a T. rex skull, the David Sole scelidosaur, the famous tail-bristled Psittacosaurus, and several ceratopsid skulls. There are also several life-sized (or half life-sized) models, some animatronic, plus there are copious panels and art. That’s a lot of stuff.

**Caption:** ceratopsid ceratopsian skulls on show at Dinosaur rEvolution. In the image at left, we see (l to r) *Diabloceratops*, *Kosmoceratops* and *Coahuilaceratops*. The photo at right shows *Coahuilaceratops* on its own. This animal is supposed to have especially massive horns relative to the rest of its skull but, as you can see, it’s not entirely obvious that this is true. Images: Darren Naish.

It was a great opportunity to hear Luis recount his personal journey of discovery with respect to feathery and filamentous dinosaur fossils. As I’ve said before, he’s one of several artists who feels vindicated by 21st century discoveries: the decision to put feathers and spiny quills on Mesozoic dinosaurs always was a good one, and those scientists who – back in the 1990s – made Luis remove feathers from his illustrations don’t look prescient today. It’s difficult to say because it often sounds targeted at individual scientists and authors, and thus rude, but the conservatism so prevalent in the popular British dinosaur-themed literature and exhibitions of the 1980s, 90s and 2000s didn’t do us any favours. We’re stuck with work from that age that was decidedly lacklustre and uninspiring when new, let alone today. I predict that history will be kind to Luis Rey for promoting an avant-garde radicalism that proved broadly correct in the end.

The Horniman has other exhibits that are worth looking at, but their natural history gallery is currently closed, meaning that we couldn’t get to see the famous walrus or the various vintage palaeontology-themed displays. The walrus is famous because the taxidermist who prepared it incorrectly made it monstrously rotund in order to smooth out the wrinkles.

**Caption:** the Horniman Museum has a connection with mermaids thanks to study of this specimen, obtained by London’s Wellcome Trust in 1919, transferred to the Horniman Museum in 1982, and studied recently by Paolo Viscardi and colleagues. Coincidentally, Paolo spoke about this specimen and his work on it (Viscardi *et al*. 2014) at **the very first TetZooCon of 2014**. This work showed that the old story about mermaids like this being fish and monkey parts stitched together is not at all correct. Image: Darren Naish.

To the south and the west! And that was that for London; we bade farewell to the capital and headed out west, our destination for the evening being Bournemouth on the south coast of Dorset. To get there, we travelled through the New Forest – a fascinating area of the British countryside that I know well – and the Great Heath that surrounds Bournemouth town. How is it that Britain includes these large and not especially young, relatively open areas given the traditional view that prehistoric Britain was thickly forested north to south, east to west? We now think that ancient Britain was more of a habitat mosaic, with parkland-like areas existing between more wooded ones. This concept will be familiar to you if you’ve read Isabella Tree’s book Rewilding (Tree 2018) or – good for you – kept up to speed with publications on Britain’s prehistoric pollen record (Fyfe et al. 2013).

**Caption:** this image – showing Acres Down near the centre of the New Forest – wasn’t taken on the tour, but it depicts the sort of mosaic, partly open habitat that characterizes this famous area of south-west England. Is this a new thing, historically speaking, or do spaces like this have a long history in the British countryside? Image: Darren Naish.

In Bournemouth, we stayed in Boscombe, an attractive, tree-lined part of the city, and at a hotel just a couple of minutes away from the headquarters of the Bournemouth Natural Science Society (or BNSS). This has a natural history museum of its own that includes a palaeontological collection with numerous Jurassic marine reptile fossils. I tried and failed to get us special out-of-hours access, oh well.

Palaeo-Mecca, the fabled town of Lyme Regis. Why stay in Bournemouth? To be within easy reach of one of the world’s most famous palaeontological and geological locations, the south Dorset, seaside town of Lyme Regis. I’m lucky enough to have visited Lyme Regis many times, but many interested people – and not just those from abroad – have never had this privilege. The town is not exactly built for coaches (it has steep hills, narrow roads and very tight corners) but we got there and disembarked in perfect weather, only a day or two after moderately heavy winds and waves had swept along the coast. That was good, since it meant that fresh erosion had occurred across the cliffs and beaches, and that’s ideal when looking for fossils.

**Caption:** the Lyme Regis coast looking east, with East Cliff Beach (where we did our fossil hunting) visible in about the middle of the shot. The tide is out, exposing the wavecut platform along the shore. The vegetated strip inshore is home to numerous insects, and I recorded several interesting species on the day. Image: Darren Naish.

Literally on stepping off the coach we were met by palaeontologist and Lyme Regis Museum engagement and collections officer Dr Natalia Jagielska who had prepared a bespoke itinerary based on the things I wanted us to see. After a tour of Lyme Regis Museum and a talk on fossils and fossil-finding, we trekked to the amazing East Cliff Beach for an hour or so of fossil finding. This was timed to match low tide, of course. A good number of fossils were found, including attractive ammonites and bivalves, and partial belemnites and crinoids. Alas, no vertebrates of any sort (that’s not a ridiculous idea. I’ve found isolated ichthyosaur vertebrae on two previous occasions).

**Caption:** Lyme Regis Museum is a small but excellent museum, a must-visit if you’re interested in the history of the town or in its geology or palaeontology. Several key Jurassic marine reptiles, fishes and other animals are on show, as is associated art and sculpture. Images: Darren Naish.

We then visited the Mary Anning statue (where obligatory group photos and such were taken) before visiting the Anning family grave, another famous photo stop for people interested in palaeontology and its history.

The Lyme Regis dinosauroid. Then it was time for Dinosaurland, properly Dinosaurland Fossil Museum. I haven’t visited Dinosaurland for many years and was surprised by how different it looked relative to my previous visit (which probably happened more than 15 years ago). Dinosaurland is a converted former congregational church (Mary Anning was baptised there in 1799, and worshipped there until her conversion to Anglicanism during the 1830s) and features room after room of fossils of all sorts: over 20,000 in all. A mezzanine is home to a series of prehistoric dioramas that depict ancient life from the Silurian (or thereabouts) to the end of the Cretaceous. As you can see, the dioramas are of that special home-made quality that somehow combines impressive craftsmanship with pure distilled nightmare fuel.

**Caption:** the front of Dinosaurland Fossil Museum in Coomb Street, Lyme Regis. The museum’s ground-floor section has a one way system that is not great if you suffer from claustrophobia, but the building is fun overall. The tree fern on the right adds a Mesozoic vibe to the exterior. Image: Darren Naish.

  **Caption:** a fossil Mesozoic marine reptile montage from Dinosaurland, showing ichthyosaurs at left and a teleosauroid thalattosuchian at right. Hey, **did I mention that I wrote a book on Mesozoic sea reptiles?** I understand that it’s sold out in a lot of places now. Images: Darren Naish.

  **Caption:** a model like this (it’s big: somewhere round about 6 m long) has various technical, anatomical inaccuracies. But imagine building a big model like this yourself, with affordable materials. I think it’s actually pretty good. It depicts the British spinosaurid *Baryonyx walkeri*. Image: Darren Naish.

  **Caption:** dinosaur-themed dioramas from the mezzanine section of Dinosaurland. At left, an Early Jurassic scene depicting what I think is a *Coelophysis* adult and juvenile. At right, a Late Jurassic scene that shows (perhaps) *Ornitholestes* (holding a lepidosaur of some sort in its right hand), with a geologically older sauropodomorph peeking in at left. The look of these models remind me of illustrations from Michael Tweedie’s 1977 book *The World of Dinosaurs*, and might be based on them. Images: Darren Naish.

I was there for one model in particular. Tucked away at the end of the dioramas and in its own glass case is ‘Saurian’, a very English version of Dale Russell and Ron Séguin’s dinosauroid model of the early 1980s. Dinosaurland’s ‘Saurian’ isn’t, technically or artistically, quite on par with the Canadian original, and an accompanying information sheet doesn’t really explain its backstory: it notes that the model depicts what “dinosaurs could have looked like if they had survived to the present day” (what… all of them?) but it doesn’t make reference to Russell and Séguin’s project nor explain how its appearance is based on that of their very excellent model.

**Caption:** Dinosaurland’s infamous ‘saurian’ model, one of several similar models present worldwide. As might be obvious from the image at right, we had fun having our photo taken nearby. Thanks to the anonymous museum-goer who kindly took this photo for me. Images: Darren Naish; anonymous museum-going woman.

It does, however, have a valid connection to the real dinosauroid in Canada because the reaction of naïve visitors to its presence is tellingly similar. As Will Tattersdill and I recounted in our 2021 article on the dinosauroid (Naish & Tattersdill 2021), people encountering the model for the first time (and unaware of its backstory) assume that it’s a misplaced alien. The similarity of the dinosauroid with certain fictional aliens is not coincidental given that Dale Russell was interested in, and very much aware of, humanoid alien stories: one of the ideas he was driving at is that humanoids were inevitable and – potentially – a widespread part of evolutionary history across the cosmos (Naish & Tattersdill 2021).

And that was that for Lyme Regis. A few members of our group took time to visit various of the shops and eateries about the town, but it would have been good to spend more time there. Alas, we only really had a few hours in Lyme Regis since arriving late and leaving early is all part of the coach tour experience (it has to be because of leaving, and getting to, hotels). I’ll keep that in mind for the future though.

**Caption:** our group at one of several Mary Anning-themed stops in the town, namely the family grave. It has become traditional to leave both flowers and fossils at the grave. Its location at the very front of the graveyard is not coincidental, and is related to the fact that she is its most famous resident. Image: Darren Naish.

I owe massive thanks to the Lyme Regis Museum staff – Natalia in particular – for doing such a phenomenal job in hosting us and enabling us to see everything I hoped we would. We left on time, and struck out east, heading this time to England’s south-east, and specifically Folkestone on the coast of Kent.

At the coast of Kent. Our second hotel stay was at the large, coast-side Grand Burstin Hotel in Folkestone. One of the concerns when leading a tour is that hotels can take a long time – more than an hour – to register all members of a large group, something that can be tedious and problematic when you’re arriving late, or behind schedule. I’m very pleased to say that we had none of that, and that our hotels were fully prepared for our arrival. A number of us stayed up too late in the bar. I don’t remember why, but I think that mekosuchines and books on sauropods were mentioned at one point or another.

**Caption:** the morning view from my room at Folkestone. For reasons, I got upgraded to an executive suite but don’t tell the others. Image: Darren Naish.

  **Caption:** Folkestone Harbour on the morning of our departure, showing the swing bridge and some attractively textured sediment. Looks great in nice weather. Image: Darren Naish.

The reason for our stay in Folkestone is that our final excursion was to Howletts Wild Animal Park – often called Howletts Zoo or just Howletts – for a day of zoo-going. There’s a lot to say about Howletts, in part because the zoo is synonymous with the late John Aspinall and a fairly notorious former safety record. I was careful on our approach to emphasize that this is nothing for a visiting tour party to worry about, since the relevant incidents have all involved keepers. Anyway, the drive from Folkestone to Howletts is not a long one and we arrived before the zoo was even open, giving us time to obtain the group photo you see here.

**Caption:** our group outside Howletts. I’m in this photo so obviously didn’t take it. The mammoth with the massive dome on its head is very distinctive and various models of this likeness are scattered about the outdoor attractions of the UK. This is the second image here that features a photobomb from a dinosaur. Image: Marc Bacon.

Animals at Howletts. Due to a technical fault that was absolutely not due to me forgetting to charge the battery, my camera was inoperative during my time at Howletts, so no good photos… oh, how that hurts. Howletts is a reasonably large zoo and I built our itinerary on the assumption that we might get to see about half of what the zoo has on show. But no, in the end we had time to do it all, despite the inconvenience of a brief, heavy rain shower.

The zoo is wholly mammal-focused (ostriches are present in one enclosure but that seems to be it for non-mammals) and is especially strong on primates, big hoofed mammals, cats and wild dogs. Beginning with African lion Panthera leo, Giant anteater Myrmecophaga tridactyla, Lowland tapir Tapirus terrestris, bongos (specifically the rare Keynan Mountain bongo Tragelaphus eurycerus isaaci), Sumatran tiger P. tigris sumatrae, Amur leopard P. pardus orientalis, Snow leopard P. uncia and more, we walked west to have lunch at the Pavilion Restaurant before heading north and back east to complete the circuit.

**Caption:** the lack of a functioning camera means that I had to rely on my phone, but the photos it takes aren’t all that bad. Left to right: Servals *Leptailurus serval* (one of which is carrying a deceased rodent in its mouth), juvenile lions (one of which had a deformed hindfoot), and East Javan langur or Javan lutung *Trachypithecus auratus*. Images: Darren Naish.

Animals seen on the second half include European wolf Canis lupus, Mainland clouded leopard Neofelis nebulosa and Caracal Caracal caracal, and we finished at the large, field-like enclosures that are home to African savannah elephant Loxodonta africana and Black rhino Diceros bicornis.

**Caption:** my photos might not be at all great, but at least a few other people on the visit did have good cameras with them. At left, a Sumatran tiger. At right, a male Western lowland gorilla *Gorilla gorilla gorilla* with an especially impressive head dome. I *think* that this is Djanhou, born at Howletts in 1993 and father to eight offspring. Images: Zach Wait.

Life-sized fossil mammals. An interesting feature we just had to look at was the outdoor Ice Age Mammals exhibit where a number of life-sized models of fossil mammals – and not all of them from the ‘Ice Age’ (meaning the Pleistocene) – are arranged in a wooded area in the north of the zoo. As you can see, some of the models are not that bad. Others are bad indeed. A concern I have is that many are equipped with full coats of simulated fur. While I’m not sure, I’m reasonably confident that this is fake and plastic, and I wonder if we have to worry about such material being shed into the environment and thereby contributing to plastic pollution.

**Caption:** models from near the end section of the outdoor fossil mammal exhibit, depicting a Woolly mammoth and the giant Asian rhino *Elasmotherium sibiricum*. I agree with arguments that *Elasmotherium* didn’t have an immense keratinous horn like this, but instead possessed a rounded, more dome-like, keratinous structure. It’s important to recognise how big *Elasmotherium* is: its skull is about 75 cm long. Images: Darren Naish.

Anyway… the exhibit included not only stalwarts like Woolly mammoth Mammuthus primigenius, Smilodon and a mega-horned version of Elasmotherium, but a dinoceratan (I think Eobasileus), the twin-horned Arsinotherium and an especially terrifying rendition of the stem-whale Ambulocetus. It turned out that we moved through the exhibit the wrong way round, since we started in the Pleistocene and ended up in the Late Cretaceous, rather than vice versa. The sole Cretaceous feature, if you’re wondering, is a Triceratops skull covered in clambering little Purgatorius.

**Caption:** models from near the middle section of the fossil mammal exhibit. The dinoceratan at left is just about the only model you can approach closely, as I’m doing here. The top of its head and shoulders had a smattering of small bloody chunks and feather tufts about them, showing that the model had recently been used as a plucking perch by a raptor (likely a sparrowhawk). At right: a scary *Ambulocetus*. I don’t think that this is a good look for the animal. I’ve reconstructed it several times myself and both the head shape and tooth configuration shown here aren’t right. Images: Zach Wait; Darren Naish.

And that was that. We had time to visit the shop and were fully boarded and ready to head back to our London drop-off point precisely on time. Traffic in London is notoriously difficult, but we made good and reached the drop-off just before 5pm, as planned. We then had a bit of trouble with our otherwise dependable wheelchair lift, but this was fixed after a little while and wasn’t disruptive in the end.

And that was the first TetZooTour. What a win! Everything ran to time, nobody got lost or injured, and we got to all of our stops and venues as planned with no disruptions or diversions. Each and every one of the tour-goers did us proud in terms of keeping to time and functioning as part of a group. I had a great time and the feedback I’ve received so far is positive.

Setting up and running this tour was a total gamble and I had no idea whether enough people would pay up and want to come along, but in the end they did, and costs were covered. It should also be obvious that the whole thing was a lot of fun and that I found it rewarding. It’s too early at this stage to be sure that a repeat performance will be happening next year (and beyond) but – right now – I consider it likely that it will, with different stops and different TetZoo-relevant attractions. Remember that, whatever happens, it’ll be part of DinoCon from 2025.

**Caption:** our shiny new coach. It served us well. Image: Darren Naish.

It only leaves me to say thanks once more to Marc (our driver and co-organizer) and South East Coaches, to Luis Rey and staff at Horniman Museum and Gardens, to Natalia and other staff at Lyme Regis Museum, and to those fine people who opted to join me. And that was the very first TetZooTour.

For previous articles on TetZooCon, see…

The events of TetZooCon 2014, July 2014
TetZooCon 2015 Is On, July 2015
The Events of TetZooCon 2015, November 2015
Coming Soon: TetZooCon 2016, September 2016
The Day After TetZooCon, October 2016
The Fourth TetZooCon, September 2017
The TetZooCon of 2017, October 2017
Reasons to Attend TetZooCon 2018, September 2018
TetZooCon 2018: Best TetZooCon So Far, October 2018
Announcing TetZooCon 2019 – the Biggest Yet, August 2019
Final Call For TetZooCon 2019, October 2019
The Sixth TetZooCon, October 2019
TetZooCon 2020 + Zoom = TETZOOMCON 2020, December 2020
TetZooMCon 2020 an Unbridled Success, December 2020
Cronch Cats, Beasts of Gévaudan, Dinosauroids, Mesozoic Art and Much More: TetZooMCon 2021 in Review, September 2021
The Ninth and Largest of the Tetrapod Zoology Conventions, December 2022
Announcing the 10th Tetrapod Zoology Convention, October 2023
At TetZooCon 2023, November 2023
TetZooCon 2023 the report, December 2023
The Last TetZooCon, October 2024

If you enjoyed this article and want to assist me in what I do — and see behind-the-scenes stuff in development — please support me at patreon. Click anywhere on this text in bold.

Refs - -

Fyfe, R. M., Twiddle, C., Sugita, S., Gaillard, M.-J., Barratt, P., Caseldine, C. J., Dodson, J., Edwards, K. J., Farrell, M., Froyd, C., Grant, M. J., Huckerby, E., Innes, J. B., Shaw, H. & Waller, M. 2013. The Holocene vegetation cover of Britain and Ireland: overcoming problems of scale and discerning patterns of openness. Quaternary Science Reviews 73, 132-148.

Naish, D. & Tattersdill, W. 2021. Art, anatomy and the stars: Russell and Séguin’s dinosauroid. Canadian Journal of Earth Sciences 58, 968-979.

Tree, I. 2018. Rewilding: The Return of Nature to a British Farm. Pan Macmillan, London.

Viscardi, P., Hollinshead, A., MacFarlane, R. & Moffatt, J. 2014. Mermaids uncovered. Journal of Museum Ethnography 27, 98-116.

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Once again, it was recently that part of the year where a good number of science enthusiasts, natural historians, wildlife artists, scientists, researchers and interested members of the public gather in London for what we call TetZooCon: the annual Tetrapod Zoology Convention…

**Caption:** at left, the front entrance of our venue... Bush House. At right: what's this... DinoCon? Images: Darren Naish; Gemma Hazeborg.

Held once more at Kings College in central London, TetZooCon 2024, the 11th of these meetings, was the biggest and best yet, and perhaps also the most tidily organized and well-run because… oh my god have we learnt from the lessons of the past.

TetZooCon 2024 ran from September 27th to 29th and was co-led by myself, palaeoartist John Conway, historian and author Chris Manias, and a team of helpers and assistants. Our venue (Bush House) provides good-sized auditoriums and lecture theatres, but this year we also won access to two massive and spectacular terrace areas, one of which (the South Terrace) has a balcony with an incredible view of the London skyline. You’ll see what I mean from the photos below.

**Caption:** the view from the South Terrace was fantastic when the weather and lighting was right. The tower to the left belongs to the (closed) St Mary le Strand church; across the Thames we see the London Eye and the Palace of Westminster at far right. Waterloo Bridge and (further away) Westminster Bridge are visible here too. Image: Darren Naish.

Things kicked off on Friday evening with a drinks reception followed by the on-stage panel event ‘Of Weird Animals, Books, and Publishing’, themed broadly around books and on being an author. I led, and was joined by zoological consultant and author Dani Rabaiotti, researcher and author Natalie Lawrence, and the aforementioned Chris Manias. And that was that for the Friday. I didn’t stay out too late.

**Caption:** our Friday evening panel event ‘Of Weird Animals, Books, and Publishing’ showing (l to r) Darren Naish, Dani Rabaiotti, Natalie Lawrence and Chris Manias. On the screen behind us are images of various of the books we’ve been involved in: each author is sat close to images of their own book or books. Image: Georgia Witton-Maclean, used with permission.

Doors opened to TetZooCon proper on Saturday morning, and here’s where most of the stalls were set up and made ready for trading. Our stalls were incredible, surely representing the very best in terms of what the UK has to offer in terms of animal- and palaeo-themed merch. Toys, models, pin badges, ornaments, art prints, posters, books and much more were on offer. I signed and sold copies of both Tetrapod Zoology: Book One and Ancient Sea Reptiles (second edition), as well as a large number of animal models.

I don’t want to start talking too much about the stalls because there were many of them and they were all good. But I’ll mention Sam St Leger and his All Yesterdays figures, Matt Dempsey and his army of 3D-printed tyrannosaurs, Carla Owens and her beautifully crafted dinosaurs, pterosaurs and other animals, Paul Glynn and his phenomenal sharks, bronze dinosaurs and ceramic whales, and of course Jed Taylor and Ruadhrí Brennan’s incredible model dromaeosaurids and replica dinosaur skulls. My god there was a lot of great stuff. You know that it’s only going to get better in future… or worse, depending on your finances.

**Caption:** Paul Glynn of **The Younger Earth** at his TetZooCon stall, showing his incredible models and sculptures of sharks, dinosaurs, dinosaur skulls, whales and more. I now own that pike and one of the bowhead whales! Image: Georgia Witton-Maclean, used with permission.

  **Caption:** a more detailed view of pieces on show at Paul Glynn's stall. Why couldn't I be born rich? I need me those things. The megalosaur here (the white one at left and the bronze one at right) also exists in beautiful painted form: see below. Image: Darren Naish.

  **Caption:** TetZooCon heroes Jed Taylor (l) and Ruadhrí Brennan, here holding Baby Bruce the Blue Goose, a life-sized juvenile *T. rex* bust who had a starring role in this year's events. He peered about London town, he consumed a few stall-holders and guests, and he pranced like a bipedal pony in the group photo. This photo was taken somewhere about half-way through TetZooCon, so some of Jed and Ru's wares had already disappeared. Look at the amazing replica skulls (including the pelagornithid or pseudotoothed bird), the magnet at far right, and the model *Megalosaurus* near the middle. Image: Georgia Witton-Maclean, used with permission.

TetZooCon talks, day 1. On talks and panel events and so on, John and I gave our short, customary welcome and here’s where John dropped – in passing – the bombshell that this was set to be the last TetZooCon. An audible gasp and several wails and sobs rippled through the auditorium. We planned to return to this point right at the end, on Sunday evening. Things then kicked off with my ‘Brian Ford and the Dinosaurs: an Unsuccessful Effort to ‘Make Dinosaurs Aquatic Again’’; the talk might be familiar, albeit entertaining, stuff if you’re followed the conversation, but this time round it’s framed to emphasize how scientists and science popularisers need to constantly work hard to push back against the anti-scientific egotists who aim to subvert society for their own ends. The good news is that quite a few good people are doing this sort of work in the public sphere. The bad news is that there aren’t enough of them in view of the sway the contrarians have and the size of the platforms they keep being given.

**Caption:** me, with the opening slide of my talk on Brian Ford’s aquatic dinosaur thing. I tweeted about the existence of this talk prior to TetZooCon 2024, and a consequence is that Mr Ford himself piped up in my twitter responses while I was on the tour. Ford’s opening gambit reads “Such sweet, old-fashioned comments. Curious that nobody is courageous enough to admit that every single scrap of scientific evidence substantiates my view”, and things went downhill from there. The relevant twitter thread is **here**, if you’re interested. Image: Georgia Witton-Maclean, used with permission.

On the subject of contrarians, some of you will know that – when giving the Ford talk at another recent event (2024’s Symposium on Vertebrate Palaeontology and Comparative Anatomy at the University of Southampton) – I was challenged, in the Q&A following my talk, by a researcher who objected to my characterisation of Alan Feduccia as a ‘contrarian’. I responded by arguing that Feduccia sure the hell is a contrarian, and that this isn’t just due to his apparent lack of familiarity with data but, more insidiously, to his wilful use of unsound argumentation. Check my review article on his book Romancing the Birds and Dinosaurs for full explanation of that.

**Caption:** at left, a signed copy of *Tetrapod Zoology: Book One*, sold at TetZooCon 2024. At right, a slide from my talk that brings attention to Tet Zoo articles I’ve published on specific contrarians. Images: Darren Naish (l); Finn Holmes-Kellett.

Palaeoart no, podcast yes. Some of you will know that palaeoartist Greg Paul was scheduled to talk at TetZooCon 2024. Greg has been a formative person in the careers of both John and myself, and it seemed right to have him attend. But, no, it didn’t work out due to illness. Once we learnt of this, we aimed to book other historically significant palaeoartists, namely Doug Henderson, Mark Hallett and Mauricio Anton. All plans fell through for one reason or another.

**Caption:** several photos taken during the podcast record have a more chaotic or active air than this sedate one, but here’s Darren Naish and John Conway talking stuff. The audience was supposed to tell me to stop when I was droning on too much but they spectacularly failed in this endeavour. Image: Georgia Witton-Maclean, used with permission.

In the end we filled the space with a live record of a podcast episode that included a bit of audience participation and some discussion of the crossover between dicynodonts, South African rock art and the contents of Cryptozoologicon Book One. The relevant episode (# 94) will be released sometime soon. Oh, for the podcast back catalogue go here.

I led one final event: a Prehistoric Planet discussion, this time focusing on the science and backstory to the Badlands episode of season 2. If you were still listening to my voice by this point… boy, are you a sucker for punishment. After lunch, some people perused the stalls some more but a bird-dominated talk session happened, as did the first of our Palaeoart Workshops. The latter were held in the grand, spacious North Terrace. What a contrast to the cramped, sometimes windowless, rooms we’ve had to occupy in the past.

**Caption:** a big surprise for me was how popular my 2010 *Tetrapod Zoology: Book One* turned out to be. All copies sold. So maybe I should buy more and repeat this. At right, I also sold cryptid-themed stickers, and they sold well too. Images: Darren Naish.

  **Caption:** over the years, I’ve grown fatter and richer by selling Invicta dinosaur models to (soon to be Dr) Matt Dempsey, and this year we both benefitted by my selling of this fine Invicta *Apatosaurus*. Matt has awarded it a name and clearly likes it very much. Image: Matt Dempsey.

The modern bird session. On to the bird talks. An excellent and inspiring talk – Curlew Conservation - A Battle for Hearts and Minds – was provided by Mary Colwell. Mary is almost certainly the most decorated person we’ve hosted as a speaker at TetZooCon and in fact was awarded an additional accolade – the ZSL Silver Medal – literally the day before speaking.

**Caption:** Mary Colwell talks curlews and her discovery of how bad things are. As explained in her 2019 book *Curlew Moon*, she found that urgent action was required. Image: Georgia Witton-Maclean, used with permission.

The plight of the Eurasian curlew Numenius arquata in the UK is depressing, but also unsurprising in view of the vulnerability of this large, ground-nesting wader to human activity, agriculture and industry. Sheep farming, arable machinery and our insistence that the world exists as a playground for pet dogs all pose serious problems for curlew persistence, and their numbers have crashed in recent decades. On realising how bad things are, Mary set about forging alliances and holding meetings, the result being the charity Curlew Action. They had a stall at TetZooCon 2024 and I hope they did well.

**Caption:** the Curlew Action stand. Books, socks, artwork, tote bags and more. Curlew Action – **their website is here** – organizes workshops and events related to curlew conservation, and merch and products are available there too. Image: Georgia Witton-Maclean, used with permission.

A second very excellent talk on modern birds was provided by Jonathan Meiburg, author, researcher and lead singer of the band Shearwater. Jonathan’s talk – The Caracaras Would Like to Meet You: Encounters with the World's Smartest and Strangest Birds of Prey – matches the contents of his book A Most Remarkable Creature (Meiburg 2021). Caracaras have fascinated and surprised people as long as they’ve known about them: these curious, intelligent, adaptable birds behave in ways that more recall corvids or parrots than falcons or other raptors. We were treated to a tour of the world’s caracara species, accompanied all the while by excellent photos.

**Caption:** a left, Jonathan Meiburg talks caracaras, with the most familiar member of the group – the Crested caracara *Caracara plancus* – on screen. At right, my personal copy of Jonathan’s 2021 book *A Most Remarkable Creature*. This is the initial, hardback edition. More recent versions of the book have a white cover with a photo of a Striated caracara *Phalcoboenus australis*. Images: Georgia Witton-Maclean, used with permission; Darren Naish.

I’ve mentioned several times how good Jonathan’s book (Meiburg 2021) is, so I was shocked to hear how it essentially sank out of trace here in the UK. In fact, it’s currently only available via print on demand! That’s not right and I want to turn this around. My review will appear here as soon as I have time to finish it. If you’re at all interested in predatory birds, or in bird biology, evolution or behaviour in general, in the history of zoological discovery, or in the keeping of animals in captivity, be sure to order this book yourself right now. You won’t regret it.

Palaeoart and book signings. As mentioned, the first of our Palaeoart Workshops also happened after lunch and in parallel to the bird talks, alas. There isn’t enough time to do everything. Indeed, I always miss the palaeoart workshops because I have to field the talk sessions. As usual though, I’ve only heard good things. Saturday’s workshops included a show and tell, and also a special event – co-led by Gemma Hazeborg, Marc Vincent and Natee Himmapaan of Love in the Time of Chasmosaurs – titled ‘An Ode to Bad Palaeoart’ and marking 15 years of LITC. I salute the team, and it’s worth noting that all LITCers (those based in the UK, anyway) are long-time TetZooCon attendees and supporters. Following the talk, attendees were invited to produced ‘bad’ palaeoart of their own. Much of it was surprisingly good… if you know what I mean.

**Caption:** this image gives some idea of how grand and spacious our palaeoart workshop was. This was taken when the LITC team (visible standing at right) were introducing their Ode to Bad Palaeoart segment. John Conway stands at left. Image: Georgia Witton-Maclean, used with permission.

  **Caption:** just some of the ‘bad palaeoart’ created by the workshop participants, spread across the stage. Pretty obvious that there’s a lot of good stuff here, and I spy what’s becoming a (Dougal Dixon-inspired) TetZooCon meme. Image: Georgia Witton-Maclean, used with permission.

  **Caption:** another image of the ‘bad palaeoart’ illustrations produced by the workshop participants. The musculoskeletal reconstruction of a tyrannosaurid on the screen was produced by John Conway, I think for use in a certain Apple TV+ series devoted to a planet that was prehistoric. Image: Darren Naish.

The latter part of Saturday involved more art: this time a proper exhibition featuring numerous original images kindly loaned from Dave Hone’s personal collection.

**Caption:** just some of the original framed art brought along to TetZooCon 2024 and put on display in the North Terrace by Dave Hone. I see original pieces here by Mark Hallett, Luis Rey and Joschua Knüppe, among others. Image: Georgia Witton-Maclean, used with permission.

Jamale Ijouiher showed up on Saturday with stock of his 2022 The Desert Bones: the Paleontology and Paleoecology of Mid-Cretaceous North Africa (Ijouiher 2022). Joschua Knüppe, who produced art that appears throughout the book, was present as well, as was James McKay, creator of the excellent cover art, this meaning that you could get the book signed by all three people who contributed to it. I’m pleased to report that Jamale shifted his entire stock. Take note authors of dinosaur-themed books.

**Caption:** at left, Joschua Knüppe (l) and Jamale Ijouiher signing copies of *The Desert Bones: the Paleontology and Paleoecology of Mid-Cretaceous North Africa*. James McKay had had to leave by the time this photo was taken. At right: Jamale once commissioned his own *Spinosaurus* model, and here it is. It was constructed by artist and sculptor Victoria Nampisano. Images: Darren Naish.

Vanishing vipers and the making of monsters. A final set of post-lunch talks happened in lecture theatre 1during Saturday afternoon. We don’t hold all talks in the main auditorium, a decision made to avoid it becoming too hot. Those of you who attended TetZooCon 2023 will know that being too hot turned out not to be such a problem. Anyway, these talks were decidedly not-dinosaur-themed. First up was Angela Julian of Amphibian and Reptile Conservation and Amphibian and Reptile Groups of the UK on ‘Revealing the Secrets of the Vanishing Viper’.

The UK’s only viper – the Adder Vipera berus – is in worrying decline, and now extinct across some section of the English midlands. Despite concerted conservation efforts and grassroots campaigns to promote feelings of good will towards these small, beautiful snakes – much of it led by Angie and her colleagues – there remain rogue individuals who kill adders on sight, and of course incessant tabloid headlines about 15 foot adders murdering dogs and so on. Adders are resilient though, and sites that are home to them today were wholly unsuitable just a few decades ago.

**Caption:** the good thing about living in a nation where there are scarcely any native (non-bird) reptile species is that an interested person can get to know all of them. At left, here's a female Adder I observed at Bossiney Cove, Cornwall, in 2018. At right, Angela Julian talks grassroots conservation efforts relevant to adders. Images: Darren Naish (l); Georgia Witton-Maclean, used with permission.

My feelings on the state of wildlife in the UK – speaking here as a natural historian who spends a lot of time looking at, and for, local wildlife – is not positive and much of our wildlife is on its last legs. That’s a point that was revisited several times during TetZooCon 2024.

Natalie Lawrence spoke next on ‘Making Monsters’, a tie-in with her new book Enchanted Creatures (Lawrence 2024). Are monsters based on misunderstanding and misinterpreting exotic animals, what role have ‘Chinese whispers’ and copying played in historical interpretations of animals like walruses and dodos, and how has the discovery of animals from the geological past played into thoughts on what monsters might be like? Natalie brought stock of Enchanted Creatures with her and we held another signing event. Again, they all sold… are you noticing a pattern here?

**Caption:** Natalie Lawrence talks making monsters. If you know about the history of monster depiction in classical literature and art, you should recognise the beast being ridden in the illustration. Image: Georgia Witton-Maclean, used with permission.

  **Caption:** at left, Natalie Lawrence discusses the recycling and re-using of images in old literature on unusual animals. The misunderstood walrus shown at left has appeared here and there in the literature as a hitherto unknown beast associated with freshwater environment. At right, Natalie and your humble author. Images: Finn Holmes-Kellett (l) and Darren Naish.

Dinosaur biology and behaviour. Sunday morning kicked off with a Mesozoic dinosaur session. Kai Caspar – a primate expert who’s also published on Mesozoic marine reptiles, spiders and dinosaurs too – led with ‘How smart was T. rex? Current debates in dinosaur neurology and cognition’. The core of this talk was coverage of the paper (Caspar et al. 2024) in which we responded to Herculano-Houzel’s (2022) argument that big Mesozoic theropods (and other fossil dinosaurs too) were primate-like in intelligence. The Tet Zoo overview of this work is here.

**Caption:** Kai Caspar talks Mesozoic dinosaur brain anatomy and what it might, and might not, mean for behaviour and biology. And yay for more *Prehistoric Planet* representation. Image: Georgia Witton-Maclean, used with permission.

But Kai covered so much more, including brain anatomy and neuron density across vertebrates in general, the history of hypotheses on animal intelligence (whatever we mean by that term), and where we’re at in our understanding of intelligence across reptiles. As I’ve emphasized in several articles here at Tetrapod Zoology, non-bird reptiles of several sorts have performed well in cognitive tests, and it’s reasonable to suggest that some of these traits could have been present in the reptiles of the Mesozoic. But none of this means that these animals were primate-like. This was my favourite talk of the whole event, I loved it.

**Caption:** one of the graphs from **Caspar *et al*. (2024)**, here being explained by Kai during his talk. As explained in the paper, we found non-bird maniraptoriforms to be more like birds in relative brain size than are other theropods.... but this wasn't true of *all* of them. Some alvarezsaurids are outliers. Image: Georgia Witton-Maclean, used with permission.

Dave Hone followed with a whistle-stop tour of what we know, and what we can sensibly infer, on Mesozoic dinosaur behaviour. This is a summarized form of what Dave covers in his brand-new book Uncovering Dinosaur Behaviour (Hone 2024). Princeton University Press had a stall at TetZooCon 2024 and – great news – were able to bring along advance stock of Dave’s book. I’m pleased to report that the books sold well and quickly, and all copies of Uncovering Dinosaur Behaviour were gone within a couple of hours. PUP also bought copies of my Dinopedia, Greg Paul’s field guides, Katrina van Grouw’s Unnatural Selection, and more.

**Caption:** the presence of Princeton University Press (PUP) meant that copies of Katrina's *Unnatural Selection* were on sale (sadly, *The Unfeathered Bird* is very much sold out and out of print), as was my *Dinopedia*. Images: Princeton University Press.

  **Caption:** me (at left) introducing Dave Hone and his TetZooCon talk. I made a joke about Dave deliberately courting controversy seeing as he's worked on spinosaurid behaviour, sexual selection theory, and tyrannosaurid ecology and body size. There's a photo of him making a very unusual expression while I was saying those words. Image: Georgia Witton-Maclean, used with permission.

  **Caption:** Dave Hone's new book features throughout illustration by the very talented Gabriel Ugueto, and here are two of them. I think that the context should be obvious, though the point remains that finding fossil dinosaurs preserved while engaging in copulation is not entirely impossible. In addition, we might also be able to find compelling evidence for the existence of grooming or preening in extinct dinosaurs (I've written articles on this). Image: Finn Holmes-Kellett.

  **Caption:** Gabriel Ugueto couldn't make the meeting in person (we asked!) but at least he was there in spirit. You can see that Dave said very nice things about him. Image: Darren Naish.

Why Dinosaurs? at TetZooCon. A dinosaur-themed, err, theme continued over lunch as we hosted a screening of the new film Why Dinosaurs?, one of just a handful of showings it’s so far had in the UK. Ellinor Michel (of Friends of Crystal Palace Dinosaurs fame) introduced the film for us, which is appropriate given that she stars in it. We owe massive thanks to Tony Pinto for permission to screen the film and for working with us in making it happen.

I wasn’t able to attend the screening (I saw the film previously at the Lyme Regis Fossil Festival earlier in the year), but I heard that the audience reaction was highly positive and involved appropriate reaction. If interested in arranging a screening of Why Dinosaurs? yourself, contact the team via their website here.

**Caption:** **Friends of Crystal Palace Dinosaurs** were once again represented at TetZooCon and I hope they did well. You can see a stack of Mark Witton and Ellinor Michell’s book *The Art and Science of the Crystal Palace Dinosaurs* in the background, but new for this year were these 3D-printed busts of several of the Crystal Palace models. It’s obvious that, were these for sale, they would sell like proverbial hotcakes. Hopefully that will be the case in the near future. Image: Darren Naish.

  **Caption:** an indication of how busy the stalls areas were at times. The Friends of Crystal Palace Dinosaurs stall is in the foreground. Image: Georgia Witton-Maclean, used with permission.

Unfeathered Bird II and Nextinction. More dinosaur-themed material occurred after lunch as Katrina van Grouw presented ‘Unfeathered Bird Update: Progress Report From the House of Bones’. I regard Katrina’s The Unfeathered Bird as one of the most interesting and important zoological books of recent years, and the good news is that it’s going to get a substantially expanded, upgraded second edition. I very much look forward to more news on that project, and Katrina’s talk made it clear that it will include so much new material relative to the first one. My old review of The Unfeathered Bird, incidentally, can be found here [UPDATE: I cannot find a working link. To be added!].

The last talk of the day was provided by conservationist, author, TV presenter, wildlife photographer and artist Chris Packham – what a coup – and was titled ‘Nextinction’. Chris is best known today as an environmental campaigner: from an introduction that explained the plight and decline of local butterflies, he discussed his thoughts on legal action against the British government, the importance of protest, and what we can do about industrial agriculture and raising awareness of the pre-eminence of environmental concerns. Things are bad, and it’s obvious that politicians, industrialists and some branches of the media would like us to ignore this and think of other things.

The subject is a bit of a downer but we cannot ignore it.

**Caption:** heartfelt thanks to the incredible Chris Packham for taking time to join us and give a talk. This photo was taken during the Sunday afternoon drinks reception and art exhibition: l to r, Darren Naish, Chris Packham, John Conway. Image: Georgia Witton-Maclean, used with permission.

  **Caption:** we finally remembered to do a group photo! We did one way back at the very first TetZooCon (in 2014) but have consistently failed to do once since. The photo doesn’t include everyone who was present, but it’s the bulk of them at least. Georgia, our official photographer, is in the photo at far left so can’t be credited as having taken this one. Image: Neil Phillips, used with permission.

The quiz and the end. As ever, we wrapped up with our famous/infamous TetZooCon quiz. 30 questions from things related in some way to the TetZooniverse, ranging from the taxonomic history of obscure cat species to the cast of Disney’s Dinosaur and the breeding history of the UK’s egret species. Prizes included books provided by Steve White, artwork by Joschua Knüppe, and a good number of animal figures and models from our friends at Everything Dinosaur. I’m told that this year’s quiz was especially fiendish, but – even so – we still had players scoring 19 out of 30 (well done Sean Hennessy and James Appleby).

**Caption:** our two cosplay winners. At left, James Appleby as E. D. Cope (James did have a plesiosaur skeleton with the head wrongly mounted on the end of the tail about his person). At right, Katrina van Grouw as a museum specimen of a Great auk *Pinguinus impennis*. Images: Neil Phillips (l), Darren Naish.

  **Caption:** at left, TetZooCon 2024 quiz winner (joint winner) Sean Hennessy with his prize. At right, Jed Taylor and Baby Bruce again. Images: Georgia Witton-Maclean, used with permission; Darren Naish.

Prizes were also given out for palaeoart winners from the Palaeoart Workshop, for our cosplay participants (James Appleby, again, and Katrina van Grouw), and also for whoever played the quiz and got the lowest score, who had attended the greater number of TetZooCons (Bob Nicholls!), and who had the highest number of dinosaur-themed tattoos! That last prize went to Franklin, who is quite the walking piece of art, let me tell you.

**Caption:** I don’t have photos of Franklin’s upper half, which features a ton of great dinosaur-themed art, but here are lower leg images showing Mary Anning’s original *Plesiosaurus*, a *Dilophosaurus* skull, a Shoebill, a phorusrhacid (its bill peeking in at the right) and more. That’s commitment to the cause right there. Images: Darren Naish.

And thus things came to an end. But no ordinary end…. the end. Because this, you see, was the last ever TetZooCon. TetZooCon is such hard work and provides such insufficient financial return for our time and effort that we just can’t do it anymore, and so John and I bid our farewells and left the stage. Only to return and announce what’s happening next. From 2025 onwards, we’re (me and others; John’s involvement is being pared down) transitioning to a larger, out-of-London event dubbed DinoCon. No, it won’t just be dinosaurs, it’s just that this name has more of a popular slant and will help attract more people. More news in time; check out the website for now.

**Caption:** more great work from Paul Glynn, this time a Jurassic diorama featuring the European tetanuran *Megalosaurus*. Paul first started showing dioramas at TetZooCon in 2019 and has had stalls at the more recent two or three of them. Image: Darren Naish.

  **Caption:** seeing as it was the last one, it was only right that we got some photos of John and I together. Despite everything, we’re still on mostly friendly terms, evidently. Images: Georgia Witton-Maclean, used with permission.

And then that was it. There was a final flurry at the stalls, a final restaurant trip, and a final pub trip. Ordinarily, here’s where I, too, would join in and be social. But not this time. For you see, I would need to be ready early the following morning for Day 1 of... the TetZooTour.

More on that later, but for now massive thanks to my co-organizers John Conway and Chris Manias, to Jenny, Chantelle and Diallo for staffing the desks and being brilliant, to Georgia Witton-Maclean for being our official photographer, to Steve White for stall management and to all the stall holders, to all the speakers, presenters, panellists and event leaders, to Hel for powerpoint help, to Toni for merch design, to Everything Dinosaur for donations, to Tony Pinto and Ellinor for help with the Why Dinosaurs? screening, and finally to everyone who bought tickets, showed up and made it what it was. That was TetZooCon 2024!

**Caption:** I’m generally first one in, last one out at these meetings. Saying goodbye to an empty hall is a slightly melancholy thing, but here we are. This is not the end. Image: Darren Naish.

A few TetZooCon 2024 reports have already appeared online…

TetZoo 2024. The Show Must Go On, by Luis Rey at the Luis V. Rey Blog
TetZooCon 2024, by Albertonykus at Raptormaniacs

For previous articles on TetZooMCon and TetZooCon, see…

The events of TetZooCon 2014, July 2014
TetZooCon 2015 Is On, July 2015
The Events of TetZooCon 2015, November 2015
Coming Soon: TetZooCon 2016, September 2016
The Day After TetZooCon, October 2016
The Fourth TetZooCon, September 2017
The TetZooCon of 2017, October 2017
Reasons to Attend TetZooCon 2018, September 2018
TetZooCon 2018: Best TetZooCon So Far, October 2018
Announcing TetZooCon 2019 – the Biggest Yet, August 2019
Final Call For TetZooCon 2019, October 2019
The Sixth TetZooCon, October 2019
TetZooCon 2020 + Zoom = TETZOOMCON 2020, December 2020
TetZooMCon 2020 an Unbridled Success, December 2020
Cronch Cats, Beasts of Gévaudan, Dinosauroids, Mesozoic Art and Much More: TetZooMCon 2021 in Review, September 2021
The Ninth and Largest of the Tetrapod Zoology Conventions, December 2022
Announcing the 10th Tetrapod Zoology Convention, October 2023
At TetZooCon 2023, November 2023
TetZooCon 2023 the report, December 2023

If you enjoyed this article and want to assist me in what I do — and see behind-the-scenes stuff in development — please support me at patreon. Click anywhere on this text in bold.

Refs - -

Herculano-Houzel, S. 2022. Theropod dinosaurs had primate-like numbers of telencephalic neurons. Journal of Comparative Neurology 531 962-974.

Hone, D. 2024. Uncovering Dinosaur Behaviour: What They Did and How We Know. Princeton University Press, Princeton and Oxford.

Ijouiher, J. 2022. The Desert Bones: the Paleontology and Paleoecology of Mid-Cretaceus North Africa. Indiana University Press, Bloomington, Indiana.

Lawrence, N. 2024. Enchanted Creatures: Our Monsters and Their Meanings. Widenfeld & Nicolson, London.

Meiburg, J. 2021. A Most Remarkable Creature: the Hidden Life and Epic Journey of the World’s Smartest Birds of Prey. Alfred A. Knopf, New York.

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Once again it’s time to continue with my slow-burn zoo review series. I’ve just returned from a trip to Tokyo, you see, and while there I visited two zoos. Today we look at the first of them: Ueno Zoological Gardens (usually just called Ueno Zoo), located in Ueno Park in Taito City, central Toyko…

**Caption:** some zoos have awesome, attractive entrances. Ueno Zoo... maybe not so much. Having said that, this isn't *the* main entrance, but the alternative front entrance to the west. Image: Darren Naish.

Ueno Zoo – generally regarded as Japan’s flagship zoological collection – is similar in size to London Zoo (about 35 acres) and is old as zoos go, having been founded in 1882. It started out as a menagerie connected to the National Museum of Natural History, built on land owned by the imperial family; it transitioned to government ownership in 1924. Over its long history, the zoo has been notable in its breeding of Giant pandas Ailuropoda melanoleuca (which have been at the zoo since 1972), Aye-aye Daubentonia madagascariensis and Pygmy hippo Choeropsis liberiensis, but like other old zoos worldwide it suffers from the fact that many of its enclosures are old-fashioned and not of satisfactory size for the animals, especially the big ones. Having said that, I didn’t see any indication that animals of any sort were poorly cared for, and indeed it was obvious that modernisation had occurred where space allowed. I will avoid talking about the zoo’s history during WWII because… oh boy, it’s not a pretty story.

Anyway… the south-west quadrant of the zoo is occupied by the enormous Shinobazu Pond, much of which was covered by lotus at the time of our visit. Islands at the edges of the pond – some connected to the mainland by walkways – are home to lemurs (more on that later) as well as to pelicans and other birds.

**Caption:** Shinobazu Pond as seen from Aesop Bridge, looking west. Obviously, it was over-run with lotus at the time of our visit. A Great egret *Ardea alba* is visible in the middle of shot but other wild birds were present in the pond at the same time. Image: Darren Naish.

  **Caption:** I was impressed with the signage at Ueno Zoo, and this extended to discussion of wild animals occurring in the grounds. This large sign at the edge of Shinobazu Pond points out which bird species are likely to be observed. Image: Darren Naish.

For reasons of location and history, Ueno Zoo today exists on either side of a large road (Dobutsen dori Street) and is thus split in two, its halves being termed the West Garden and East Garden. Visitors have to cross a high bridge (Aesop Bridge) to get from the western half to the eastern one. In the discussion that follows here, I’m going to talk about exhibits and their animals as I encountered them on my walk through the zoo, rather than in phylogenetic order or anything like that.

**Caption:** zoos should, where possible, have sections that look like 'natural' environments, meaning that we need plants, rock faces and natural waterways. I'm not pretending that this is anything other than simulated, but we're simple animals and the simulation is good enough to work. This rock garden is located close to Tiger Forest. Warning signs noted that Japanese honeybees were nesting in this area. Image: Darren Naish.

On entering Benten Gate at the far south of the West Garden (Shinobazu Pond being to your left), you immediately pass a children’s zoo with domestic rabbits, guinea-pigs and so on. I and my companions entered via the West Garden, but the main entrance is actually in the East Garden. The pond is used by numerous locally occurring wildfowl and other waterbirds and good signage alerts you to what species you might see. Only Eastern spot billed duck Anas zonorhyncha, Great egret and Great cormorant Phalacrocorax carbo were visible during my visit, though the massive amount of vegetation covering the pond meant that ducks, grebes and rails would have been mostly invisible.

**Caption**: select animals from the small house near Benten Gate. Left to right: Asian swamp eel *Monopterus albus* (swamp eels aren't eels at all but a very odd group of acanthomorphs), Reeve’s turtle *Mauremys reevesii* and Chinese softshell turtle *Pelodiscus sinensis*. Captive softshells are often perpetually moving and thus hard to photograph well; this juvenile (carapace length about 8 cm; in an adult it's over 30 cm) was co-operative enough to pause and stick its head and neck out of the water at least once. Images: Darren Naish.

A small educational facility nearby houses tanks holding swamp eel, Reeve’s turtle, Chinese softshell turtle and various impressive insects, including mantids. Several birds are also on show in this part of the zoo, including Emu Dromaius novaehollandiae, Japanese night heron Gorsachius goisagi, Oriental white stork Ciconia boyciana and – on the far side of the pond – Cackling geese Branta hutchinsii.

**Caption:** an avian montage. At left, Emu (the pixely look isn't because of a camera malfunction but because of the screen-like mesh around the enclosure). Upper right: Japanese night heron, one of two extant *Gorsachius* species (and which are no longer included in *Nycticorax*). Lower right: a view through two different enclosures at the same time, showing Cackling goose at left, crane in the middle, and Oriental white stork at right. Images: Darren Naish.

To the pandas! Pretty soon, you approach Panda Forest, a recently constructed building complex housing Giant panda, where you have to join a queue. People are allowed into the building in groups of about 15 at a time and the staff and signage inform you that you’re not allowed to linger at one spot for more than a few minutes. The great joke about Giant pandas at zoo is that you might queue for an age only to then catch a glimpse of the top of the back of a motionless, sleeping panda that’s hardly visible behind the throng of people in front of you.

**Caption:** one of the buildings housing the zoo's two Giant panda, as seen from the middle of the queue. You enter through large, 'traditional' (maybe Edo style) wooden doors at the side. The now defunct monorail is overhead. Image: Darren Naish.

  **Caption:** Shin Shin, a female at Ueno Zoo, doing what a Giant panda does. That's mostly eating and sitting. Here's your regular reminder that the Giant panda isn't 'the panda': that honour goes to the Red or Lesser panda, the panda that became known to science and to Europeans first. Hence *Giant* panda. And, yes, it's a bear, not a close relative of actual pandas, or raccoons, as was considered correct during the mid 20th century. Images: Darren Naish.

This was very much not the case here. We only queued for about 20 minutes and the number of people allowed in meant that we could get close to the glass. The pandas were readily visible, as you can see from my photos. The zoo currently has two: Shin Shin, a female, and Ri Ri, a male. They’re bred on several occasions. Both arrived from China in 2011. I’ve seen Giant pandas at three zoos now, most notably at Chengdu Research Base of Giant Panda Breeding (aka PandaBase) in Sichuan, China. I’m of the opinion that bears in general are not built for life in captivity. They require access to massive space and constant behavioural occupation, and confinement very quickly leads to depression and psychosis. But I don’t think that this is necessarily the case for Giant pandas, mostly because they spend so much of the day sitting in the same place, eating bamboo.

**Caption:** we take it for granted that what are among the weirdest and most specialized of birds are, actually, quite easy to keep, rear and breed in captivity. Maybe this reflects how tough and adaptable flamingos are. Flamingos can live to 50 years or more in captivity, so you're likely to see the same bird(s) if you visit the same zoo more than once within your lifetime. Image: Darren Naish.

The hallowed Shoebill. Nearby, an old African, Cape or Jackass penguin Spheniscus demersus exhibit was empty of actual penguins and American flamingo Phoenicopterus ruber were on show. But the centre of the western half is dedicated to an animal that Ueno Zoo is now quite famous for: the Shoebill Balaeniceps rex. Don’t call it ‘Shoebill stork’, since it’s not a stork nor closely allied to them.

**Caption:** Shoebill 1, here photographed as it moved its head about and slowly moved its legs and feet. It did eventually walk left and into its house. Shoebills are long-legged kin of pelicans, despite old arguments that they might be storks. Images: Darren Naish.

  **Caption:** at left, a close-up of Shoebill 1. At right, the rather smaller Shoebill 3. Like all modern animals, the Shoebill is no more 'prehistoric' than you are, but of incidental interest is that some claimed 'pterodactyl' and monster bird sightings from tropical Africa might have been misinterpreted Shoebill observations. Images: Darren Naish.

  **Caption:** Shoebill 2 as seen from behind, or obliquely so. Note the way the feathers grow horizontally backwards to form a midline dorsal ridge beneath the crest, and how the pupil is still visible even from this angle (meaning that the bird potentially has a field of vision well exceeding 180 degrees). Images: Darren Naish.

  **Caption:** Shoebill 2, a large individual that spent a bit of time moving its head around while I was watching it. Shoebills can reach 1.5 m in standing height and have a wingspan of 2.6 m. A big male might weigh over 5 kg. Images: Darren Naish.

The zoo houses three Shoebills and all were showing during the time of my visit. They weren’t doing much apart from standing around and occasionally taking the odd step, but one did shake its head from side to side in an interesting manner at one point. I don’t think I’ve ever seen live Shoebills before and I took numerous photos of all three, sometimes from angles I haven’t seen before (have you ever actually looked at the back of a Shoebill’s head?). The Shoebill enclosures are spacious and well planted, but don’t include the sort of massive papyrus stands that Shoebills favour in the wild. If they did, I don’t think that we’d get to see much of the birds.

**Caption:** the enclosure belonging to Shoebill 1. I think it looks pretty good, and is appropriately spacious for the animal. It's said of raptors that they'll happily sit on a perch for 20 hours a day and be apparently happy so long as they get those short bouts of flying and eating. I think that the same might be true of quite a few birds. Not parrots, not corvids. Image: Darren Naish.

Ueno Zoo knows what a big deal Shoebills are, and numerous Shoebill-related products were on sale. More on that matter later. Shoebills are officially listed as ‘Vulnerable’ in terms of conservation status and things are worrying: there are likely less than 5000 individuals in existence and a long list of issues affect them, including illegal trade, hunting, human disturbance, habitat deterioration and drought.

**Caption:** none of the Shoebills were especially close to the fences when I was at the zoo, but I can buy that they might have a decent bite. I like that little cartoon. Image: Darren Naish.

At the time of writing, I don’t know what the deal is with respect to captive breeding but I do know that a nest protection and reintroduction programme exists in the Bangweulu Wetlands of Zambia. Shoebills produce two eggs but one of the chicks always dies or is killed by its sibling, so a scheme exists whereby the ‘waste’ chick is rescued, raised to fledging, and released. Read about this scheme and adopt a chick yourself here!

**Caption:** nice art on the walls of the Small Mammal House. Bats (vesper bats of some sort, I think) and a tree squirrel.... Images: Darren Naish.

  **Caption:** ..... and flying squirrels in action! I'm not sure if the zoo currently has flying squirrels of any sort, but this wasn't the only flying squirrel imagery at the Small Mammal House, so I'm pretty sure they've had them in the past at least. Image: Darren Naish.

‘Small Mammals’: marsupials, aardvarks, echindas, shrews and more. Moving now away from the Giant pandas, Red pandas Ailurus fulgens – the actual pandas – are on show too, and are adjacent to a very impressive Small Mammal House. I like bass reliefs and other aspects of zoo architecture that reflect a zoo’s history and occupants, and this building is one of the best examples, its outsides being decorated with grand artwork depicting bats, squirrels and fishing owls.

**Caption:** Common marmoset at left, Naked mole-rat exhibit at right. There's some material in the Tet Zoo archives on mole-rats (naked and otherwise) but good luck finding a non-ruined version. Naked mole-rats were conventionally argued to be the most specialized (and, presumably, geologically youngest) of mole-rats but fossil and molecular data shows them to be one of the oldest lineages within the group (apparently diverging from other mole-rats in the Late Eocene, over 34 million years ago). Images: Darren Naish.

Indoors, a daylight section included Common marmoset Callithrix jacchus, a number of Pallas’s cat Otocolobus manul (there’s total inconsistency on whether its name should be written Pallas’s, Pallas’ or Pallas; my favourite solution is to give up and use the Kyrgyz name Manul), a Naked mole-rat Heterocephalus glaber colony (not a first for me, since I’ve also seen them in Oregon Zoo), what I think were Common degu Octodon degus, and Southern three-banded armadillo Tolypeutes matacus.

**Caption:** a Pallas’s cat montage, showing playful and highly active kittens as well as at least one of their more sedate parents (the animal sat on the box). Three kittens were born during April this year, so we were lucky to see them at this stage of life. Images: Darren Naish.

  **Caption:** a relaxing degu group. I'm assuming – I hope correctly – that these are Common degu, one of five species within the genus *Octodon*. These are mostly animals of Chile, but Ricardo Ojeda's degu *O. ricardojeda* (only recognized as a distinct species in 2020) has a range that extends into western Argentina as well. Image: Darren Naish.

More interesting was the night-time section of the house. Low, red tint lighting and my lack of a good low-light camera, combined with constant movement from the animals themselves, means that my photos are poor to terrible, but there were a great many animals here that I was very happy to see, mostly for the first time. They include Asian house shrew Suncus murinus (surprisingly big if you’ve only seen ordinary, European shrew species before), Short-beaked echidna Tachyglossus aculeatus, Six-banded armadillo Euphractus sexcinctus, Aardvark Orycteropus afer, Woylie or Brush-tailed rat kangaroo Bettongia penicillata, Greater Egyptian jerboa Jaculus orientalis, Prince Demidoff’s bushbaby Galagoides demidoff (though still included in the old, inclusive version of the genus Galago according to the signage) and Spectral tarsier Tarsius tarsier. That’s an impressive collection, and of note is that all the animals here were out and about, and active.

**Caption:** smaller mammals from the nocturnal section of the Small Mammal House, House shrew at left and Brush-tailed rat kangaroo, Woylie, or Brush-tailed bettong at right. There can't be many zoos that have shrews of any sort on show. Images: Darren Naish.

  **Captions:** at least three aardvarks were on show, all in two connected rooms. I have to say that this was a bland, clinical exhibit. Aardvarks are highly variable across their range and multiple subspecies have been recognized historically. Images: Darren Naish.

  **Caption:** more badly photographed mammals from the nocturnal section of the Small Mammal House, namely Greater Egyptian jerboa and very blurry Short-beaked echidna. The echidna was constantly on the move and this is the best of my photos, ha. Images: Darren Naish.

African megamammals. Moving now to the far north-west, the western edge of the zoo is home to hippos of both species as well as Eastern black rhino Diceros bicornis michaeli (pretty unusual to have Black rhinos in a zoo identified to subspecies). Rothschild’s giraffe Giraffa camelopardalis reticulata / G. reticulata was present too. A noteworthy aspect of the zoo’s history with respect to giraffes is that it was home to two spotless giraffes, one called Ryoko and one Toshiko, born in 1967 and 1972. I didn’t get to see the hippos for time reasons. Jiro, a bull Common hippo Hippopotamus amphibius, died at the zoo in 2022 at a very respectable 38 years old. An Okapi Okapia johnstoni was formerly on show next to the giraffes but died in 2023 and the enclosure was empty at the time of my visit.

**Caption:** Ueno Zoo currently has two Black rhinos, Argo the female (here at left) and Maro the male. Maro has lots of black staining around the top and sides of his head; I don't know what this represents but wonder if it comes from rubbing his head against objects in the enclosure. The kind of wear that rhinos have on their horns is also interesting and says lots about their behaviour and personal habits. Images: Darren Naish.

  **Caption:** more views of Maro the male Black rhino. He was born in 2000; Argo was born in 1995. Captive black rhinos live into their fourth, fifth or sixth decades (the oldest on record died at age 57), well beyond the 18 considered average for wild ones. Images: Darren Naish.

While all of the enclosures for these animals looked well designed, with pools, planted areas, trees and rocky faces, they were – as mentioned earlier – very much on the small size given the size of their denizens. I didn’t get the impression that the animals were morose or especially bored (if anything, the contrary), but it did definitely seem that their spaces were not sufficiently big.

**Caption:** Argo the rhino in her enclosure. The pool in the foreground is empty; that in Maro's enclosure was not and he spent some time walking in and out and sloshing around in the water. Image: Darren Naish.

I was excited to learn that a reptile house – labelled Vivarium – is present nearby… but it was closed! The building did have rather brutalist overtones, this perhaps meant to superficially convey the rough form of its tenants (it’s not old, having been built in 1999). A Komodo dragon and giant tortoise statue were nearby.

**Caption:** the vivarium, though sadly closed. Frustrating, as I'm pretty sure it would have housed reptile and amphibian species unfamiliar to a visitor of European zoos. Image: Darren Naish.

  **Caption:** imagery on the outside of the vivarium, I think giving some idea of what should be in there. I haven't tried to work out what species of crocodile is shown at left, but I think it's a Nile croc *Crocodylus niloticus*. A giant salamander is at lower right. Image: Darren Naish.

  **Caption:** the life-sized Komodo dragon statue near the Vivarium. It has some issues with its forelimbs but is fun nonetheless. Tokyo is sufficiently hot and sunny in the summer that metal statues can become very warm to the touch, hence the 'don't touch' sign on the cone. Then again, this can happen even here in the UK: the Pygmy hippo statue at my local zoo (Marwell) can get hot in the sun as well. Image: Darren Naish.

Madagascar ahoy. Finally for the West Garden – and taking us back to the edge of Shinobazu Pond – we come to Aye-aye Forest, a section dedicated to the endemic wildlife of Madagascar. Enclosures (some on islands, as mentioned earlier) are home to Aye-aye, Black-and-white ruffed lemur Varecia variegata, Black lemur Eulemur macaco and Ring-tailed lemur Lemur catta. Radiated tortoise Astrochelys radiata are present too. I understand that a Fossa Cryptoprocta ferox was in the collection until recently (perhaps this year), but has died.

**Caption:** Radiated tortoise at left, male Black lemur at right. The Radiated tortoise is mid-sized (carapace length is typically around 30 cm) and individuals have exceeded 180 years in age. Black lemurs are unique to Madagascar's north-west (and adjacent islands) and are famously dimorphic; females are brown or orange-brown with white ear tufts. Images: Darren Naish.

  **Caption:** Shinobazu Pond again, this time showing part of the Madagascar exhibit at its eastern edge, hence the model baobab. Ring-tailed lemurs are present on the island we see here, but I don't think you can see them in the photo. Image: Darren Naish.

What impressed me most about the exhibit was its inclusion of features relevant to Madagascar’s unique biological heritage and history. A life-sized model of a baobab (albeit not a very big one) stands on the island with the Ring-tailed lemurs, and a slightly oversized bronze statue of an aepyornithid – ‘elephant bird’ or vorompatra – and its massive egg stands proud nearby. Animal statues are good, I always appreciate them.

**Caption:** the zoo's gigantic aepyornithid statue as seen from several views. I like it, though I'm not sure about that bushy head crest. This a fascinating group of birds and we still know so little about them. Images: Darren Naish/Em Naish.

Also nice were two very artful wooden pillars on either side of the entrance to the Madagacar section. As you can see, they feature Malagasy endemics (chameleons, lemurs, tenrecs) but also depict the location of Madagascar within Gondwana and show a symbolic lemur rafting across the Mozambique Channel from the African mainland. That’s a lot of information to convey via carved wooden posts, and what a great job!

**Caption:** educational, but also artistic, Madagascar-themed wooden poles! Images: Darren Naish.

  **Caption:** more installations from the Madagascar section, showcasing the zoo's history of keeping and breeding lemurs. I grew up imagining aye-ayes as squirrel-sized (meaning about 30 cm in total) so was shocked when I first saw a live one (this was at Bristol Zoo): in total length, they're around 80 cm. Images: Darren Naish.

To the east. Late in the day, we walked across Aesop Bridge and entered the zoo’s East Garden. It’s about similar in size to the western half and contains a greater assortment of larger animals, including bears, big cats, elephants and primates. A massive, jagged mock mountain is the centrepiece of a Japanese macaque Macaca fuscata exhibit. Thanks to the good signage, I learnt that this mountain was built in 1932, making it one of the oldest structures in the zoo, and that zoos with similar features were inspired by this one. In addition, the specific Japanese macaques here are from the Shimokita Peninsula of Honshū and are thus the most northerly occurring of all non-human primate species.

**Caption:** the majority of the zoo's Japanese macaques spend a lot of their daytime sitting and walking on the sides of their enclosure's artificial mountain, but some individuals can be seen in the moat surrounding it too. Not shown in the photos here are the concrete pools at the side of the mountain. I reckon that someone decided to include heated pools so that the monkeys might choose to sit in the water, as is so famously seen in the Jigokudani 'snow monkey' population. Alas, that's a learned bit of culture specific to that population, not a species-wide bit of behaviour. Images: Darren Naish.

Asian elephants Elephas maximus are housed in Elephant Forest nearby. Due to a number of constraints – timing, the extreme heat at the time of our visit, our initial misunderstanding of the zoo’s layout, and slow walking on my part due to an injury – there’s quite a bit of the zoo that we didn’t get to before it started to shut down at 4pm (zoos in Japan close off certain parts of attractions about an hour before final closing, it seems).

**Caption:** the Asian elephant house and enclosure are ok, but nothing to write home about. You should be able to make out the trunk, and part of the head and body of the elephant in the house at left. Images: Darren Naish.

As a result, I never got to the Polar Bear and Seal Oceans section or Bird House in the northern part of the East Garden, nor to the section in the south-west devoted to Japanese animals, or the tropical monkeys, bison and prairie-dogs nearby. It seems that I missed a lot: I wasn’t able to get to the area where the bears were, but the zoo’s website lists an amazing four species being on show as of this year (Polar bear, Hokkaido brown bear, Sun bear, and Japanese black bear).

The eastern edge of the East Garden is occupied by Gorilla Woods and Tiger Forest. I didn’t get to the gorillas before the exhibit closed but did see a tiger. I wondered what sort of tiger the zoo might have, given that the majority on show at home in the UK are Siberian Pantheria tigris altaica. It turned out to be a Sumatran tiger P. t. sumatrensis (I later saw a Siberian tiger during my visit to Tama Zoological Park, more on that in future).

**Caption:** I photographed two captive tigers while in Japan (the other one was at Tama Zoological Gardens), and both made direct eye contact with me, or with my camera at least. In the UK, tigers and other big cats seem to avoid eye contact with cameras... or, that's my impression anyway. This makes me wonder if the zoo tigers of Japan don't have people pointing cameras at them so often. Or am I completely off base here? Image: Darren Naish.

On show nearby are Eurasian otter Lutra lutra, Edward’s pheasant Lophura edwardsi, Australian brushturkey Alectura lathami, Golden-breasted starling Lamprotornis regius, Red-headed wood pigeon Columba janthina, Leschenault’s rousette Rousettus leschenaultii and Brazilian or Lowland tapir Tapirus terrestris. Dhole Cuon alpinus are on show too, or were until recent years.

A notable feature of the zoo, encountered at this point, is the Animal Cenotaph, a monument dedicated to animals that have died at the zoo over its history. Signage explained how a monument was initially built in 1931 close to Gorilla Woods and Tiger Forest but that the present one was constructed in 1975. I also saw an animal memorial in a later visit to Tama Zoological Park and wonder if this is a normal feature of eastern zoological parks. It’s a nice and touching feature that gives pause for thought. Remember that many animals kept in captivity – of all sorts, from lizards and molluscs and fish to charismatic big lizards, birds and mammals – become ‘known’ individuals that keepers and visitors form bond with.

**Caption:** the zoo's modern animal cenotaph. Note the fresh flowers at left and the well-kept condition of the hedges. Image: Darren Naish.

Raptors, owls, cranes. A large aviary section on the south-east edge of the zoo was mostly devoted to owls and raptors, and I spent some time here as they had some spectacular animals on show. They included several Indian white-backed vulture Gyps bengalensis, Andean condor Vultur gryphus and Steller’s sea eagle Haliaeetus pelagicus of both sexes, Harris hawk Parabuteo unicinctus and Mountain hawk-eagle Nisaetus nipalensis. Owls on show included Japanese scops Otus semitorques, Snowy owl Bubo scandiaca and Ural owl Strix uralensis.

**Caption:** at left, White-backed vulture with what looks like a full crop. At right, Mountain hawk-eagle. Old World hawk-eagles like this were included in same genus as the American *Spizaetus* species until around 2005 but molecular data has shown that they don't belong in the same clade, instead being close to the Black eagle *Ictinaetus malaiensis*. Images: Darren Naish.

  **Caption:** spectacular female Steller's sea eagle, a large raptor of north-east Asia's Pacific coastline. A big female can have a wingspan of 2.5 m and weigh 9.5 kg, making this species generally bigger than the Harpy *Harpia harpyja*. Images: Darren Naish.

  **Caption:** male Andean Condor, at left, and Harris hawk at right. Condors are incredible, and yet another bird group where so many parts of their anatomy have been co-opted for display and communication. The Harris hawk is a familiar bird to Europeans and Americans, but maybe it feels more exotic if you live in eastern Asia. Images: Darren Naish.

  **Caption:** a view of the raptor and owl aviary section. A number of vultures were kept together in the big enclosure on the left; owls, eagles, hawks and condors were on the right. Image: Darren Naish.

Also impressive was the crane display, which featured three species in adjacent enclosures (so, more cranes together than I think I’ve seen before): Wattled crane Bugeranus carunculatus, Black-necked crane Grus nigricollis and Red-crowned crane G. japonensis. A Secretary bird Sagittarius serpentarius pair were also here and I got to see them being fed. Southern bald ibis Geronticus calvus were present nearby, as were Hamerkop Scopus umbretta but they weren’t showing. I should add here that some – perhaps many – birds kept by the zoo weren’t seen on my visit, either because they were in aviaries that I didn’t get to, or because they were off-show due to current concerns about H5N1 bird flu.

**Caption:** a crane montage, showing (clockwise from top left) Black-necked crane, Red-crowned crane (in side view and when looking right at me), and Wattled crane. Some cranes have really gone to town on the developing of epidermal lumps and nodules: the red crown of a Red-crowned crane is formed of pointed, red papillae, not feathers as you might guess. Images: Darren Naish.

  **Caption:** a possibly disturbing photo of a Secretary bird scoffing down a chick, whole. Of interest is the timing of nictitating membrane moment and that the rictus (the web of tissue at the corner of the mouth) is similar in colour to the face, not a wholly separate and differently pigmented sheet of tissue as we tend to expect. Image: Darren Naish.

And that about wraps up all my animal viewings of the zoo. What about those aspects of the zoo that don’t concern the animals on show? An important thing (for European tourists at least) in the Tokyo region concerns access to shade and cold drinks, because boy was it warm and humid during our time there. The zoo was great on these things, there being numerous rest stops and vending machines. We also found the restaurant to be conveniently placed, as well as good, efficient and reasonably priced.

Signs, and statues and statues. Signage at the zoo was very good. All enclosures were well labelled, and big, attractive panels and display boards did a good job of informing you about what was on show. An important feature of zoos for me is the general look of the place, and the buildings, artwork and installations present throughout the zoo grounds. Ueno Zoo does well on those fronts, being attractive and very well landscaped throughout. The walking disability that affected me during my visit made me acutely aware of stairs and steep slopes, and I’m pleased to say that Ueno was good and convenient in terms of access.

**Caption:** another indication of how good the signage is at the zoo. I consider this pretty impressive as an effort to educate the public (though getting them to actually read it is a different matter). Our fascination with the enigmatic Giant panda has resulted in numerous public descriptions of its skull and dentition, and of its famous 'sixth finger' or 'thumb' (actually a modified pisiform bone adjacent to the thumb). Image: Darren Naish.

I also like statues at zoos. If they’re not especially accurate in terms of anatomy or proportions… well, that’s ok because they’re abstract. If they are accurate… well, that’s great too because they’ve succeeded in depicting the form or feel or size of the subject. The sculptures at Ueno were all great: there were the Komodo dragon, giant tortoise and vorompatra already mentioned, but others I saw included an Asian elephant and one devoted to a famous, and famously endangered, east Asian species specially associated with Japan: the Crested ibis Nipponia nippon.

**Caption:** the Crested or Japanese ibis is a conservation icon. In Japan, the last wild bird died in 2003 following gradual decline caused by overhunting, habitat loss and other causes. Wild populations persist in Shaanxi Province, China; something only appreciated in 1981. Captive breeding has since resulted in the wild release of birds in Japan and South Korea too. Images: Darren Naish.

On shops and things to buy. Finally… here in the UK I am constantly disappointed by the state of zoo shops. They have a few souvenirs that adults might be interested in – like hats, towels and fridge magnets – but otherwise they don’t cater for anyone seriously interested in zoos, in animals, or in natural history.

**Caption:** it was obvious from signage, products and things on sale that Ueno Zoo was proud of its Shoebills. I resisted the urge to buy a cuddly one (though I did buy a plush Malayan tapir; how could I not). Images: Darren Naish.

  **Caption:** your zoo has PALLAS'S CATS so, surely, you stock and sell Pallas's cat merch, toys, souvenirs and so on, right? In Japan, the answer is a confident YES. Image: Darren Naish.

Well, Japan has very much the opposite problem. They know exactly what they’re doing and have phenomenally good stuff in their shops, catering very much to an interested adult audience. A zoo that features an especially remarkable and charismatic animal – like Shoebill or Manul or Tasmanian devil – will have a whole section devoted to merch and products featuring that animal, it’s great. High-quality toys, models and figures are present, and (as a collector of such) I was simply thrilled at what was available. Kids can buy and enjoy these things, for sure, but they’re not for kids only.

**Caption:** as a dedicated and pretty serious collector of animal figures, Japan was *lethal*. All the animal-themed attractions I went to stocked excellent selections of animal toys and models. The larger figures at the bottom are by Safari and Schleich, the smaller ones that come in the rectangular boxes are by Colorata. Image: Darren Naish.

And that is where we end my look at Ueno Zoo, I hope you enjoyed it. As ever, we finish with my wholly subjective scoring system…

Selection of species: 7 out of 10
Zoo nerd highlights: Giant panda, Shoebill, Short-beaked echidna, House shrew, Aye-aye, Naked mole-rat, selection of cranes together
Quality of signage: 9 out of 10
Value for money: 9 out of 10
Overall worthiness: 8 out of 10

And for previous articles in my zoo reviews series, see…

Tet Zoo Reviews Zoos: Edinburgh Zoo, August 2016
Tet Zoo Reviews Zoos: Colchester Zoo, October 2017
TetZoo Reviews Zoos: the Isle of Wight Zoo, February 2020
TetZoo Reviews Zoos: ZSL Whipsnade Zoo, June 2021
TetZoo Reviews Zoos: Bristol Zoo, July 2021
TetZoo Reviews Zoos: Tripoli Zoo in Libya, January 2023
Tet Zoo Reviews Zoos: New Forest Wildlife Park, April 2024

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It's time once again to look at squamates…

**Caption:** I've mentioned before how inspirational the wonderful illustrations of Alan Male (from Philip Whitfield’s *Reptiles and Amphibians: An Authoritative and Illustrated Guide* of 1983) were to a young me, and here are the several cordylids he illustrated. It's sometimes obvious that Male only had access to photos, and photos that weren't especially useful at that, and hence made mistakes (the head of the *Chamaesaura* here is inaccurately short, for example). Images: Alan Male, from Whitfield (1983).

That’s snakes and lizards… though, given that snakes are a specialized group of lizards, maybe I should stop using ‘snakes and lizards’ as a descriptor. In fact, maybe we should get into the habit of calling all squamates ‘lizards’ and hope eventually that tree-based thinking wins the day. Ha, yeah right.

Here, we’re going to embark on a whistlestop tour of the cordylids, or girdled lizards: a diurnal, endemic African group, mostly associated with dry rocky places and hilly slopes, and well known for the armour-plated, sometimes spiky, covering of many of its species. The use of burrows and crevices is common across the group, and the majority are viviparous, giving birth to between one and twelve babies. These animals are generally between about 12 and 40 cm long in total.

**Caption:** some of the books I consulted while compiling this article. Even in the age of the internet, books are essential and you still have to do everything you can to accrue them. Remind me why I have no money. Image: Darren Naish.

Within squamate phylogeny as a whole, cordylids appear part of Scincoidea, the large clade that also includes the hyperdiverse skinks, night lizards (Xantusiidae) and plated lizards (Gerrhosauridae). To use the most general generalization possible, scincoids are mostly smooth-scaled, insectivorous, ancestrally terrestrial lizards that often have small bony plates (osteoderms) embedded in the skin and use their well-clawed limbs to clamber, run and dig. Within Scincoidea, both anatomical similarities and molecular data show that cordylids and gerrhosaurids are close, the two forming the clade Cordyliformes (Lang 1991). An old-fashioned view is that gerrhosaurids should be subsumed into Cordylidae. That’s a bad idea (come on, they’re easily ‘distinct enough’ to warrant ‘family’ status) and it’s not currently in fashion.

How, then, to discuss this moderately (but not especially) speciose group, about 70 extant species of which are currently recognized? Our coverage of cordylids here will revolve around taxonomy, for reasons that’ll be obvious once we get into it. For girdled lizards, you see, were, until recently, grouped into four genera: the extremely flat-bodied flat lizards Platysaurus, the flattish crag lizards Pseudocordylus, the deeper-bodied, often spiny, girdled lizards, zonures or sungazers Cordylus and the serpentiform grass lizards, snake lizards or ground lizards Chamaesaura. Let’s get to it, starting with flat lizards.

**Caption:** the ‘original’ girdled lizard… the Cape girdled lizard *Cordylus cordylus*, type species of the genus *Cordylus* and thus the groups Cordylinae and Cordylidae. This individual was photographed in Stony Point Nature Reserve, Western Cape, South Africa. Image: Bernard Dupont, CC BY-SA 2.0 (**original here**).

Flat lizards are very special. As might be obvious, flat lizards are extremely flattened, both in the head and body and also the thighs and tail base. 16 species are currently recognized, the most recently named of which is Attenborough’s flat lizard P. attenboroughi Whiting et al., 2015 of southern Namibia (yup, species number 42 or something named after Sir David).

Bright colours and bold patterns are present across Platysaurus species, with blues, oranges, yellows and reds being present on the head, flanks, limbs and especially the undersides of the head and belly. The lizards flash these ventral markings by tilting or lifting themselves, and their ventral distribution of course means that they aren’t obvious at all times. These colours are typical of adult males; females across most species are generally blackish-brown with three white longitudinal stripes. However, the females in some species are brightly coloured too.

**Caption:** a montage that gives some idea of how variably coloured and patterned, and how spectacular, some flat lizards are. From left to right, we’re seeing a Common flat lizard *P. intermedius* in Zimbabwe, a Broadley’s flat lizard *P. broadleyi* in Augrabies National Park, South Africa, and an Imperial flat lizard *P. imperator* in Zimbabwe. These three species represent three of the four main clades within *Platysaurus*. Images: Sputniktilt, CC BY-SA 3.0 (**original here**); Bernard Dupont, CC BY-SA 2.0 (**original here**); © i\_c\_riddell, CC BY 4.0 (**original here**).

Flat lizards are mostly insectivorous, and climb, leap and chase in their efforts to catch flies, butterflies and other insects. They’ve even been seen to dive into pools and grab insects from below the water (Alexander & Marais 2007). Some amount of herbivory and frugivory is present too, some species eating flowers, leaves, seeds, berries and – in the Augrabies flat lizard P. broadleyi – figs (Pianka & Vitt 2003, Alexander & Marais 2007). The lizards mostly eat ripe figs on the ground but will also climb intro trees to get them, and will “use birds as cues to find fruiting fig trees” (Pianka & Vitt 2003, p. 229). P. broadleyi, incidentally, is another recently named species (it was only scientifically named in 1997, having previously been included within the Cape or Namaqua flat lizard P. capensis). Herbivory in lizards is generally associated with large size, but it appears that these lizards break that rule.

**Caption:** flat lizard be flat. This view of a Broadley’s flat lizard gives an excellent indication of what platysaurine proportions are like. Numerous individuals of this species are readily visible in Augrabies National Park in South Africa where they can be observed leaping to catch black flies. Image: Bernard Dupont, CC BY-SA 2.0 (**original here**).

We’ve seen already that cordylids are overwhelmingly viviparous. Platysaurus is oviparous, females producing two elongate, soft-shelled eggs. The remarkable flat bodies of these lizards have apparently resulted in the evolution of low clutch size and unusual egg shape (Pianka & Vitt 2003). Flat lizards are highly gregarious, tens of individuals typically feeding and hanging out in the same rocky areas, and sharing the same crevices after sundown. Their nesting is often communal too.

Wherefore art though, Platysaurus? The general view across much of the 20th century was that flat lizards are nestled somewhere in among the cordylid assemblage, in which case their lack of both osteoderms and spininess would be advanced traits. This hypothesized phylogenetic position (Lang 1991) also required that ovipary re-evolved in these lizards from viviparous ancestors… which is unlikely albeit not impossible (I’ve discussed this phenomenon a few times before, most recently in my 2020 article on the reproductive habits of Mesozoic mammals).

**Caption:** this is cordyliform phylogeny as hypothesized by Lang (1991). Note that the serpentine chamaesaurs are outside of Cordylinae, so that the primary split within Cordylidae is between Chamaesaurinae and Cordylinae. That was a 'traditional' view that had existed since the 1930s. Within Cordylinae, flat lizards are highly derived (this term has now been ruined, since some take it to mean 'very recently evolved within a group', and others to mean 'highly modified relative to the ancestral condition'... here, it can mean either or both). Molecular studies have shown these key propositions to be wrong. It should be added that Lang (1991) is a good example of a morphology-based phylogenetic study done well: Mathias Lang described and illustrated relevant character states, explained and described his methodology and approach, considered alternative hypotheses, and went to considerable trouble to obtain and observe relevant specimens. Image: Lang (1991).

The hypothesis that flat lizards – which, in some ways, seem ‘more typical’ than other cordylids – are the most modified (‘most derived’) cordylids always did seem a bit odd… couldn’t it be that things were the other way round? Molecular work published this century appears to confirm exactly this, since Platysaurus has more recently been recovered as the sister-group to the rest of Cordylidae. This, combined with anatomical distinctiveness, means that Cordylidae is now generally considered to consist of the two clades Platysaurinae and Cordylinae (Pyron et al. 2014).

**Caption:** a few cordylid anatomical characters of special interest, as illustrated by Lang (1991). At left, we're looking at the left side of the palatal surface of the skull of *Cordylus cordylus*, the unusual feature being the subdental shelf: the hollowed-out area, lingual to the teeth, and marked at its edge by the crista dentalis. The function of this shelf is unknown but it might have a role in holding food items in place while they are crushed. At right, a montage showing osteoderm distribution in (A) *Cordylus*, ventral (left) and dorsal views, (B) *Pseudocordylus*, ventral (left) and dorsal views, (C) *Platysaurus*, dorsal view and (D) *Chamaesaura*, dorsal view. All from Lang (1991).

Cordylus, the break-up. Within Cordylinae specifically, molecular work published in the 1990s and early years of the 21st century indicated that Pseudocordylus (the crag lizards) is not monophyletic, with different species being in different places among Cordylus (Frost et al. 2001). In addition, Cordylus seemed paraphyletic not just to Pseudocordylus, but to the snake-like Chamaesaura species too. One solution to this problem might be to sink Pseudocordylus and Chamaesaura into Cordylus (Frost et al. 2001, du Toit et al. 2004). A criticism of the relevant studies, however, is that they didn’t sample a sufficiently broad range of species to really get to grips with their phylogeny (and hence taxonomy).

Stanley et al. (2011) opted to sort things out via much more representative sampling. They found that several clades traditionally included within Cordylus are actually outside a clade that included Chamaesaura, Pseudocordylus and ‘core’ Cordylus (the type species is the Cape girdled lizard Co. cordylus (Linnaeus, 1758) of South Africa). A solution could have been to give up on the genus-level distinctions and just lump everything together, but Stanley et al. (2011) instead opted to devise a new taxonomy that reflected this phylogeny.

**Caption:** a fairly messy effort to depict cordylid phylogeny as recovered by Stanley *et al*. (2011), with photos depicting genus-level diversity within the group. It should be obvious immediately that *Smaug* is big and very spiky, that *Ninurta* is small and delicate, that *Chamaesaura* is serpentine, that *Pseudocordylus* lacks spines and big, plate-like scales, that *Ouroborus* curls up into a ring, that *Karusasaurus* and *Namazonurus* are broad-bodied, that *Hemicordylus* is a bit generic, and that *Cordylus* in the strict, modern sense is speciose. Images: *Smaug*: Eric Johnston, CC BY-SA 3.0 (**original here**); *Ninurta*: Amada44, public domain (**original here**); *Chamaesaura*: Alandmanson, CC BY-SA 4.0 (**original here**); *Pseudocordylus*: Bernard Dupont, CC BY-SA 2.0 (**original here**); *Ouroborus*: Handré Basson, CC BY-SA 3.0 (**original here**); *Karusasaurus*: Tony Rebelo, CC BY-SA 4.0 (**original here**); *Namazonurus*: © Tim Brammer, CC BY-NC 4.0 (**original here**); *Hemicordylus*: Abu Shawka, CC0 (**original here**); *Cordylus*: Bernard Dupont, CC BY SA-2.0 (**original here**).

Describing everything they did would take too long to explain, but the takehome is that those ‘Cordylus’ clades recovered outside of ‘core Cordylus’ required new names. So, hello Smaug, Ninurta, Ouroborus, Karusasaurus and Namazonurus, and hello again to a resurrected Hemicordylus (originally named in 1838). A ‘restricted’ version of Pseudocordylus (‘core’ Pseudocordylus = the clade that includes the type species P. montanus and its closest relatives) is now in use.

This new taxonomy makes it easiest to talk about cordyline diversity, since the members of the different clades mostly look obviously different and also do different things. It won’t be lost on you that some of these names are recognizable: Ouroborus was named for the mythical symbol whereby a dragon-like reptile is consuming its own tail, while Smaug was named for the dragon of Tolkien’s 1937 book The Hobbit. By the way, this new taxonomy corresponds quite closely to distribution and morphology (Stanley et al. 2011).

**Caption:** name an animal – any animal, I think – after a famous character from popular fiction, and you’re gonna win some coverage in the popular media. That’s exactly what happened when *Smaug* the cordylid was published in 2011. Nine *Smaug* species are currently recognized, the best known of which is the Giant girdled lizard *S. giganteus*, shown here. Images: Unwin Paperbacks; Wilfried Berns, CC BY 2.5 (**original here**).

Giant, terricolous Smaug, ring-forming Ouroborus. The Smaug species are extremely large (SVL can be 20 cm and total length 40 cm), robustly built, have an especially spiny tail, and are associated more with burrows and crevices in soil than rock (they’re terricolous, rather than rupicolous). The burrows of the Giant girdled lizard or Sungazer S. giganteus, excavated in fine soil, can be at least 1.8 m long and are sometimes home to a mother as well as her young (Branch 1988). They’re notably deep-headed relative to rupicolous cordylines.

**Caption:** a slightly sketchy effort to depict a Giant girdled lizard in life, and which depicts some of the remarkable features of this large lizard (it can reach 40 cm in total length). The tail spines are especially large, and also slightly curved, and the spines around the back of the head are especially prominent. Image: Darren Naish.

Ouroborus contains the somewhat Smaug-like Armadillo girdled lizard O. cataphractus, the spiny species famous for grabbing hold of its tail in its mouth and forming a spiky ring. While most cordylines are (apparently) solitary, this species is gregarious, with groups of up to 30 sharing the same refuge site and colonies consisting of hundreds of individuals (Alexander & Marais 2007). Loners exist though and tend to be adult males; these differ from females in reaching larger sizes and in having proportionally larger heads and longer tails (Mouton et al. 2006).

**Caption:** the Armadillo girdled lizard is another incredibly charismatic, handsome animal… its machine-perfect plates give it such an awesome look. And of course it’s famous for rolling into a defensive circle. This species has been over-collected for the pet trade in the past, though this is now illegal. Images: Kevin Murray, CC BY 4.0 (**original here**); Handré Basson, CC BY-SA 3.0 (**original here**).

The Pseudocordylus crag lizards are flat-headed, rock-dwelling animals with spiny tail whorls but a head, neck and body mostly covered in granular, plate-like or slightly convex scales. They mostly eat arthropods but small lizards and leaves, flowers and berries are eaten by some species too (Alexander & Marais 2007). They’re mostly associated with the Drakensburg and Cape Fold Mountains of South Africa, Swaziland and Lesotho, and some (like the Drakensburg crag lizard P. subviridis) live at sufficient altitude that they can sometimes be subjected to freezing temperatures. So long as they’re sufficiently deep in crevices and cavities, they avoid exposure and are ok.

**Caption:** another fine reminder of what spectacular beasts some of these cordylids are. This is a Drakensburg crag lizard *Pseudocordylus subviridis* of South Africa and Lesotho. An obvious feature of these lizards is that the head and tail possess osteoderms whereas the body and limbs do not. Populations included within this species are variable and there's some controversy over how their taxonomy should be resolved. Image: Amada44, CC BY 3.0 (**original here**).

Namazonurus, Karusasaurus, Ninurta. Moving now to more of the new genera named by Stanley et al. (2011), we come to Karusasaurus, a group of just two species that occur across South Africa and southern Namibia. The best known of them is the widely distributed Karoo girdled lizard K. polyzonus (the other is the Namibian or Jordan’s girdled lizard K. jordani). Karusasaurus species recall Smaug and Ouroborus in having spiky tail whorls but their bodies are notably broad and flattened and mostly covered in small, smooth scales arranged in transverse rows. Karoo girdled lizards are mostly mottled in brown and cream but melanistic populations occur on the cool Atlantic coast of South Africa (Broadley & Branch 2002, Mouton et al. 2002). Namazonurus is Karusasaurus-like, also relatively flat-bodied, but is evidently more closely related to Cordylus (Stanley et al. 2011, Pyron et al. 2014). Both genera (Namazonurus and Karusasaurus) share a transparent lower eyelid, a feature not present in Cordylus.

**Caption:** Karoo girdled lizard, exemplar of *Karusasaurus*. The generic name incorporates the Khoisan word *karusa*, meaning dry, barren, thirstland. Image: Tony Rebelo, CC BY-SA 4.0 (**original here**).

Ninurta contains only N. coeruleopunctatus of South African’s southern Cape region, a mid-sized (SVL 8 cm), lightly built cordylid with a flattish head. It grouped close to the Chamaesaura species in Stanley et al.’s (2011) phylogeny but a position close to Hemicordylus has been found elsewhere (Pyron et al. 2014).

**Caption:** a Blue-spotted girdled lizard, sole representative of the genus *Ninurta*. The general appearance of this lizard – it’s small, skinny, long-bodied and with small scales across its body that aren’t big and plate-like – is consistent with phylogenetic studies that find it close to the ancestry of the long-bodied *Chamaesaura* species. Image: Amada44, public domain (**original here**).

Finally, Cordylus itself – even with several lineages now removed and placed in their own genera – is still a large clade with a complex phylogenetic structure. This is the most widely distributed cordylid clade, occurring across a large part of southern and eastern Africa as far north as Ethiopia. There are dwarf species within this group (like the Rooiberg girdled lizard C. imkeae and its relatives; SVL can be 4 cm, total length 10-12 cm) and arboreal ones too (like the Tropical girdled lizard C. tropidosternum). These live in hollow trees and stumps and hide under loose bark and dead leaves.

**Caption:** a montage of species still included in *Cordylus*. At left, the strikingly attractive Black girdled lizard *Co. niger*, here photographed at Cape of Good Hope Reserve, Western Cape, South Africa. At right, Cape girdled lizard *Co. cordylus* – the ‘original’, or OG, cordylid – photographed in Mountain Zebra National Park, Eastern Cape, South Africa. Images: Bernard Dupont, CC BY SA-2.0 (originals **here** and **here**).

Snake-like cordylids! Most of us who know even a bit about squamates are aware that a snake-like form evolved multiple times within these animals. Dibamids, pygopodids, amphisbaenians, skinks (several times), anguids…. But poorly known even among specialists is that cordylids include snake-like species too, and here I’m taking about the Chamaesaura species, the five or so species of which are variously dubbed grass lizards, snake lizards or ground lizards. ‘Grass lizard’ is a not especially useful name, given that it's used around the world for various entirely different types of lizard.

These lizards are so weird compared to other cordylids – come on, the idea of snake-like cordylids is wild! – that they have, at times, been considered the sister-group to the remainder of the group (Lang 1991). They’re widespread across Africa’s southern half, occurring along the coastal strip of South Africa, throughout Angola, and (on the eastern side of the continent) from Zimbabwe and Mozambique to Rwanda, Tanzania and Kenya.

**Caption:** at left, *Chamaesaura* skulls illustrated by Lang (1991). (A) *Ch. macrolepis* in dorsal view, (B) *Ch. anguina* in right lateral view, and (C) the lower jaw of *Ch. anguina* in lingual (above) and labial views. These skulls are shallow at the snout end (as is typical for rupiculous cordylines) and more pointed (in dorsal view) than is the case in many other cordylids. At right, a range map for the *Chamaesaura* species as depicted by Lang (1991). If you’ve thought of these animals as mostly limited to the south of the continent, it’s obvious that they’ve actually staged a decent invasion of the continent’s southern half. Images: Lang (1991).

They’re mid-sized to long (with SVLs of between 14 and 17 cm, and total lengths of as much as 50 cm), extremely long-tailed (the tail might be three or four times the body length), and snake-like in movement. While they do have a ‘cordylid look’ to their integument (the body scales are sharply keeled), they differ from most other cordylids in that osteoderms are restricted to the head, a condition which has convergently arisen in Pseudocordylus. The head is also flattened, this being consistent with phylogenies that find Chamaesaura within a section of the tree where the surrounding lineages are rupicolous (Stanley et al. 2011).

The Chamaesaura species are further interesting in that they seem to represent a continuum, with one species (Ch. aenea) having five clawed digits on all four limbs, another (Ch. anguina) having only one or two clawed digits on its much reduced limbs, and a third (Ch. macrolepis) lacking forelimbs entirely, and having only a single clawed digit on its tiny, spike-like hindlimbs (Branch 1988). Patterns of variation like this are seen in some other reduced-limbed, serpentiform lizard groups (like certain skinks). Broadley & Branch (2002) noted that the Ukinga girdled lizard Co. ukingensis approaches the Chamaesaura species anatomically in having a reduced osteoderm compliment and in lacking the lateral groove that cordylines normally have along the side of the body. However, molecular work finds it to be part of Cordylus in the new, restrictive sense, and thus not phylogenetically close to Chamaesaura. Any similarities are therefore convergent.

**Caption:** the snake-like *Chamaesaura* species are often larger than I'd assumed previously (reaching 50 cm in total), and I think that that's conveyed in this photo. They aren't specialized climbers but, as shown here, are obviously capable of clambering in shrubs, bushes and grass clumps. This is the Cape grass lizard *Ch. anguina*, and specifically part of the type population *Ch. a. anguina* (the subspecies-level taxonomy of this species is controversial and some studies have raised the subspecies to species level). Image: Chris Vynbos, CC BY-SA 4.0 (original here).

As suggested by one of their common names (‘grass lizard’), the Chamaesaura species are strongly associated with grassy areas, mostly in hillside and plateau environments, and they don’t regularly use rocky retreats as their cousins do. They pursue grasshoppers, beetles and other insects in these habitats and ‘swim’ rapidly through grass via horizontal undulation. Alexander & Marais (2007) noted that their reliance on grass patches makes them vulnerable both to isolation (since grassland fragmentation and the construction of roads, walls and channels is a problem across the region), and to the loss or deterioration of these places by fire. Both issues could worsen across time, especially in a warmer, less climatically predictable world.

**Caption:** this fairly remarkable photo of a mother and baby Armadillo girdled lizard creates the impression that giving birth to a proportionally big, scale-covered baby might be a traumatic, perhaps painful event. The photo is (c) Gary Fogel and used to be present on a page dedicated to these lizards. It’s no longer findable online.

We’ve mentioned viviparity a few times now. The Chamaesaura species are no exception and can have among the largest litters – up to 12 babies – of any cordylid. Viviparity in squamates has often been linked with evolution in cool environments. We might not think of southern Africa as cool, but there are climatic models in which it was when cordylines initially diversified, this being the Oligocene (between about 35 and 25 million years ago) (Mouton & Van Wyk 1997, Stanley et al. 2011). Some studies have therefore proposed that cordylines “are a cool-adapted lineage and that viviparity developed in the family during cold conditions” (Stanley et al. 2011, p. 68).

And that’s where things must come to an end. This article is another rescued and revamped one from the archives, the original version appearing back in 2008 (a wayback machine version is here). More squamates are coming!

**Caption:** screengrabs of the original version of this article, from 2008. I managed to obtain and use some different images of *Chamaesaura* back then.

For previous Tet Zoo articles on squamates, see…

In Quest of Anguids, May 2006
Pompey and Steepo, the World-Record-Holding Champion Slow-Worms, May 2007
Cambodia: now with dibamids!, May 2011
The Tet Zoo Guide to Mastigures, August 2018
The Remarkable Basilisks, May 2023
Do Lizards Really Have ‘Mite Pockets’?, March 2024
Meeting Lake Zacapu’s Garter Snake, March 2024
Ray Hoser, Number 1 Taxonomic Vandal, May 2024
The Mysterious Dibamids, May 2024
Side-Stabbing Stiletto Snakes, Redux for 2024, May 2024
The Rehabilitation of Günther’s Black Cameroonian Snake, May 2024
Ikaheka and Other ‘Palatine Draggers’, Cryptozoic Elapid Snakes of Melanesia, June 2024
Arboreal Alligator Lizards of Mesoamerica... and Beyond!, July 2024
Grayiids: Africa Has Water Snakes Too, July 2024

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

Refs - -

Alexander, G. & Marais, J. 2007. A Guide to the Reptiles of Southern Africa. Struik Publishers, Cape Town.

Branch, B. 1988. Field Guide to the Snakes and Other Reptiles of Southern Africa. New Holland, London.

Broadley, D. G. & Branch, W. R. 2002. A review of the small east African Cordylus (Sauria: Cordylidae), with the description of a new species. African Journal of Herpetology 51, 9-34.

du Toit, A., Mouton, P. le F. N. & Flemming, A. F. 2004. Aseasonal reproduction and high fecundity in the Cape grass lizard, Cordylus anguinus, in a fire-prone habitat. Amphiphia-Reptilia 24, 471-482.

Frost, D., Janies, D., Mouton, P. le F. N. & Titus, T. 2001. A molecular perspective on the phylogeny of the girdled lizards (Cordylidae, Squamata). American Museum Novitates 3310, 1-10.

Lang, M. 1991. Generic relationships within Cordyliformes (Reptilia: Squamata). Bulletin de l'Institut Royal des Sciences Naturelles de Belgique, Biologie 61, 121-188.

Mouton, P. le F. N., Flemming, A. F.& Kanga, E. M. 2006. Grouping behaviour, tail-biting behaviour and sexual dimorphism in the armadillo lizard (Cordylus cataphractus) from South Africa. Journal of Zoology 249, 1-10.

Mouton, P. le F. N., Nieuwoudt, C. J., Badenhorst, N. C. & Flemming, A. F. 2002. Cordylus polyzonus (Sauria: Cordylidae) populations in the Western Cape, South Africa: relics or ecotypes? Journal of Herpetology 36, 526-531.

Mouton, P. LeF. N. & Van Wyk, J. H. 1997. Adaptive radiation in cordyliform lizards: an overview. African Journal of Herpetology 46, 78-88.

Pianka, E. R. & Vitt, L. J. 2003. Lizards: Windows to the Evolution of Diversity. University of California Press, Berkeley.

Pyron, R. A., Burbrink, F. T. & Wiens, J. J. 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13, 93.

Stanley, E. L., Bauer, A. M., Jackman, T. R., Branch, W. R. & Mouton, P. LeF. N. 2011. Between a rock and a hard polytomy: rapid radiation in the rupicolous girdled lizards (Squamata: Cordylidae). Molecular Phylogenetics and Evolution 58, 53-70.

Whitfield, P. 1983. Reptiles and Amphibians: An Authoritative and Illustrated Guide. Longman Group Ltd, Harlow, UK.

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As remarkable as it might sound, we’re presently in the midst of a ‘modern werewolf’ craze…

**Caption:** werewolves in modern day America. From left to right: Linda Godfrey illustration of a creature seen kneeling at the roadside in Elkhorn, Wisconsin, as reportedly seen by Lori Endrizzi in 1989; the 1936 ‘gadarrah’ creature described by Mark Schackelman from near Jefferson, Wisconsin; and Tina Cole’s sketch of a creature she recalls seeing in Michigan in 2001, when she was eight years old.

I don’t mean with respect to erotic fanfiction or fursona representation at conventions, but to claimed real-life encounters with living, breathing, dog-headed humanoids. Such is current interest in these werewolf monsters – we’re talking about an entity most often dubbed dogman – that they’re among the most frequently discussed of all creatures within the mystery beast canon.

The dogman craze got off the ground following the early 1990s collecting of reports from the vicinity of Elkhorn, Wisconsin, and indeed Wisconsin and adjacent Michigan are the areas most associated with dogman accounts. These involve the claimed observation of creatures today known as the Beast of Bray Road or Bray Road Beast – Bray Road, Wisconsin, being the place where several of the initial reports occurred – or the Michigan dogman.

**Caption:** Linda Godfrey was a skilled artist who illustrated dogman / the Beast of Bray Road several times. One of her illustrations – it featured in her 2003 book *The Beast of Bray Road* – is shown here; the photo of Linda shown here appeared in several interview articles on her.

One person above all others got dogman widely known and catalogued in the literature, and that’s Wisconsin-based journalist, author and researcher Linda Godfrey. Godfrey died of Parkinson’s Disease in 2022, aged 71, and probably all of the many obituaries devoted to her note her influence in bringing the Beast of Bray Road to wide attention. She was very much the ‘go to’ person on the subject after uncovering the creature’s alleged existence in 1991.

**Caption:** the small town of Elkhorn, Wisconsin, is about 100 km to the west of Lake Michigan, about the same distance to the south-west of Milwaukee, and about 80 km to the north-west of Chicago. It’s in the extreme south of Wisconsin and was home to around 10,000 people in a 2010 census. Image: Google maps.

  **Caption:** Bray Road – marked with an arrow – as it looks from the air. The rural, agricultural nature of the surrounds is obvious. The entire location is fairly open farmland with some tree cover. Image: Google maps.

The sightings. As relayed in her 2003 book The Beast of Bray Road, Godfrey owed her initial awareness of these accounts to an unnamed local freelance journalist who, in turn, had heard accounts relayed by a bus driver called Pat Lester. Lester’s daughter, Lori Endrizzi, had seen “a wolf or large dog with human characteristics” in the autumn of 1989, and it turned out that a teenage neighbour – Doris Gipson – had seen it too. Endrizzi’s sighting described the animal kneeling at the roadside at 1.30 in the morning, apparently holding roadkill. Gipson’s account, dating to Halloween 1991 (yes, Halloween), involved her hitting an animal with her car, stopping to check out the damage, and then having to escape at speed from the pursuing creature… which she then saw again on the same night, and had it attack the back of her car (Godfrey 2003).

**Caption:** images of Bray Road as it looks in 2024. Images courtesy of Tyler Stone, used with permission.

A December 1990 account, reported by a witness who was only 10 or 11 at the time, was also regarded as part of the same sighting flap, as were additional observations from other children in the same group (Godfrey 2003, p. 12). Clawed footprints suspected to be from the creature had been mentioned to the director of the Lakeland Animal Shelter, and at least some local people apparently took to using the nickname ‘Pottsy’ for the creature, presumably after Potter’s Road where the tracks were found (Godfrey 2003, p. 12).

Also during 1990, Mike Etten was driving along Bray Road late at night when he observed an animal bigger than a wolf, “sitting like a raccoon sits” and eating something held in its forepaws. A high school student called Tom Brichta is credited with two 1992 sightings of a shaggy-coated humanoid creature, the first of which involves the creature partly colliding with his car, the second – this one making reference to the dog-like head – describing how the animal was seen standing among 6-ft-tall corn (Godfrey 2003, pp. 38-41).

**Caption:** it might be obvious that the people of the Bray Road area are having fun with the dogman phenomenon today; an image taken during the Summer of 2024. Image courtesy of Tyler Stone, used with permission.

Some of the accounts that Godfrey uncovered long pre-dated the 1990s, in particular one recalled from 1936 that took place close to Jefferson, Wisconsin. This came to Godfrey via author and editor Joe Schackelman, who was himself relaying what his father, Mark, told him in 1958 (Coleman 1998, Godfrey 2003, DeMello 2023). A large, human-sized animal had been observed, kneeling on and clawing at an indigenous burial mound created by the Hopewell culture (DeMello 2023). What was supposedly the same creature was seen again on a second night, this time in a more extended encounter. 6 foot tall and with visible fangs and pointed ears, it made eye contact and an utterance described as “gadarrah”… more on that below. Joe Schakelman’s drawing, based on his interpretation of his father’s description, shows an erect-bodied, tailless, dog-headed creature with extended arms and folded wrists. What were interpreted as its first and fifth fingers were curled up and “shrivelled” (Coleman 1998, Godfrey 2003, pp. 26-27). Also of interest is that the illustration shows the animal as plantigrade, meaning that it was placing the heel of its foot on the ground.

**Caption:** the Mark Schackelman dogman/werewolf, as illustrated by his son, Joe, decades after the sighting is said to have happened. This drawing has appeared several times in the literature (Coleman 1998, Godfrey 2003, 2012). Things to note include the plantigrade feet, the bent wrists, and interesting curled structures on the inner and outer parts of the hands. Image: Joe Schackelman, from Godfrey (2003).

Other alleged dogman encounters have been reported and recorded since Godfrey amassed this selection in the early 2000s, but these are the foundational reports that got the phenomenon off the ground. They hint at several interesting points worth commenting on. The first is that a tradition of werewolf belief, local to the Bray Road area, had arisen among people there, especially the young ones. The suspicion when such things happen is that a sort of snowball effect emerges, either because other people want in on the fun, or because the phenomenon becomes a source of local identity that others seek to reinforce.

**Caption:** Godfrey, as shown in the 2024 Small Town Monsters movie, holding open the issue of *The Week* that contains her article ‘Tracking down ‘The beast of Bray Road’’. The illustration shows the kneeling dogman, a piece of carrion in its hand, described by Lori Endrizzi. Image: (c) Small Town Monsters.

The second is that some of the accounts that Godfrey collected don’t seem at all reliable, especially those involving children and what sound like tall tales from Halloween. And I want to emphasise that last point in particular: the initial, foundational dogman accounts are all highly suspicious, mostly unrealistic, and really make it seem that this was a cultural event whereby a bunch of locals were telling tall tales and fun, scary stories. I think it at least plausible that some observations of real animals – Etten’s roadside animal that sat like a raccoon could actually have been a raccoon, for example – got mixed in with the whole thing, but… otherwise, we really should be very sceptical of the inference that there was anything ‘real’ here in the first place. Are any of the dogman sightings reported in more recent years really descriptions of encounters with real animals? I’ve assumed that some are, but we really are talking about a handful of reports here.

Indeed, an issue obvious throughout Godfrey’s writings is that she collected and published any and all stories and anecdotes describing unusual hairy animals, the result being that what might look like a substantial roster of dogman reports is diluted by a number of probably irrelevant cases. Her 2003 book The Beast of Bray Road includes reports of big, shaggy, dog or wolf-life quadrupeds (pp. 31-34, 58-59), wildman or bigfoot-type creatures (pp. 34-38), and quadrupedal canids described as recalling giant coyotes or German shepherds (pp. 51-52).

**Caption:** I’m not sure how relevant it is to the whole dogman/American werewolf phenomenon, but it’s worth noting that certain domestic dogs can look surprisingly odd – disturbing even – in some circumstances. I like this photo of a large pet dog (it looks to be an Irish wolfhound), since it makes the point. Imagine how a person might react to seeing an animal like that standing bipedally for a moment, glimpsed among crops or foliage. I saved this image after seeing it on Facebook and haven’t been able to find a proper source.

Indeed, obvious throughout Godfrey’s writings is that she was forever aiming to bulk up the exceedingly slim dogman dossier through reference to any and all connected things, however tenuous the link. A fair amount of her The Beast of Bray Road describes local media coverage, Godfrey’s own experiences as an interviewee and media personality, and a huge quantity of musings concerning whatever she could dig up on werewolves and lycanthropy (Godfrey 2003).

**Caption:** Steve Cook’s song *The Legend* was first released on the radio (on April Fools Day, 1987), and later sold on CD and DVD. It is a work of fiction. Local TV station WTCM held an art competition, inviting viewers to send in their depictions of what the dogman might look like. The charcoal illustration (at right) of an especially monstrous dogman – by Brian Rosinski – became regarded as the best and most chilling of these. Rosinski was 23 years old at the time and without artistic training. The image is still associated with *The Legend* at some locations online.

Incidentally, a song – Steve Cook’s The Legend, first played on public radio in 1987 and then available for sale on CD and, later, DVD – relates a number of real-sounding Michigan dogman encounters and thereby creates the impression that the phenomenon has a pre-Godfrey tradition. But… it’s a song; essentially a work of creative fiction. Godfrey’s interpretation of its contents after discussing it with Cook was that the tales it recounts reminded her of dogman stories she knew from Wisconsin (Godfrey 2003, p. 64), and Cook’s song did help solidify the idea that dogman might be real, and with an attested history in the region. But Cook’s song was played as an April Fools Day gag (Godfrey 2012, p. 288). It was never meant to be relaying ‘real’ history of any sort.

**Caption:** Godfrey’s two original books. 2003’s *The Beast of Bray Road* was self-published. She produced others later on, including *American Monsters: A History of Monster Lore, Legends, and Sightings in America* in 2014 and *Monsters Among Us: An Exploration of Otherworldly Bigfoots, Wolfmen, Portals, Phantoms, and Odd Phenomena* in 2016.

The Gable dogman film. Another piece of creative media was also used, for a time, to bolster the case for dogman: a supposed vintage home movie, dubbed the Gable dogman footage. It did the rounds in 2007 and was uploaded to YouTube in 2009. You can watch it yourself here. Grainy, and revealing clothing and vehicles suggestive of its being set in the 1970s, it depicts countryside scenes from rural North America – Michigan, apparently – as a Mr Aaron Gable is filmed by one of his sons. Gable junior, filming from a car window, captures a glimpse of a human-sized, dog-like quadruped standing on a rise at the side of the road. The creature bounds on all fours towards the car and briefly disappears from view before its toothy maw flashes in front of the camera just before the film ends. A second, separate segment of film shows part of the police investigation as detectives survey a scene of death and dismemberment.

**Caption:** stills from the Gable film. At left, the creature begins to bound towards the camera operator. At right: shock horror, the creature leaps up at the window and we see its open mouth for a second or two… then the footage ends.

Despite its name, the Gable film – even if interpreted as genuine – never should have been considered part of the same phenomenon as Linda Godfrey’s Beast of Bray Road seeing as the creature it depicts is a bounding quadruped, not a humanoid. It was shown to be a hoax in 2010, and in fact a staged ‘unveiling’ of the whole thing, performed in co-operation with the hoaxer himself – Mike Agrusam – formed the focus of the final episode (titled America’s Wolfman, broadcast in March 2010) of the History Channel TV series MonsterQuest. Agrusam used an 8mm camera to create the impression of age and had worn a ghillie suit, and bounded on all fours while wearing it, when playing the creature.

**Caption:** another still from the Gable film. The camera operator, a young person with what looks like a hand-operated Nikon Super Zoom 8, is glimpsed briefly in the wing mirror of the car. A nice touch!

It turned out that Steve Cook – the musician behind The Legend song – was responsible for this piece of media too and had devised the film as a consequence of flagging sales of The Legend CD. He worked together with Agrusam and edited out some comical moments that Agrusam had initially included (Godfrey 2012). Cook’s cover story was that the film, stored in old cannisters labelled ‘Gable’, had been obtained via an estate sale. He sent copies to Godfrey and other researchers (Godfrey 2012) but obviously kept them in the dark as to the film’s fraudulent nature.

Dogman right now. Today, dogman accounts are not infrequently recounted in venues devoted to bigfoot and allied phenomena, and my rough impression is that something like 50% of people who endorse the existence of bigfoot also regard dogman as a genuine phenomenon, rather than as just a piece of folklore. At this point we come to a philosophical impasse of the sort not uncommon in the world of cryptozoological claims. To people who view the universe from a critical, evidence-based perspective – and this includes the majority of naturalists, scientists and authors with even a passing interest in cryptozoology – dogman is not real, cannot be real, and all reported accounts can only be hoaxes, tall tales, old folkloric accounts that are recast to sound as if they’re set in the modern day, or mistakes.

**Caption:** if you believe in bigfoot, chances are high that you accept the existence of dogman too. Twenty years ago, this would have been a ridiculous thing to say. These days…. well, not so much. Here are different images of ‘Patty’, the supposed bigfoot that features in the 1967 Patterson-Gimlin film. Images: Darren Naish.

But to a number of people who talk about dogman on podcasts, YouTube and the like, the implication is that we really are hearing here about a phenomenon unexplained by science, concerning either a mystery flesh-and-blood animal – a new species or some ungodly hybrid – or an entity from another realm that somehow, sometimes, manifests in our own. Here’s where we come back to Schackelman’s ‘gadarrah’ creature. Gadarrah is a word that has connections with Biblical Christianity. Also spelt Gadara, it’s the name of a region of Judea close to the Sea of Galilee, and famous as the place where Christ performed an exorcism.

**Caption:** the story of Jesus performing an exorcism and casting out demons that then possessed swine (that then leapt into the river and drowned) is a well known bit of Christian lore. It has been associated with places variously termed Gerasa, Gadara, or Gergesa. This 6th century AD mosaic, depicting the scene, is at the Basilica of Sant'Apollinare Nuovo in Ravenna, Italy. Image: public domain (from **here**).

Godfrey (2003, 2012) and others heaped significance on this point, the idea being that this ‘links’ dogman with a spiritual view of the world, perhaps one involving black magic and the occult. Maybe – according to this view – dogman itself is ‘demonic’. In talks she gave, Godfrey emphasized the fact that Schackelman’s encounter was associated with a burial mound, the implication being that dogman has a connection with the spiritual, satanic or demonic (J. Card, pers. comm. 2024). The 2024 Small Town Monsters movie The Bray Road Beast explains that Endrizi, one of the initial dogman witnesses, regarded dogman as “satanic in origin: a demonic entity or, perhaps, the Dark Prince himself”, and suggestions that dogman might be evil have been made by others too.

To clarify my own position in case it’s not already clear… there are hoaxes and fabrications in the dogman canon, but I also think that there are misinterpreted or embellished accounts involving bears, wolves, coyotes and other animals too. Perhaps most interesting is that we appear to be seeing the transmission of deliberately frightening stories that form part of a modern, evolving folklore. It’s not a coincidence that dogman accounts are mostly relayed via podcasts and YouTube channels devoted to the scary and the weird, the very places many of us go to learn about frightening phenomena, places where ‘authenticity’ is ambiguous and sometimes unimportant. The humanoid werewolf did not die out in the 1700s or earlier. It is in fact very much alive and even undergoing a renaissance.

**Caption:** until recently, the image of werewolves most familiar in popular culture was the human-faced, human-shaped monster of the sort shown here. At left, Lon Chaney as *The Wolfman*, from the 1941 movie of the same name. At right, a rendition of the same sort of creature from Usborne’s *All About Monsters* (**which I wrote about here**). Images: Universal (**original here**); Usborne/Miller 1977.

Origins of the humanoid werewolf. Having mentioned ‘humanoid werewolves’… the impression prevalent in the literature on monsters is that ‘werewolves’ as believed in during the Middle Ages and beyond were meant to look like large but anatomically normal wolves (Woodward 1991). Of those eyewitness accounts purportedly describing observations of actual werewolves recounted by Baring-Gould (1995) – originally published in 1865 and still regarded as the primary reference on the subject – all describe wolves, not hairy, dog-faced humanoids.

**Caption:** originally published in 1865, Baring-Gould’s *The Book of Werewolves* is still the standard reference on werewolves. It interprets accounts as folkloric transmissions (and connected to the long history of ideas that humans might, at times, undergo transformation into the form of another animal), mixed with accounts involving mental illness, and has what feels like a 'modern' approach.

The humanoid werewolf, supposedly, is a 20th century invention connected to cinema, and some authors have implied that it owes its origins to this art form alone (Woodward 1984). The thing is, though: that’s not true. In fact, artwork from the 1600s onwards shows that at least some people were indeed imagining werewolves to be bipedal, and to essentially be dog-headed humanoids. An engraving by George Jacob Schneider, made during the 1680s and connected to a spate of claimed werewolf attacks from Eschenbach in Germany, shows a bipedal werewolf wearing a ragged sheet (Ruickbie 2016). A much-reproduced 18th century engraving – it appears in most 20th century books that discuss the subject of werewolves (e.g., Farson 1975, Bradbury 1981, Grant 1992) – shows a ‘dogman’ werewolf carrying a woman in its jaws.

And a famous illustrating showing werewolves lining up for a nighttime jumping contest – produced for Maurice Sand’s 1858 book Légendes Rustiques – also depicts werewolves of a sort consistent with the modern, pop-culture vision of the creature. This illustration is also a staple of books that cover monsters and related phenomena (e.g., Bradbury 1981).

**Caption:** at left, a very famous 18th century image showing a werewolf carrying off an unfortunate victim. The point is made that her Christian faith did not save her. At right, werewolves lining up against a cemetery wall in order to compete in a jumping competition: a famous illustration that first appeared in Maurice Sand’s *Légendes Rustiques* of 1858.

It can be argued that these images were inaccurate in view of what werewolves were really ‘meant’ to be like (Baring-Gould 1995), but the point remains that these humanoid depictions are, and have long been, a staple of the literature. The idea that the humanoid werewolf originated in cinema is, therefore, not true, and we’ve been subjected to this template for some hundreds of years. I put it that this template has persisted to modern times.

The Beast of Seven Chutes. For all the dogman reports that exist, there are no photos or pieces of film. Or, at least, none worth spending any time discussing. But there’s one exception: the June 2005 image that’s become known as the ‘Beast of Seven Chutes’ photo. Taken in Quebec at Le Parc des Sept Chutes (chutes = waterfalls) by a semi-anonymous, French-speaking dump-truck worker named Lary, the photo depicts woodland surrounding a large river. Partly hidden by foliage and shadow at far right, and unnoticed by the photographer at the time, is a large and vaguely humanoid figure standing in a small clearing adjacent to a path.

**Caption:** Le Parc des Sept Chutes, also known as Sept-Chutes Regional Park or Parc régional des Sept-Chutes, in Lanaudière in the south of Quebec. The eponymous seven waterfalls are located along a 17 km stretch of the Noire River. Image: Google maps.

The figure looks superficially like a dog-faced biped, covered in brown fur and with a whiteish nape and red crown. To its left, adjacent to the lower part of its chest, is what appears to be a distinct, pale object, and if we look really hard we can see that this object has projections that look like legs and a tail. It would appear that the dogman has abducted, and is carrying, a small dog (though other suggestions include a pig, a chunk of meat or a plush toy), a point arguably consistent with eyewitness accounts that describe dogmen holding carrion.

**Caption:** at left, a close-up shot of the mystery creature, captured accidentally in the photo by semi-anonymous Lary. At right, an extreme blow-up of what is taken to be its face. Images: these are taken from Rob Gaudet’s now defunct website, still findable **here** via wayback machine.

In current, online cryptozoological theorising (at Cryptid Wiki, for example), there’s the claim that this creature is one and the same as the gugwe, this being a word that comes from Mi’kmaq lore of the Atlantic provinces of Canada. This name is, in fact, more associated with the Seven Chutes photo than ‘dogman’, and I presume that those endorsing this proposal regard gugwe and dogman as the same thing. This is a blatant effort to give dogman’s reality tacit support via supposed connection to an indigenous source, and I don’t see much reason to take it seriously.

**Caption:** interpretations of the Seven Chutes creature. At left, an impressive effort to visualize the animal’s face as it might look if it were a real animal. A giant, dog-faced primate might be terrifying, but this vision makes it look almost friendly. At right, an effort to show the outlines of the beast if it’s interpreted as a humanoid holding a dog-shaped object. Images: again, I only know these from reddit and have not found their original sources. The artistic reconstruction is credited to someone with a name similar to David A. Cleara. Any help with proper credit is appreciated.

When photos of alleged monsters exist, you must never assume that they’ve gone un-investigated or been deemed of little value or interest. On the contrary, a massive amount of discussion and analysis surrounds these images, and at least some of it has value if you’re interested in knowing what the images really represent. Such is the case here. As documented at a now defunct website and 2023 discussion thread at reddit, author, entrepreneur and humanitarian worker Rob Gaudet looked in detail at this case, made contact with the photographer (initially buying the photo off him, since he was selling it on ebay!), and obtained additional photos and video footage taken at what is undoubtedly the same location.

**Caption:** Lary’s original photo of Le Parc des Sept Chutes, presumably taken because he was enjoying the look of the white water. Submerged in shadow and vegetation at lower right is a browny-greyish object that looks like a human-sized creature. Image: Lary/Rob Gaudet.

  **Caption:** the same photo but with ‘the creature’ circled in red. Image: Lary/Rob Gaudet.

While the semi-anonymous nature of Lary the photographer and the venue of the discussion – an inactive webpage and a reddit thread – might throw up some red flags, I don’t see a good reason to doubt the veracity of Gaudet’s work and I think that he really did succeed in contacting the photographer as stated.

Based on the interview data that Gaudet obtained, Lary the photographer was alone in the park when the photo was taken, and only noticed the creature when checking his images later on. He was sufficiently intrigued that he later visited the site on three additional occasions, one time taking his girlfriend so that she could stand in the relevant spot to provide a sense of scale. She wasn’t able to stand in the exact same spot due to new vegetation that had grown in the interval, but it’s still evident that ‘the beast’ was approximately human-sized. It also appears that it probably can’t be explained as a misidentified rock, tree stump, fallen log or pile of vegetation since none of those things can or did exist in the specific spot. I think it really must have been an animal of some kind.

**Caption:** Lary’s photos of the specific location in Le Parc des Sept Chutes where the beast was standing. At left, the original image, with the beast in place. At right, the same location with a person standing – as best as possible – in the same spot. The horizontal lines show how the creature and a person are similar in size. Images: Lary/Rob Gaudet.

Could it be a bear? That doesn’t look right based on its size, shape and posture, since it looks too erect and too broad-chested, and also in the possession of shoulders. These points also rule out wolf, coyote and other known species that might be encountered in the region. What about a non-native, escapee animal? A specific photo of a squatting gorilla has been noted by some as looking similar to ‘the beast’ but, as you can see from the overlay shown below, the similarities are superficial and not impressive. The idea that a gorilla might have been living in the wild in Quebec is unlikely anyway, we have no reason to think that one was in the area at the time the photo was taken, and there’s nothing in the photo that makes it look as if an image of a gorilla has been composited in at some point.

**Caption:** the photo of a captive Western lowland gorilla *Gorilla gorilla gorilla* shown here has been linked by some with the Seven Chutes photo. But an overlay of the two shows that the similarity is coincidental since there’s insufficient similarity of shape. The original gorilla image has been flipped, since the source image (published at Wikipedia) shows the animal facing to the right. Image: Greg Hume, CC BY-SA 3.0 (**original here**); Lary/Rob Gaudet.

What about the most familiar animal of them all, by which I mean… the human. We tend not to see a human-like shape in ‘the beast’, but that’s because we forget (or fail to account for) objects that often accompany humans when they’re out walking in natural areas, like hats, bags and cameras. A sensible and clever interpretation that I find worthy of consideration is that ‘the beast’ is actually a hiker seen from behind, and that what looks like the head and muzzle of ‘the beast’ is a large camera, held upwards because the person was aiming it at an object up in a tree or in the sky.

A claim often made when reinterpretations like this are on offer is that the revisionary ‘gotcha’ is just as speculative and pareidolic as the original monster claim (here, that it’s a giant, dog-headed, furry humanoid carrying an abducted pet dog), but… come on, is it? I feel it likely that, alas, there is no ‘Beast of Seven Chutes’ in the photo, and that we are indeed mis-reading an image of a person.

**Caption:** at left, another zoom-in showing the beast with, at right, a suggestion interpretation in which the animal is reinterpreted as a person holding a camera aloft. The pale patch is their hair (or headgear), the upper arms, shoulders and sides of the head are the arms and hands of the person, and the muzzle and reddish forehead is a camera body and lens. I do find this interpretation pretty plausible. Images: Lary/Rob Gaudet; I only know the reinterpretation from **this reddit page** and have been unable to find out who created it originally.

A final piece of alleged dogman evidence is worth mentioning, this being an audio recording where two people – supposedly involved in a late-night collision of their vehicle with an unknown large animal – are attacked by what’s presumed to be the same creature. It’s a noisy, growling beast and, just before the recording cuts out, we hear the desperate screams of its unlucky victims. I certainly haven’t investigated this case in appropriate depth, but I submit that it can be ruled out as an amusing bit of monster theatre.

Explaining dogman. Can there be any rational, real-world, flesh-and-blood interpretation of dogman? Could ‘dogman’ be exactly what some people evidently hope it is: a humanoid canid? That is, a genuine member of the dog family that has evolved a humanoid physique, an obligatorily bipedal stance, and giant size relative to other canids? Linda Godfrey hinted at this idea at times, suggesting that adaptation to a North American habitat dominated by tall grasses had encouraged the evolution of a new, bipedal configuration in wolves. According to this view, people would be seeing a recently evolved, erect-bodied wolf adept at using its forelimbs in grabbing and carrying.

**Caption:** dogman as envisioned by Tyler Stone, representing a vision of this creature now endorsed by believers. An erect-bodied canid with digitigrade hindlimbs but hominid-like pectoral and forelimb adaptations. Image: Tyler Stone, used with permission.

Is this view scientifically reasonable? The short answer is no. The long answer is that we have no reason to think from fossils, from the functional anatomy of canids living and extinct, or from the role that canids play in ecology (or played, across their whole evolutionary history) that bipedal, semi-bipedal or facultatively bipedal canids might have evolved. Wholly extinct canid groups existed in the past, including the archaic hesperocyonines and the diverse borophagines, the latter sometimes termed bone-crushing dogs. Members of those groups were different from the sole living canid group (Caninae) in some respects, but not so different that we see anything like the evolutionary potential for a ‘non dog’ body plan.

Godfrey’s model also requires that this evolutionary event occurred at breakneck speed within (at most) the last few thousand years. And wolves haven’t been present in North America for more than 30,000 years anyway… a very short time, geologically speaking. We know that profound evolutionary changes can occur in a short timeframe – examples are demonstrated by the cichlid fishes in the great lakes of the African Rift Valley – but no such event is known from the evolutionary history of mammals. Our own species, to take one example, is less than 500,000 years, yet species extremely similar to use in shape and proportions were in existence for more than a million years prior to this.

**Caption:** we’re basically at a point where those promoting the existence of dogman are endorsing the existence of animals like this… the humanoid werewolf of popular media. These particular figures represent the Victorian werewolf from a 2006 episode of *Dr Who*, and a 2017 werewolf figure made by Safari Ltd. Image: Darren Naish.

Actual bipedal dogs. Here's a second ‘flesh and blood’ possibility. Despite what I just said, we do, actually, know of fully bipedal canids. The caveat is that I’m referring to malformed individuals of the domestic dog Canis familiaris born without forelimbs, or unlucky enough to have had their forelimbs removed or damaged by accident. Such individuals have had no choice but to walk bipedally. A famous example is Faith, a pet dog who lived in Oklahoma and then Indiana, USA, between 2002 and 2014. We know from other examples that quadrupedal mammals can become bipeds when malformation demands it. The best-known case is that of the bipedal goat described by Everhard J. Slijper in 1942. This animal lacked forelimbs and special attention was paid to its flattened, humanoid chest, modified pelvis and enlarged back muscles (Slijper 1942).

**Caption:** Faith the bipedal dog (2002-2014) was something of a celebrity, and appeared numerous times on TV and in online articles. She was not swift or agile, but was very much able to walk unassisted. She was born with three limbs, but a deformed forelimb was amputated at age seven months. Images: Mike Matney, CC BY-SA 2.0 (**original here**); (c) Shirley Ann Dennis.

Is it worth considering the possibility that a mutant wolf (or coyote, coywolf, wolf-dog hybrid, or whatever), affected by the same developmental issues, might occur in the wild and hence explain at least some dogman accounts? Two problems affect this hypothesis. The first is that dogman accounts generally make a point of describing large, well-developed forelimbs, so ‘forced’ bipeds these are not. The second is that any hypothetically deformed wild canid is going to be a one-off or, at least, an extremely rare occurrence. There won’t be a number of them spread across a wide area, and there certainly won’t be a self-sustaining population. So… no, I don’t think that malformed or disabled wild-living canids have any contribution to the dogman phenomenon.

The most novel of hybrids. The notion that canids and primates might have hybridized and that dogman is the result – yes, this idea is also hinted at in discussions about dogman – is also a non-starter. Hybridization among mammal species is rampant in the natural world, but it only works when the parent species are sufficiently close behaviourally, physically and genetically. Distantly related species might be compatible when it comes to the relevant physical acts, but hybrid babies are not the result. Canids, for example, cannot produce hybrids with bears, seals, skunks or cats, yet those animals are close relatives of canids compared to primates.

**Caption:** the idea that bigfoot has been genetically sequenced and has a hybrid origin that involves humans as well as an unknown, mysterious, additional being surely can’t be connected to views that werewolves are real…. can it? Hold my beer. Images: screengrabs from **this CFI article** on Melba Ketchum.

With that in mind, the notion that anonymous geneticists working in a secret lab might have spliced dog and human DNA together is, for sure, an entertaining or disturbing notion, but also a sci-fi one outside our current abilities. Why any such chimaeric novelty (which would have been extremely expensive to produce and nurture) might be released into the farms and woodlands of the American mid-west is also a question that would need answering. I don’t think that this idea can be taken seriously but I mention it for want of completion. It was promoted by Melba Ketchum (of ‘bigfoot DNA reveals hybrid origin’ fame) in a Coast to Coast interview.

Of mystery kangaroos and devil monkeys. Another idea is that these creatures are not canids with primate-like bodies, but primates with canid-like faces. We know of dog-faced monkeys (drills, baboons and kin). Could there also be big, ape-like, dog-faced primates that await scientific recognition? Again, I think that the short answer is no. However…

**Caption:** so-called dog-faced monkeys are remarkable animals with a striking and sometimes shocking appearance. Some researchers have speculated that North American primates of this sort actually exist and await recognition. Mandrill *Mandrillus sphinx* at left, Drill *M. leucophaeus* at right. Images: Darren Naish.

The notion that North America (and elsewhere) might be inhabited by a large, bipedal or semi-bipedal, dog-faced primate is not new, having first been proposed in the 1970s by cryptozoological investigators seeking to explain animals identified by witnesses as ‘kangaroos’ or ‘devil monkeys’ (Coleman 1998, Coleman & Huyghe 1999). The accounts concerned are vague with little to tie them together. My opinion is that they include observations of escapee wallabies and kangaroos, and that there isn’t a valid category of cryptid here. Perhaps worth noting is that these ‘devil monkeys’ are supposed to be smaller than a person, typically less than 1 m tall, whereas dogman is a giant, at least as big as a human adult if not taller. In other words, the Beast of Bray Road just isn’t at all like the creatures described in alleged ‘devil monkey’ accounts.

**Caption:** my effort to reconstruct the ‘devil monkey’/’American kangaroo’ cryptid endorsed in parts of the cryptozoological literature (Coleman & Huyghe 1999) and suggested therein to be a kind of dog-faced, bipedal giant monkey. Coleman & Huyghe (1999) proposed that *nalusa falaya* of the Choctaw of Louisiana represents the same sort of animal, and that these creatures exist “from Alaska to New Brunswick, with a concentration of contemporary sightings in the Mid-west” (p. 60). Image: Darren Naish.

Other cryptozoologists have also suggested a primate identity for dogman, this time that it might be a variant of bigfoot that has a superficially dog-like muzzle and enlarged canines. The ‘gugwe’ term that we saw earlier is applied by some researchers to this alleged bigfoot variant.

My bias – though I don’t think ‘bias’ is the right word – is that I can’t take seriously the hypothesis that North America might be inhabited by a novel mammal species of this size or sort. That’s a view shared with the majority of biologists, naturalists and ecologists.

The bear hypothesis. If we suppose that dogman observations really do describe encounters with real animals, could these be garbled accounts of bears? Bears are not ordinarily bipedal, but they will stand on two legs for a while and some individuals will even walk this way for a while.

**Caption:** you can decide for yourself how seriously you take this image. The dogman that people are claiming to see is generally supposed to be very distinct from a bipedally-standing bear in a number of important respects. Image: Darren Naish.

An especially bold proponent of dogman might argue that bipedal bears differ from dogmen in that the latter reportedly have prominent shoulders and human-like arms and hands, a most unbear-like anatomical configuration. In addition, bears are plantigrade, meaning that they plant the whole hindfoot on the ground and don’t have ‘hocks’ (bony ankle joints that are raised well up off the ground). That last feature has been mentioned quite specifically by some dogman witnesses and their insistence that the animal they saw was digitigrade and not plantigrade is a problematic contention for the bear hypothesis.

However, that’s not the case across the board. You’ll recall me noting earlier that Mark Schackelman’s Michigan dogman from 1936 apparently had plantigrade feet. In fact, the Schackelman dogman has a few other traits that could be consistent with a bear identity: its strongly flexed wrists (which resulted in the hands hanging down at a 90° angle to the forearm) very much recall those of bipedal bears, and a rounded projection on the outer edge of its wrist looks like the pisiform pad that bears have in the same location (though a dogman that’s a speculative bipedal carnivoran could well have this same feature too).

**Caption:** underweight, mange-infected Black bears *Ursus americanus* can look very odd, and with more than a passing similarity to werewolf-type creatures described in certain of the reports. Could witnesses have actually observed bears like the individual shown at left? We also know that bears – some bears, anyway – can walk bipedally. The photo at right shows Pedals, a famous individual from New Jersey that was shot dead in 2016. Note the forelimb pose here and compare it with that of the Schackelman animal shown above. Images: Pennsylvania Game Commission; (c) Ron Cronk.

It’s also worth noting that bears with mange could be relevant to dogman accounts. These unfortunate animals have a skinny appearance where the neck looks thinner than is the case in normal bears and the head looks more clearly demarcated. Parts of the arm, chest and back can be unfurred and exposed, and could create the impression of a pseudo-human body shape. I’m far from the first person to propose this and recall Pat Spain (of the Nat Geo Beast Hunter TV series, and much else besides) suggesting it within recent years.

Does this mean that all dogman accounts are confused descriptions of encounters with bipedally walking bears suffering from mange? No. But some might be.

What, then, to do with dogman? From a sceptical, scientific perspective, there’s no reason to think that there’s a genuine zoological phenomenon here. There are no bones or other remains, and no worthwhile photographic evidence. There are no otherwise-unexplained DNA, hair or scat samples, audio recordings, or tracks or bite marks that require the serious consideration of dogman as a real animal. It’s a pop-culture, folkloric phenomenon, the ‘evidence’ being a tainted pool of dodgy accounts collected by people who’ve been specifically looking for weird tales to relate in popular books and articles, and in podcasts.

**Caption:** yes, that’s a Linda Godfrey action figure, photographed by Loren Coleman and part of the collection on show at the **International Cryptozoology Museum**. Part of me disapproves of the lionization, even canonization, gifted to investigators of cryptozoology. They’re often not heroic at all and are sometimes in it for self-aggrandizement, not philanthropy! But another part of me likes action figures and geeky humour. Image: Loren Coleman.

Hypotheses that dogman might be explained by a recently evolved, novel form of canid or developmentally aberrant wild-living canid specimens don’t withstand scrutiny and shouldn’t be endorsed, and notions of a hybrid or lab-born origin are pseudoscience pushed by unreliable narrators.

Dogman, beloved of anti-science types. My personal opinion is that this phenomenon persists for two reasons. The first is that some people – often, those who promote the idea that the scientific consensus fails to explain the real nature of the universe – very much want inexplicable creatures of this sort to exist. It fits within a “science can’t explain everything, the world is stranger than we know, scientists are only in it for the money” worldview aligned with conspiracy theory and anti-science feelings, and a cynical take could be that it’s symptomatic of the poor science literacy and aggressive, right-leaning libertarianism prevalent in parts of the USA. I’ve seen this anti-scientific view pushed on pro-cryptozoology podcasts (thinking in particular of Wes Germer’s Sasquatch Chronicles, sorry Wes), and also by researchers who promote anti-scientific views (like Melba Ketchum).

**Caption:** discussions and stories relating to dogman and American werewolves do appear in printed media, but their primary theatre is the internet, YouTube and the podcasting world in particular. Move in the right circles, and you will be *constantly* subjected to dogman lore and discussion.

I have some fondness and respect for Linda Godfrey, but like many writers who make or made a living from the generation of content on weird phenomena, she was guilty of mystery-mongering: of maintaining and building interest in dogman through what amounts to constant promotion.

The second reason is the sociocultural one I’ve pushed before with respect to monster sightings and belief (Naish 2016, 2023). Dogman witnesses are not reporting their claimed observations as encounters with an entirely new, unfamiliar creature, but – on the contrary – are describing something deeply familiar and very much ‘known’; the dog-headed, humanoid werewolf of cinema, literature and mythology. As others have noted before me, dogman reports have been coming in at about the same time as humanoid werewolves have been making a minor resurgence in the cinema: the Canadian movie Ginger Snaps saw release in 2000, and the very successful UK horror flick Dog Soldiers was outed in 2002. Creatures of this sort were very much in people’s minds.

**Caption:** the dog-headed humanoid is a very familiar image these days, thanks in no small part to memorable creatures from cinema. At left, a werewolf as portrayed in *Ginger Snaps* of 2000. At right, a Scottish werewolf as seen in *Dog Soldiers* of 2002.

Is dogman ‘real’? It’s real and alive in modern culture, for sure, and in fact this is a fascinating case of evolving folklore, very much worthy of analysis and investigation. But is it a flesh-and-blood creature that walks the land and awaits scientific recognition? No.

For previous Tet Zoo article on monsters, cryptozoology, folklore and connected issues, see…

Bigfoot’s Genitals: What Do We Know?, August 2018
Books on the Loch Ness Monster 1: Ronald Binns’s The Loch Ness Mystery Reloaded, March 2019
Books on the Loch Ness Monster 2: Gareth Williams’s A Monstrous Commotion, March 2019
Usborne’s All About Monsters, April 2019
Sea Monster Sightings and the ‘Plesiosaur Effect’, April 2019
Books on the Loch Ness Monster 3: The Man Who Filmed Nessie: Tim Dinsdale and the Enigma of Loch Ness, August 2019
Lore of the Loveland Frog, January 2020
The 1972 Loch Ness Monster Flipper Photos, August 2020
Monsters of the Deep, a Ground-Breaking Exhibition at the National Maritime Museum, Cornwall, October 2020
The Lake Dakataua ‘Migo’ Lake Monster Footage of 1994, February 2021
What Was the Montauk Monster? A Look Back to 2008, October 2021
Santa Cruz’s Duck-Billed Elephant Monster, Definitively Identified, November 2021
Legend of the Black Dog, August 2022

Acknowledgments. This article was first published at my patreon and I benefitted from comments added by patrons. Thanks also to Sharon Hill and Jeb Card for comments and corrections, and Tyler Stone for kind use of images.

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

Refs - -

Baring-Gould, S. 1995. The Book of Werewolves. Senate, London.

Bradbury, W. 1981. Into the Unknown. The Reader’s Digest Association, Pleasantville, New York/Montreal.

Coleman, L. 1998. The Wisconsin werewolf. Fortean Times 108, 47.

Coleman, L. & Huyghe, P. 1999. The Field Guide to Bigfoot, Yeti, and Other Mystery Primates Worldwide. Avon Books, New York.

DeMello, M. 2023. Bigfoot to Mothman: A Global Encyclopedia of Legendary Beasts and Monsters. Bloomsbury Academic, New York.

Farson, D. 1975. Vampires, Zombies, and Monster Men. Aldus Books, London.

Godfrey, L. 2003. The Beast of Bray Road: Tailing Wisconsin’s Werewolf. Linda Godfrey.

Godfrey, L. 2012. Real Wolfmen: True Encounters in Modern America. Tarcher/Penguin, New York.

Grant, J. 1992. Monster Mysteries. The Apple Press, London.

Naish, D. 2016. Hunting Monsters: Cryptozoology and the Reality Behind the Myths. Arcturus, London.

Naish, D. 2022. A cultural phenomenon. The Biologist 69 (3), 16-21.

Ruickbie, L. 2016. The Impossible Zoo: An Encyclopedia of Fabulous Beasts and Mythical Monsters. Robinson, London.

Slijper E. J. 1942. Biologic-anatomical investigations on the bipedal gait and upright posture in mammals, with special reference to a little goat, born without forelegs. Proceedings of the Koninklijke Nederlandse Akademie Van Wetenschappen 45, 288-295, 407-415.

Woodward, I. 1991. Delusions and transformations. In Brookesmith, P. (ed) Creatures from Elsewhere: Weird Animals That No-One Can Explain. Macdonald & Co, London, pp. 81-84.

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You’ll recall that my aim for 2024 is to rescue and revamp a good deal of old squamate-themed material from the Tet Zoo archives….

**Caption:** Smith’s African snake *Grayia smythii* (…. or should that be ‘*smithii*’?), photographed in the wild. When encountered by people, these animals will hiss and open the mouth in threatening fashion. The scales are smooth and the pupils are round. Image: Marius Burger, public domain (**original here**).

In previous articles – including those on garter snakes and house snakes and kin – we’ve looked at snakes conventionally lumped together within the old-fashioned, super-inclusive version of Colubridae. For the latter part of the 20th century at least, this group was interpreted as including basically all caenophidian snakes that weren’t filesnakes, viperids or elapids. I like Gower et al*.’s (2023) characterisation of this situation…

“Snake biologists understood that this was unsatisfactory, but they were somewhat overwhelmed by the morphological and ecological diversity of these many hundreds of species, and so put up with the situation while continuing their research” (p. 160).

Caenophidia = so-called ‘higher snakes’, the vast assemblage that contains file snakes and Colubroides and excludes archaic Mesozoic snakes, the weird worm, thread and blind snakes, and the pythons and boas.

Caption: massively simplified phylogeny of crown snakes to show that Caenophidia is the giant clade that includes the youngest major snake groups. Elsewhere in the tree, studies continues to disagree on whether boas and kin form a clade with pythons and kin. The monophyly of Scolecophidia has also been challenged. Images: as ever, these are for the textbook. More on that here.

But as our understanding of caenophidian phylogeny, diversity and anatomy has improved (and as views on how taxonomy should reflect phylogeny have changed too), it’s become increasingly realized that ‘Colubridae’ of tradition should be split up. In part this is because it was polyphyletic, various of its constituent groups being closer to elapids (cobras, mambas and kin) than to the Eastern racer Coluber constrictor, the ‘core’ animal of Colubridae. And it’s also because the groups are sufficiently diverse – in all measures – to be considered ‘families’ all their own.

Introducing the grayiids. Here, we look at a poorly known group previously included within Colubridae as a ‘subfamily’. They’re often termed African water snakes (or watersnakes) and all species (five have been named to date) are included in the genus Grayia. Regarded as a subfamily within Colubridae, the group is thus Grayiinae. In taxonomic systems where Colubridae is restricted to the ‘Coluber group’ alone (and where what used to be Colubridae is Colubroidea), they’re Grayiidae, the grayiids.

**Caption:** big grayiids are said to be sluggish and even somewhat clumsy on land (the idea that they move “ponderously” was stated by Stephen Spawls and colleagues in 2002), whereas juveniles look to be agile, faster-moving animals. This juvenile *G. tholloni* was photographed in Togo; this species maintains banding into adulthood, even though the bands fade with age. Image: toganim, CC BY-NC (**original here**).

What’s with the origin of the scientific name? Grayia was first reported scientifically in 1818 when G. smythii was described from Democratic Republic of the Congo (DRC) as a species of Coluber – the concept of that genus was, at the time, substantially broader that it is today – by British zoologist William Elford Leach (1791-1836). That specimen had been collected in DRC on an expedition led by Captain J. K. Tuckey, and then studied by Leach back at the British Museum’s Natural History Department in London. Perhaps the most memorable detail of that expedition’s history, at least with respect to zoological specimens, is that three lion cubs were given to Tuckey and his men by local people. These cubs were “kept alive three days and fed on soaked bread, which doubtless caused their death” (Cranch in Tuckey & Christen 1818, p. 405). Anyway, you might know Leach best thanks to the giant New Caledonian gecko Rhacodactylus leachianus, named in his honour by Georges Cuvier in 1829, or by Leach’s storm petrel Hydrobates leucorhous.

**Caption:** taxa prominently associated with William Leach, rightly or not. At left, New Caledonian or Grande Terre giant gecko *Rhacodactylus leachianus*. At right, Leach’s storm petrel *Hydrobates leucorhous*. Images: Lennart Hudel, CC BY 4.0 (**original here**); Alexis Lours, CC BY 4.0 (**original here**).

In 1858, German-born British zoologist Albert Günther named a new snake that he regarded as worthy of its own genus: Grayia silurophaga, ‘Gray’s catfish eater’. This later proved synonymous with Coluber smythii… which thus became Grayia smythii. ‘Smythii’, incidentally, almost certainly honours Norwegian botanist Christen Smith (1785-1816) – he collected the type specimen on Tuckey’s expedition and somehow died on the same trip – and thus should really be ‘smithii’. Annoyingly, ‘smythii’ and ‘smithii’ are both used in the literature on this snake, as are their variants ‘smythi’ and ‘smithi’ (I’m sticking with smythii here as I still see it as the name that’s most prevalent in the literature). And, yes, it’s no secret that the reptiles (and other animals) of the global tropics sure are named after a whole lot of dead northern European white guys.

**Caption:** a hatchling *G. smythii*, photographed in the wild in eastern Congo. Juveniles of this species are conspicuously banded; adults are generally yellowish-brown, olive, or black with a mottled appearance where the cross-bars of juveniles have merged and ‘filled in’ the lighter areas. Image: (c) Kate Jackson.

So, what’s with the name ‘Grayia’? The British zoologist John Edward Gray (1800-1875) has been mentioned on numerous previous occasions here at Tet Zoo, often because he named the animals I write about (examples include arboreal alligator lizards Abronia and the whale Kogia). Well, Günther named Grayia specifically in his honour. Additional Grayia species were named during the 1860s and 90s: Caesar’s African water snake G. caesar by Günther in 1863, the Ornate African water snake G. ornatus by José Vicente Barbosa du Bocage in 1866, and Tholloni’s African water snake G. tholloni* by François Mocquard in 1897.

Actually, he named it Xenurophis caesar. George Boulenger, in 1910, transferred it to Grayia.

Caption: John Edward Gray sure was a hard-working individual when it came to describing and naming animals that fell within his broad sphere of interest, which basically included all animals. He initially joined the British Museum’s Zoology Department to help catalogue reptiles, but ended up becoming Keeper of Zoology. At right are just two of the taxa he named: Kogia the whale and Abronia the lizard. Images (clockwise from left): public domain; Robert Pitman, in public domain (original here); Ethan Kocak, used with permission.

A phylogenetic analysis published this year (Chaney et al. 2024) found G. smythii and G. ornata to be sister taxa, and G. caesar and G. tholloni to also be sister taxa. They also found that these two clades diverged during the middle of the Oligocene (around 27 million years ago). That’s a deep divergence and this, combined with anatomical differences, led them to suggest resurrection of the name Xenurophis (originally applied to G. caesar), albeit as a subgenus within Grayia. Chaney et al. (2024) also found that Grayia included a hitherto overlooked distinct lineage that appeared to be cryptic species, and named G. obscura for a population from the Upper and Middle reaches of the Congo River and nearby. Spawls et al. (2018, p. 536) made reference to “one undescribed species” known from Cameroon, though I don’t know of its current status.

Incidentally, the common name ‘Caesar’s African water snake’ for G. caesar is fairly inappropriate since it seems that Günther named this snake in possible recognition of its magnificent, regal form (Beolens et al. 2011) (‘caesar’ being a name used for Imperial things thanks to its original association with the Roman emperor). A more accurate vernacular name would thus be ‘Magnificent African water snake’ or ‘Regal African water snake’ or such. I should also add that I find ‘African water snake’ clumsy and unlikeable – it’s just too generic – and would prefer it if we adopted an African name for the group. The local name dibomina is used for these snakes in many places, but one problem is that its plural is mabomina, which would be inconsistent with English conventions.

**Caption:** unfortunately, I don’t have any images of *G. caesar* that show how attractive and impressive it is in life, and this image of a dead one – presumably squished on a road – hardly does it justice. Image: (c) M. Cristina Carboni (**original here**).

  **Caption:** Ornate African water snake *G. ornata* photographed in the wild. A prominently banded pattern is typical for this species and explains its vernacular English name. I find this snake to have a natricid-like demeanour and would suspect it to be a member of that group if I didn’t know better. Image: Marius Burger, CC0 (**original here**).

Some biology and natural history. Grayia snakes are robust and large, reaching 1.7 m and even 2.5 m in a few places in western Africa (Spawls et al. 2004). Relatively little is known about their natural history, they’re regarded as elusive, and they inhabit places – seasonal swamps and watercourses – where they can be hard to see and find. But as suggested by the common name, African water snakes are semi-aquatic and eat fish, frogs and tadpoles. A study of G. smythii in Nigeria demonstrated Tilapia and the catfish Clarias to be the main fish prey, with Xenopus tropicalis – tadpoles as well as metamorphs – being the frog the snakes ate the most (Godfrey & Luiselli 2001). This diet appears typical of G. smythii, at least, across its range.

**Caption:** everyone’s heard of *Xenopus laevis*, but less familiar (unless you’re a massive herp nerd) is the Western or Tropical clawed frog *X. tropicalis*. It’s generally smaller as an adult than X. laevis (with a snout to vent length of 3-5 cm) and often darker on its dorsal surface. Image: Václav Gvoždík, CC BY-SA 3.0 (**original here**).

Egg-laying happens during the winter dry season, and an unusual behavioural trait is that their eggs are deposited in several separate batches at two or three separate sites, an aspect of behaviour unknown for other snakes (Godfrey & Luiselli 2001). Nests are among leaf litter gathered between buttress roots and not located in an aquatic setting (I know that that would be highly unusual but it’s worth commenting on). Males and females are similar in size with males having proportionally longer tails, as is typical for snakes.

Until recently, I’d wrongly thought that these were animals of the Congo region alone. But they’re actually hugely widespread across continental Africa, with a range extending from Niger in the north to Angola in the south-west, and Kenya and Tanzania in the east. They occur, then, across a vast portion of one of the world’s greatest continents, albeit not in the far north or south, and not in the deserts.

**Caption:** a map of Africa, showing the countries from which four of the living grayiid species have been reported (so, it’s not really a range map, and it doesn’t include data from all recognized species). It’s messy given that some countries are inhabited by two or even three (like Cameroon, Angola and Democratic Republic of the Congo) grayiid species. With occurrences in nations like Chad, Ethiopia and Senegal, this group is not ‘Congolese’ only, even employing the most generous use of that term. Image: public domain.

Ethnozoological knowledge. A common complaint made about animals of all sorts is that it can be really hard, if not impossible, to find what local people – not explorers, naturalists or scientists from foreign, typically European, lands – know or think about them. The good news for Grayia is that a published study dedicated to local knowledge exists, albeit pertaining only to the Chaillu Massif of southern Gabon, and only to the Ornate African water snake G. ornata (Pauwels et al. 2002).

Known to Loumbou, Massango, Pounou and Nzebi people there as the dibomina (other local names exist too), the snake is regarded as non-venomous and as the “grandfather of all the other snakes” (Pauwels et al. 2002, p. 139). It’s widely eaten and regarded as a valued food item. Being aquatic, the snakes are most often captured in fishing nets but they’re also caught by hand in submerged burrows otherwise being investigated for catfish. In some places, G. ornata is also used to provide medical assistance to women in labour: the snake’s dried head is kept in a safe location, and water poured through it is drunk by the mother. Pauwels et al. (2002) explained that this connects the snake’s use of water as a refuge with the belief that the baby’s head will emerge faster than otherwise. Somewhat more magical beliefs in some places connect the application of fat from the snake with the attaining of improved swimming and fishing abilities.

**Caption:** it’s 2024, but we’re still at the point where you have to consult actual books made of paper to get good info on obscure animals like the snakes discussed here. Here are some (but not all) of the snake-themed books I checked while preparing this article. Image: Darren Naish.

On natural history, Grayia is known to be highly aquatic but also to be a good climber that drops into the water from overhanging branches when alarmed. It’s said to hunt underwater at night and a belief encountered in the Lunda area of Angola is that the snakes hunt in pairs (Pauwels et al. 2002). This is of special interest given suggestions that social hunting might be present in certain snake species. Otters are reported to eat Grayia trapped in nets, and the crocodiles Mecistops and Osteolaemus prey on them too (Pauwels et al. 2002). This is an impressive amount of information and an illustration of how much ethnozoological data can be collected if only researchers go to some trouble to collate and record it.

**Caption:** African crocodiles that can and do predate on grayiid snakes. At top, the slender snouted *Mecistops* (this individual, photographed in Tanzania, is supposedly *M. leptorhynchus*). Below, two different captive *Osteolaemus*, one formerly on show at Bristol Zoo and one formerly at Marwell Zoo. *Osteolaemus* is a species complex and working out which species captive specimens belong to is hard. Images: Leyo, CC BY-SA 3.0 (**original here**); Darren Naish.

Outside of ethnozoological data, Godfrey & Luiselli (2001) reported that herons and Nile monitors Varanus niloticus are also predators of these snakes and it’s been suggested that cobras – some of which forage aquatically – might also predate on them.

Where in phylogeny? Grayia has always been difficult to place phylogenetically. Though (as discussed above) conventionally lumped into the great, sprawling, traditional version of Colubridae and regarded as somewhat nondescript, it’s become more obvious as more evidence has come in that it’s unusual and distinctive. These are big and robust snakes relative to standard ‘colubrids’ and their semi-aquatic habits are consistent with views that aquatic habits might be archaic – more ‘ancestral’ – for colubroids. Hemipenis anatomy in snakes provides a great deal of phylogenetically important information and the hemipenis of Grayia differs importantly from that of most other colubroids: in many, the hemipenis is asymmetrical whereas it’s symmetrical and has a forked sulcus spermaticus in Grayia.

Add all of this together, and we have the idea of a distinct group that perhaps diverged early in evolutionary history from the lineage that includes most other colubroids. What has molecular data said on all this? The following area is a bit difficult to discuss given the competing taxonomic schemes used by different authors, so keep in mind that – from hereon – I’m using the system preferred by Zaher et al. (2009), where Colubridae is restricted to the Coluber clade, and several other supposed ‘colubrid subfamilies’ are elevated in rank.

One of the first studies to analyse caenophidian snake phylogeny using genetics – they used one nuclear and three mitochondrial genes – was published by Vidal & Hedges (2002). And… newsflash: Grayia was the sister-taxon to virtually the whole of the rest of Colubroidea (the mudsnakes or Indo-Australian water snakes – the homalopsids – were shown diverging one node further down the tree), this implying that Grayia really should be imagined as the earliest-diverging, arguably ‘most archaic’, lineage within the whole group.

**Caption:** one of the preferred consensus phylogenetic trees from Vidal & Hedges (2002). Taxa ‘traditionally’ lumped into the highly inclusive late 20th century version of Colubridae are in disparate positions about the tree (they’re marked with the image of *Coluber*, the Eastern racer), and because viperids (marked with an image of an Adder *Viperus berus*), elapids (marked with a *Naja* cobra) and atractaspidids (marked with a photo of a burrowing asp/stiletto snake) are nested *within* this version of Colubridae, that version of Colubridae is paraphyletic, and here is part of the reason for its dissolution. Within the clade that includes colubrids in the most restrictive sense (shown here as colubrines), you can see that *Grayia* (red arrow) is sister to everything else. Image: Vidal & Hedges (2002); Darren Naish.

Kelly et al. (2003), just a little later, found Grayia to be an early-diverging lineage within a clade (let’s call it Clade X) that also included calamariids (reed snakes) and colubrids (Eastern racer and kin). This isn’t totally consistent with Grayia being an ‘early diverging colubroid’, however, since Clade X was sister to a Clade Y that included natricids (keelbacks, Natrix water snakes and kin) and xenodontids (reed snakes). These clades (X and Y) need names, by the way… hint hint. In the several molecular studies incorporating Grayia following that one, Grayia was specifically found to be part of the colubrid lineage (Pinou et al. 2004), the Asian vine snake + colubrid clade (Lawson et al. 2005) or Natricidae (Kelly et al*. 2009).

This group has been called both Ahaetuliinae and Chrysopeleinae, with the latter having priority.

These diverse results are variable enough that, by around 2009-ish, there wasn’t a pinned down position on Grayia within Colubroidea, and it might have been best to regard it as incertae sedis. Zaher et al. (2009) thought that the balance of evidence made a position within Colubridae (remember: in the strict, narrowest sense) most likely. This emphatically refutes the idea that Grayia is ‘early diverging’ within Colubroidea as a whole. It’s deeply nested within the group, and in fact close to a clade generally regarded as one of the youngest within the whole assemblage.

A similar result was found by Pyron et al. (2011): a big Clade X that includes natricids and dipsadids is sister to Clade Y, and Grayia is again close to the base of Clade Y. But in a later and more comprehensive study, Pyron et al. (2013) found Grayia to be sister to the clade that includes Asian vine snakes and kin and Colubridae (again, meaning Eastern racer and kin). A similar position was also discovered by Figueroa et al. (2016) and Zaher et al. (2019).

**Caption:** a substantially simplified phylogenetic tree of colubrids and kin, based on the results of **Figueroa *et al*. (2016)**, but using (where applicable) the taxonomy proposed by **Zaher *et al*. (2009)**… though this doesn’t quite work, since those two studies find very different positions for some of the relevant lineages (for **Zaher *et al*. (2009)**, for example, hinge-toothed snakes or sibynophiines are within Colubrinae). The main point here is that grayiids are close to the restricted version of Colubridae. Images: Sibynophiinae: **Thomas Brown**, CC BY 2.0 (**original here**); Natricidae: **Orchi**, CC BY-SA 3.0 (**original here**); Pseudoxenodontidae: **Umeshsrinivasan**, CC BY-SA 3.0 (**original here**); Dipsadidae: **Geoff Gallice**, CC BY 2.0 (**original here**); Grayiidae: Kate Jackson, used with permission; Calamariidae: in public domain; Chrysopeleinae/Ahaetuliinae: **Rushenb**, CC BY-SA 4.0 (**original here**); Colubrinae: **Dawson**, CC BY-SA 2.5 (**original here**).

The conclusion has to be that, despite early indications (Vidal & Hedges 2002), Grayia isn’t especially ancient or ‘early diverging’ within Colubroidea after all. It’s instead part of a clade that includes the ‘core’ members of the whole lot, and any features that make it seem at all archaic are secondarily so and probably the result of specialisation for an unusual lifestyle.

And that about wraps everything up. A group of snakes that have long been obscure and enigmatic turn out to be reasonably well studied once you seek out the literature on them. And what were so often a sort of ‘footnote’ group that only get the briefest of mentions in discussions of colubroids can, in review texts of the future, now receive maybe a little more coverage.

For previous Tet Zoo posts on other squamates see…

In Quest of Anguids, May 2006
Pompey and Steepo, the World-Record-Holding Champion Slow-Worms, May 2007
Cambodia: now with dibamids!, May 2011
The Tet Zoo Guide to Mastigures, August 2018
The Remarkable Basilisks, May 2023
Do Lizards Really Have ‘Mite Pockets’?, March 2024
Meeting Lake Zacapu’s Garter Snake, March 2024
Ray Hoser, Number 1 Taxonomic Vandal, May 2024
The Mysterious Dibamids, May 2024
Side-Stabbing Stiletto Snakes, Redux for 2024, May 2024
The Rehabilitation of Günther’s Black Cameroonian Snake, May 2024
Ikaheka and Other ‘Palatine Draggers’, Cryptozoic Elapid Snakes of Melanesia, June 2024
Arboreal Alligator Lizards of Mesoamerica... and Beyond!, July 2024

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

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Beolens, B., Watkins, M. & Grayson, M. 2011. The Eponym Dictionary of Reptiles. Johns Hopkins University Press, Baltimore.

Chaney, T., Pauwels, O. S. G., Nagy, Z. T., Gvoždík, V., Kusamba, C., Badjedjea, G., Masudi, F. M., Akuboy, J. B., Ernst, R., Trape, J.-F., Chirio, L., Conradie, W., Keates, C., Wallach, V., Zassi-Boulou, A.-G., Vaughan, E. R. & Greenbaum, E. 2024. Phylogenetics and Integrative Taxonomy of African Water Snakes (Squamata: Colubridae: Grayia). Herpetological Monographs 38, 1-52.

Cranch, J. 1818. Appendix No. IV. A general notice of the animals taken by Mr John Cranch, during the expedition to explore the source of the River Zaire. In Tuckey, J. H. & Christen, S. Narrative of an Expedition to Explore the River Zaire, Usually Called the Congo, in South Africa, in 1816. J. Murray, London, pp. 407-419.

Figueroa, A., McKelvy, A. D., Grismer, L. L., Bell, C. D. & Lailvaux, S. P. 2016. A species-level phylogeny of extant snakes with description of a new colubrid subfamily and genus. PLoS ONE 11, e0161070.

Godfrey, G. C. & Luiselli, L. 2001. Ecological studies on a population of the water snake Grayia smythii in a rainforest swamp of the Niger Delta, Nigeria. Contributions to Zoology 70, 139-146.

Gower, D., Garrett, K. & Maddock, S. 2023. Snakes: Their Diversity, Ecology and Behaviour. Natural History Museum, London.

Kelly, C. M. R., Barker, N, P. & Villet, M. H. 2003. Phylogenetics of advanced snakes (Caenophidia) based on four mitochondrial genes. Systematic Biology 52, 439-459.

Kelly, C. M. R., Barker, N. P., Villet, M. H. & Broadley, D. G. 2009. Phylogeny, biogeography and classification of the snake Superfamily Elapoidea: a rapid radiation in the late Eocene. Cladistics 25, 38-63.

Lawson, R., Slowinski, J. B., Crother, B. I. & Burbrink, F. T. 2005. Phylogeny of the Colubroidea (Serpentes): new evidence from mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 37, 581-601.

Pauwels O. S., Toham, A. K. & Mamonekene, V. 2002. Ethnozoology of the dibomina (Serpentes: Colubridae: Grayia ornata) in the Massif du Chaillu, Gabon. Hamadryad 27, 136-141.

Pinou, T., Vicario, S., Marschner, M. & Caccone, A. 2004. Relict snakes of North America and their relationships within Caenophidia, using likelihood-based Bayesian methods on mitochondrial sequences. Molecular Phylogenetics and Evolution 32, 563-574.

Pyron, R. A., Burbrink, F. T., Colli, G. R., Montes de Oca, A. N., Vitt, L. J., Kuczynski, C. A. & Wiens, J. J. 2011. The phylogeny of advanced snakes (Colubroidea), with discovery of a new subfamily and comparison of support methods for likelihood trees. Molecular Phylogenetics and Evolution 58, 329-342.

Pyron, R. A., Burbrink, F. T. & Wiens, J. J. 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13, 93.

Spawls, S., Howell, K., Drewes, R. & Ashe, J. 2004. A Field Guide to the Reptiles of East Africa. A & C Black, London.

Vidal, N. & Hedges, S. B. 2002. Higher-level relationships of snakes inferred from four nuclear and mitochondrial genes. C. R. Biologies 325, 977-985.

Zaher, H., Grazziotin, F. G., Cadle, J. E., Murphy, R. W., Cesar de Moura-Leite, J. & Bonatto, S. L. 2009. Molecular phylogeny of advanced snakes (Serpentes, Caenophidia) with an emphasis on South American xenodontines: a revised classification and descriptions of new taxa. Papeis Avulsos de Zoologia 49, 115-153.

Zaher, H., Murphy, R. W., Arredondo, J. C., Graboski, R., Machado-Filho, P. R., Mahlow, K., Montingelli, G. G., Quadros, A. B., Orlov, N. L., Wilkinson, M., Zhang, Y.-P. & Grazziotin, F. G. 2019. Large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (Squamata: Serpentes). PLoS ONE 14, e0216148.

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Ever keen to cover more of squamate diversity – Squamata = snakes and lizards – we here look at a really interesting group of mostly Mexican lizards. They’ve led us on a merry chase with respect to their diversity, taxonomy, phylogeny and historical biogeography…

**Caption:** some *Abronia* species are strikingly coloured, as is obvious from these images. Both show Green, Mexican, Sierra de Tehuacan, or Terrestrial arboreal alligator lizards – yes, a name that’s internally contradictory – *A. graminea*. This species occurs in Veracruz, Puebla and Oaxaca in eastern and southern Mexico. Images: María Eugenia Mendiola González, CC BY-SA 4.0 (**original here**); Ismael EPM, CC BY-SA 4.0 (**original here**).

One of the several major branches on the lizard family tree is Anguimorpha, the big group that includes the fantastic monitor lizards and the spectacular gila monsters and kin. The ‘core’ group of Anguimorpha is Anguidae, the family that includes slow-worms, glass lizards and alligator lizards (and, maybe, no longer galliwasps… more on that point later).

**Caption:** I’ve mentioned in previous squamate-themed articles that the art of Alan Male was highly influential to my nascent interest in squamates and herpetology in general. Here are Male’s anguid illustrations, as featured in Philip Whitfield’s *Reptiles and Amphibians: An Authoritative and Illustrated Guide* of 1983. Galliwasps (diploglossines) have conventionally been included within Anguidae but this is now controversial, since several recent phylogenetic studies have excluded them from this group. *Ophisaurus* and *Anguis* are conventionally regarded as close relatives within the anguid clade Anguinae, but some studies find *Ophisaurus* to be paraphyletic with respect to *Anguis*. **UPDATE:** I’d missed that this is no longer an issue given that the sheltopusik is now *Pseudopus*. See comments! Images: Alan Male, from Whitfield (1983).

And, in turn, the ‘core’ anguid is Anguis, the slow worms. These are superficially snake-like, limbless anguids of Eurasia and north Africa (the Western Palaearctic), and my familiarity with the European slow worm Anguis fragilis in particular means that I’ve written about them a fair bit at Tet Zoo over the years. Slow worms and their close relatives are sorta typical of Western Palaearctic reptiles in that they’re not colourful or striking in appearance, and their biology and natural history is (comparatively) well known and substantially recorded in the literature.

**Caption:** in recent years, I’ve photographed every *Anguis* I’ve encountered. Here are a few. This is the only anguid I’ve seen in the wild, and indeed the only one we have here on the Atlantic fringes of western Europe, a place of such depauperate herpetofauna. See the links below for more on this animal. Images: Darren Naish.

But anguids aren’t just animals of the Palaearctic. They occur throughout the subtropics and tropics too, and in fact this is where the bulk of species occur… as ever, our reliance on Palaearctic forms as anchors in nomenclature and natural history knowledge is a quirk of our Eurocentrically-biased scientific history, sorry rest of humanity. Tropical anguids often are colourful and striking, and – in contrast to northerners like Anguis – their natural history and biology is typically poorly known and poorly documented.

In this article (another one rescued from the archives, and now updated and modified), we look at a very beautiful, sometimes bright green, group of mostly arboreal, mostly high-altitude anguids from Mexico and Central America: the Abronia species, sometimes dubbed arboreal alligator lizards. They’re pretty special animals, and – as ever – we don’t know as much about them as we might like.

**Caption:** Monte Cristo arboreal alligator lizard *Abronia montecristoi*, a poorly known, endangered cloud forest species from El Salvador and Honduras. It’s mostly brownish; parts of its body were described as ‘cinnamon’ by its describers. Image: Josiah Townsend, CC BY-SA 4.0 (**original here**).

A brief scientific history. Abronia has been known to science since 1828 when Deppe’s arboreal alligator lizard A. deppii (also called the Guerreran arboreal alligator lizard) was reported from Mexico by German zoologist Arend F. A. Wiegmann (1802-1841), though Wiegmann misidentified this species as part of the alligator lizard genus Gerrhonotus. ‘Deppe’ is German explorer, artist and naturalist Ferdinand Deppe (1795-1861) who collected the specimen on one of his several Mexican expeditions of the late 1820s. In 1838, British zoologist John Edward Gray (1800-1875) realized that Deppe’s species was unusual enough for its own genus and hence the name Abronia was published. Gray coined innumerable generic names across his long career but tended not to explain the etymologies behind them, and even today many have mysterious or unresolved origins. In the case of Abronia, it’s thought that he was commemorating the ‘delicate’ nature of the tail, since that’s what ‘abros’ means in Greek. ‘Delicate’ in this context was presumably a reference to its length and slenderness.

**Caption:** *Abronia deppii* photographed in the wild in 2018. These images were uploaded to iNaturalist and an important point worth making here is that records of endangered species – especially of attractive reptiles that are popular in the exotic animal trade – need to have their locality data protected, since there’s a history of unethical collectors visiting specific places to collect live animals. Image: (c) Eusebio Roldán Félix, CC BY-NC (**originals here**).

During the 1860s, 70s and 80s, new Abronia species were described from additional locations in southern Mexico in addition to Guatemala, Costa Rica and Panama. These showed both that these lizards were moderately speciose – ten had been named by the end of the 19th century – and present across essentially the whole length of Central America. Since then, additional species have come in on a fairly regular basis (the species count right now is 41) and they’ve turned out to include more than half of all species within Gerrhonotinae, the alligator lizard clade. This fact seemingly establishes Mexico and Central America as the pulsating hub of gerrhonotine evolution and diversity, with the USA being a ‘fringe’ area inhabited by immigrants that have moved north. But read on…

**Caption:** the distribution of *Abronia* as depicted by Campbell & Smith (1993). Since 1993, additional records demonstrate the presence of *Abronia* in additional locations in the far west of Honduras. In addition, the inclusion of *Mesaspis* within *Abronia* (see main text) means that the group is also present in northern Nicaragua, through part of inland Costa Rica, and in northern Panama.

Goodbye Mesaspis! During the 1990s, an effort to better understand the evolutionary history of these lizards resulted in a taxonomy whereby the more than 20 species known at the time were grouped into six subgenera, namely Abaculabronia, Abronia, Aenigmabronia, Auriculabronia, Lissabronia and Scopaeabronia (Campbell & Frost 1993). The Central American alligator lizards (Mesaspis), which are smaller, browner, more terrestrial and generally less impressive in appearance than typical Abronia species, were generally regarded as the Abronia sister-group.

**Caption:** Morelet’s alligator lizard *Abronia moreletii*, but formerly *Mesaspis moreletii*, photographed in Guatemala. Obviously, the species formerly included in *Mesaspis* look quite distinct from the most familiar members of *Abronia* (like bright green, rugose *A. graminea*), so it’s not surprising that they were previously considered generically distinct. Image: Todd Pierson, CC BY-SA 2.0 (**original here**).

However, molecular results published in 2013 and more recently have shown – a bit surprisingly – that Mesaspis and Abronia are both non-monophyletic (Pyron et al. 2013), and that taxa previously grouped in Mesaspis are scattered about the Abronia tree, grouping specifically with clades of locally occurring Abronia species (Gutiérrez-Rodríguez et al. 2020). A full discussion of what this means and what its implications are is beyond the scope of this article (as ever, I started compiling this intending it to be a very brief review), but the outcome is that the former Mesaspis species are currently subsumed into Abronia, and Mesaspis is no longer in use. Additionally, certain of those supposed Abronia subgenera are not clades (Gutiérrez-Rodríguez et al. 2020).

**Caption:** the Green, Mexican or Terrestrial arboreal alligator lizard *A. graminea* is now quite frequently encountered in the pet trade. Its popularity is not surprising in view of how amazing it looks: a local pet reptile establishment near me even has a giant photo of one on their front door. However, it’s endangered and on the IUCN Red List, and collecting for the trade has been one of the contributory factors. Seems pretty wild that you can own an endangered species as a pet…. Image: Ethan Kocak, used with permission.

Appearance and anatomy. Abronia species are often striking in appearance. Like other gerrhonotines, they usually have prominent, rugose dorsal scales on top of similarly rugose osteoderms (the osteoderms are the bony plates beneath the skin; some species have a strongly reduced osteoderm compliment), and a broad and flattened head. Some (like A. aurita and A. anzuetoi, both from Guatemala) possess short spikes around the ears. The limbs of Abronia species are ‘normal’ in proportion, as expected for animals that regularly climb, and not proportionally small as they are in other gerrhonotines. The tail is prehensile, but like other anguids they’re still able to autotomize it if necessary, with regenerated tails being shorter and less functionally effective.

   **Caption:** anatomical illustrations of the heads of various *Abronia* species, in dorsal and left lateral views, from Campbell & Smith (1993). Note how variable they are with respect to depth of the head, the rear part in particular, and in the distribution, size and number of spike-like scales dorsal to the ear opening. Three of the species here were described and named by Campbell & Smith (1993).

They’re mostly green. Some are bluish or turquoise, some are brown or red, and some have areas of yellow, orange or black on their bodies. Adding further difficulty to efforts to generalize, captive individuals of some species have been observed to change colour over a period of a few months. In some species the pigmentation forms patterns recalling patches of lichen or moss. All Abronia species are viviparous, with some giving birth to a single young, while others produce litters of up to 12 babies. Some authors describe Abronia as ovoviviparous, this being a kind of viviparity where the babies don’t emerge from shelled eggs but aren’t nourished by a placenta. Authors are still frustratingly lazy on the terminological distinction here, and I’m not sure which way we should go (I disagree with the idea that ‘viviparity’ requires the presence of a placenta).

**Caption:** captive *A. graminea* providing good views of the rugose dorsal scales and vivid green colour. *Abronia* is like many anguids in possessing a distinct ‘lateral fold’ that runs along the lower part of the body and separates the scales of the flank from those of the belly. The fold in *Abronia* is weakly developed compared to that of some other anguids, and its extent on the neck varies across species. Image: Ethan Kocak, used with permission.

Natural history notes. Field data shows that many Abronia species spend most of their lives among epiphytes, and they take shelter in bromeliads and tree holes. Some species live in tree-tops 40 m above the ground, whereas others are encountered beneath logs and rocks on the forest floor. Bogert & Porter (1967) noted that, where arboreal Abronia species occur in Oaxaca, Mexico, the sympatric Barisia and Gerrhonotus are strictly terrestrial while, in Arizona and California (where there are no Abronia species), Gerrhonotus can be found in trees. That might suggest competitive exclusion and maybe ecological competition between these taxa. Unexpectedly, an individual of A. fimbriata in Guatemala was reported swimming in a stream and repeatedly diving to the bottom. This is a reminder of the adage that animals do what they damn well please… or, anatomy is not destiny, take your pick.

**Caption:** representatives of two other gerrhonotine anguid genera that live in sympatry with *Abronia* in some places. At top: *Barisia*, specifically Chihuahuan alligator lizard *B. levicollis* photographed in Chihuahua, Mexico. At bottom: *Gerrhonotus*, specifically Pygmy alligator lizard G. *parvus* photographed in Cumbres de Monterrey National Park, Mexico. Images: Marisa Ishimatsu, from Lemos-Espinal *et al*. (2017), CC BY-SA 4.0 (**original here**); Michael Price, CC BY-NC-ND (**original here**).

As ever with unusual tropical squamates, it’s hard to recommend a single go-to literary source on this group. The most useful and comprehensive review is a 1993 paper by Campbell & Frost (1993). 23 species were known to them at the time of their review, and they grouped these into six subgenera, namely Abaculabronia, Abronia, Aenigmabronia, Auriculabronia, Lissabronia and Scopaeabronia. They noted their awareness of several additional species that were awaiting publication, and it was clear that diversity within the group was set to increase a fair bit over coming decades. As noted earlier, the number of recognized species right now is 41, recently named species including such Mexican endemics as the Sierra Morena arboreal alligator lizard A. morenica Clause et al., 2020, A. zongolica García-Vázquez et al., 2022 from Veracruz, and the Coapilla arboreal alligator lizard A. cunemica Clause et al., 2024 from Chiapas.

**Caption:** at left, portrait of Sierra Morena alligator lizard *Abronia morenica*, a species endemic to southern Mexico’s Sierra Madre de Chiapas, named in 2020. At right, depiction of *A. zongolica* from the Sierra de Zongolica in Mexico’s south-east, named in 2022. These illustrations do a good job of showing the sculpted surface texture of the scales present across the head, plus much of the body as well. Images: AMANTEDESAURIOS CC BY-SA 4.0 (originals **here** and **here**).

Sympatry among Abronia species is unusual and rare, and reported cases have proved controversial and either erroneous, or just about impossible to verify (Campbell & Frost 1993, Townsend Peterson & Nieto-Montes 1996, Pianka & Vitt 2003).

The bad news is that the very localized distribution of many Abronia species makes them vulnerable to extinction, and Campbell & Frost (1993) estimated that perhaps 13 species will become extinct in the next few decades. Indeed, some species are known from just a single specimen, or have gone undocumented for several decades (e.g., A. montecristoi from El Salvador, named in 1983), meaning that we have no information on their current status.

Incidentally, when I published the previous version of this article back in 2007, I listed A. mitchelli from Oaxaca (named in 1982) as among those species known from a single specimen. Since then, a claim has been made that at least one additional individual has been encountered in the wild, leading some authors to list it as ‘rediscovered’ (Lindken et al. 2024). A. ochoterenai from Chiapas, a bright red species named by Rafael Martín del Campo in 1939 but then considered lost, has also recently been rediscovered alive in the wild (Lindken et al. 2024).

**Caption:** *Abronia ochoterenai* was named in 1939 by Mexican herpetologist Rafael Martín del Campo y Sánchez, initially as a subspecies of an anguid regarded at the time as a *Gerrhonotus* alligator lizard (this was *G. vasconcelosii*, which was itself transferred to *Abronia* later on). Unfortunately, del Campo only gave the type locality as "Santa Rosa, Comitan", Chiapas, Mexico and at least 16 villages in the region have this name. Substantial discussion has surrounded where additional specimens might be found, and things were confused and unresolved until recently. As reported by the **HERP.MX** team in 2019, additional specimens have now been discovered, specifically at an unnamed sierra on the Atlantic slopes of southeastern Chiapas. As you can see, it’s a spectacular animal. Image: **HERP.MX**.

Complications provided by fossils. Back when I first wrote about Abronia (the aforementioned 2007), the genus didn’t have a fossil record. It was appropriate, however, to note that a few fossils appeared close to it and might ultimately prove relevant to its ancestry. Gauthier (1982) and Estes (1983) noted that Gerrhonotus mungerorum from the Miocene and Pliocene of Nebraska and Kansas – known only from its frontal bone and sometimes called Munger’s alligator lizard – resembles Abronia and might be close to it (though they also noted that it resembles the Mexican gerrhonotine Barisia). And then there’s Paragerrhonotus ricardensis, supposedly the closest relative of Abronia, from California (with some questionable specimens from Nebraska).

**Caption:** the *Gerrhonotus mungerorum* holotype frontal (anterior to the right; scale bar = 1 mm) as figured by J. Alan Holman in 1975, but here taken from the **Kansas Herpetofaunal Atlas**. What herpetologists have conventionally called ‘the frontal’ in squamates is actually the two frontal bones fused into a single unit, but people are nothing if not inconsistent across research groups.

If these fossils are indeed close to Abronia, maybe they demonstrate that it originated in the USA before moving southwards during the Pliocene, perhaps as a result of cooling conditions. Relevant here is that old cautionary tale about neglecting fossil species when looking at problems of this sort. Macey et al. (1999) produced a biogeographical analysis of gerrhonotines based on extant taxa and – because ‘Mesaspis’, Abronia, Gerrhonotus and Barisia all have distributions centred around Mexico, Texas and Central America – their area cladogram makes it look as if Mexico (or thereabouts) was the place of origin for this clade. Throw in the fossils though, and doubt arises.

Post-2007, we have a definitive fossil Abronia, a very nice skull from the Miocene Caliente Formation of California (and thus about 11 or 12 million years old), named A. cuyama by Scarpetta & Ledesma (2023). The specimen was collected in the Cuyama Valley Badlands in southern California back in the 1950s but misidentified as a Gerrhonotus. It’s part of an assemblage of small animal fossils that appear to represent an accumulation of owl pellets. Owls, we’ve learnt, are the friends of the vertebrate palaeontologist, species worldwide having contributed massively to our knowledge of small animals. Subjected to CT-scanning and described in detail, the A. cuyama skull has the wide frontal and heavily sculpted osteoderms, with a ‘vermiculate’ ornamentation, unique to Abronia, and it has osteoderm characters unique to itself and not present in any other Abronia species.

**Caption:** at left, the *Abronia cuyama* fossil (in both colour and black and white; in right lateral, left lateral, and dorsal views), a partial skull with intact cranial osteoderms. At right, map from Scarpetta & Ledesma (2023) showing how distant the collection locality of *A. cuyama* is from the modern range of *Abronia*. Does this mean that *Abronia* previously occurred right across the south-western USA and northern and western parts of Mexico? Images: Scarpetta & Ledesma (2023).

What does its presence in California mean for the prehistoric distribution and origin of this group? That’s not entirely clear, mostly because – surprisingly – the specimen was found to be deeply nested within Abronia when included within a phylogenetic analysis. It’s not an outgroup to the rest of Abronia, and thus can’t be used as evidence for more northerly origins. Instead, maybe it shows that Abronia was ancestrally present throughout southern California and, presumably, across the Sierra Madre Occidental of Mexico’s north before becoming extinct in both regions. If that’s true, habitats like those inhabited by Abronia today were presumably more widespread during the later parts of the Miocene. Another possibility is that A. cuyama represents a singular expansion of the group northwards. We can’t really say in the absence of additional fossils. The fact that an owl transported the specimen could mean that the animal wasn’t local when alive, but this appears contradicted by the idea that the sort of owl responsible for the pellets was most likely a ground-hunting, relatively sedentary species (Scarpetta & Ledesma 2023).

**Caption:** so that’s another squamate-themed article from the Tet Zoo archives updated and rescued. This one originally appeared at Tet Zoo ver 2 (the ScienceBlogs years) in 2007. As you can see from these screengrabs, the illustrations originally included were very different and I previously included a short diversion on phylogeneticist Jacques Gauthier, his squamate-themed research, and the books visible in the background of the photo I’d featured. I couldn’t do that this time around as I couldn’t find a version of the photo that’s of sufficient resolution.

Goodbye, Abronia. And that about sums up everything I wanted to say. Anguids, and anguimorphs as a whole, are among the most charismatic and popular of lizards and there’s lots more about them I need to publish… my galliwasp writings, promised since about 2007, are now waaay overdue. Abronia specifically is now a bit of a superstar and also, increasingly, a flagship genus in terms of conservation interest, so I’m pleased to have covered it again. These lizards are beautiful and fascinating, still so mysterious and enigmatic, and perpetually at risk from the destruction and deterioration of their habitat.

For previous Tet Zoo posts on other squamates see…

In Quest of Anguids, May 2006
Pompey and Steepo, the World-Record-Holding Champion Slow-Worms, May 2007
Cambodia: now with dibamids!, May 2011
The Tet Zoo Guide to Mastigures, August 2018
The Remarkable Basilisks, May 2023
Do Lizards Really Have ‘Mite Pockets’?, March 2024
Meeting Lake Zacapu’s Garter Snake, March 2024
Ray Hoser, Number 1 Taxonomic Vandal, May 2024
The Mysterious Dibamids, May 2024
Side-Stabbing Stiletto Snakes, Redux for 2024, May 2024
The Rehabilitation of Günther’s Black Cameroonian Snake, May 2024
Ikaheka and Other ‘Palatine Draggers’, Cryptozoic Elapid Snakes of Melanesia, June 2024

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

Refs - -

Bogert, C. M., & Porter, A. P. 1967. A new species of Abronia (Sauria, Anguidae) from the Sierra Madre del Sur of Oaxaca, Mexico. American Museum Novitates 2279, 1-21.

Campbell, J. A. & Frost, D. R. 1993. Anguid lizards of the genus Abronia: revisionary notes, descriptions of four new species, a phylogenetic analysis, and key. Bulletin of the American Museum of Natural History 216, 1-121.

Estes, R. 1983. Handbuch der Paläoherpetologie. Part 10A. Sauria Terrestria, Amphisbaenia. Gustav Fischer Verlag, Stuttgart, New York.

Gauthier, J. A. 1982. Fossil xenosaurid and anguid lizards from the Early Eocene Wasatch Formation, southeast Wyoming, and a revision of the Anguioidea. Contribution to Geology of the University of Wyoming 21, 7-54.

Gutiérrez-Rodríguez, J., Zaldívar-Riverón, A., Solano-Zavaleta, I., Campbell, J. A., Nelsi Meza-Lázaro, R., Flores-Villela, O., Nieto-Montes de Oca, A. 2021. Phylogenomics of the Mesoamerican alligator-lizard genera Abronia and Mesaspis (Anguidae: Gerrhonotinae) reveals multiple independent clades of arboreal and terrestrial species. Molecular Phylogenetics and Evolution 154, 106973.

Lindken T., Anderson, C. V., Ariano-Sánchez, D., Barki, G., Biggs, C., Bowles, P., Chaitanya, R., Cronin, D. T., Jähnig, S. C., Jeschke, J. M., Kennerley, R. J., Lacher, T. E., Luedtke, J. A., Liu, C., Long, B., Mallon, D., Martin, G. M., Meiri, S., Pasachnik, S. A., Reynoso, V. H., Stanford, C. B., Stephenson, P. J., Tolley, K. A., Torres-Carvajal, O., Waldien, D. L., Woinarski, J. C. Z. & Evans, T. 2024. What factors influence the rediscovery of lost tetrapod species? Global Change Biology 30, 1-18.

Macey, J. R., Schulte, J. A., Larson, A. Tuniyev, B. S., Orlov, N. & Papenfuss, T. J. 1999. Molecular phylogenetics, tRNA evolution, and historical biogeography in anguid lizards and related taxonomic families. Molecular Phylogenetics and Evolution 12, 250-272.

Pianka, E. R. & Vitt, L. J. 2003. Lizards: Windows to the Evolution of Diversity. University of California Press, Berkeley.

Pyron, R. A., Burbrink, F. T. & Wiens, J. J. 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13, 93.

Scarpetta, S. G. & Ledesma, D. T. 2023. A strikingly ornamented fossil alligator lizard (Squamata: Abronia) from the Miocene of California. Zoological Journal of the Linnean Society 197, 752-767.

Townsend Peterson, A. & Nieto-Montes, A. 1996. Sympatry in Abronia (Squamata: Anguidae) and the problem of Mario del Toro Aviles’ specimens. Journal of Herpetology 30, 260-262.

Whitfield, P. 1983. Reptiles and Amphibians: An Authoritative and Illustrated Guide. Longman Group Ltd, Harlow, UK.

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It’s time to start preparing for TetZooCon – our annual Tetrapod Zoology Convention – and oh boy is a lot of stuff happening this year…

Once again, TetZooCon occurs at Bush House, Kings College, London (though in different parts of the building from previous events). This year, TetZooCon is a late September thing. We kick off with an evening discussion on Friday September 27th, and then continue across the weekend of September 28th and 29th. Doors open at 10am on both days and we’re generally done by 6pm. **Tickets and further info can be found here.**

As ever, we have a busy schedule of talks and events. I should note up-front that Mesozoic dinosaurs will be a prominent theme this year. Stalls will be present throughout, items on sale including art, stickers, plushies, animal figures, models, books and more… frankly, a frightening amount of merch, palaeoart and wildlife art. We’ll also have book-signing events – I, for one, am selling copies of Ancient Sea Reptiles and a few of my other books – and a silent auction will be happening too. We’re also hosting a palaeoart-themed art exhibition, and of course our patented* palaeoart workshop occurs on and off throughout the weekend.

I use the term loosely.

Caption: if you’ve attended TetZooCon before, you’ll know that – oh boy – a lot of amazing stuff is on sale at the numerous stalls. The montage here includes (upper left) models, fridge magnets and more from Jed Taylor and Ruadhrí Brennan’s stall; prints and more on sale at Lee Brown’s stall (Lee runs DailyDinoSketch, and sells art, stickers, rock art and more here at Etsy); and some of the All Yestersdays figures made by Sam St Leger and available from the Splendid Edition shop on Etsy. Images: Ruadhrí Brennan, George Lathouris, Darren Naish.

We also hope to show a movie across lunchtime on Sunday, specifically the feature-length documentary Why Dinosaurs?, produced by Tony Pinto. Why Dinosaurs? has, so far in the UK, only been screened at special events (in London, Bristol and Sheffield, and in Lyme Regis for the 2024 Fossil Festival), so it’s a big deal that we’re able to show it.

Speakers 1. Talks kick off on Saturday morning with me; I’ll be discussing an unusual episode in the history of dinosaur research. We then welcome pioneering palaeoartist, author and researcher Greg Paul. Yes, Greg Paul in person. For those of you who don’t know, Greg was instrumental in constructing the view of dinosaur life appearance we have today, his technical, high-fidelity skeletal and muscular illustrations being among the first widely available images to properly depict dinosaurian anatomy. He has coupled this tradition of rigorous anatomical reconstruction with efforts to imagine Mesozoic dinosaurs in life, and his black-and-white drawings and colour paintings include some of the most influential illustrations in the entire history of palaeoart.

**Caption:** author, researcher, artist and publishing scientist Greg Paul will be speaking at **TetZooCon 2024**, and will also be participating in discussion events and more. Images: (c) Greg Paul, from **The Science and Art of Gregory S. Paul**.

  **Caption:** Greg Paul’s books include some of the most influential volumes yet published on dinosaurs; I count 1988’s *Predatory Dinosaurs of the World* as among the formative of works I’ve read. In the 21st century, Greg has published a series of field guides through Princeton University Press. We hope to have these works on sale at TetZooCon 2024.

Later on Saturday, we welcome Natalie Lawrence, who’ll be talking about ‘Making Monsters’. Natalie’s book Enchanted Creatures: Our Monsters and Their Meanings is set to appear later this year, though I’m fairly confident that copies won’t be on hand in time for her TetZooCon talk, alas. Angela Julian, coordinator of Amphibian and Reptile Groups of the UK, will also join us for her talk ‘Revealing the Secrets of the Vanishing Viper’.

**Caption:** we welcome author and researcher Natalie Lawrence as a first-time speaker at **TetZooCon 2024**. Natalie’s previous books include (with Paco Calvo) *Planta Sapiens: Unmasking Plant Intelligence* (2022) and *Feathers and Eggshells, The Bird Journal of a Young London Girl* (2005). Images: (c) **Natalie Lawrence**.

A lunchtime discussion revolving around the Apple TV+ / BBC Studios series Prehistoric Planet will also happen, as will an on-stage discussion relating to dinosaur life appearance and palaeoart, involving Greg Paul, John Conway, and myself.

Speakers 2. Moving to Sunday, our Mesozoic dinosaur theme remains apparent thanks to talks from Kai Caspar (‘How Smart was T. rex? Current Debates in Dinosaur Neurology and Cognition’) and David Hone, who’ll be speaking about dinosaur behaviour in lieu of his soon-to-be-published Princeton University Press book. Princeton themselves will have a stall at the meeting and we hope – though this can’t yet be confirmed – that they’ll have advance copies of Dave’s book on hand. It’s illustrated throughout by Gabriel Ugueto (whose other engagements mean that he’s unfortunately unable to attend. We tried to get him!). Princeton have also published Greg Paul’s recent books as well as my own Dinopedia, so it’s likely that those works will be present. If so, we’ll arrange signing events.

**Caption:** dinosaur and pterosaur research Dr David Hone will be speaking at **TetZooCon 2024**; his talk is titled ‘Uncovering Dinosaur Behaviour’ and is connected to his soon-to-be-published book of the same name. Images: Dave Hone/Princeton University Press.

Later on Sunday, Katrina van Grouw (The Unfeathered Bird, Unnatural Selection) will be talking about progress on The Unfeathered Bird II. If you’ve seen any of the new illustrations that Katrina has been preparing, you’ll know that her plans involve extending her thoughts well beyond the limits of Aves as we know the group today…

**Caption:** artist, author and researcher Katrina van Grouw is probably best known today for her spectacular *The Unfeathered Bird* of 2013 (which I reviewed **here**, though I can’t see that article as it’s paywalled to me). *TUB* is out of print today and only available at extremely high prices. So it’s great news that a second edition is currently in prepration — come to **TetZooCon 2024** and hear how it’s coming along! Images: (c) Katrina van Grouw.

I’m very pleased to report that we’ll also be joined by TV presenter, photographer, author, artist and conservationist Chris Packham, who’ll be talking about Nextinction. Chris is an outstanding naturalist and natural historian and – for those of you outside the UK who might not know this – he’s one of the most prominent and respected people working in the promotion of conservation and natural history today.

**Caption:** Chris Packham CBE is best known to British people of a certain age for his association with the BBC’s *The Really Wild Show* (1986-1995). Today, he co-presents the BBC Natural History Unit’s flagship series *Springwatch*. Chris also has associations with a large number of conservation and wildlife organisations, including the RSPB, Butterfly Conservation, the Wildlife Trusts, Hampshire Ornithological Society and others. Image: (c) Jo Charlesworth.

Also on Sunday, we’ll once again be hosting cosplay and an on-stage competition will be led by my daughter Hel. Sunday evening will end with our regular quiz, featuring – as usual – a range of excellent prizes. So, that’s a lot of stuff; we hope that attendees enjoy it.

TetZooTour! One thing above all others will make TetZooCon 2024 unusual relative to its predecessors, and that’s that – for the first time ever – we’re leading a coach tour, a TetZooTour, of select sites about southern England. This is a separately ticketed event from TetZooCon proper and its cost includes hotel accommodation, morning and evening meals, and entry to the respective venues. I’ll be going along as a guide, as will other organisers. I’ve worked as a tour guide in the past and will be providing commentary on the journey.

We board our coach at 10am on Monday September 30th (in Surrey Street, adjacent to Bush House) before disembarking for the Horniman Museum in Forest Hill, London, for a special visit to the Dinosaur rEvolution exhibit before it goes off show. The images and models featured in Dinosaur rEvolution are based on the art and imagery of TetZooCon regular Luis Rey, and I’m happy to say that Luis will be accompanying us during our time at the Horniman.

We then set off to the English south-west, first staying overnight in Bournemouth, Dorset, before driving on Tuesday October 1st to historic Lyme Regis. Here, we’ll spend time at Lyme Regis Museum – built on the site formerly occupied by Mary Anning’s house – and will also visit the shore for fossil collecting. We’ll be guided by Lyme Regis Museum staff Natalia Jagielska, Paul Davis and Kieran Satchell. At this point in the year, it’s difficult to know what conditions will be like in early October, but the general forecast is that it’s due to be somewhere around 14° C and with a possible chance of rain. Keep this in mind.

**Caption:** Lyme Regis remains ground zero for British palaeontological research, and it’s a beautiful place with a unique charm. Lyme Regis Museum is an unusual building with a good amount of interesting local material on show, and it’ll be great to visit it during our tour. I can’t guarantee that we’ll have time to visit the Mary Anning statue, shown here during May 2024, but we’ll see. Images: Darren Naish.

  **Caption:** East Beach, Lyme Regis, during May 2024. We won’t be exploring the shore directly beneath the cliffs – that’s dangerous! – but will be visiting at least part of the shore, time and conditions permitting. Image: Darren Naish.

Once done at Lyme Regis, we head east for a second overnight stay, this time at Folkestone in Kent, and we then disembark on Wednesday October 2nd for Howletts Wild Animal Park, near Canterbury. Howletts is home to African elephants, rhinos, tapirs, gorillas, African wild dogs, dholes, European wolves, spectacled bears, silver-cheeked gibbons and much, much more. And that’s the end: we leave at 3pm, and then return to our drop-off point in central London, hopefully by 5pm. And that’s it. Hopefully, things will work out and it’ll be a great trip.

**Caption:** images from a 2014 trip I took to Howletts. Their selection of bovids, canids and primates is really impressive, and they hold the largest herd of African elephants (all *Loxodonta africana*) in the UK and possibly in Europe. Images: Darren Naish.

Tickets for TetZooCon 2024, including for the tour, are already selling fast. GO HERE to buy tickets now (there are no physical tickets: you’re added to a list and everything is electronic). It should be a great event with a great crowd, and we look forward to seeing you in London for what might be the biggest and best TetZooCon so far. As ever, we encourage sharing of things as they happen on social media, and a full report of everything will appear here as and when time allows.

For previous articles on other TetZooCons, see…

The events of TetZooCon 2014, July 2014
TetZooCon 2015 Is On, July 2015
The Events of TetZooCon 2015, November 2015
Coming Soon: TetZooCon 2016, September 2016
The Day After TetZooCon, October 2016
The Fourth TetZooCon, September 2017
The TetZooCon of 2017, October 2017
Reasons to Attend TetZooCon 2018, September 2018
TetZooCon 2018: Best TetZooCon So Far, October 2018
Announcing TetZooCon 2019 – the Biggest Yet, August 2019
Final Call For TetZooCon 2019, October 2019
The Sixth TetZooCon, October 2019
TetZooCon 2020 + Zoom = TETZOOMCON 2020, December 2020
TetZooMCon 2020 an Unbridled Success, December 2020
Cronch Cats, Beasts of Gévaudan, Dinosauroids, Mesozoic Art and Much More: TetZooMCon 2021 in Review, September 2021
The Ninth and Largest of the Tetrapod Zoology Conventions, December 2022
Announcing the 10th Tetrapod Zoology Convention, October 2023
At TetZooCon 2023, November 2023
TetZooCon 2023 the report, November 2023

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It’s time once more to visit the amazing world of squamates, and again we’re looking at snakes. Today: the extremely obscure Small-eyed or Ikaheka snake of New Guinea and some of the surrounding islands. What’s the deal with this unusual animal?

**Caption:** a very glossy, clean Small-eyed snake photographed in Karkar Island, New Guinea in 2010. There’s an almost iridescent sheen to some of the scales. Image: Wolfgang Wüster, used with permission.

Known technically as Micropechis ikaheka, the Small-eyed snake is stocky and medium-sized, reaching 2 m in total. Its unusually small eyes are among its most distinctive features, and indeed it’s sufficiently unusual in anatomy that it’s the only (currently) recognized member of its genus. Incidentally, the name ‘Small-eyed snake’ is also sometimes used for Cryptophis nigrescens, an east Australian species more commonly called the Eastern small-eyed snake to remove confusion.

Little has been published on Micropechis in total but – unlike many of the snakes covered at Tetrapod Zoology in the past – comparatively little has been published on it since I first wrote about, that being back in 2010 when it was covered at ver 2 (the wayback version of that article is here).

**Caption:** yes, this is another ‘old’ Tet Zoo article – again from ver 2, the ScienceBlogs years – that I’ve now rescued and resuscitated. At right: I’m a big fan of the work and writings of Dr Mark O’Shea; all of his books are excellent. This is the cover of the second, 2011 edition of his *Venomous Snakes of the World*, and… yes it does include a section on *Micropechis*. As ever, good images of the cover online are hard to find, so this is a photo of my own (signed) copy.

Some history and some natural history. Micropechis had its first outing in the scientific literature in 1830 when French surgeon, naturalist and zoologist René Primevère Lesson (1794-1849) described it as a new species of Coluber, initially using the spelling Coluber ikaheca. Lesson was describing specimens collected on his global voyage of 1822-1825 aboard the sailing ship La Coquille, and his visit to New Guinea and the nearby islands is best known for his observations of birds-of-paradise, for he was seemingly the first European scientist to see these birds alive in their natural habitat.

**Caption:** the original plate depicting this species, produced to accompany Lesson’s initial description of 1830. I haven’t translated the description’s text from the original French, but it looks like this particular specimen was plain in colour and markings relative to others.

The specific name for this snake (that is, the second part of its scientific binomial) – ‘Ikaheka’ – means ‘land eel’ in one of the local Papuan dialects, and this apparently refers to the fact that it’s sometimes associated with streams and other damp habitats (O’Shea 2011). After Lesson’s description, there existed disagreement on what sort of snake the Ikaheka really was, some herpetologists of the late 1800s realizing that its front-fanged condition made it no Coluber, but instead perhaps a viper or sort of cobra. In 1896, George Boulenger realized that it was unusual enough to warrant its own genus, and thus Micropechis was coined. It has been asserted that this must be pronounced ‘microp-echis’, and not ‘micro-pechis’ (Warrell et al. 1996).

**Caption:** live *Micropechis*, encountered in the field in Karkar Island, New Guinea, in 2010. One of the reasons that snakes (and other reptiles) have such clean, glossy scales most of the time is that their scales have a self-cleaning, dirt-shedding micro-ornamentation. Image: Wolfgang Wüster, used with permission.

Micropechis is smooth-scaled, light brown overall, and marked with transverse reddish bands. It’s secretive, nocturnal or crepuscular, and hunts for reptiles, frogs and mammals in cluttered rainforest and wetland habitats. O’Shea (2011) noted that it often hides in piles of discarded coconut husks, and that snakes including Candoia ground boas and other Micropechis individuals are among its prey. It’s oviparous. It’s also regarded as dangerous to humans and several fatalities are on record. Most recorded bites have involved local swelling, myalgia, systemic bleeding and the passing of dark urine (Warrell et al. 1996).

The other Micropechis. There’s a reason why I said that M. ikaheka is the only currently recognized member of its genus. This is because Boulenger named a second species – M. elapoides – in the above-mentioned text of 1896. This second species was from the Solomon Islands and had originally been included in the genus Hoplocephalus (today restricted to the Australian Broad-headed snake H. bungaroides and kin). Some later authors went further in terms of making this species an ally of M. ikaheka, even regarding it as a subspecies of M. ikaheka (Loveridge 1948).

**Caption:** a live Solomons coral snake *Salomonelaps par* photographed in the field in 2015. This photo was uploaded to (and taken from) iNaturalist, which I encourage people to use themselves (I do, albeit not as regularly as I should). Image: jqrichmond, CC BY-NC (**original here**).

However, the Solomon Islands snake differs notably from Micropechis proper in the anatomy of its teeth, palatine bones, scalation and hemipenes and – according to McDowell (1970) – appears more closely related to the Solomons coral snake Salomonelaps par and the Australian bandy-bandy snakes (Vermicella). For those reasons, McDowell gave it its own genus – Loveridgelaps – in 1970, this name honouring British zoologist Arthur Loveridge, best known for his work on snakes and lizards. The species is sometimes called the Solomon or Solomons small-eyed snake and is even more obscure than the one we’re mostly looking at here. Micropechis and Salomonelaps* both remain poorly studied, but the view that they’re distinct at the generic level has been followed in the post-McDowell literature.

And yes, the name is indeed spelt ‘Salomonelaps’, not ‘Solomonelaps’. McDowell (1970) only said of this spelling that it referred to “the geographic region to which the genus is confined” (p. 146), so we must assume that he was referring to the historic ‘Islas Salomón’ term for the Solomon Islands.

Whence within Elapidae? Micropechis is certainly an elapid, possessing the proteroglyphous (front-fanged) anatomy of this group as well as the characteristic palatal and facial bone configuration, head scalation and much more. Within this large group, its anatomy, distribution and molecular traits show that it’s part of the clade that includes the Australasian taipans, tiger snakes and kin in additional to the viviparous or true sea snakes (as opposed to the laticaudine sea kraits, which are less specialized for marine life). This group is generally termed Hydrophiinae, though Sanders & Lee (2008) argued that Oyxuraninae might be used for it instead.

**Caption:** McDowell’s 1970 diagrams of a skinned head and skull of museum specimens of *Micropechis* in the collections of the American Museum of Natural History in New York. A few things are of interest. The venom gland (vg) is the large, convex area covering the cheek region; both the premaxillary salivary gland (pg) and nasal gland (ng) are visible on the snout. Image: McDowell (1970).

A diversion on palatine erection vs palatine dragging. In a moderately influential article of 1970, Samuel B. McDowell of Rutgers University in New Jersey argued that elapids possess two distinct ways of erecting their fangs. In one group – it includes all African, Asian and American elapids in addition to laticaudine sea snakes and Parapistocalamus of Bougainville Island – McDowell proposed that the vaguely triangular palatine bone is pulled vertically as the maxilla (the bone bearing the fang) is erected to deploy the fang. The palatine thus (so McDowell thought) ends up with a strong anterodorsal inclination during fang erection (meaning that it projects forwards and upwards at a diagonal slant). He termed the snakes that do this the ‘palatine erectors’ (McDowell 1970).

**Caption:** at left, from top to bottom, (A) the palatine and pterygoid bones (with the palatine, the more anterior of the two, at right) of a cobra, both in medial (inner) view, (B) the left palatine and anterior part of the pterygoid of a taipan in medial and (C) lateral view, and likewise for a death adder in (D) medial and (E) lateral views. The difference in shape of the cobra palatine from those of the taipan and death adder is obvious. At right, a ‘palatine erector’ (a cobra) at the top, and ‘palatine dragger’ (a taipan) below, these diagrams showing palatine movement as hypothesized by McDowell (1970). The palatine is the bone inboard of the fang-bearing maxilla (mx). It’s raising up into a diagonal position in the ‘erector’ and is pointing forwards at semi-horizontal orientation in the ‘dragger’. But be sure to check the main text! Images: Deuful & Cundall (2010).

Something different, McDowell proposed, happens in hydrophiines. In these animals, the palatine is bar-like, rather than subtriangular. As the maxilla is erected to deploy the fang, the palatine – McDowell thought – remains subhorizontal and in line with the similarly-shaped pterygoid to its posterior. Connection between the maxilla and palatine then meant that the latter is dragged anteriorly, such that its tip ended up being closer to the snout tip than is the leading edge of the maxilla (McDowell 1970). He termed these snakes the ‘palatine draggers’.

**Caption:** lest we forget, we’re actually here to talk about *Micropechis*. Here’s an especially useful, detailed shot of the face. The small, dark eyes are a familiar feature, but note also the slightly downturned snout (suitable for a snake that burrows in leaf litter, under logs and so on) and convex cheek regions (that are in keeping with moderately large venom glands). Image: Wolfgang Wüster, used with permission.

But here’s the problem. It turns out that the bones of live animals don’t behave or operate in the way you might think from museum specimens alone, dry skulls especially. McDowell didn’t precisely describe how he examined cranial kinesis in elapids, but it seems from the context of his work that he was manipulating skulls and/or skinned specimens, and not looking at the performance of live snakes. In a series of recent studies, Alexandra Deufel and David Cundall examined elapid skull function by filming live snakes as they feed, and then used these data to work out what’s happening with the skull bones (Deuful & Cundall 2003, 2006, 2010).

The bad news is that McDowell’s hypotheses of ‘palatine erection’ and ‘palatine dragging’ don’t really stand up, in part because the palatine and other bones don’t act as linked ‘pullers’ and ‘pushers’ of other bones. Instead, it’s the contractions of the protractor and levator pterygoidei muscles that are the main drivers of movement here: as the maxilla is protracted, the palatine moves anterolaterally (forwards and sideways). For McDowell’s ‘palatine erectors’, the palatine is indeed elevated as he proposed (Deuful & Cundall 2003), albeit not only because it’s attached to the maxilla as McDowell thought. In McDowell’s supposed ‘palatine draggers’, a strong ligamentous connection between the maxilla and palatine makes the palatine flex laterally (outwards) at its joint with the pterygoid, and ventrally (downwards) as the maxilla is protracted (Deuful & Cundall 2010). The palatine isn’t ‘dragged’ anteriorly as McDowell thought. Both ‘erectors’ and ‘draggers’ end up with a ventrally convex palatine-pterygoid bar during protraction.

**Caption:** diagram from Deuful & Cundall (2003), showing a ‘palatine erector’ elapid undergoing protraction of its right maxilla and palatine-pterygoid bar, the long axes of the palatine and pterygoid being shown via the intersecting lines. One thing that I haven’t discussed in this article is that the palatine rotates about its long axis. Hey, I can’t do everything. Image: Deuful & Cundall (2003).

What this means, ultimately, is that so-called palatine erectors and draggers don’t have the substantial differences in palatine movement that McDowell proposed. In any case, the palatine isn’t all that important in prey apprehension and swallowing, since it’s the pterygoid to its posterior that does the bulk of the work in ratcheting back and forth and ‘walking’ objects toward the throat. And if ‘erectors’ and ‘draggers’ aren’t performing the fundamentally different actions that McDowell thought they were, we really should give up on those terms (Deuful & Cundall 2010). I’ve continued to use them here because I’m discussing these hypotheses within a historical context.

But… come on, whence Micropechis? Anyway, the whole reason for this long discussion about palatine erectors and palatine draggers is to say that Micropechis has, historically, been considered an archaic member of the ‘palatine dragger’ assemblage. DNA-based studies that include Micropechis have appeared in print ever since the late 1990s, and they mostly find what was already suspected on the basis of anatomy: Micropechis is an early-diverging hydrophiine, outside the clade that includes Australasian taipans, tiger snakes and kin, and the viviparous or true sea snakes (Sanders & Lee 2008, Sanders et al. 2008, Pyron et al. 2013, Figueroa et al. 2016, Strickland et al. 2016).

**Caption:** highly simplified elapid cladogram (based mostly on **Pyron *et al*. (2013)**), shown to emphasize that *Micropechis* is closer to Australian viviparous elapids and true sea snakes than are sea kraits and Old World elapids like cobras. Images: *Micrurus*, public domain; *Naja*, kalmalnv, CC BY 3.0 (**original here**); *Laticauda*, Jens Petersen, CC BY-SA 3.0 (**original here**); *Micropechis*, Wolfgang Wüster (used with permission); *Pseudechis*, Smacdonald, CC BY 3.0 (**original here**); *Oyxuranus*, AllenMcC., CC BY-SA 3.0 (**original here**); *Hydrophis* Rasmussen *et al*. (2011), CC BY 2.5 (**original here**).

However, variations in results include the grouping of Micropechis with the whipsnake genus Demansia (Keogh 1998), and the recovery of both genera in a clade that also includes taipans (Oxyuranus), brown snakes (Pseudonaja) and black snakes (Pseudechis) (Scanlon & Lee 2004). Most recently, Micropechis has tended to group with whipsnakes and the New Guinean Toxicocalamus (Pyron et al. 2013, Figueroa et al. 2016). Incidentally, some Toxicocalamus species look superficially much like Micropechis, so much so that certain Toxicocalamus specimens were initially misidentified as individuals of Micropechis (O’Shea et al. 2018).

**Caption:** *Toxicocalamus* specimens can look quite similar to *Micropechis*. This montage, from O’Shea *et al*. (2018), shows museum specimens of (A, A') *T. ernstmayri*, (B, B') *T. grandis* and (C, C') a pale individual of *Micropechis ikaheka*. The colour coding of the head scales shows that *Micropechis* is similar to *Toxicocalamus* in having six supralabials (orange), one anterior temporal (yellow), and two posterior temporals (blue), but differs from it in possessing a temporolabial (red). Image: O’Shea *et al*. (2018).

One thing worth emphasizing here is that several of the hydrophiines mentioned or discussed so far are not Australian like good, classic terrestrial hydrophiines, but are Melanesian: that is, they inhabit New Guinea, the Solomon Islands, Fiji and other islands of the region. That goes for the aforementioned Micropechis, Salomonelaps, Loveridgelaps and Toxicocalamus, but also for the Fiji snake Ogmodon vitianus and the New Guinean crowned snakes (Aspidomorphus). In addition, these are generally all cryptozoic: that is, they spend a lot of time in leaf litter, and under logs and rocks.

**Caption:** Melanesia – originally named on the basis of its dark-skinned people – includes New Guinea in the west, Fiji in the east, and also the Solomon Islands, New Caledonia, Vanuatu and Nauru. The region has a fascinating reptile fauna that includes unusual elapids, giant geckos, far-flung iguanians and remarkable skinks. Image: Oceania UN Geoscheme, CC BY 3.0 (**original here**).

What’s implied by the distribution and anatomy of these snakes is that they represent a sort of ‘ancestral grade’ for hydrophiines, and that the Australian terrestrial lineages and true sea snakes evolved from snakes of this sort. If this is correct, hydrophiines have their roots in the wet tropical forests of Melanesia, in cryptozoic snakes that are quite different ecologically from classic Australian hydrophiines (let alone sea snakes), and that they moved into Australia from the north. Here I want to mention in passing that the idea of hydrophiines – yes, all of them – originating from marine forms has been mentioned more than once.

At the moment, I think this this scenario (of Melanesian, cryptozoic ancestry) still appears broadly supported, though works finding whipsnakes and such to be within that ‘ancestral grade’ do suggest that things are more complex. Maybe there were multiple independent invasions of Australia, or maybe there was repeated exchange between Melanesia and Australia. Exactly that was promoted by Strickland et al. (2016). Also on the subject of hydrophiine evolution, molecular clock estimates indicate that the entire radiation is young, the divergence between Laticauda sea kraits and hydrophiines happening about 12 million years ago, in the middle of the Miocene. The 140 or so oviparous and viviparous terrestrial hydrophiines and 64 or so species of sea snake all emerged very rapidly between about 10 and 6 million years ago (Sanders et al. 2008), in the late Miocene. This is a geologically young, explosively successful radiation, and there’s a lot to say about it.

**Caption:** two different *Micropechis* specimens from Karkar Island off the north-east coast of Papua New Guinea, showing some of the variation in striping and colour present in this species. Warrell *et al*. (1996) noted that the name ‘tiger snake’ is used for the species in their study region. The animal at left was 1.5 m long; the one at right 1 m long. An especially darkly pigmented head is a common feature of terrestrial hydrophiines. Images: Warrell et al. (1996).

For now, we end here, and I’m pleased to have ‘rescued’ another Tet Zoo squamate-themed article from the archives. For previous Tet Zoo posts on snakes and other squamates see…

Cambodia: now with dibamids!, May 2011
The Tet Zoo Guide to Mastigures, August 2018
The Remarkable Basilisks, May 2023
Do Lizards Really Have ‘Mite Pockets’?, March 2024
Meeting Lake Zacapu’s Garter Snake, March 2024
Ray Hoser, Number 1 Taxonomic Vandal, May 2024
The Mysterious Dibamids, May 2024
Side-Stabbing Stiletto Snakes, Redux for 2024, May 2024
The Rehabilitation of Günther’s Black Cameroonian Snake, May 2024

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

Refs – -

Deufel, A. & Cundall, D. 2003. Prey transport in “palatine-erecting” elapid snakes. Journal of Morphology 258, 358-375.

Deufel, A. & Cundall, D. 2006. Functional plasticity of the venom delivery system in snakes with a focus on the poststrike prey release behavior. Zoologischer Anzeiger 245, 249-267.

Deufel, A. & Cundall, D. 2010. Functional morphology of the palato-maxillary apparatus in “palatine dragging” snakes (Serpentes: Elapidae: Acanthophis, Oxyuranus). Journal of Morphology 271, 73-85.

Keogh, J. S. 1998. Molecular phylogeny of elapid snakes and a consideration of their biogeographic history. Biological Journal of the Linnean Society 63, 177-203.

Loveridge, A. 1948. New Guinean reptiles and amphibians in the Museum of Comparative Zoology and United States National Museum. Bulletin of the Museum Comparative Zoology, Harvard 101, 305-430.

McDowell, S. B. 1970. On the status and relationships of the Solomon Island elapid snakes. Journal of Zoology 161, 145-190.

O’Shea M. 2011. Venomous Snakes of the World. New Holland Publishers, London.

O’Shea, M., Herlihy, B., Paivu, B., Parker, F., Richards, S. J. & Kaiser, H. 2018. Rediscovery of the rare Star Mountains Worm-eating Snake, Toxicocalamus ernstmayri O’Shea et al., 2015 (Serpentes: Elapidae: Hydrophiinae) with the description of its coloration in life. Amphibian & Reptile Conservation 12, 27-34.

Pyron, R. A., Burbrink, F. T. & Wiens, J. J. 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13, 93.

Scanlon, J. D. & Lee, M. S. Y. 2004. Phylogeny of Australasian venomous snakes (Colubroidea, Elapidae, Hydrophiinae) based on phenotypic and molecular evidence. Zoologica Scripta 33, 335-366.

Sanders, K. L. & Lee, M. S. Y. 2008. Molecular evidence for a rapid late-Miocene radiation of Australasian venomous snakes (Elapidae, Colubroidea). Molecular Phylogenetics and Evolution 46, 1165-1173.

Sanders, K. L., Lee, M. S. Y., Foster, R. & Keogh, J. S. 2008. Molecular phylogeny and divergence dates for Australasian elapids and sea snakes (hydrophiinae): evidence from seven genes for rapid evolutionary radiations. Journal of Evolutionary Biology 21, 682-695.

Strickland, J. L., Carter, S., Kraus, F. & Parkinson, C. L. 2016. Snake evolution in Melanesia: origin of the Hydrophiinae (Serpentes, Elapidae), and the evolutionary history of the enigmatic New Guinean elapid Toxicocalamus. Zoological Journal of the Linnean Society 178, 663-678.

Warrell, D. A., Hudson, B. J., Lalloo, D. G., Trevett, A. J., Whitehead, P., Bamler, P. R., Mamy Ranaivoson, Wiyono, A., Richie, T. L., Fryauff, D. J., O’Shea, M. T., Richards, A. M. & Theakston, R. D. G. 1996. The emerging syndrome of envenoming by the New Guinea small-eyed snake Micropechis ikaheka. Quarterly Journal of Medicine 89, 523-530.

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Within recent days, the world has learnt of the passing of American writer, author, artist and natural historian Richard Ellis (1938-2024), best known for his many works on marine animals and their environment...

**Caption:** Richard Ellis in 1978, a portrait provided by the Ellis family and used in **this obituary at *The New York Times***. Image: (c) Ellis family.

Richard and his work had a major and formative impact on myself, and I suspect on quite a few of you reading this article. I was lucky enough to have corresponded with him from the late 1990s onwards and thus had a modicum of insider info on his projects and thoughts. In view of all this, I felt it appropriate to pen some words.

**Caption:** I don’t (yet) own all of Richard’s books, but here are some in my collection here…

Growing up in New York and with a connection to the sea from an early age, he graduated from the University of Pennsylvania in 1959 before working at the American Museum of Natural History (AMNH) as an exhibition designer. Richard ultimately gained enviable field experience with sharks, marine mammals and other marine wildlife worldwide, became an experienced diver, and underwent scuba and cage diving, including with Great whites Carcharodon carcharias. Even by the time he published his first book (1975, when he was in his early 40s) he was extremely well travelled.

I essentially know nothing of what led Richard to become a painter or writer, but it was a 1972 job for Encyclopedia Britannica (for their 1974 edition) that made him a professional illustrator. He did the sharks for that work and also the whales and assorted other animals too, turning in an incredible 40 paintings a month (Ellis 1975, p. 13). From these beginnings, he later had work published in Reader’s Digest, GEO, Sea Frontiers, Scientific American, Science Digest, Audubon and elsewhere, and exhibited his paintings in numerous galleries and museums. By the 1980s, he held associations with the Society of Vertebrate Paleontology, American Society of Mammalogists, New York Academy of Sciences and the Explorers Club, and had a research affiliation with the AMNH.

An emphasis Richard made is that painting sharks from references isn’t easy since images (speaking here of a 1970s perspective) generally show dead, landed animals. You need to know how they functioned and fitted into the water. The following words will echo feelings familiar to many of you…

“I liked the way sharks looked. They seemed frighteningly efficient, and they reminded me of the fighter planes of World War II. I grew up in the era of the P-38, P-40, and P-41; as a boy in the early 1940s, I had an almost total preoccupation with the Mustang, Spitfire, ME-109, and the Zero. Eventually I grew out of this phase, but it was replaced by a more interesting group of streamlined, efficient “machines”, the sharks.” (Ellis 1975, p. 15).

**Caption:** this painting does appear in *The Book of Sharks*, but small and in black and white, not in the colour plate section. The version here was sent by Richard on Christmas cards, this one specifically from 2011. Image: (c) Richard Ellis.

He goes on in that section of his 1975 book – The Book of Sharks – to describe his total immersion in the world of sharks, both in terms of looking at them as live animals, learning about their three-dimensional form, and attending conferences, speaking to researchers, shark fishermen and trophy hunters, and shark keepers at aquariums and oceanariums. It was obvious that his level of experience was vast and considerable, even in the mid-70s.

**Caption:** one of the most striking features of Richard’s paintings is how vibrant the colours are. That’s especially obvious in the leaping Common dolphin *Delphinus delphis* at left, a version of which appears in the plate section of his 1982 *Dolphins and Porpoises*. I can’t get that same look to come across in the photos shown here (if I increase the contrast, the background blue just looks over-saturated). At right, a North Atlantic right whale *Balaena glacialis*, from 1980’s *The Book of Whales*. This is a really interesting painting because the perspective and crystal-clear view (not possible for the human eye to obtain) makes the whale look small. In reality, this animal is about 18 m long. Images: (c) Richard Ellis.

Helping to build life-sized whales and squid. To go back to that 1960s stint at New York’s AMNH, we know that Richard helped create the life-sized Blue whale Balaenoptera musculus model on display in the museum’s Hall of Ocean Life. Its mid-1960s construction is itself related to the existence of an older and slightly smaller 1930s-era Blue whale model at London’s Natural History Museum (the construction of which I wrote about here).

Richard maintained connections with museum exhibition work later in his career, since he painted the life-sized giant squid on show at the National Museum of Scotland in Edinburgh in 1997 (Ellis 1998). I’m not sure if he was invited because of his (by then) respected status as an author-artist specializing on marine life, or if the museum really was in quest of someone with relevant knowledge and experience… either way, it’s a great connection.

**Caption:** the life-sized giant squid *Architeuthis dux* model on show at the National Museum of Scotland, Edinburgh, where it hangs in the main hall, surrounded by models, taxiderm specimens and mounted skeletons of other swimming animals. The position of the squid means that it’s a bit difficult to photograph. The cetacean making a guest appearance is a Pygmy killer *Feresa attenuata*. Richard painted this model squid. Image: Darren Naish.

Influential books on sharks and cetaceans. Richard’s 1975 The Book of Sharks was followed by the similar-format The Book of Whales in 1980 and Dolphins and Porpoises in 1982. All three are large and combine a plate section that features his spectacular, vibrant paintings with species-by-species accounts that cover natural history, ecology, behaviour and more. Naturally, The Book of Sharks is not intended to be a thorough coverage of all species (somewhere over 510 are recognized at the time of writing, and less than 300 were regarded as valid when Richard was writing) but it still serves as an excellent introduction. And it was a commercial success, with reprints occurring in 1976,1983 and 1989.

Dolphins and Porpoises was my first introduction to Richard’s work, and I well remember it being my staple reading for some chunk of the early 1990s. I adore its style, arrangement and picture-to-text ratio (very much not a trivial thing when it comes to book design). A major strength of the book is the presence of tidy diagrams and drawings that illustrate aspects of anatomy discussed in the text. When pondering living things and the way they look, we mostly think that pigmentation, ornamentation and shape is adaptive, and linked to behavioural syndromes and ecological specializations. That sounds like an obvious proposition, but it takes a lot of effort to learn about ‘form-function’ correlations of this sort and point them out, and it’s done well far less frequently than you might think.

**Caption:** I adore the black and white illustrations that appear throughout Richard’s two whale-themed books of the early 1980s. At upper right, the pale fields and patches on the body of *Orcinus*. Lower left, a juvenile Risso’s dolphin *Grampus griseus*, showing the prominent pale folds often present on the bodies of very young cetaceans. At right, various ‘blackfish’ (albeit not all of them) drawn to scale with a person. Image: Ellis (1982).

It’s actually quite funny that The Book of Whales and Dolphins and Porpoises turned out the way they did because, as Richard wrote in the preface to The Book of Whales (Ellis 1980, p. xi), the original intention was for there to be a single book, and for it to be a small pocket guide. I’m very happy that this plan went so disastrously awry.

**Caption:** more great images from Richard’s cetacean books, again showing interesting aspects of anatomy that generally hadn’t been shown in books before. Those here show (clockwise from upper left) different ventral pigmentation patterns in cephalorhynchine dolphins, the remarkable degree of sexual dimorphism in *Physeter*, the distinct dorsal and ventral sonar fields of *Platanista* hypothesized by Pilleri in 1979, and the steep hump-backed diving posture of *Inia*. Images: Elllis (1980, 1982).

It was thanks to cetaceans that I first got to know Richard. I never met him in person, but – like many people of my age (I’m somewhere in my late 40s) – I got to know a great many people thanks to the internet mailing lists and message boards of the late 20th century. Sometime around 1996, I was an erstwhile participant in MARMAM, the Marine Mammals Mailing List, still in existence today and operated from the University of Victoria. I was especially interested at that time in whales that people claimed to have observed, but which don’t match officially recognized species. Crypto-whales, if you like, a topic a younger version of me explored in some number of published articles. I’m pretty sure that’s why I first corresponded with Richard, probably because he was telling me to avoid paying attention to sources generally considered dubious. And on that note…

**Caption:** Richard’s first three books showcase many of his paintings in the largest format possible (err, excluding gatefolds), but a drawback is that many of them then cross the book’s spine. Such is the case in the painting at top left, showing a group of Sowerby’s beaked whale *Mesoplodon bidens*. At right, the spectacular cover of his 1980 *The Book of Whales*. Another thing that these books demonstrate is Richard’s versatility as an artist: pencil drawings, black and white diagrams, colour paintings, skeletal diagrams… At lower left, we see his drawing of a Heaviside’s dolphin *Cephalorhynchus heavisidii*. What a contrast in style to the colour paintings. Images: Ellis (1980, 1982).

On monsters and cryptozoology. Any writer interested in poorly known creatures of the seas and the history of human knowledge pertaining to them will, inevitably, develop a familiarity with cryptozoology. Richard touched on this subject in several of his books – his shark-themed works discuss claims that the megatooth shark Otodus megalodon might be alive today – and ultimately gathered enough material for his 1996 Monsters of the Sea (Ellis 1996). It’s fondly remembered and remains one of few technically well produced, mainstream books on the topic. Richard once scolded myself and my co-authors for failing to cite it (Woodley et al. 2011), a guilty oversight on our part.

**Caption:** it can be surprisingly difficult to find useable images of book covers online. They’re often at a resolution too low to be useful, or badly photographed at a weird angle. I’ve done my best to photograph some covers here but haven’t done an especially good job. Whatever, here’s the cover of the hardback first edition of Richard’s *Monsters of the Sea* (Ellis 1996)… a signed copy! Image: Darren Naish.

On cryptozoology, Richard was an avowed sceptic and was critical of cryptozoological claims and evaluations, and indeed of cryptozoologists themselves. A complaint one can make about cryptozoologists is that their statements about non-cryptozoological topics are often wrong or naïve. It often seems trite to point this out… but Richard liked to do it. My favourite, albeit trivial, example: in In The Wake of the Sea-Serpents, Bernard Heuvelmans says eels are “immensely powerful constrictors” (Heuvelmans 1968, p. 264), to which Richard responds “… they certainly are not” (Ellis 1996, p. 240).

Richard sometimes seemed a bit prickly when commenting on cryptozoological issues. In a TV show devoted to the examination of supposed unexplained phenomena, Richard was shown evaluating some bigfoot footage. “This is awful!”, says Richard in what sounds like a genuinely annoyed tone as a person in a costume – I assume – strides through a cornfield on the TV screen he’s viewing.

**Caption:** in case you’re wondering, the to-and-fro about the constricting abilities of eels has its origins here… the Pauline sea monster account of 1875, wherein a sperm whale off Brazil was seen in supposed battle with a long object twisted around its body. Heuvelmans (1968) thought that the long object was the gigantic crypto-eel – *super eel* – that he endorsed in his book. More recent suggestions include that the witnesses saw, and misinterpreted, the whale’s large, pale penis (Paxton *et al*. 2005), or that this was a whale entangled in rope (France 2019).

When preparing his 2019 book Disentangled: Ethnozoology and Environmental Explanation of the Gloucester Sea Serpent, Robert France phoned Richard to talk about tuna (Richard was a tuna expert, having written the 2008 Tuna: Love, Death and Mercury). Richard “abruptly hung up the phone when thrice I contacted him” (France 2019, p. 248). I’d love to know what it was that made Richard so irascible. I put it that saying “Richard Ellis once slammed the phone down on me” is something of an accolade. A bit like being punched in the face by a famous boxer.

Richard and the discovery of the Megamouth. Having mentioned sharks, I have to recount another Ellis-themed anecdote. It’s a story that I’ve seen recounted several times online but not in print.

We know from comments made in 1991’s Great White Shark, a book that Richard co-authored with ichthyologist John McCosker, that Richard was aware of the discovery of the Megamouth Megachasma pelagios right from the off in November 1976. None other than Peter Benchley had phoned to tell him… though Benchley had wrongly referred to the new shark as a Megalodon, oopsie (Ellis & McCosker 1991, p. 44).

**Caption:** at left, the Megamouth holotype specimen as it looked after being brought to the surface following entanglement with the parachute sea anchor of US Navy vessel AFB-14 on November 15th 1976, at the Kaneohe Bay facility of the Naval Undersea Center. At right, the specimen on show at the Los Angeles County Museum, photographed in 2009. It’s in a giant, glass-topped case which makes photography (with a normal camera) difficult. Images: Leighton Taylor; Darren Naish.

A common occurrence in the history of science is that people recognize or discover a thing, but then take years, even decades, to get it into print. Why are they taking so long, others say, despite knowing that just about everyone in science is overworked, underpaid, and perpetually on the brink of physical or emotional collapse. Such was the precise complaint of Richard and John when they visited Leighton Taylor at Honolulu in 1980. Taylor was supposed to be leading the Megamouth’s long-awaited description, yet it had so far failed to materialize. Richard and John hatched a plan.

One day, a surprised Leighton Taylor received an article, published in a Japanese-language ichthyological journal, describing a new species of remarkable shark that was one and the same as the Megamouth. He had been beaten to it by a rival team, the sort of thing that can happen any time a scientist or group of scientists are preparing work for publication. But to cut right to the chase, it was a spoof paper, cobbled together from random text on rhinos in captivity and the history of the domestic cat in Japanese art and featuring a pretend binomial (I’d like to know what that binomial was. I’ve been told in the past but have since forgotten). A few clues gave the game away, and it’s this adventure that led to Richard and John being credited for “preparation of a preliminary manuscript which was of great help in the production of this final paper” in the technical description of the Megamouth (Taylor et al. 1983, p. 110). A fuller version of events can be found here at the late R. Aidan Martin’s elasmo-research site.

**Caption:** the Megamouth type description and official naming occurred in volume 43 of *Proceedings of the California Academy of Sciences*, published in 1983. Its second page features a full-page colour painting by Richard Ellis, part of which is shown here. Images: (c) Richard Ellis; Taylor *et al*. (1983).

Taylor et al. (1983) features on its second page Richard’s colour painting of a live Megachasma, so his place in the history of the species has been assured, practical joke or not.

Richard and Sea Dragons. At some point during the late 1990s or early 2000s, Richard jokingly told me that he was running out of animals to write about and thus had finally decided to tackle the great sea reptiles of the Mesozoic. Thanks mostly to my adventures at the dinosaur mailing list or DML – yes, another 20th century discussion board – Richard was aware of my interest in ichthyosaurs, plesiosaurs and their contemporaries (I’d also published a few articles on them by this time). Would I be able to review the manuscript for his book on these animals? Because I’m an idiot who’s always said yes to non-paying work, I said yes.

**Caption:** it’s hard to say from *Sea Dragons* whether Richard liked ichthyosaurs, plesiosaurs or mosasaurs more. Maybe the cover demonstrates a bias for mosasaurs. Richard did his illustrations of these animals before it became widely known that they had vertical tail flukes.

As a consequence, I get extremely fair mention in the acknowledgements (p. xi) of 2003’s Sea Dragons: Predators of the Prehistoric Oceans and am also name-checked a few times in the text itself (Ellis 2003). It remains one of comparatively few books devoted to Mesozoic marine reptiles, and of course I don’t need to say that I’ve contributed to that select number myself in recent years (Naish 2022).

As revealed in the preface to Sea Dragons, Richard’s plans to write about fossil marine reptiles extended back to the 1970s when he worked with Robert Bakker and first learnt about these creatures. It turns out that the two even considered co-operating on such a venture at one point. We all think of Bakker as a dinosaur guy, but in fact his influence on marine reptile studies has been profound too and – as I argued in my book (Naish 2022) – much of what’s happened in plesiosaur research over the past few decades can be pinned on an article that Bakker published in 1992.

**Caption:** a large sticker can be found on the inside cover of my copy of *Sea Dragons*, making it unique and highly valuable. Thank you, Richard.

Legacy: all the ocean’s animals. Richard’s quip that he was “running out of animals” was hardly true. Post-2003, he published additional books devoted to tuna (Tuna: Love, Death and Mercury, 2008), sperm whales (The Great Sperm Whale: A Natural History of the Ocean’s Most Magnificent and Mysterious Creature, 2011), swordfish (Swordfish: A Biograph of the Ocean Gladiator, 2013) and beaked whales (Beaked Whales. A Complete Guide to Their Biology and Conservation, 2017; co-authored with James Mead). Additional works covered the global marine environment, climate change and the evolution of life in the sea. There’s both a satisfying arc to the publication chronology of this body of work, and a bewildering, terrifying productivity that will hardly be matched by any living natural history writer.

**Caption:** two of Richard’s several 21st century books, these both devoted to cetaceans. I still don’t own the beaked whale volume, for shame, mostly because I have yet to see it appear at reasonable price. It’s very expensive.

And there ends my thoughts. Richard Ellis died on May 21st this year, aged 86, and my thoughts go to those who knew and loved him. Obviously, Richard and his books were and are personally important to me, but it’s no exaggeration to say that he has to be considered one of the most prominent, productive, respected and influential of writers on natural history in the modern age.

I don’t know that this was made clear to him during his own lifetime, but dearly hope that it was.

For previous articles relevant to the subjects mentioned here, see…

Sea Monster Sightings and the ‘Plesiosaur Effect’, April 2019
Remembering Lyall Watson’s Whales of the World, June 2019
Whale Watching in the Bay of Biscay, August 2019
Extreme Cetaceans, Part 1, September 2019
Extreme Cetaceans, Part 2, September 2019
A Review of Robert L. France’s Disentangled: Ethnozoology and Environmental Explanation of the Gloucester Sea Serpent, November 2019
Extreme Cetaceans, Part 3, December 2019
Monsters of the Deep, a Ground-Breaking Exhibition at the National Maritime Museum, Cornwall, October 2020
Kogia, Shark-Mouthed Horror, May 2023
News from the World of Ancient Sea Reptiles, May 2023

You can support this blog and my other projects here at Patreon

Refs - -

Ellis, R. 1975. The Book of Sharks. Alfred Knopf, New York.

Ellis, R. 1982. The Book of Whales. Alfred Knopf, New York.

Ellis, R. 1983. Dolphins and Porpoises. Robert Hale, London.

Ellis, R. 1996. Monsters of the Sea. Alfred A. Knopf, New York.

Ellis, R. 2003. Sea Dragons: Predators of the Prehistoric Oceans. University Press of Kansas, Lawrence, Kansas.

Ellis, R. & McCosker, J. E. 1991. Great White Shark. Stanford University Press, Stanford, California.

France, R. L. 2019. Disentangled: Ethnozoology and Environmental Explanation of the Gloucester Sea Serpent. Wageningen Academic Publishers, Wageningen, The Netherlands.

Heuvelmans, B. 1968. In the Wake of the Sea-Serpents. Hill and Wang, New York.

Naish, D. 2022. Ancient Sea Reptiles. Natural History Museum, London.

Paxton, C., Knatterud, E. & Hedley, S. L. 2005. Cetaceans, sex and sea serpents: an analysis of the Egede accounts of a “most dreadful monster” seen off the coast of Greenland in 1734. Archives of Natural History 32, 1-9.

Taylor, L. R., Compagno, L. J. V. & Struhsaker, P. J. 1983. Megamouth – a new species, genus, and family of lamnoid shark (Megachasma pelagios, family Megachasmidae) from the Hawaiian Islands. Proceedings of the California Academy of Sciences 43, 87-110.

Woodley, M. A., Naish, D., & McCormick, C. A. 2011. A baby sea-serpent no more: reinterpreting Hagelund’s juvenile “cadborosaur” report. Journal of Scientific Exploration 25, 495-512.

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It’s time once again to look at a very interesting bunch of snakes….

**Caption:** we’ll be seeing this illustration again… I really like it. Image: G. H. Ford, in the public domain.

In my continuing efforts to rescue and rehabilitate material now essentially lost due to the destruction of Tetrapod Zoology versions 2 and 3, here’s another article on obscure snakes. And it has some degree of connection to the other snake-themed articles published here recently, which is good. The article here was originally published at ver 2 back in September 2010 and covers Bothrolycus ater*, a relatively obscure African snake (go here for a wayback version of the original). At that time, I was super excited to learn that herpetologist and African snake expert Kate Jackson of Whitman College, Washington, had photographed this snake in living state, something that possibly hadn’t been done before. Dr Jackson very kindly gave permission for my use of these photos, and here we are. Below, please find an updated and tidied version of that 2010 article, and if you read it first time around... well, 2010 is a long time ago so you may as well read it again.

Your reminder that the dead internet problem is a real and present danger, and yet another item on the list of bad things we need to worry about, sorry.

Caption: Bothrolycus in the flesh, photographed in the Republic of the Congo in 2010. Note the slightly reddish tint to the head, the shiny, smooth-scaled look, and the white spots around the jaws. Image: © K. Jackson.

Yes, this is Bothrolycus ater, one of so many snakes conventionally treated as a sort of ‘miscellaneous colubrid’. Colubridae, you’ll recall, is the snake group name mostly associated with racers and (rightly or not…) garter snakes, water snakes and kin. One of the first books I check when wanting to know more about obscure snakes is Chris Mattison’s The Encyclopedia of Snakes (Mattison 1998). Of this animal, Mattison lists it in his unsorted colubrid section, and states that it’s “A small snake about which almost nothing appears to be known” (p. 211). That’s it. Not encouraging. Incidentally, 2010 was the year during which I discovered – by complete coincidence – that Chris Mattison used to live about five minutes away from my house. Anyway… after a bit of research I discovered that – despite its obscure status – Bothrolycus isn’t that poorly known, and in fact should certainly be more familiar than it is.

**Caption:** at left, screengrab of the 2010 ancestor of the article you’re reading now. Yeah, the original article title was way wordier than the one I’ve gone for this time around. At right, Mattison (1998). It has served me well over the years.

  **Caption:** this image of *B. ater*, by George Henry Ford, is from Günther’s original description. The chameleons are Spectral pygmy chameleons *Rhampholeon spectrum*, also known as the Western pygmy chameleon or Cameroon stumptail chameleon. I really like this entire illustration and wish that a clean white copy was available. I only have access to the very shady scan provided by the Biodiversity Heritage Library, which I’ve lightened as much as I can. Image in public domain.

The name Günther’s black snake is sometimes used for this species. Named by hyper-prolific German-born British zoologist Albert Günther in 1874, it’s from central Africa, its range including Cameroon, Gabon, Equatorial Guinea and the Republic of the Congo (Trape 1985, Lasso et al. 2002, Pauwels et al. 2006); Günther (1874) originally described it from Cameroon. The individual you see in the photos here was encountered by Kate in the Republic of the Congo in 2010, sadly in an area about to be strip-mined for iron ore.

In terms of habitat preference, Bothrolycus is an inhabitant of montane and sub-montane forest, and some studies indicate that it may be moderately common in some places. An interesting statement made by Georges Boulenger is that it’s semi-aquatic, which would be unusual for snakes of its sort. I haven’t seen this confirmed by other sources. It’s smooth-scaled (Moore & Jackson 2010) and can reach just over 41 cm. In adults, the blackish head is flecked with white, but in juveniles the top of the head and nape is cream-coloured (Loveridge 1936).

**Caption:** check out some of the great details in the George Ford illustration from above. The chameleon is totally not happy about seeing the snake, and the snake is beautifully portrayed, with an excellently portrayed amount of sheen across its scales. Ford (1808-1876) joined the British Museum in 1837 and illustrated works on human anatomy, fishes, amphibians and reptiles. Images in the public domain.

Notable sexual dimorphism. I was surprised to learn that Bothrolycus has sometimes been cited as one of the only snakes where sexual dimorphism is obvious. Schmidt (1923) regarded the difference in size between the sexes as being “unusually pronounced” and “quite exceptional” while Davis (1936) noted that it was (to his knowledge) the only snake where males and females consistently exhibit different scale counts: the former exhibiting 17 and the latter 19 on the “anterior part of the body”. Boulenger (1919) was the first author to bring attention to this presence of obvious dimorphism.

A snake now regarded as a junior synonym of B. ater – that’s Pseudoboodon albopunctatus Anderson, 1901 – might owe its initial mis-description as a distinct taxon due to this sexual dimorphism, since I think (but am not 100% sure) that its holotype represents the opposite sex from the B. ater holotype. It’s moderately well known that the males and females of various animal species were initially described as different species… well, here’s an apparent case among snakes.

**Caption:** image of the *Pseudoboodon albopunctatus* holotype, from Anderson (1901). Compare it with the images of *B. ater*: note the pale spots on the labial scales and the pit in front of the eye. *P. albopunctatus*, by the way, was only one of several *Pseudoboodon* species. The genus (named by Mario Giacinto Peracca in 1897) is still a valid boaedontin taxon (four species are currently recognized). Image in public domain.

Anyway, since the above comments of the early 1900s were published, sexual dimorphism in snakes has become better known and is now understood to be quite widespread, including in size, mass and relative head size (Shine 1991). Nevertheless, given the apparently distinct – and historically significant (if you like) – sexual dimorphism of Bothrolycus, I’m surprised that the species is as little-mentioned as it is.

By the way, it’s obvious how virtually everything published about this snake comes to us from European scientists, often writing about museum specimens brought to European institutions by colonial explorers. What I’m getting at is that we essentially never hear about whatever it is that local people knew or know, or thought about or think about this snake. A striking animal such as this surely had some reputation to the people who lived and live alongside it. If such information is recorded in the literature, I’d love to know more.

What sort of snake is Bothrolycus? When I first wrote about this snake in 2010, the idea most familiar thanks to then-current literature was that Bothrolycus was part of the colubrid group Boodontinae (spelt Boaedontinae by some authors) (Dowling 1969, McDowell 1987, Zaher 1999, Lawson et al. 2005). This inclusion was historically based on hemipenial morphology: like other members of the group, each hemipenis in Bothrolycus is slightly bilobed and has centrolineal, bifurcating sulci spermatici and longitudinal rows of medium-sized spines that are connected by wavy, spinulate ridges (Zaher 1999). Yes, to be an expert on snake phylogeny and diversity, you have to be very familiar with male genitalia.

**Caption:** an Olive house snake *Boaedon olivaceus*, standing in as an exemplar for the lamprophiine genus *Boaedon*. *Boaedon* was mostly regarded as synonymous with *Lamprophis* – though the members of the two genera sure have a complex taxonomic history – until it was resurrected as valid by Kelly *et al*. (2009). Image: Erik Paterson, CC BY 2.0 (**original here**).

Since about 2010, Boodontinae/Boaeodontinae has undergone a name change: its type genus – Boaedon (the Cape house snake B. capensis and kin) – is almost definitely closely related to Lamprophis, meaning that the correct name for this group is Lamprophiinae. A version of this name (though written Lamprophes) was first published by Austrian zoologist Leopard Fitzinger in 1843, whereas Boodontinae wasn’t published until 1893 (by E. D. Cope) (Kelly et al. 2009).

**Caption:** this illustration, by Joseph Smit and published in 1887, depicts Fisk’s house snake *Lamprophis fiskii* and accompanied Boulenger’s original description of this species. Under the current, restricted use of the name, *Lamprophis* only includes three extant species. Image in public domain.

And as you’ll know if you read the other recent snake articles here, both Lamprophiinae and Lamprophiidae are now in general use in the snake literature, recent studies agreeing that this is an important clade within Elapoidea: it is not close to Colubridae, but instead to Elapidae (the cobras, kraits, sea snakes and so on) (Pinou et al. 2004, Lawson et al. 2005, Kelly et al. 2009, Pyron et al. 2011, 2013, Zaher et al. 2009, 2019, Figueroa et al. 2016, Das et al. 2023). If Bothrolycus is a lamprophiine… well, that puts an end to our need to refer to it as a ‘colubrid’. By the way, lamprophiines/lamprophiids have sometimes been vernacularly termed ‘house snakes’ (since that name is attached to the Boaedon and Lamprophis species that form the ‘core’ of the group) but ‘African nocturnal snakes’ was suggested by Kelly et al. (2009).

**Caption:** here’s that very simplified cladogram again, showing relationships among the main endoglyptodont groups. Elapoidea includes numerous lineages and views differ on which should and should not be included within Lamprophiidae. The topology shown here is consistent with several recent studies, but the taxonomy specifically follows that of **Zaher *et al*. (2009)**. Image: this uses images created for the textbook I’m putting together. **More on that on patreon.**

And… is Bothrolycus a lamprophiine? Zaher et al. (2009) supported this on the basis of hemipenial morphology, and it’s since been supported in molecular studies. Specifically, Kelly et al. (2011), Pyron et al. (2011, 2013), Figueroa et al. (2016), Portillo et al. (2019), Zaher et al. (2019) and Tiutenko et al. (2022) all found Bothrolycus to be close to Lamprophis and Boaedon, and thus unambiguously within Lamprophiinae even in the very strictest sense of that name. All of these studies found a Bothrolycus + Bothrophthalmus clade as the sister-group to a Lamprophis + Boaedon clade, this whole lot corresponding to Boaedontini as used by Dowling (1969).

Bothrophthalmus, if you’re wondering, includes the Red-striped black snake Bo. lineatus of tropical Africa, its range extending from Guinea in the west to Uganda in the east and Angola in the south. Conventionally, there’s only one Bothrophthalmus species. But that might be wrong. I should also mention that Tiutenko et al. (2022) recognized the so-called Ethiopian house snake – conventionally Boodon erlangeri – as a boaedontin close to the Bothrolycus + Bothrophthalmus clade and gave it the new generic name Bofa. I tell you, caenophidian snake phylogenetics really is where it’s at.

The big deal about Bothrolycus.One thing makes Bothrolycus a particularly special snake, and it’s not its sexual dimorphism or its position within phylogeny. It’s the presence of unusual openings on the side of the face, located just in front of the eyes and referred to by Anderson (1901) as elongate deep loreal pits. You should be able to make them out in Kate’s photos.

**Caption:** that live *Bothrolycus* individual again, photographed in the Republic of the Congo in 2010. An interesting anatomical detail here is that the maxillary tooth tips are partly visible, perhaps because the snake has exposed these as a threat display. I owe thanks to snake expert John Scanlon for the following observation: “And those tips form two distinct groups: long fang-like teeth at the front, and shorter ones more closely spaced posteriorly. All those teeth are on the maxilla; this pattern is rather like elapid dentition, and probably not by coincidence. And the ‘wolf’ reference in the name is presumably due to the large canine-like anterior teeth, as in several other snake genera with ‘lyco’ names”. Image: © K. Jackson.

If you know snakes, you’ll no doubt be thinking of the facial pits present in vipers, pythons and boas. These house thermoreceptors and allow the snakes to detect heat sources, these typically being warm-bodied prey animals. So… do the pits in Bothrolycus also contain thermoreceptors? Does this African elapoid also use heat-detection to find prey? I don’t know the answers to these questions and, so far as I can tell, nor does anyone. In fact, I haven’t seen any information on the diet or feeding behaviour of this snake. If Bothrolycus does possess heat-detecting pits, then these remarkable structures must have evolved in yet another snake group.

**Caption:** closeup of the snake’s right side, this time showing the loreal pit and also the white facial spots in detail. The latter are restricted to the labial scales and are arranged approximately symmetrically on both upper and lower jaws. Image: © K. Jackson.

I started this article by noting how obscure Bothrolycus is, and you might recall that I quoted certain sources as stating that nothing, essentially, is known about this snake. But now that we’re at the end of an article that includes over 1620 words, we may ask: does the fact that it’s obscure mean that there’s nothing to say about it? I think not.

I need to finish by expressing massive thanks to Dr Kate Jackson for supplying the Bothrolycus images in the first place (visit Kate’s webpage). For previous articles on squamates, see…

Cambodia: now with dibamids!, May 2011
The Tet Zoo Guide to Mastigures, August 2018
The Remarkable Basilisks, May 2023
Do Lizards Really Have ‘Mite Pockets’?, March 2024
Meeting Lake Zacapu’s Garter Snake, March 2024
Ray Hoser, Number 1 Taxonomic Vandal, May 2024
The Mysterious Dibamids, May 2024
Side-Stabbing Stiletto Snakes, Redux for 2024, May 2024

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

Refs – -

Anderson, L. G. 1901. Some new species of snakes from Cameroon and South America, belonging to the collections of the Royal Museum in Stockholm. Bihang Till Köngl. Svenska Vetenskäps-Akadeillens Handlingar 4, 3-26.

Boulenger, G. A. 1919. Un cas interessant de dimorphisme sexuel chez un serpent africain (Bothrolycus ater Günther). Comptes Rendu de l’Academie des Sciences, Paris168, 666-669.

Das, S., Greenbaum, E., Meiri, S., Bauer, A. M., Burbrink, F. T., Raxworthy, C. J., Weinell, J. L., Brown, R. M., Brecko, J., Pauwels, O. S. G., Rabibisoa, N., Raselimanana, A. P. & Merilä, J. 2023. Ultraconserved elements-based phylogenomic systematics of the snake superfamily Elapoidea, with the description of a new Afro-Asian family. Molecular Phylogenetics and Evolution 180, doi.org/10.1016/j.ympev.2022.107700

Davis, D. D. 1936. Courtship and mating behaviour in snakes. Zoological Series of Field Museum of Natural History 22, 257-290.

Dowling, H. G. 1969. Relations of some African colubrid snakes. Copeia 1969, 234-242.

Günther, A. 1874. Descriptions of some new or imperfectly known species of reptiles from the Camaroon [sic] Mountains. Proceedings of the Zoological Society of London 42, 442-445.

Kelly, C. M. R., Branch, W. R., Broadley, D. G., Barker, N. P. & Villet, M. H. 2011. Molecular systematics of the African snake family Lamprophiidae Fitzinger, 1843 (Serpentes: Elapoidea), with particular focus on the genera Lamprophis Fitzinger 1843 and Mehelya Csiki 1903. Molecular Phylogenetics and Evolution 58, 415-426.

Lasso, C. A., Rial, A. I., Castroviejo, J. & De la Riva, I. 2002. Herpetofauna del Parque Nacional de Monte Alén (Río Muni, Guinea Ecuatorial). Graellsia 58, 21-34.

Loveridge, A. 1936. African Reptiles and amphibians in field museum of natural history. Field Museum of Natural History – Zoology 22, 1-111.

Mattison, C. 1998. The Encyclopedia of Snakes. Blandford, London.

McDowell, S. B. 1987. Systematics. In Seigel, R. A., Collins, J. T. & Novak, S. S. (eds) Snakes: Ecology & Evolutionary Biology. Macmillan (New York), pp. 3-49.

Moore, K. & Jackson, K. 2010. A quantitative analysis of two scale characters in snakes. Amphibia-Reptilia 31, 175-182.

Pauwels, O. S. G. Burger, M., Branch, W. R., Tobi, E., Yoga, J.-A., & Mikolo, E.-N. 2006. Reptiles of the Gamba Complex of Protected Areas, Southwestern Gabon. Bulletin of the Biological Society of Washington 12, 309-318.

Portillo, F., Stanley, E. L., Branch, W. R., Conradie, W., Rödel, M. O., Penner, J., Barej, M. F., Kusamba, C., Muninga, W. M., Aristote, M. M., Bauer, A. M., Trape, J. F., Nagy, Z. T., Carlino, P., Pauwels, O. S. G., Menegon, M., Ineich, I., Burger, M., Zassi-Boulou, A. G., Mazuch, T., Jackson, K., Hughes, D. F., Behangana, M. & Greenbaum, E. 2019. Evolutionary history of burrowing asps (Lamprophiidae: Atractaspidinae) with emphasis on fang evolution and prey selection. PLoS ONE 14, e0214889.

Pyron, R. A., Burbrink, F. T. & Wiens, J. J. 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13, 93.

Schmidt, K. P. 1923. Contributions to the herpetology of the Belgian Congo based on the collection of the American Museum Congo Expedition, 1909-1915. Part II. Snakes. Bulletin of the American Museum of Natural History 49, 1-146.

Shine, R. 1991. Intersexual dietary divergence and the evolution of sexual dimorphism in snakes. The American Naturalist 138 103-122.

Tiutenko, A., Koch, C., Pabijan, M. & Zinenko, O. 2022. Generic affinities of African house snakes revised: a new genus for Boodon erlangeri (Serpentes: Elapoidea: Lamprophiidae: Lamprophiinae). Salamandra 58, 235-262.

Trape, J. F. 1985. Les serpents de la région de Dimonika (Mayombe, République Populaire du Congo). Revue De Zoologie Africaine 99, 135-140.

Zaher, H. 1999. Hemipenial morphology of the South American xenodontine snakes, with a proposal for a monophyletic Xenodontinae and a reappraisal of colubroid hemipenes. Bulletin of the American Museum of Natural History 240, 1-168.

Zaher, H., Grazziotin, F. G., Cadle, J. E., Murphy, R. W., Cesar de Moura-Leite, J. & Bonatto, S. L. 2009. Molecular phylogeny of advanced snakes (Serpentes, Caenophidia) with an emphasis on South American Xenodontines: a revised classification and descriptions of new taxa. Papeis Avulsos de Zoologia 49, 115-153.

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An updated look at a very special group of remarkable burrowing snakes…

**Caption:** a fairly familiar portrait of a burrowing asp / stiletto snake that has appeared several times in the literature, and showing the right maxillary fang protruding while the mouth is closed. Image: I’m not sure of the origin of this image and will add credit info when I find it! I think it first featured in Underwood & Kovcha (1993).

Tet Zoo, the Squamate Years. In case it isn’t already obvious, one of my aims for 2024 is to release a lot of squamate-themed content here at Tet Zoo ver 4, and in part this involves rescuing and updating material previously published at ver 2 and ver 3. Today we’re going to look at a really fascinating group of caenophidian snakes called the burrowing asps or mole vipers.

The bulk of the text here was originally published at ver 2 (the ScienceBlogs years) back in May 2008, but – like all Sb material from that time – it’s since been removed or ruined. A wayback machine version does exist (here). Interesting discoveries have been made about burrowing asps since 2008, so I’ve made updates to the text where appropriate, and I’ve also updated it overall. Ok, let’s get to it.

**Caption:** if you’ve been following Tetrapod Zoology for a while, you might remember this very popular article from 2008. I know I go on about this a lot, but it’s a real source of frustration and disappointment to me that so much of my old Tet Zoo content is now unfindable *except at* wayback machine (which comparatively few people use or even know about). I see this as part of the larger, ‘dead internet’ problem that we now face as a society.

It goes without saying that most predatory animals need to open their mouths when they want to stab or bite potential prey items. But – get this – there’s a group of snakes that can erect their teeth and stab prey with a closed mouth. And that’s not all that’s interesting about these snakes. There are an awful lot of weird snakes, and one of my favourite groups is the atractaspidids (or atractaspids), and in particular the genus Atractaspis. These snakes are variously referred to as mole vipers, burrowing asps, burrowing adders, stiletto snakes or side-stabbing snakes. I’m going to (mostly) be referring to them as burrowing asps.

Burrowing asps are specialized for fossoriality (burrowing), with shiny-scaled, cylindrical bodies, small heads, a countersunk lower jaw, indistinct neck, short tail, and small eyes. Specialized fangs, a sophisticated venom apparatus and other characters show that they’re part of Colubriformes (sensu Zaher et al. 2009), the huge snake clade that includes viperids, elapids and that massive number of lineages traditionally grouped together as Colubridae.

**Caption:** good photos of burrowing asps in life aren’t all that common. Here’s a good one, showing a Southern stiletto snake *A. bibronii*, the type species for the genus. It occurs widely across the southern half of Africa. Note the protective or defensive pose, with the head held low and the tail tip raised high. Image: Ryan van Huyssteen, CC BY-SA 4.0 (**original here**).

Burrowing asps occur throughout much of Africa (with the exception of the north and south-west), and also the Sinai and Arabian peninsulas as far north as Israel. Around 22 species are currently recognized, but suspicion persists that cryptic species – those that look similar to others but are genetically very distinct from them – exist and await recognition. The type species for the genus is Bibron’s or the Southern stiletto snake A. bibronii, named by British zoologist Andrew Smith (who became Sir Andrew Smith in 1858) in 1849, the scientific name honouring his French colleague Gabriel Bibron (1806-1848). Species have been described at a fairly steady rate since then, the newest at the time of writing being Branch’s stiletto snake A. branchi of Liberia, named in 2019 (Rödel et al. 2019). These snakes are generally between 30 and 50 cm long. Oh, and they have a distinctive aromatic smell (Branch 1988). No-one knows why.

How to be a ‘fang stabber’. Burrowing asps have a formidable venom delivery apparatus. The maxillary fangs are massive, being about one-third as long as the whole skull, and the venom glands are enormous, extending into the neck and – in some species – continuing for approximating 20% of total body length (Underwood & Kochva 1993, Wollberg et al. 1998).

**Caption:** the burrowing asp skull – the lower of the two shown here – really is one of the most modified in all of Squamata. As is obvious if you compare it with the viperid skull shown at the top, the burrowing asp skull has a modified prefrontal (pf) and maxilla (mx), a startling lack of teeth, and a much pared down, reduced anatomy overall. Image: Deufel & Cundall (2006).

The rest of the dentition is highly reduced, there being just two short, gently curved dentary teeth and a couple of very small palatine teeth. The maxillary fangs (there are two in each maxilla, one of which is a replacement tooth kept in reserve) are huge compared to the short, block-like maxilla, and virtually its entire length is occupied by the transversely arranged fang sockets. The maxilla articulates with the relatively immobile prefrontal by way of a saddle-shaped joint, and this allows the maxilla to rotate. This is quite different from the condition present in viperids, where the articulatory surfaces between the maxilla and prefrontal are flat.

**Caption:** fang erection mechanism in *Atractaspis*, as illustrated by Deufel & Cundall (2003b). At left, it should be obvious that the articulation between the prefrontal (pf) and maxilla (mx), allow substantial rotation of the maxilla (and hence the giant fang(s) it houses). At right, the extreme shortness of the maxilla means that it can be rotated through a much larger angle than it can in more ‘normal’ snakes, like the *Xenocalamus* shown below. Figure from Deufel & Cundall (2003b).

Because both the maxillae and the fangs are directed posteriorly when at rest, and because the prefrontal doesn’t move much relative to the braincase, the maxilla-prefrontal unit can’t be thrown forwards to project the fangs anteroventrally (as happens in viperids and elapids).

The rotation of the maxilla is assisted by musculature attached to the slender, rod-like pterygoid-ectopterygoid unit. Usually in snakes, the pterygoid is attached to the palatine, but in burrowing asps the two are widely separated and only have a ligamentous connection (Underwood & Kochva 1993, Deufel & Cundall 2003a, b). This allows the pterygoid-ectopterygoid unit to swing antero-posteriorly without interference from the palatine. This has some implications, as we’ll see.

**Caption:** viperid, burrowing asp, and elapid palatal bones compared, from Deufel & Cundall (2006). The massive size of the burrowing asp fang (f) is obvious, as is the small size of the maxilla (mx). Note the slender, toothless pterygoid (pt) in the burrowing asp, and the lack of a bony connection (there’s a ligament there instead) between the pterygoid and palatine (pal). The reduced, bar-like form of the ectopterygoid (ec) in the burrowing asp is notable too. See Deufel & Cundall (2006) for a fuller explanation.

As the maxilla rotates ventrally, it opens up a slit along the mouth-line, providing enough space for the fang to protrude out of the mouth. The projecting tooth is then stabbed into prey with a swift posteroventral (down and backwards) jerk of the head. The snake might erect either the left-side fang or the right-side one: they don’t seem to deploy fangs from both sides at the same time (even though they probably could). When grasped behind the head in what would normally be regarded as a safe handling posture, a burrowing asp can – without opening its mouth – erect one of its super-long fangs and stab the hand of the person holding it. Kurnik et al. (1998) reported a case in which a herpetologist – specifically, one of the authors of the paper – was bitten on the finger by an Ein-Geddi burrowing asp A. engaddensis. “Local effects, oedema, erythema and numbness appeared within minutes, followed by systemic effects, including general weakness, sweating, pallor, fluctuations in the level of consciousness, vomiting and watery non-bloody diarrhoea. Gross oedema of the hand developed and extended up to the forearm” (Kurnik et al. 1998, p. 223). While the local effects healed within a few weeks, “some discoloration and tenderness remained even 10 months after”. Yikes. Don’t get bitten by a burrowing asp, that’s my advice.

And burrowing asp fangs aren’t just hollow cones, but (in all but two species) both canaliculate (that is, they house a tubular canal) and keeled along the posterior edges of their tips. This keel cuts into tissue when the snake stabs, presumably increasing the size of the wound and hence aiding the absorption of venom (Golani & Kochva 1988). It’s also been suggested that the keel helps the snake to yank its fangs out of its prey: while most long-fanged snakes strike at prey from a distance and only briefly engage with the prey, burrowing asps get right up close to prey before stabbing, sometimes stabbing several times. For this reason, Deufel & Cundall (2003b) recommended that a burrowing asp attack shouldn’t be referred to as a ‘strike’, but as a ‘fang stab’. When confronted with several prey items, burrowing asps have been reported to stab and envenomate several individuals before beginning to feed.

Specialized – but specialized for what? Given that burrowing asps do their fang stabbing in burrows and other confined spaces, you’d guess that all of these morphological and behavioural specializations have evolved to allow attacks where relatively little room for manoeuvring is possible. Who, or what, is getting stabbed? Atractaspis species prey on nestling mammals (mostly murids and shrews), and Deufel & Cundall (2003b) proposed that a reliance on such prey shaped their evolution, suggesting that “the success of [Atractaspis and relatives] is partly attributable to the use of the envenomation apparatus on mammals” (p. 58).

**Caption:** at left, another image showing how the maxillary fangs can be erected in a manner that’s very unusual relative to what’s more familiar. This is a preserved specimen of a Fat burrowing asp *A. corpulenta*. The snake is small, so that fang is only 4 mm long. At right, a photo from a sequence provided by Deufel & Cundall (2003b) where a Bibron’s burrowing asp *A. bibronii* is manipulating a rodent baby in preparation for swallowing. The snake has hooked a maxillary fang into the rodent and is using it as a gaff. Images: Deufel & Cundall (2003b); Tilbury & Verster (2016).

However, Shine et al. (2006) argued that mammals make up less than 25% of the diet of Atractaspis and drew attention to data showing that elongate fossorial squamates were the most important items in their diets. Attacking burrowing skinks and amphisbaenians within their burrows poses a problem, as the tails of these animals are similar in diameter to their bodies, making it difficult for an attacking snake to move past the tail and grab the body (you don’t want to grab the tail as both skinks and amphisbaenians are capable of autotomy).

**Caption:** amphisbaenians are one of my favourite groups of animals. We know for sure that snakes related to burrowing asps, like quill-snouted snakes (*Xenocalamus*, read on), are occasional predators of these dedicated burrowers. It’s thought that burrowing asps predate on them as well. The amphisbaenian portraits here show (at left) *Amphisbaena* of South America (an animal that’s therefore not relevant to burrowing asp predation) and Zarudny’s worm lizard *Diplometopon zarudnyi* of Iran, Iraq, the Arabian Peninsula and nearby. Images: Darren Naish.

Some burrowing asp specializations might, therefore, allow these snakes to push past the tail and envenomate or seize the prey’s body. Incidentally, Shine et al. (2006) noted that the ability to autotomise the tail among fossorial squamates might be an anti-atractaspidid adaptation, as a shed tail might both block the burrow and prevent venom injected into the tail from reaching the body. More study is needed to test this intriguing idea.

Being a good burrower makes a snake a poor swallower. The reduced palatal dentition and ligamentous linkage between the pterygoid and palatine raises an issue: how do these snakes transport prey within the mouth? Most snakes employ ‘pterygoid walking’, engaging the maxillary and pterygoid teeth on the left side with the prey and dragging it toward the throat, disengaging, and then doing the same with the maxillary and pterygoid teeth on the right, and so on. Burrowing asps can’t do that, since they sacrificed the ability to employ pterygoid walking when they de-coupled the palatine from the pterygoid and evolved a toothless pterygoid whose only function is to aid erection of the rotating maxilla and its fang. What, then, is a burrowing asp to do?

**Caption:** at left, Western Forest stiletto snake *Atractaspis aterrima*, quite possibly not in living state. At right, montage showing *A. branchi*, the most recently named member of the group (it was published in 2019). Images: Violette Dérozier, CC BY 4.0 (**original here**); **Rödel *et al*. (2019)**, CC BY 4.0 (**original here**).

Deufel & Cundall (2003b) looked at this question. Firstly, burrowing asps sometimes used their super-long fangs as gaffs to manipulate prey into swallowing position. Movements of the maxilla and/or pterygoid are indeed not used in transporting prey, but by shifting the lower jaw posteroventrally, bending the anterior trunk region from side to side, and compressing and extending the neck, the snakes are able to move the mouth over the prey. They aren’t very good at it though, and take a long time to successful ingest prey (the account here is very much simplified: for the full story see Deufel & Cundall 2003b and Cundall & Deufel 2006).

The question of how burrowing asps transport prey within the mouth illustrates the point that these snakes have made a sort of trade-off in view of competing evolutionary pressures. To be a good burrower, and to function as a specialized fang stabber, Atractaspis has lost or modified some of the kinetic zones ordinarily present in the colubriform skull: the burrowing asp snout is relatively immobile relative to the braincase, for example, and the palatine doesn’t move with the pterygoid-ectopterygoid unit. But these modifications mean that burrowing asps have had to find other solutions to the problems posed by feeding, and in fact their solution is convergently similar to that evolved by some other fossorial snakes, like pipe snakes (Cylindrophis).

For more on this whole fascinating subject, be sure to check out the excellent papers published by Alexandra Deufel and David Cundall (Deufel & Cundall 2003a, b, 2006, Cundall & Deufel 2006). It should be obvious that I relied on them heavily in my discussion here.

**Caption:** the image of live burrowing asps we’ve seen so far make them look dark brown or nearly or entirely black. As this image of a Beaked burrowing asp *A. duerdeni* (of Namibia, Botswana and South Africa) shows, they’re sometimes marked with light, pinkish tones too. Image: Joubert Heymans, CC BY 4.0 (**original here**).

A controversial radiation. Finally, where do burrowing asps fit in phylogenetic terms? Their large venom glands, long, posteriorly inclined quadrates, vestigial or absent left lung, absent pelvis and many other details show that they’re part of the grand radiation formerly termed Colubroidea but better named Colubriformes (McDowell 1987, Lee & Scanlon 2002). I’m now following Zaher et al. (2009) in thinking that the term Colubroidea should be restricted to what was formerly ‘Colubridae’.

Burrowing asps were long regarded as viperids. However, Bourgeois (1961) proposed that they might be particularly closely related to another group of colubriforms, the aparallactines. Aparallactines are small-headed, fossorial snakes, often back-fanged. Some are blunt-snouted, others sharp-snouted. The ‘core’ member of this group is Aparallactus (the centipede-eaters), but it also includes the quill-snouted snakes (Xenocalamus), harlequin snakes (Homoroselaps) and purple-glossed snakes (Amblyodipsas). Molecular studies published within recent years have generally supported an atractaspidid – aparallactine alliance (Kraus & Brown 1998, Pyron et al. 2011, 2013, Figueroa et al. 2016, Zaher et al. 2019).

**Caption:** aparallactines are elusive, poorly known snakes, though there are certainly more images of them online today than there were when I first needed them back in 2008! Here’s a species that’s relatively well known within the group, the Slender quill-snouted snake *Xenocalamus bicolor*, one of five species within this endemic African genus. Image: Ryan van Huyssteen, CC BY-SA 4.0 (**original here**).

Molecular studies have also found this atractaspidid + aparallactine clade to be close to elapids (Kraus & Brown 1998, Pyron et al. 2011, 2013, Figueroa et al. 2016, Zaher et al. 2009, 2019). This atractaspidid, elapid and kin clade in turn belongs within Endoglyptodonta, and – within that – Elapoidea (Zaher et al. 2009). This accords with general thinking on where burrowing asps might belong based on anatomy. However, studies also find a number of other lineages to be even closer to burrowing asps than elapids are, among them the African house snakes and kin, or lamprophiines. Some studies have thus applied the name Lamprophiidae to the clade that includes lamprophiines, burrowing asps and various related group (e.g., Pyron et al. 2011, 2013, Figueroa et al. 2016). If this is followed, I shouldn’t be referring to ‘atractaspidids’… they are, instead, astractaspidine lamprophiids.

Other studies, however, do not find burrowing asps and kin to be so clearly nestled among lineages that might be united within that same version of Lamprophiidae, and instead have used a taxonomy where Atractaspididae is maintained as a distinct ‘family’-level group within Elapoidea (e.g., Zaher et al. 2009, 2019). This has implications for the taxonomy of various other, related snake groups too, but that’ll have to be a subject for another time. As ever, remember that the taxonomy we superimpose on top of the phylogeny is often up for debate, and still somehow subjective at some level.

**Caption:** a massively simplified depiction of phylogeny within the colubriform clade Endoglyptodonta. Burrowing asps are certainly elapoids, and within this clade are close to or part of Lamprophiidae. The topology shown here is consistent with several recent studies, but the taxonomy specifically follows that of Zaher *et al*. (2009). Image: this uses images created for the textbook I’m putting together. **More on that on patreon.**

That’s where we’ll end things for now. However… if you have any memory of the initial publication of this article at Tetrapod Zoology ver 2 back in 2008, you might recall that it resulted in a follow-up article discussing physical encounters – yes, I mean bites – with these snakes. It should republish that here as well, stay tuned…

For previous articles on squamates, see…

Cambodia: now with dibamids!, May 2011
The Tet Zoo Guide to Mastigures, August 2018
The Remarkable Basilisks, May 2023
Do Lizards Really Have ‘Mite Pockets’?, March 2024
Meeting Lake Zacapu’s Garter Snake, March 2024
Ray Hoser, Number 1 Taxonomic Vandal, May 2024
The Mysterious Dibamids, May 2024

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

Refs – -

Bourgeois, M. 1961. Atractaspis – a misfit among the Viperidae? News Bulletin of the Zoological Society of South Africa 3, 29.

Branch, B. 1988. Field Guide to the Snakes and Other Reptiles of Southern Africa. New Holland, London.

Cundall, D. & Deufel, A. 2006. Influence of the venom delivery system on intraoral prey transport in snakes Zoologischer Anzeiger – A Journal of Comparative Zoology 245, 193-210.

Deufel, A. & Cundall, D. 2003a. Prey transport in “palatine-erecting” elapid snakes. Journal of Morphology 258, 358-375.

Deufel, A. & Cundall, D. 2003b. Feeding in Atractaspis (Serpentes: Atractaspididae): a study in conflicting functional constraints. Zoology 106, 43-61.

Deufel, A. & Cundall D. 2006. Functional plasticity of the venom delivery system in snakes with a focus on the post-strike prey release behavior. Zoologischer Anzeiger 245, 249-267.

Golani, I. & Kochva, E. 1988. Striking and other offensive and defensive behavior patterns in Atractaspis engaddensis (Ophidia, Atractaspididae). Copeia 1988, 792-797.

Kraus, F. & Brown, W. M. 1998. Phylogenetic relationships of colubroid snakes based on mitochondrial DNA sequences. Zoological Journal of the Linnean Society 122, 455-487.

Kurnik, D., Haviv, Y. & Kochva, E. 1999. A snake bite by the burrowing asp, Atractaspis engaddensis. Toxicon 37, 223-227.

Lee, M. S. Y. & Scanlon, J. D. 2002. Snake phylogeny based on osteology, soft anatomy and ecology. Biological Reviews 77, 333-401.

McDowell, S. B. 1987. Systematics. In Seigel, R. A., Collins, J. T. & Novak, S. S. (eds) Snakes: Ecology & Evolutionary Biology. Macmillan (New York), pp. 3-49.

Pyron, R. A., Burbrink, F. T. & Wiens, J. J. 2013. A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13, 93.

Rödel, M., Kucharzewski, C., Mahlow, K., Chirio, L., Pauwels, O. S. G., Carlino, P., Sambolah, G. & Glos, J. 2019. A new stiletto snake (Lamprophiidae, Atractaspidinae, Atractaspis) from Liberia and Guinea, West Africa. Zoosystematics and Evolution 95, 107-123.

Shine, R., Branch, W. R., Harlow, P. S., Webb, J. K. & Shine, T. 2006. Biology of burrowing asps (Atractaspididae) from southern Africa. Copeia 2006, 103-115.

Tilbury, C. & Verster, J. 2016. A fatal bite from the burrowing asp Atractaspis corpulenta (Hallowell 1854). Toxicon 118, 21-26.

Underwood, G. & Kovcha, E. 1993. On the affinities of the burrowing asps Atractaspis (Serpentes: Atractaspididae). Zoological Journal of the Linnean Society 107, 3-64.

Wollberg, M., Kochva, E. & Underwood, G. 1998. On the rictal glands of some atractaspid snakes. Herpetological Journal 8, 137-143.

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Among the most poorly known of all squamate groups are the dibamids…

**Caption:** a dibamid exemplar. Note the smooth-scaled, shiny overall look, the essentially eyeless head, and the pale patches on the snout and scattered across the body. This specimen, from the Philippines, was identified as *Dibamus* cf *leucurus*, so hadn’t been pinned down to species when published. Image: **Brown *et al*. (2016)**, CC BY 4.0 (**original here**).

… a group so obscure that they don’t really have a common or vernacular name (though read on). Little information on the group is available, and in this article I aim to cover essentially everything about them that I consider worth saying…

What are dibamids? They’re near-limbless, fossorial, snake-shaped, near-blind, insectivorous squamates mostly occurring in southern and south-eastern Asia (including the Wallacean region as far east as New Guinea). A single species – the Mexican blind lizard Anelytropsis papillosus – occurs in eastern Mexico. Excluding Anelytropsis, all the other species are included within the genus Dibamus. Members of both genera have been known to science for some time: Dibamus was named by French zoologists André Marie Constant Duméril and Gabriel Bibron in 1839 for D. novaeguineae (originally described from New Guinea, in case you’re guessing) while Anelytropsis* was first published by Edward Cope in 1885.

Within recent decades, the number of dibamid species has increased quite markedly. Books I recall from childhood (e.g., Whitfield 1983) said that there were just three species, a 1985 review increased this to nine (Greer 1985), and the steady and gradual description of new species since then has put us at 26 species as of May 2024. It’s not unlikely that the count will double within the next few decades.

**Caption:** the holotype specimen of *Dibamus dalaiensis*, discovered in the southwestern Cardamom Mountains, Cambodia, and described by Neang *et al*. (2011). It confirmed the presence of this group in Cambodia. Those patches that you might assume are areas where the skin is sloughing are actually pale grey areas, and a normal feature of the pigmentation. Note also the slight iridescent sheen on the dorsal reflective areas. Image: Neang *et al*. (2011).

A notorious taxonomic vandal, whose work cannot be taken seriously and whose taxonomic proposals should be ignored, has proposed several genus-level names for clades within Dibamus. For reasons discussed below, it’s almost inevitable that Dibamus will indeed undergo taxonomic revision at some point. When that happens, however, we won’t be using the name proposed by the taxonomic vandal alluded to here (see Kaiser et al. 2013, Rhodin et al. 2015).

All dibamid species are less than 25cm long, and a curious feature is that the tail is proportionally short, usually being less than one-third SVL (snout to vent length) and less than 10% of SVL in some Dibamus species (Greer 1985). Snakes are short-tailed relative to SVL, so this is a snake-like character.

**Caption:** I’ve said several times in older articles that my very first introduction to a good number of amphibian and reptile species came via Philip Whitfield’s *Reptiles and Amphibians: An Authoritative and Illustrated Guide* of 1983, and more specifically to the astonishing art by Alan Male. In cases, it was obvious that Male only had access to vague or poor reference photos or other images, and I think we can say that this was the case with his dibamid illustration, shown in the book on the same plate as anguids and anniellids. When the text was written, only three dibamid species were recognized. Image: Alan Male, from Whitfield (1983).

As ever with obscure reptiles, good luck if you want to find a comprehensive source that says much about these animals. Most books that cover squamate diversity mention dibamids in passing and don’t dwell on the things that make them interesting. The standard and best reference on them remains Allen Greer’s Journal of Herpetology paper from 1985, and I relied on it heavily here. It is fair to say, however, that dibamid skull anatomy is sufficiently unusual that many people interested in squamates have had reason to investigate it and ponder on what it might mean for squamate biology and evolution more broadly (Rieppel 1984, Greer 1985, Hallermann 1998).

Having said that dibamids don’t have a common name, both ‘blind lizard’ and ‘blind skink’ have been used. But both are less than ideal: ‘blind lizard’ is similar to ‘blind worm lizard’ used for some amphisbaenians (which are a very separate group, not closely related), and ‘blind skink’ is not good either since dibamids are not skinks nor close to them either.

**Caption:** at left, the head of *D. dalaiensis* from Cambodia in right lateral view, showing the prominence of the rostral and mental pads. At right, a montage showing the heads of (left to right) the *D. greeri* holotype, the *D. greeri* paratype, and the *D. smithi* holotype. The heads are shown in dorsal (upper row), left lateral (middle row) and ventral (lower row) views. The scale bar is 1 mm. Various of the scales have been labelled, including ocular (o), postocular (po), frontal (f) and mental (m) (there’s an error: some lower jaw scales labelled ‘im’ should be labelled ‘if’ for infralabial). All the main scales on the squamate head have names, and its typical to refer to their form and placement in descriptive discussions. Images: Neang *et al*. (2011), Darevsky (1992).

Dibamid anatomy, a primer. What are dibamids like in anatomical terms? Live dibamids are blunt-snouted, blunt-tailed, shiny-scaled squamates that range from purplish to dark brown, the underside being paler. The tongue is not bifid, but broad and fleshy. Pale patches are present across the body in some species. A peculiarity of some species – like D. greeri from Vietnam (named after the aforementioned Allen Greer) – is the presence of bright blue rings on the body. Darevsky (1992) noted that these can’t have any role in intraspecific signalling (given that dibamids are nearly blind, and subterranean too), so might represent mimicry of sympatric megascolecid earthworms, some of which possess unusually coloured body rings that are apparently toxic. Birds including spurfowl, peafowl and junglefowl might, Darevsky (1992) speculated, therefore avoid predating on dibamids… or, on some dibamids, anyway, and Darevsky (1992, p. 8) did bring attention to the un-tested and speculative nature of this hypothesis.

**Caption:** a beautifully detailed photo and diagram of *Dibamus manadotuaensis*, a species from Manado Tua, a small island off the northern coast of Sulawesi, Indonesia named in 2019. Note the tiny size and the small sensory papillae scattered across the surfaces of the large face plates. Extra points if you think this looks like the 2014 Gareth Edwards Godzilla, since it totally does. Image: Koppetsch *et al*. (2019).

Thick-skinned, milky-coloured plates occur across the anterior halves of the dibamid snout and lower jaw. Greer (1985) termed these the rostral and mental pad, respectively. They aren’t homogenous across their surfaces since they’re broken up by sutures (the distribution of which varies from one species to the next), and a “large number of evenly distributed sensory papillae” (Greer 1985, p. 121) also occur across their surfaces. The implication is that these structures have some kind of sensory role, but one that’s uninvestigated so far as I can tell. A pale area corresponding to the eye is present in some species but not all.

The dibamid skull is rigid (as is typical for burrowing squamates), has closed fenestrae (again, as typical for burrowers), and is tiny at just 5-7 mm in length. Numerous details are unusual relative to the condition present in other squamates. The bony palate is one of the most ‘complete’ within Squamata, extending as far posteriorly as the anterior edge of the braincase, albeit with a midline furrow at its rear margin. This is occupied by spongy, possible glandular, tissue in life (Greer 1985). Within the snout chamber, scroll-like structures and bony laminae enclose the nasal passages (Greer 1985, Hallermann 1998). Similar configurations are seen in the fossorial acontine skinks and the unusual feylinines (Greer 1985) – there are some similar laminae in uropeltid snakes as well – so presumably this is something to do with breathing while burrowing through sediment.

**Caption:** it’s not often that you get to see the cartilaginous nasal capsule within the snout of a squamate. This diagram – from Hallermann (1998) – was based on reassembled serial sections. The nasal capsule – the sinuous structure closest to us in the diagram – is paired, and each one is separated along the midline by a septum (nas. sept.). Each capsule has a complex relationship at the snout’s tip with the septum, the capsule forming a curled anterior nasal cupola (a. nas. cup.). Hallermann (1998, p. 389) suggested that this complex anterior anatomy might be “a consolidation in adaptation to burrowing habits”.

The lower jaw is simplified (being formed of just three bones, the most posterior of which looks to be a compound amalgamation of the ancestrally separate five), and has a low tooth count, there being 8-10 teeth (on one side) depending on the species. The tooth count in the upper jaw is harder to discuss. Why? Because the conjoined premaxillae (the bones at the front of the upper jaw) sometimes have an odd number of teeth (7) due to the presence of a midline tooth. This feature is not unique within squamates since it also occurs within some geckos – remember this point!

**Caption:** dibamids can’t be described as limbless. At left, the hindlimb and pelvis of male *Dibamus* specimens, as figured by Greer (1985). A and B show bones from a male *D. taylori* (with fibula and tibia); C and D show bones from a female *D. novaeguineae*. The scale bars are 1 mm. The pelvic girdle is mostly formed of a large ilium (il), with a tiny pubis (pu) and ischium (is) persisting as small ossicles visible in B. At right, the flipper-like hindlimbs of a male *D. floweri* from peninsular Malaysia, a species described in 2017. I don’t think it’s typical for the hindlimbs to be crossed like this. Images: Greer (1985), Quah *et al*. (2017).

Another anatomical area of interest concerns limbs, or their absence. Forelimbs are wholly lacking and only a relictual, splint-like pectoral girdle remains. On the issue of hindlimb presence or absence, dibamids are ambiguous, one reason being that they’re sexually dimorphic on this point: females lack limbs entirely, but males have small, flap-like hindlimbs that they use to grip females while mating. On bones, males have a femur, tibia and fibula (plus some attendant cartilage caps) while females just have a femur. Anelytropsis is variable with respect to all this, some males having a tibia and fibula and others being like females in lacking them (Greer 1985). One final thing on skeletal anatomy: it’s been stated that osteoderms are present in some dibamids but this was regarded as a mistake by Greer (1985).

**Caption:** the complete skeleton of the recently named Vietnamese dibamid *Dibamus deimontis*, specifically the male paratype. It’s 15.3 cm long in total. Note the relatively well developed hindlimb (though lacking a foot) and pelvis, and the relatively short length of the tail. Image: Kliukin *et al*. (2024).

Reduced eyes, lone lungs and erectile scales. Dibamid eyes are reduced (no sclerotic ring, no lens, no iris, no vitreous body), covered by scales, and presumably only have a role in light sensing. Both external ears and a middle ear cavity are absent.

On organs, dibamids are like some amphisbaenians in only having a single lung: on the left side, it’s anterior to the heart, but on the right side it’s posterior to the heart. This is different from the condition in snakes, since in them both lungs are generally present and it’s the right lung that’s larger and more important than the left. Alveoli are apparently absent in the dibamid lung (Greer 1985), as is the case in snakes. Another snake-like feature is that the heart is situated far back in the body (relative to what we mammals are familiar with).

An especially notable dibamid features concerns the ability of at least some species to elevate their body scales such that they project almost perpendicularly from the body. This gives them a rugose or wrinkled appearance: one suggestion is that this is another convergence with megascolecid earthworms that inhabit the same environment, since some of those are bristle-covered (and presumably toxic) (Diaz et al. 2004, Kliukin et al. 2023). Prior to reading about this, I wasn’t aware that the scales of any squamates were mobile in this manner, and the fact that they are in at least some groups raises all kinds of questions…. and possibilities if you’re interested in speculative evolution.

**Caption:** some preserved dibamid specimens are almost orange. However, this is almost certainly because they’ve been sitting in preservative fluid for years or decades, and it likely doesn’t reflect their colour in life. This is the holotype of *Dibamus tebal* from Pulau Simeuleu (off the south-west coast of Sumatra), named as a new species in 2016 but collected at some point in the 1910s (Das & Lim 2009). The scale bar is in millimetres; the total length is 158 mm. Image: Das & Lim (2009).

Distribution and habitat, part 1. There’s a lot to say about dibamid distribution and what it means for our knowledge of the group. First things first, their disjunct distribution – they occur in eastern Asia, western Australasia and Mexico and nowhere in between – obviously requires an explanation. If this group has a history extending back to the Paleogene or earlier (as it likely does), then maybe this distribution reflects an older, continuous one whereby species formerly occurred across Beringia and North America. If so, we predict the discovery of fossil species in North America and perhaps elsewhere.

On that note, if this group evolved especially early in squamate history, it might even be that they were substantially more widespread in the distant past. An Early Cretaceous or Jurassic origin, for example, could mean a pan-Laurasian distribution, perhaps even a pan-Pangaean one.

An alternative explanation of dibamid distribution is that over-water dispersal occurred, and of course small burrowing reptiles of this sort could conceivably travel out to sea in sediment associated with big floating root masses or clusters of plants. Townsend et al. (2011) examined dibamid distribution and proposed that dibamids crossed the Beringian landbridge during the Paleocene or Eocene, but did also consider the possibility of trans-Pacific rafting. Townsend et al. (2011) also found Dibamus to be paraphyletic, with one set of species being closer to Anelytropsis than to other Dibamus species. This implies that Anelytropsis is geologically young within the group, that eastern Asia is the group’s ‘centre of origin’, and that the movement that explains their disjunct distribution was eastwards in relation to the Pacific.

**Caption:** the area of Nui Chua National Park, Ninh Thuan Province, southern Vietnam, in which *Dibamus tropcentr* was discovered. This is a hot and dry location, and not the sort of habitat that you might ordinarily associate with dibamids. The holotype was specifically found in a rotten log, in association with termites. Image: Kliukin *et al*. (2023).

If dibamids have an evolutionary history going back at least 40 million years, we might speculate that they include more diversity – in ecology, behaviour and physiology – than generally thought. I’d thought from prior reading on the group that they were all animals of tropical forest floors, but this is not so: the Asian/Australasian species occur in tropical and subtropical forest environments while the Mexican one appears to be a habitat generalist, occurring in cloud forest, brush-covered, arid mountains, deciduous woodland, arid scrubby plains and elsewhere. And having said that the Asian/Australasian species “occur in tropical and subtropical forest environments”, it’s important to note the diversity of habitats they occur in, since there are species in lowland arid habitats (including dry maritime evergreen forest) and upland forests too.

Incidentally, Darevsky (1992) reported that the holotype of D. greeri was discovered within a mass of soil, overgrown with epiphytes attached to a moss-covered tree, that fell to the ground from a height of about 3 metres. So, this dibamid was living up in a tree. Presumably, the animal had travelled up the tree by climbing through and under the thick moss layer. Worms and fossorial arthropods are also sometimes found in soil attached to trees in the tropics, so this is not a wholly surprising observation.

**Caption:** Nui Chua Mountain in Nui Chua National Park in southern Vietnam, the type locality for *D. deimontis*, described this year (Kliukin *et al*. 2024). The arrow marks the discovery spot. This is an area where mixed montane evergreen forest occurs alongside grassy areas. There are also streams that contain large, root- and moss-covered granite boulders. The dibamids were collected from the boulder surfaces where they were associated with the mosses and roots. Image: Kliukin *et al*. (2024).

Distribution and habitat, part 2. There’s one more thing I want to say on distribution and habitat, and this is that our scant knowledge of these animals – most species are known from only a handful of specimens – means that the map is quite patchy in terms of where we know they occur. On mainland Asia, the group’s most northerly occurrence is Guangxi Province in southern China, with Hong Kong being their most easterly occurrence. In Southeast Asia, they’re known from the north and south of Vietnam as well as its narrow central section, from south-eastern as well as extreme southern, peninsular Thailand, western and probably northern Cambodia, and the northern half of Malaysia.

As you can see from the map I created (see caption for caveats!), large gaps are present in confirmed dibamid distribution. These gaps suggest that dibamids await discovery in numerous places, including central and southern Vietnam, much of Cambodia, some of eastern and southern Thailand, and probably southern Laos at least. The presence of the group in Tioman Island, adjacent to eastern mainland Malayasia, hints at a presence in southern Malaysia. Certain of the mainland locations from which dibamids are known are (relatively) ‘recent adds’: the 2011 naming of D. dalaiensis from Cambodia, for example, was significant enough that I covered it at Tet Zoo.

**Caption:** by combing the primary literature on dibamids, I extracted as much locality data on Asian/Australasian dibamids as I could and depicted it on this map. I marked ‘small island’ locations with a spot or oval but attempted to depict ‘large island’ and mainland locations with an extrapolated range. Some caveats are necessary: when several records are known from the same approximate area (e.g., southern Vietnam), I extrapolated a rough range area by ‘connecting the dots’. Records from Sumatra and Sulawesi suggest that dibamids occur across the better part of these islands, but this likely isn’t correct and a more detailed examination would result in far patchier ranges for these regions. Anyway… the key point here is that dibamids remain unknown or scarcely known from many places across the region, and could await discovery in many additional localities.

Islands are important in dibamid distribution. The group’s most westerly Asian occurrence is on the Nicobar Islands, which – politically – are part of India. On the Indonesian islands, dibamids have a known presence (going from west to east) in Sumatra and some surrounding islands, northern Borneo, much of Sulawesi, various of the Lesser Sunda Islands, and Seram. They’re present on some of the southern islands of the Philippines and western New Guinea too. Again, there are lots of gaps here, and I think it’s reasonable to suggest that dibamids might prove more widely distributed in Borneo and might also await discovery on quite a few additional Indonesian islands, including Taliabu, Buru and numerous other of the Maluku Islands, Timor, and Java.

At the moment, the majority of Asian dibamid species occur on islands. But the fact that several new ones have been named from the Asian mainland in recent years indicates that their continental diversity has been unappreciated and Kliukin et al. (2023, p. 318) recently stated that “it is highly likely that the territory of the mainland Indochina, including Vietnam, cradles a comparable if not higher diversity of Dibamus lizards”. They ended their text by emphasizing that more sampling is needed in appropriate habitats in nations like Vietnam.

**Caption:** maps from recently published descriptions of Asian dibamid species, showing locations relevant to the discovery of species from eastern Southeast Asia. The map on the left is from Kliukin *et al*. (2023), the description of *Dibamus tropcentr* from eastern Vietnam; that on the right is from the description of *D. deimontis* Kliukin *et al*., 2024 from the same area. The publication of *D. deimontis* takes us to 26 recognized species. Images: Kliukin *et al*. (2023), Kliukin *et al*. (2024).

While we’re here… similar comments could be made about the distribution of Anelytropsis in Mexico. Its current distribution is also patchy, there being a large gap between the type locality in central Veracruz and its other occurrences in southern Tamaulipas, San Luis Potosí, and northern Guanajuato and Querétaro. Based on its ecological flexibility, some authors of decades past suggest that it might await discovery further west and south. When Axtell (1958) reported the most northerly known occurrence – at Ciudad Victoria in Tamaulipas – he noted that this might be indicative of “a more widespread range at a time (early Pleistocene?) when humid conditions were more pronounced” (Axtell 1958, p. 189). More record records indicate that this view of a wider Mexican distribution is likely correct: Valdez-Vilavicencio et al. (2016) reported occurrences in Oaxaca in southern Mexico, and Montiel-Veranza et al. (2022) described the presence of the species in Comapa in Veracruz, which is further to the east in Veracruz than previous records.

**Caption:** images of *Anelytropsis* from recently published range extension records. At left, an orange specimen from Oaxaca in southern Mexico, reported by Valdez-Vilavicencio *et al*. (2016) (again, the specimen had been kept in preservative for some years prior to 2016, so this colour might not represent the condition in life); at right, a pinker one from Comapa in Veracruz, reported by Montiel-Veranza *et al*. (2022).

And… where do dibamids fit in phylogeny? Moving now away from distribution and biogeography, where do dibamids fit in the squamate family tree? I was planning to cover that issue in this article but there’s enough to say about it that I’m going to postpone coverage and return to it later. The spoiler is that dibamids have proved enigmatic, with different authors proposing quite different phylogenetic positions for the group, and with several pushing the hypothesis that dibamids might link amphisbaenians and snakes and prove important in models of snake origins (this being one of the most contentious issues within the field of squamate evolution). Some authors regard dibamids as a highly distinct, ancient lineage within Squamata, and it’s mostly for that reason that the name Dibamia – it implies a ‘higher’ taxonomic status than the ‘family-level’ Dibamidae – has been used by some authors. We will return to these topics in time…

Incidentally, I promised back in 2011 that “much more on dibamids” would appear at “some other time”, so never let it be said that I don’t keep my promises. Sorta.

For previous articles on squamates, see…

Cambodia: now with dibamids!, May 2011
The Tet Zoo Guide to Mastigures, August 2018
The Remarkable Basilisks, May 2023
Do Lizards Really Have ‘Mite Pockets’?, March 2024
Meeting Lake Zacapu’s Garter Snake, March 2024

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

Refs - -

Axtel, R. W. 1958. A northward range extension for the lizard Anelytropsis papillosis, with notes on the distribution and habits of several other Mexican lizards. Herpetologica 14, 189-191.

Brown, R. M., Sanguila, M. B., Cobb, K. A., Siler, C. D., Diesmos, A. C., Alcala, A. C. & Brown, R. M. 2016. The amphibians and reptiles of Mindanao Island, southern Philippines, II: the herpetofauna of northeast Mindanao and adjacent islands. ZooKeys 624, 1-132.

Darevsky, I. S. 1992. Two new species of the worm-like lizard Dibamus (Sauria: Dibamidae) with remarks on the distribution and ecology of Dibamus in Vietnam. Asiatic Herpetological Research 4, 1-12.

Das, I. & Lim, K. K. 2009. A new species of Dibamus (Squamata: Dibamidae) from Pulau Simeuleu, Mentawai Archipelago, Indonesia. Zootaxa 2088, 15-23.

Diaz, R. E., Leong, T. M., Grismer, L. L. & Yaakob, N. S. 2004. A new species of Dibamus (Squamata: Dibamidae) from West Malaysia. Asiatic Herpetological Research 10, 1-7.

Greer, A. E. 1982. 1985. The relationships of the lizard genera Anelytropsis and Dibamus. Journal of Herpetology 19, 116-156.

Hallermann, J. 1998. The ethmoidal region of Dibamus taylori (Squamata: Dibamidae), with a phylogenetic hypothesis on dibamid relationships within Squamata. Zoological Journal of the Linnean Society 122, 385-426.

Kliukin, N. S., Bragin, A. M., Nguyen, T. V., Le, S. X., Tran, T. T. V., Gorin, V. A. & Poyarkov, N. A. 2024. Another new species of Dibamus Duméril & Bibron, 1839 (Squamata: Dibamidae) from Nui Chua National Park, Ninh Thuan Province, Vietnam. Zootaxa 5406, 87-104.

Kliukin, N. S., Nguyen, T. V., Le, S. X., Bragin, A. M., Tran, V. T., Gorin, V. A. & Poyarkov, N. A. 2023. A new species of the genus Dibamus Duméril & Bibron, 1839 (Squamata: Dibamidae) from the driest and hottest place of Vietnam. Zootaxa 5380, 301-320.

Koppetsch, T., Böhme, W. & Koch, A. 2019. A new species of Dibamus Duméril & Bibron, 1839 (Squamata: Dibamidae) from Pulau Manado Tua, Northern Sulawesi, Indonesia. Zootaxa 4555, 331-345.

Montiel-Veranza, J. A., Vásquez-Cruz, V. & Morales-Leal, I. 2022. New records of Anelytropsis papillosus Cope, 1885 (Squamata: Dibamidae) for the Municipality of Comapa, in the central-western of Veracruz, Mexico. Revista Latinoamericana de Herpeología 5, 139-141.

Quah, E. S. H., Shahrul Anuar M. S., Grismer, L. L. & Grassby-Lewis, R. 2017. A new species of Dibamus Duméril & Bibron 1839 (Squamata: Dibamidae) from a hill station in Peninsular Malaysia. Raffles Bulletin of Zoology 65, 681-690.

Rhodin, A. G. J., Kaiser, H., van Dijk, P. P., Wüster, W., O’Shea, M., Archer, M., Auliya, M., Boitani, L., Bour, R., Clausnitzer, V., Contreras-MacBeath, T., Crother, B. I., Daza, J. M., Driscoll, C. A., Flores-Villela, O., Frazier, J., Fritz, U., Gardner, A., Gascon, C., Georges, A., Glaw, F., Grazziotin, F. G., Groves, C. P., Haszprunar, G., Havaš, P., Hero, J. M., Hoffmann, M., Hoogmoed, M. S., Horne, B. D., Iverson, J. B., Jäch, M., Jenkins, C. L., Jenkins, R. K. B., Kiester, A. R., Keogh, J. S., Lacher Jr., T. E., Lovich, J. E., Luiselli, L., Mahler, D. L., Mallon, D., Mast, R., Mcdiarmid, R. W., Measey, J., Mittermeier, R. A., Molur, S., Mossbrugger, V., Murphy, R., Naish, D., Niekisch, M., Ota, J., Parham, J. F., Parr, M. J., Pilcher, N. J., Pine, R. H., Rylands, A. B., Sanderson, J. G., Savage, J., Schleip, W., Scrocchi, G. J., Shaffer, H. B., Smith, E. N., Sprackland, R., Stuart, S. N., Vetter, H., Vitt, L. J., Waller, T., Webb, G., Wilson, E. O., Zaher, H. & Thomson, S. 2015. Comment on Spracklandus Hoser, 2009 (Reptilia, Serpentes, ELAPIDAE): request for confirmation of the availability of the generic name and for the nomenclatural validation of the journal in which it was published. (Case 3601; see BZN 70: 234-237; 71: 30-38, 133-135, 181-182, 252-253). Bulletin of Zoological Nomenclature 72 (1): 65-78.

Rieppel, O. 1984. The cranial morphology of the fossorial lizard genus Dibamus Dumeril & Bibron, with a consideration of its phylogenetic relationships. Journal of Zoology 204, 289-322.

Townsend, T. M., Leavitt, D. H. & Reeder, T. W. 2011. Intercontinental dispersal by a microendemic burrowing reptile (Dibamidae). Proceedings of the Royal Society B: Biological Science 278, 2568-2574.

Valdez-Villavicencio, J., Garcia-Padilla, E. & Mata-Silva, V. 2016. Anelytropsis papillosus Cope, 1885 (Squamata: Dibamidae), an overlooked species in the state of Oaxaca, Mexico. Mesoamerican Herpetology 3, 178-180.

Whitfield, P. 1983. Reptiles and Amphibians: An Authoritative and Illustrated Guide. Longman Group Ltd, Harlow, UK.

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For some years now, a prolific amateur herpetologist has published an absolutely extraordinary number of new taxonomic names for snakes, lizards and other reptiles…

**Caption:** lest we forget, the world is full of amazing snakes. Top row, left to right: Prairie or Western rattlesnake *Crotalus viridis*, Bornean keeled green pitviper *Tropidolaemus subannulatus* and Mole snake *Pseudaspis cana*. Lower row, left to right: Rock rattlesnake *Crotalus lepidus*, Rhinoceros viper *Bitis nasicornis* and Smooth-scaled death adder *Acanthophis laevis*. All photos by Wolfgang Wüster and used with permission.

In addition to naming well over 100 supposedly new snake and lizard genera, this individual has also produced taxonomic revisions of the world’s cobras, burrowing asps, vipers, rattlesnakes, water snakes, blindsnakes, pythons, crocodiles and so on. But, alas, his work is not of the careful, methodical, conservative and respected sort that you might associate with a specialized, dedicated amateur. Instead, his articles appear in his own, in-house, un-reviewed, decidedly non-technical publications. These are notoriously unscientific in style and content, and his taxonomic recommendations have been demonstrated to be problematic, frequently erroneous and often ridiculous. He has, for example, named many new taxa after his pet dogs.

In short, the new (and really terribly formulated) taxonomic names that this individual throws out at the global herpetological community represent what’s known as TAXONOMIC VANDALISM. We’re expected to use these names, and – indeed – they’re supposedly officially valid according to the letter of the law, yet they besmirch the field, litter the taxonomic registry with monstrosities, and cause working herpetologists to waste valuable time clearing up unnecessary messes.

**Caption:** Egyptian cobra *Naja haje*: included within the *Naja* subgenus *Uraeus* by **Wallach *et al*. (2009)** but given the new genus ‘Wellsus’ by Hoser. Hoser’s proposed name honours Richard Wells (on which, read on). Image: (c) Wolfgang Wüster, used with permission.

I am of course talking about Australian researcher and snake hobbyist Raymond Hoser. The charges against him are many. I’ve mentioned Hoser on a few previous occasions at Tet Zoo, most notably in the article on Australian freshwater crocodiles. It’s time to explore the issue in more depth. UPDATE: the article you’re reading here was originally published at Tetrapod Zoology ver 3 (the Scientific American years) in 2013 and includes several post-2013 updates.

A bit of required background: on taxonomic freedom and the Principle of Priority. One of the key principles of zoological taxonomy – the practice and science of naming animals – is what’s known as taxonomic freedom. It’s acknowledged that not all experts agree on how animals should be classified when it comes to where the boundaries are. Are a group of individuals all part of the same species, or do some actually represent a distinct, second species? Debates over species boundaries and how taxonomy should reflect those boundaries are commonplace and it can take a lot of work to sort them out, typically via statistical analyses of large numbers of individuals, molecular phylogenetics and so on.

But, whatever, if one or more authors do regard individuals within a group as a distinct taxon – here, we’re mostly interested in debates at the species level – they’re within their rights to act on this within the literature. ‘Taxonomic freedom’ allows this.

**Caption:** the African crocodile *Crocodylus suchus*, placed in the new genus ‘Oxycrocodylus’ by Hoser. What’s the etymology? The names honours Hoser’s dog, Oxyuranus (itself named after the Australian snake). Image: stuart Burns [sic], CC BY 2.0 (**original here**).

A well-established rule of the taxonomic naming system is the so-called Principle of Priority. Because people sometimes name the same organism more than once (sometimes because they don’t know about the work of their predecessors, sometimes because they think they’re dealing with a new taxon when they actually aren’t, and sometimes because they’re deliberately trying to beat other authors into print), it’s agreed that the very first name given to an organism is the one we have to stick with, even if that first name is deemed poorly chosen in some way.

There are special cases where a name can be overturned but, by and large, the Principle of Priority is pretty important and more or less guarantees a namer’s ‘place in history’. Remember that the full scientific name of an organism includes more than the organism’s name alone: Homo sapiens, for example, is properly Homo sapiens Linnaeus, 1758.

**Caption:** the Principle of Priority makes sense, but it’s not always our friend. We’re stuck, for example, with the name *Basilosaurus* for this extinct whale. Image: Tim, CC BY-SA 2.0 (**original here**).

So, if you encounter an animal that you, personally, regard as worthy of distinct taxonomic recognition, you are – thanks to taxonomic freedom – within your rights to name it as such, and – thanks to the Principle of Priority – you’ve earned a place in history if you’re the first person to name the taxon in question. It’s assumed that you’ve correctly followed the rules set out in the ICZN (= International Code of Zoological Nomenclature) by the ICZN (= International Commission on Zoological Nomenclature). Yeah, same acronym. Once your new name is published, it’s essentially forever burned into history.

**Caption:** Hugh, what have you done? Hugh Edwin Strickland (1811-1853), as a young man of 26 at left (illustration by Francis William Wilkin, dating to 1858), and as an… older man at some later date. However, he died at age 42, so never got to be ‘old’ at all. In 1842, Strickland worked with others in the British Association to establish the Principle of Priority. Images in public domain.

What are these “rules set out in the ICZN”? As you can see for yourself at the ICZN site, a new name has to be published in a permanent, duplicable form that’s available to others, it has to be clearly stated as a new name, it has to be published within the context of the binomial (or binominal) system, and it must be established on a type specimen (a key reference specimen). Notably, many of the key ideas that we typically associate with the publication of scientific research – like standards of practice, an appropriate level of scholarship, and peer review – are, actually, not required by the ICZN. Perhaps surprisingly, the ICZN is actually quite lax about those venues that past muster when it comes to the publishing of new taxonomic names.

In other words, individuals can work within the mandates of the International Code of Zoological Nomenclature even if their conclusions, proposals and work in general is problematic and unsatisfactory. They can still name new species that, technically speaking, are valid, available and (in theory) fixed due to the Principle of Priority.

Mega-prolific Raymond Hoser: one of the greatest herpetologists of all time! Back to Hoser. I don’t necessarily mean to denigrate Hoser’s research abilities, experience with snakes and other reptiles, or intelligence. As others before me have said, it’s obvious that he does have extensive, impressive, detailed knowledge of reptile diversity, anatomy and biology. But the fact is that he is very obviously ‘cheating’ his way through zoological nomenclature. Yes, he’s naming, and publishing umpteen new herpetological names. If you want some figures: between 2000 and September 2012 alone he named 89 tribes and subtribes, 113 genera, 64 subgenera, 25 species and 53 subspecies. That’s 76% of all new genera and subgenera named worldwide during that period (Kaiser et al. 2013). If these taxonomic recommendations and proposals were valid, they would make Hoser a more significant taxonomic force than most of the great explorer-herpetologists of the 19th and 20th centuries.

Unlike the respected work produced by the experts of the past, however, Hoser’s is amazingly slapdash.

**Caption:** George Boulenger (1858-1937), Belgian-British zoologist and prolific describer of new amphibians, reptiles and other organisms. In all, he named over 2000 new species (556 of which are amphibians and 872 of which are reptiles). His impact on our knowledge has been enormous. Just a few of the species he named are featured here. Top row, left to right: African dwarf frog *Hymenochirus boettgeri*; New Caledonian gecko *Rhacodactylus trachycephalus*. Middle row, left to right: Stripe-tailed goanna *Varanus caudolineatus*; Sierra Leone water frog or Sabre-toothed frog *Odontobatrachus natator*; Blanford’s flying lizard *Draco blanfordii*. Lower right: Java grass lizard *Takydromus khasiensis*. Images: Boulenger image in public domain; James Gathany, public domain (**original here**); Lennart Hudel (CC BY 4.0, **original here**); Barej *et al*. (2015) (CC BY 4.0, **original here**); Haplochromis (CC BY-SA 3.0, **original here**); Rushenb (CC BY-SA 4.0, **original here**); Rohit Naniwadekar (CC BY-SA 4.0, **original here**).

In fact, the impression you get from Hoser’s articles – all appearing in his self-published Australasian Journal of Herpetology, and before that in several amateur publications including Litteratura Serpentium and The Monitor – is that they’re written as much to piss off working herpetologists and to vent his own spleen as anything else. They are shockingly and hilariously unscientific. Many include long rants directed at officials and employees of local government as well as at qualified researchers. There are in fact so many cases of this non-scientific – in fact, truly amateurish, if not childish – practice in his articles that there are too many to recount.

Over the years, Hoser has had at least a bit of appropriate criticism (e.g., Aplin 1999, Wüster et al. 2001, Williams et al. 2006, Borrell 2007). He refers to those qualified herpetologists who criticize him and his work as “the truth haters”: the obvious implication being that he’s on the side of ‘The Truth’. Jeez, what is it with people on the fringes and their adherence to the notion that only they are seeing The Truth? Incidentally, Hoser frequently charges those of us who criticize him as ‘plagiarists’. He directed that specific charge at me after I wrote (unfavourably) about his crocodile article. I can only conclude that he doesn’t know what that word really means.

**Caption:** Australia sure has some amazing elapids. Given that Hoser claims to have the interests of the animals at heart, it’s bizarre that he defaces their taxonomy with horrible names that never honour the animals themselves (read on for examples). This is *Oxyuranus microlepidotus*, the Fierce snake or Inland taipan. Image: XLerate, CC BY-SA 3.0 (**original here**).

If you’re curious about the technical shortcomings of Hoser’s taxonomic proposals, let’s look at a few of them. In order to make a claim for the distinctive nature of an alleged new taxon, you need to state those features that make it distinctive. In other words, you need to diagnose it. Hoser’s diagnoses are typically inadequate, contradictory, vague, or erroneous, sometimes referring to features that aren’t unique (example: the black labial markings in the proposed death adder species 'Acanthophis crotalusei'), and sometimes referring vaguely to work on scale counts, overall appearance or DNA that hasn’t been documented or published anywhere (Wüster et al. 2001, Kaiser et al. 2013).

**Caption:** death adders are incredible snakes. This is the Smooth-scaled death adder *Acanthophis laevis* of New Guinea and several islands to its west, including Ceram. In articles of 1998 and 2002, Hoser claimed the recognition of several additional taxa for populations otherwise considered part of this species. Their recognition was mostly rejected by **Kaiser *et al*. (2013)**. Additional work is required to resolve the taxonomy of these snakes. Image: Petra Karstedt, CC BY-SA 2.0 (**original here**).

Alleged new taxa have also been based on specimens where the ‘diagnostic’ characters clearly represent post-mortem distortion (see examples discussed in Kaiser et al. 2013). To add insult to injury, Hoser has sometimes named the same alleged new taxa on more than one occasion. 'Leiopython albertisi barkeri' Hoser, 2000 is the same as 'L. a. barkerorum' Hoser, 2009 which was then redescribed as if it were new in 2012. Similarly, 'Oxyuranus scutellatus barringeri' Hoser, 2002 is the same as 'O. s. andrewwilsoni' Hoser, 2009 (Kaiser et al. 2013). Many of the new names Hoser creates are incorrectly formulated: see Wüster et al. (2001) for a list and their amendments.

Slapping names on cladograms: it’s quick, it’s cheap, it’s dirty. Another thing that Hoser does is look at published cladograms, note cases where genera or species are shown as being non-monophyletic, and then name (in cursory, name-grabbing fashion) those lineages that don’t group with the type species of the given genus. This is known as nomenclatural harvesting (Denzer & Kaiser 2023).

**Caption:** Keeled slug snake *Pareas carinatus*. Close relatives of this species – conventionally included within *Pareas* – were put into the new genus 'Katrinahoserserpenea' by Hoser. *Pareas* is one of several genera within Pareidae, a poorly known and mostly tropical Asian colubroid snake group specialized for eating snails. Image: W. A. Djatmiko, CC BY-SA 4.0 (**original here**).

There are numerous examples of this; they explain why Hoser has published such names as 'Katrinahoserserpenea' for certain Oriental slug-eating snakes (Hoser 2012a), 'Katrinahoserea' for the Green ratsnake (Hoser 2012b), 'Swileserpens' for the Pale-headed forest snake (Hoser 2012c), 'Michaelnicholsus' for members of the Madagascan hognosed snake group (Hoser 2012c), 'Lukefabaserpens' and 'Ginafabaserpenae' for some of the cat-eyed snakes (Hoser 2012d), 'Gregwedoshus' and 'Neilsonnemanus' for certain garter snakes (Hoser 2012e), 'Jackyhosernatrix' for certain natricine water snakes (Hoser 2012f), 'Sharonhoserea' for the Southern smooth snake (Hoser 2012f), and so on and on and on.

Note the terrible, terrible names that Hoser comes up with. Other notable word-monsters of his include 'Adelynhoserserpenae', 'Charlespiersonserpens', 'Euanedwardsserpens', 'Moseselfakharikukri', 'Trioanotyphlops' and 'Martinwellstyphlops'. If this doesn’t qualify as taxonomic vandalism, it’s difficult to know what does. Most (maybe all?) of Hoser’s taxonomic names are patronyms: names that honour people. That’s fine, but there has to be a limit to this sort of behaviour, especially when the namer is repeatedly naming things after the members of their own family and after their pets. As I said earlier, Hoser has named several taxa after his pet dogs, explaining at length how these noble canines have contributed more to herpetology than have the majority of the world’s researching academics (e.g., Hoser 2012g). Again, this is transparently taxonomic vandalism. A deliberate mockery of the field and the work of other researchers.

**Caption:** Hoser frequently points to cladograms (such as this one: this is the natricine section of Pyron *et al*.’s (2011) giant colubrid phylogeny) to support the taxonomic splits and renamings that he proposes. Yes, non-monophyly abounds in studies like this. But is it right to jump all over the cladogram and get to work slapping names all over the place? These things take time and a lot of work to sort out and do properly.

We do, of course, all know of cases where long-standing genera and/or species do indeed warrant revision. However, how are researchers meant to act when they spot these sorts of problems? My suggestion: once such a problem has been identified, good practice is to compile and run your own analysis, not to rush out a brief, non-illustrated article, the only purpose of which is to slap a name on a given lineage. If a researcher played the name-bagging name once in their career they might be forgiven (as I said, we all know of cases where new names are needed and people are just waiting for someone to come along and publish an update). But if they did this as a matter of course, again and again and again, typically naming new taxa after their family members and such, it would be pretty clear that they were deliberately and desperately ‘name-bagging’ in the hope for taxonomic immortality.

Incidentally, if, at this stage, you’re thinking that the taxonomic names we apply to snakes and other reptiles don’t really matter, think again. The whole reason we give names to things is so that we can talk about those things with other people. Confusion and disagreement are the opposite of useful when we’re dealing with conservation and summoning up the political and social will to protect animals and their environments. Furthermore, venomous snakes are a special case since a stable nomenclature known to people in the healthcare profession is a must; or it is, at least, if you want people to get the right antivenom after they get bitten.

**Caption:** the extremely variable Timber, Canebrake or Banded rattlesnake *Crotalus horridus* of eastern North America. This species can be regarded as ‘the original’ rattlesnake, the first one that European colonizers got to know and name scientifically. In Hoser’s proposed taxonomy, this is the only species retained within the subgenus *Crotalus* of his restricted version of the genus *Crotalus*. Images (clockwise from left): Tad Arensmeier, CC BY-SA 2.0 (**original here**); Tanner Smida, CC BY-SA 4.0 (**original here**); Glenn Bartolotti, CC BY-SA 4.0 (**original here**).

Has anybody, actually, yet become confused by the fact that spurious and problematic name changes have been suggested for any of these animals? After all, most working herpetologists have deliberately ignored and not used Hoser names. However, the Brazilian Society of Herpetologists adopted Hoser’s (2009) taxonomic arrangement for rattlesnakes, and this had a knock-on effect in the Brazilian literature. As argued by Wüster & Bérnils (2011), Hoser’s suggestions for rattlesnakes were redundant in the first place (they mostly involve subjectively sub-dividing an already monophyletic entity, namely Crotalus), are inconsistent with some published phylogenetic work, and are based on the assumption that certain parts of the phylogeny are resolved and ‘fixed’ for the foreseeable future. These are the technical problems; there are the additional ones related to the standing of Hoser’s articles in the first place (Wüster & Bérnils 2011).

**Caption:** the awesome Western diamondback rattlesnake *Crotalus atrox*, a species endemic to the southwestern USA and Mexico and among the largest rattlesnakes of them all (giant specimens can exceed 1.8 m). Hoser wants *C. atrox* to be recognized as the type species for his new genus ‘Hoserea’ (named for his wife). Image: Gary Stolz, United States Fish and Wildlife Service in public domain (**original here**).

What to do? Quality control should be integral to taxonomic publications. What can we actually do about taxonomic vandalism of this scale? Even the most lenient of liberal libertarians will agree that there’s a problem here: we clearly have an individual who isn’t using the same standards – or anything close to them – when publishing new names, and is insanely prolific to boot. Given what I said above about taxonomic freedom and the Principle of Priority, it’s essentially impossible to use the ICZN to discount or dismiss or ignore or strike off names that have been published and which meet the basic criteria discussed above. Or is it?

In 2013, a group of working herpetologists published a concise and very readable point-of-view piece on the subject in Herpetological Review (Kaiser et al. 2013). Note that the article is open access. Just to prove what a professional, ethical individual he is, Hoser somehow got hold of a version of this article before it was published and published it himself, in full (Hoser 2012h).

**Caption:** as we’ve seen, the ICZN is surprisingly lax about what is and what is not permissible when it comes to the acceptability of taxonomic names. That would be fine and good if all people publishing taxonomic names were doing work of a high, respectable standard. But they’re not. Communities of workers must therefore take a stand and state that there has to be a limit to what we can and will accept. Such is stated here in the title of **Kaiser *et al*. (2013)**.

Predictably enough, Hoser (2013) later published another article in which, over more than 60 pages, he responded in characteristic fashion to Kaiser et al. (2013), referring to them as “alleged scientists” and “serial liars” throughout; he even (for reasons best known to himself) kept calling their article a “blog” (a blog is an updated, diary-style website: the word is not synonymous with ‘article’). Hoser’s 2013 article included a full reprinting of the final formatted version of Kaiser et al. (2013) from Herpetological Review. Hm, something tells me you’re not allowed to do that. You’ll be pleased to hear that I get a brief mention: I’m referred to as a “serial spammer”, as “a close friend of [Mark] O’Shea” and also as someone guilty of promoting the Kaiser et al. article on Twitter (Hoser 2013). Yup, guilty as charged, and proud of it (bar the erroneous “spammer” claim... again, does he know what that word actually means?).

**Caption:** here are the first three pages of the six-page-long table of Hoser names compiled by **Kaiser *et al*. (2013)**. See the paper for yourself: it’s open access. **Kaiser *et al*. (2013)** was – obviously – published more than ten years ago, and Hoser has since published numerous additional names. All should be ignored.

Kaiser et al. (2013) was not specifically about Hoser, since there have been (and are) several other authors who also self-publish taxonomic revisions where there’s little to no evidence of appropriate scholarship. Kaiser et al. (2013) also covered Richard Wells who, since 2000, has named over 25 new genera and numerous taxa in another self-published publication called Australian Biodiversity Record. Wells is notorious in the world of Australian herpetology for publishing two lengthy catalogues (co-authored with Ross Wellington) that made an enormous number of taxonomic recommendations for Australian reptiles and amphibians, few if any justified or supported in the way that’s normal for new systematic decisions (Wells & Wellington 1983, 1985).

An attempt by a group of over 150 Australian herpetologists to get the ICZN to suppress the names published by Wells and Wellington was unsuccessful: to repeat a point I made earlier, the ICZN more or less says that groups of researchers, not the Commission itself, have to police such problem areas themselves. The Wells and Wellington case is fairly well known and has been summarized and discussed numerous times in the literature (Grigg & Shine 1985, King & Miller 1985, Tyler 1985, 1988, Thulborn 1986, Ingram & Covacevich 1988, Hutchinson 1988, Iverson et al. 2001, Williams et al. 2006).

**Caption:** it might not surprise you to learn that Hoser behaves in a decidedly unhinged fashion on social media. He has numerous different accounts on Twitter/X (or, did last time I bothered to check) and is in the habit of reposting the same comment numerous times. He also makes a habit of smearing those who’ve criticized him in libellous fashion. For the record, I have no association with “fake science and fraud”, have no association with any “cohort found guilty of various criminal offences”, do not “control plenty of dodgy twitter accounts” and so on. These claims look an awful lot like projection.

With all of this in mind, Kaiser et al. (2013) argued that a measure of quality control is required if we’re to stop the literature being flooded with problem names appearing in unsatisfactory publications. The good news is that we already have such a system: namely, peer review. It makes perfect sense that new taxa should only be named in those published works that make it through the normal scientific channels and Kaiser et al. (2013) recommended that we introduce such a way of assessing the merits, or otherwise, of publications that include new taxonomic names. All published recommendations, as we’ve seen, are not created equal.

What about the Principle of Priority we looked at earlier? It’s well known that, in special cases, the ICZN will indeed rule against the use of certain names; the ICZN likes stability and the use of its rules, but it doesn’t like frivolity nor does it approve of new names that appear in non-technical publications. Kaiser et al. (2013) provided a long table that listed all the names that Hoser had published as of 2013, together with the recommended names that working herpetologists should use for the taxa in question.

**Caption:** the Black-necked spitting cobra *Naja nigricollis*, a species that’s partly served as the flashpoint for a call for action from the ICZN. *N. nigricollis* and its kin – the African spitting cobras – were given the subgeneric name *Afronaja* by Wallach *et al*. (2009), and ‘Spracklandus’ by Hoser in 2009. According to some interpretations, ‘Spracklandus’ was published first (Kaiser 2014), and thus should win out if the Principle of Priority were all that mattered. The point that’s now been made many times is that it *isn’t* all that should matter. It’s also not clear that ‘Spracklandus’ was published first anyway (Wüster *et al*. 2014). Image: Warren Klein, CC BY-SA 3.0 (**original here**).

Boycotting Hoser names. In short, it’s advised that herpetologists boycott Hoser’s names, and it’s hoped that the ICZN will eventually rule against their use entirely. In a 2015 appeal made to the ICZN, Rhodin et al. (2015) emphasized that none of Hoser’s works follow best practice, that his works basically do everything they can to undermine stability in herpetological taxonomy, and that numerous workers and organizations already boycott Hoser names. Ergo, the ICZN was asked to use its “plenary power to declare the Australasian Journal of Herpetology (AJH) Volumes 1-24 unavailable for nomenclatural purposes” (p. 77), and to make a statement asking that authors should adhere to the ICZN’s Code of Ethics (Rhodin et al. 2015).

The good news is that the ICZN did eventually publish an opinion on this (ICZN 2021). The bad news it that, par for the course, they declined to act, stating that “the Commission has declined to use its powers to confirm what is obvious” (ICZN 2021, p. 42). Again, this means that they bowed out of making a decision… though, that sure is some weird wording (“to confirm what is obvious”?).

**Caption:** a newspaper article from May 2013 that discusses some of the response to Hoser taxonomy, and notes some of the other charges against him. **Full size version here.**

Hoser’s other adventures. Elsewhere in his life, Hoser’s constant battles with local law enforcement – a subject I’m not interested in here – have led to his being found guilty of (and fined for) “scandalising the court”.

Then there’s the fact that he’s developed a technique of pinning down unanaesthetised venomous snakes on a table and cutting out their venom ducts (Hoser 2004). These snakes have been extensively handled (often in front of crowds) and Hoser is more than happy to let the snakes bite his daughters in order to demonstrate how safe they are. As you’ll find out if you check the wikipedia page on Hoser, he’s been convicted and fined for demonstrating with venomous snakes in close proximity to the public and had his commercial wildlife demonstrator license suspended. Hoser has also put himself forward as a candidate for local and state government on several occasions, including as recently as 2023, and has sued Australian companies for using the term ‘snake man’, which Hoser thinks he owns.

**Caption:** Hoser has appeared in the popular press a lot, for various reasons. This article from July 2012 discusses Hoser’s naming of the alleged new crocodile species ’Oopholis jackyhoserae’. It includes a few choice quotes from crocodile expert Professor Grahame Webb, including "The guy's a f*****g idiot". **You can see the full article here.**

Hard-working amateurs should be encouraged to contribute to science, not shunned or admonished, and it’s been stated on every occasion that unaffiliated researchers have frequently done, and do, sterling work. Fortunately, individuals like Hoser are rare and their research efforts are mostly recognized as the unsatisfactory, non-technical and bizarrely idiosyncratic contributions that they are. Nevertheless, the issue of taxonomic vandalism needs to be appreciated as widely as possible, and hopefully curtailed altogether.

UPDATE 1: this article was edited in March 2014 such that Hoser's names were taken out of italics and put in quote marks. This was done since otherwise it looked as if the Hoser names were proper scientific names on par with those used by others.

UPDATE 2: this article was originally published at Tetrapod Zoology ver 3 (the Scientific American years), but the current version of the article there is stripped of its images and paywalled. In order to make the article as widely accessible as possible, I’ve republished it here, with updates. An archived version of the 2013 article can be viewed here.

Hoser Taxonomy has been mentioned or discussed on a few previous occasions on Tet Zoo. See…

Scolecophidians: seriously strange serpents, May 2008
The Freshie: Australian crocodile, seemingly from the north (crocodiles part V), July 2012
Tetrapod Zoology enters its 8th year of operation, January 2013

Refs - -

Aplin, K. P. 1999. “Amateur” taxonomy in Australian herpetology – help or hindrance? Monitor 10 (2/3), 104-109.

Borrell, B. 2007. Linnaeus at 300: the big name hunters. Nature 446, 253-255.

Denzer, W. & Kaiser, H. 2023. Naming and gaming: The illicit taxonomic practice of ‘nomenclatural harvesting’ and how to avoid it. Journal of Zoology 320, 161-168.

Grigg, G. C. & Shine, R. 1985. An open letter to all herpetologists. Herpetological Review 16, 96-97.

Hoser, R. 2004. Surgical removal of venom glands in Australian elapid snakes: the creation of venomoids. The Herptile 29 (1), 36-52.

Hoser, R. 2009. A reclassification of the rattlesnakes; species formerly exclusively referred to the genera Crotalus and Sistrurus. Australasian Journal of Herpetology 6, 1-21.

Hoser, R.. 2012a. A new genus of Asian snail-eating snake (Serpentes: Pareatidae). Australasian Journal of Herpetology 12, 12-14.

Hoser, R. 2012b. The dissolution of the genus Rhadinophis Vogt, 1922 (Serpentes: Colubrinae). Australasian Journal of Herpetology 12, 16-17.

Hoser, R. 2012c. A new genus and new subgenus of snakes from the South African region (Serpentes: Colubridae). Australasian Journal of Herpetology 12, 23-25.

Hoser, R. 2012d. A review of the South American snake genera Leptodeira and Imantodes including three new genera and two new subgenera (Serpentes: Dipsadidae: Imantodini). Australasian Journal of Herpetology 12, 40-47.

Hoser, R. 2012e. A review of the North American garter snakes genus Thamnophis Fitzinger, 1843 (Serpentes: Colubridae). Australasian Journal of Herpetology 12, 48-53.

Hoser, R. 2012f. A review of the taxonomy of the European colubrid snake genera Natrix and Coronella, with the creation of three new monotypic genera (Serpentes: Colubridae). Australasian Journal of Herpetology 12, 58-62.

Hoser, R. 2012g. A review of the taxonomy of the living crocodiles including the description of three new tribes, a new genus, and two new species. Australasian Journal of Herpetology 14, 9-16.

Hoser, R.. 2012h. Robust taxonomy and nomenclature based on good science escapes harsh fact-based criticism, but remains unable to escape an attack of lies and deception. Australasian Journal of Herpetology 14, 37-64.

Hoser, R.. 2013. The science of herpetology is built on evidence, ethics, quality publications and strict compliance with the rules of nomenclature. Australasian Journal of Herpetology 18, 2-79.

Hutchinson, M. N. 1988. Comments on the proposed suppression for nomenclature of three works by R. W. Wells and C. R. Wellington. Bulletin of Zoological Nomenclature 45, 145.

Ingram, G. J. & Covacevich, J. 1988. Comments on the proposed suppression for nomenclature of three works by R. W. Wells and C. R. Wellington. Bulletin of Zoological Nomenclature 45, 52.

International Commission on Zoological Nomenclature. 2021. Opinion 2468 (Case 3601) – Spracklandus Hoser, 2009 (Reptilia, Serpentes, Elapidae) and Australasian Journal of Herpetology issues 1–24: confirmation of availability declined; Appendix A (Code of Ethics): not adopted as a formal criterion for ruling on Cases. The Bulletin of Zoological Nomenclature 78, 42-45.

Iverson, J. B., Thomson, S. A. & Georges, A. 2001. Validity of taxonomic changes for turtles proposed by Wells and Wellington. Journal of Herpetology 35, 361-368.

King, M. & Miller, J. 1985. Letter to the editor. Herpetological Review 16, 4-5.

Thulborn, T. 1986. Taxonomic tangles from Australia. Nature 321, 13-14.

Tyler, M. J. 1985. Nomenclature of the Australian herpetofauna: anarchy rules OK. Herpetological Review 16, 69.

Tyler, M. J. 1988. Comments on the proposed suppression for nomenclature of three works by R. W. Wells and C. R. Wellington. Bulletin of Zoological Nomenclature 45, 152.

Wallach, V. & Wüster, W., Broadley, D. G. 2009. In praise of subgenera: taxonomic status of cobras of the genus Naja Laurenti (Serpentes: Elapidae). Zootaxa 2236, 26-36.

Wells, R. W. & Wellington, C. R. 1983. A synopsis of the Class Reptilia in Australia. Australian Journal of Herpetology 1, 73-129.

Wells, R. W. & Wellington, C. R. 1985. A classification of the Amphibia and Reptilia of Australia. Australian Journal of Herpetology, Suppl. Ser. 1, 1-61.

Williams, D., Wüster, W. & Fry, B. G. 2006. The good, the bad and the ugly: Australian snake taxonomists and a history of the taxonomy of Australia’s venomous snakes. Toxicon 48, 919-930.

Wüster, W. & Bérnils, R. S. 2011. On the generic classification of the rattlesnakes, with special reference to the Neotropical Crotalus durissus complex (Squamata: Viperidae). Zoologia 28, 417-419.

Wüster, W., Broadley, D. G. & Wallach, V. 2014. Comments on Spracklandus Hoser, 2009 (Reptilia, Serpentes, ELAPIDAE): request for confirmation of the availability of the generic name and for the nomenclatural validation of the journal in which it was published (Case 3601; see BZN 70: 234–237). Bulletin of Zoological Nomenclature 71, 37-38.

Wüster, W., Bush, B., Keogh, J. S., O’Shea, M. & Shine, R. 2001. Taxonomic contributions in the “amateur” literature: comments on recent descriptions of new genera and species by Raymond Hoser. Litteratura Serpentium 21, 67-79, 86-91.

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The intelligence of non-bird dinosaurs is one of the most-asked questions about their biology...

**Caption:** a recent claim posits that *T. rex* and dinosaurs like it were as intelligent and cerebrally complex as anthropoid primates, and that tool use was a plausible aspect of their behaviour. In this reconstruction, two tyrannosaur individual co-operate, all while using a tree trunk as a tool, in capturing and subduing an ankylosaur. Credit: Andrew S. Minniear.

On the one hand, this interest stems from the well-known fact that most non-bird dinosaurs had proportionally small brains, certainly so relative to mammals like us. On the other, it reflects interest in what extinct animals were like when alive, and how complex, or otherwise, their behaviour might or could have been. The issue of how we determine ‘intelligence’ in any animal (living or extinct) is complex, and what to do when we only have empty endocranial cavities and rare brain endocasts to go on anyway? As a proxy for intelligence, we use approximate brain size – the primary measure being a brain to body size ratio termed encephalisation quotient or EQ (Jerison 1973) – and also what we know about brain structure and complexity.

**Caption:** a simplified cladogram depicting the relationships of the relevant reptile groups, with diagrammatic representations of their brains at far right. The animals of special interest to our study are shown in the shaded box. On the brains, the pallium (homologous to the mammal cerebral cortex) is shown in green. Note that theropods like tyrannosaurids are not tremendously different in brain form from crocodylomorphs and lepidosaurs. Image: **Caspar *et al*. (2024)**.

The general consensus on non-bird dinosaurs is that they were most likely on par in cognitive terms with turtles, lizards and crocodylians, though bird-like maniraptorans were likely more similar to birds like emus and ostriches. However, it’s widely recognised that EQ and other measures are only extremely rough guides to intelligence (e.g., Paulina-Carabajal et al. 2023).

On that note, an interesting caveat is the discovery that animal groups differ profoundly in the number of neurons they possess within a given volume of nervous tissue. In a mammal, turtle, and passerine bird of similar brain mass, for example, the total number of neurons contained within the brain varies from 58.8 million in the mammal, to 14.3 million in the turtle, to 164 million in the bird (Olkowicz et al. 2016, Kverková et al. 2022). Within birds, it’s worth noting that members of the clade Telluraves (birds of prey, rollers, songbirds and so on) have a much greater number of neurons than do other bird groups.

**Caption:** a montage of telluravian birds. This is the clade that includes rollers, hornbills, woodpeckers and kin, all predatory birds including owls, as well as parrots and songbirds. All individuals shown here were photographed in captivity except for the Song thrush *Turdus philomelos* at lower right. Images: Darren Naish.

Dinosaurs with primate-like neuron counts. What if an increased neuronal density was true of non-bird dinosaurs too? Could they possess unusually ‘neuron-dense’ brains and thus, potentially, possess much greater intelligence than conventionally argued? We were interested specifically in the number of neurons within the telencephalon, this being that part of the brain that contains the olfactory bulbs and tracts as well as the pallium, the region dealing with higher cognitive functions. It’s equivalent to the cerebral cortex in mammals.

Exactly this question was asked and examined by Brazilian neuroscientist Suzana Herculano-Houzel (2022). After working out brain size and volume from fossil endocranial chambers for 29 dinosaur and pterosaur taxa, Herculano-Houzel (2022) used these data to estimate telencephalic neuronal numbers based on established scaling rules. It’s important to note that her conclusions on dinosaurs have not emerged from out of the blue, but are built on previous studies that looked at brain size and neuron count in fossil vertebrates, and on neuron scaling relationships observed across animal groups (e.g., Herculano-Houzel 2017, 2019).

**Caption:** this nice piece of art, used to present one of Herculano-Houzel’s (2022) primary contentions about dinosaur intelligence, accompanies a YouTube video made specially to promote the research. **You can watch it here.**

The results: non-bird dinosaurs had exceptionally high telencephalic neuron counts. Big theropods like Acrocanthosaurus and Tyrannosaurus possessed over 2 billion telencephalic neurons, and thus exceeded corvids and were at the anthropoid primate-level (Herculano-Houzel 2022). She went on to suggest that these dinosaurs “had the biological capability to use and craft tools, and develop a culture, like modern birds and primates” (Herculano-Houzel 2022, p. 11), and also – based on previous work correlating neuron density with physiology and longevity – that (to use just one animal she examined as an example) Tyrannosaurus was endothermic, reached sexual maturity at age 5 and lived to between 42 and 49.

The decision to respond. Herculano-Houzel’s study received a large amount of coverage in the global media, and it could be argued that her statements have the potential to influence popular and scientific thought. A group of us – a team interested in the biology of extinct animals, in brain anatomy, and in inferring behaviours like tool use from the fossil record – decided that an academic response to these claims was warranted. It should be noted that – unbeknownst to us – neurobiologist Anton Reiner also decided to publish a response, and it appeared prior to the publication of our own study (Reiner 2023) and definitely compliments it in scope and conclusions.

**Caption:** Herculano-Houzel’s 2022 article got quite a substantial amount of media coverage, most of it treating her suggestions as fair and scientifically reasonable. Those shown here are from *The Daily Mail* (left) and *The Washington Post*.

And thus today sees the publication of a multi-authored response in The Anatomical Record. Titled ‘How smart was T. rex? Testing claims of exceptional cognition in dinosaurs and the application of neuron count estimates in palaeontological research’ (Caspar et al. 2024), the paper was led by Kai Caspar of Heinrich Heine University, Düsseldorf, Cristian Gutierrez-Ibanez of the University of Alberta, Edmonton, and Grant Hurlburt of the Royal Ontario Museum in Toronto. Our authorship includes tyrannosaurid specialists (Thomas R. Holtz and Thomas Carr) and animal tool-use experts (Jennifer Colbourne).

Dinosaur brain size, revisited. One of Herculano-Houzel’s (2022) primary assumptions was that brain size in non-bird theropods was similar, in proportional terms, to that of birds. Because non-bird theropods include animals only distantly related to birds as well as those extremely similar to archaic birds, it follows that (non-bird) theropod anatomy includes innumerable forms of ‘intermediacy’ with respect to a ‘reptilian’ condition versus an ‘avian’ one. In other words… yes, some non-bird theropods were bird-like in brain size. But those less close to birds were not.

**Caption:** how do non-bird dinosaurs fair in brain-to-body-size ratios, relative to other reptiles? This graph – plotted using brain sizes and body sizes that we consider plausible (Caspar *et al*. 2024) – shows that non-bird dinosaurs are mostly on the same approximate regression line as living non-bird reptiles ('“Non-avian Sauropsida”), though some are below it. *T. rex* (black circle) is above it, but still below the avian regression line (“Non-telluravian birds”). Some bird-like non-birds, however, are close to or on the avian regression line. Image: **Caspar *et al*. (2024)**.

We analysed relative brain size in dinosaurs using established methods that determine how well data points match phylogeny (Garland & Ives 2000), and one of our key conclusions was that non-bird theropods have a predicted brain size about typical for non-bird reptiles, and below that typical for modern birds. Other dinosaur groups (sauropodomorphs and certain ornithischians) have smaller brains than theropods, and – if we go with lower-end estimates – their brains were proportionally smaller than those of living reptiles (Caspar et al. 2024). Hadrosaurs, by the way, overlap similar-sized theropods in brain size.

**Caption:** when incorporating dinosaur brain volume data, we mostly used data previously compiled and verified by members of our team, Grant Hurlburt especially. For a few specimens, uncertainty over volume meant that we calculated new ones using graphic double integration (GDI). This diagram depicts how GDI was calculated (the example here being the endocast of the *T. rex* specimen AMNH 5029). Image: **Caspar *et al*. (2024)**.

Put more succinctly: we found that Herculano-Houzel (2022) substantially over-estimated brain size in non-bird dinosaurs, and the more conservative values found by previous authors (Hurlburt et al. 2013, Morhardt 2016) are more likely to be correct. This is primarily because she assumed that the brain occupied essentially the whole of the endocranial space, yet this is not true for the majority of dinosaur groups (Jerison 1973, Hurlburt et al. 2013, Caspar et al. 2024).

High neuron counts, revisited. What about those estimates of exceptionally higher neuron densities? Herculano-Houzel (2022) assumed that non-bird theropod brains should be similar to those of extant birds in telencephalic neuron density. She then scaled up the neuron counts from the brains of birds to conform to the much larger (in literal terms) brains of extinct dinosaurs, the results being extremely high telencephalic neuronal counts, sometimes in the low billions.

Two things emerge here. Firstly, we’ve seen that Herculano-Houzel (2022) miscalculated brain size. Wouldn’t such a miscalculation result in an artificially inflated neuron count? We re-analysed neuron count relative to estimated brain size, using scaling relationships derived from birds, non-bird reptiles, and mammals… and got very different results. For T. rex (which serves as an exemplar here because its brain anatomy is relatively well known), ‘reptilian scaling’ results in 360 million telencephalic neurons under some volume estimates, and 1.7 billion using ‘avian scaling’ under some volume estimates (Caspar et al. 2024). The estimates based on ‘reptilian scaling’ – which are the ones we should be relying on more than those based on ‘avian scaling’ – don’t put T. rex in an exceptional category at or above the corvid range.

**Caption:** this scatter plot shows predicted neuron count (in the cognitive part of the brain) as estimated by Herculano-Houzel (2022) (in red) and us (in green). Look at the values predicted for the *T. rex* specimen AMNH 5029 in particular. Reconstructed with a bird-like neuron count (green circles), our estimates do put *T. rex* at over 1 billion neurons. But when reconstructed with a non-bird-like neuron count (green squares), we find *T. rex* to be below 1 billion. Morhardt (2016) estimated neuron counts for the telencephalon alone and her predicted values, shown at lower right, also put *T. rex* at less than one billion neurons when ‘reptilian scaling’ was used. Image: **Caspar *et al*. (2024)**.

Secondly, are we sure that high telencephalic neuron counts – on par with those of anthropoid primates – are indicative of behavioural complexity, high intelligence and so on anyway? At this point, I think that many of us have heard about the claimed link between neuron count and intelligence.

What we argue in the paper is that data from across tetrapod diversity reveals that any such proposed link is on shaky ground. Especially high cerebral neuron counts – exceeding those of corvids and most primates – are present in giraffes (1.7 billion) but there’s no indication so far that this is correlated with tool use, complex culture, or anything else linked with intelligence. Dolphins including pilot whales and orcas have absurdly high neocortical neuron counts (exceeding 30 billion) – we humans have 15-20 billion – but there is, as yet, no evidence that they exceed us in whatever intelligence metric you wish to devise (Caspar et al. 2024). You might be wondering at this point if certain animals have high numbers of neurons due to the fact that… well, that they’re big. It has, indeed, generally been thought that this is the case. However, Herculano-Houzel (2011) has pushed back against this, her argument being that it’s absolute brain size and absolute neuron count that’s important. Under this logic, giraffes are secretly super-smart, I guess we just have yet to document it.

**Caption:** certain living animals have extremely high numbers of neurons in those parts of the brain associated with cognition. In cetaceans, like the orca shown here (I’ll avoid the species-level taxonomy for now…), we can say that this high neuron density is indeed correlated with cultural and behavioural complexity. But what about animals like giraffes? Turns out that they have a really high cerebral neuron count as well. Images: Darren Naish.

Being ‘reptile smart’… even ‘fish smart’… is not a bad thing. In the end, Herculano-Houzel’s (2022) suggestions and speculations, while fascinating, just don’t match the data. The idea that dinosaurs like T. rex might have been capable of cultural transmission or tool manufacture and use – that they could have been ‘monkey smart’, on par with macaques and baboons – should not be considered likely (Caspar et al. 2024).

The evidence from dinosaur brain size shows that non-bird dinosaurs were ‘reptile smart’ – on par with lizards and crocodylians – and not in the same league as big-brained birds or mammals, and not at the primate level. In fact, the especially small brains of sauropodomorphs and certain ornithischians – they’re smaller-brained than living lizards or crocs (Caspar et al. 2024) – might imply that they were cognitively on par with amphibians and fishes. Finally, there’s no reason to think that non-bird dinosaurs of any sort had exceptionally high telencephalic neuron counts exceeding 2 or 3 billion… and even if they did it’s not clear that this would demonstrate behaviour complexity or unusual intelligence (Caspar et al. 2024).

**Caption:** projects like *Prehistoric Planet* – hadrosaur-themed images from seasons 1 and 2 of that series are shown here – depict non-bird dinosaurs living socially, indulging in parental care, and interacting with other members of their social group. If you think that animals need to be similar in intelligence to telluravian birds or primates to exhibit such behaviours… you’re wrong. Images: Prehistoric Planet / Apple TV.

Speaking as someone who’s been involved in several efforts to reconstruct and portray the intelligence of non-bird dinosaurs – most recently for the Apple TV series Prehistoric Planet – I want to point out that comparing dinosaurs to animals like turtles, lizards or crocodylians, or even amphibians or (shudder) fishes, must never be considered a ‘bad thing’. As far as I see it, it makes little to no difference in terms of how we depict or imagine them: such aspects of behaviour as foraging, drinking, nest building, post-hatching parental care, sexual display, combat, herding behaviour, and the pursuing, subduing, killing and dismembering of prey all play out in the same way no matter what ‘intelligence’ you imagine these animals to be equipped with. I don’t watch amphibians or fish engaging in sexual display, intraspecific combat, parental care or predation and find myself thinking how disappointing it is that they don’t exhibit telluravian- or primate-like intelligence.

Granted, ‘reptile smart’ dinosaurs are unlikely to mourn or pine, exhibit a theory of mind, teach their youngsters, or pay attention to celestial events. But I think we can agree that suggestions that extinct dinosaurs might have behaved in such ways are more to do with science fiction than what we know about the behaviour of real animals.

**Caption:** your reminder that crocodiles, alligators, various lizards, turtles and so on can all be highly trainable, and exhibit many traits that we associate with ‘intelligence’. This image shows a trained Saltwater crocodile *Crocodylus porosus* at Australia Zoo. Image: Australia Zoo, CC BY-SA 2.0 (**original here**).

T. rex and similar animals were not on par with avian or mammalian tool-users, they were not big-brained, and they didn’t possess billions of neurons within the telencephalon of the brain. Does this make them any less interesting, or any less sophisticated or complex, than we’ve conventionally imagined them? Definitely not.

For previous Tetrapod Zoology articles on dinosaur brains, biology and connected issues, see…

The Sensitive Face of a Big Predatory Dinosaur, June 2017
Could We Domesticate (Non-Bird) Dinosaurs?, August 2018
Postcranial Palaeoneurology and the Lifestyles of Pterosaurs, August 2018
Alternative Timeline Dinosaurs, the View From 2019 (Part 3): the Dinosauroid and its Chums, December 2019
Did Dinosaurs and Pterosaurs 'Glow'? Extinct Archosaurs and the Capacity for Photoluminescent Visual Displays, March 2020
Humanoid Dinosaurs Revisited Again: Russell and Séguin’s Dinosauroid at (Nearly) 40 Years Old, August 2021
A brain for Baryonyx: using CT-scanning to examine British spinosaurid brains, February 2023

Refs - -

Caspar, K., Gutiérrez-Ibáñez,C., Ornella, B. C., Carr, T., Colbourne, J., Erb, A., Hady, G., Holtz, T. R., Naish, D., Wylie, D. R. & Hurlburt, G. R. 2024. How smart was T. rex? Testing claims of exceptional cognition in dinosaurs and the application of neuron count estimates in palaeontological research. The Anatomical Record 2024, doi 10.1002/ar.25459.

Garland, T., & Ives, A. R. 2000. Using the past to predict the present: confidence intervals for regression equations in phylogenetic comparative methods. The American Naturalist 155, 346-364.

Herculano-Houzel, S. 2011. Brains matter, bodies maybe not: The case for examining neuron numbers irrespective of body size. Annals of the new York Academy of Sciences 1225, 191-199.

Herculano-Houzel, S. 2017. Numbers of neurons as biological correlates of cognitive capability. Current Opinion in Behavioral Sciences 16, 1-7.

Herculano-Houzel, S. 2019. Longevity and sexual maturity vary across species with number of cortical neurons, and humans are no exception. Journal of Comparative Neurology 527, 1689-1705.

Herculano-Houzel, S. 2022. Theropod dinosaurs had primate-like numbers of telencephalic neurons. Journal of Comparative Neurology 531 962-974.

Hurlburt, G. R., Ridgely, R. C. & Witmer, L. M. 2013. Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain-endocast quantitative relationships in extant alligators. In Parrish, J. M., Molnar, R. E., Currie, P. J. & Koppelhus, E. B. (eds), Tyrannosaurid Paleobiology. Indiana University Press, Bloomington, pp. 134-154.

Jerison, H. J. 1973. Evolution of the Brain and Intelligence. Academic Press, New York.

Kverková, K., Marhounová, L., Polonyiová, A., Kocourek, M., Zhang, Y., Olkowicz, S., Straková, B., Pavelková, Z., Vodička, R., Frynta, D. & Němec, P. 2022. The evolution of brain neuron numbers in amniotes. Proceedings of the National Academy of Sciences 119, e2121624119.

Morhardt, A. C. 2016. Gross Anatomical Brain Region Approximation (GABRA): Assessing Brain Size, Structure, and Evolution in Extinct Archosaurs. PhD thesis, Ohio University.

Olkowicz, S., Kocourek, M., Lučan, R. K., Porteš, M., Fitch, W. T., Herculano-Houzel, S. & Němec, P. 2016. Birds have primate-like numbers of neurons in the forebrain. Proceedings of the National Academy of Sciences 113, 7255–7260.

Paulina-Carabajal, A., Bronzati, M. & Cruzado-Caballero, P. 2023. Paleoneurology of non-avian dinosaurs: an overview. In Dozo, M. T., Paulina-Carabajal, A., Macrini, T. E. & Walsh, S. (eds) Paleoneurology of Amniotes: New Directions in the Study of Fossil Endocasts.Springer International Publishing, Cham, pp. 267-332.

Reiner, A. 2023. Could theropod dinosaurs have evolved to a human level of intelligence? The Journal of Comparative Neurology doi 10.1002/cne.25458.

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Regular readers will be aware of my intermittent, occasional articles on the zoos and other animal-based visitor centres of the world, published as and when I remember to do them...

**Caption:** a montage of carnivorans kept at the New Forest Wildlife Park. Images: Darren Naish.

Over the weekend I visited the New Forest Wildlife Park, a visitor attraction not more than an hour’s drive from where I live, and in an effort to take advantage of what’s still live and active in my short-term memory, now is a good time to write a review.

**Caption:** the entrance sign as it looked in 2016 (at left) and as it looks in 2024. A great many visitor attractions were hit hard by the Covid pandemic and of course many didn’t survive at all. I have no idea how NFWP made it through that time, but I wonder if the greater visibility of the current sign is related to this time of hardship. Images: Darren Naish.

The New Forest Wildlife Park – NFWP from hereon – is located in Ashurst on the eastern side of New Forest National Park and has existed as an animal-based visitor attraction since 1981. At that time it was known as the New Forest Butterfly Farm and basically consisted of a large heated glasshouse with some adjacent outbuildings. By 1995, it had changed to New Forest Nature Quest, an innovative site devoted to British wildlife and priding itself on the design of its enclosures and on the fact that any observations of its animals would capture what it’s like to see them in the wild. A 1996 Independent article that explains Nature Quest and its ethos is here.

Place of otters and owls. A change of hands in 1997 saw the site’s wildlife collection expand as otters, owls, lynx and boar were added, and it then became the New Forest Otter, Owl and Wildlife Park. This explains the preponderance of owls and otters still there today, and in fact few places are home to as many otter species as this one, the current list on show including Giant otter Pteronura brasiliensis, Eurasian otter Lutra lutra, Smooth-coated otter Lutrogale perspicillata and Asian short-clawed otter Aonyx cinereus. North American river otter Lontra canadensis was there in the recent past.

**Caption:** the Giant otter occurs across a huge swathe of northern South America that historically extended from north-eastern Argentina and eastern Uruguay to Venezuela in the north, though it’s now extinct across part of this range and endangered overall. As is obvious from these photos, its short muzzle, tall forehead, bulging eyes and small ears give it a very unusual appearance that some people find disturbing. Images: Darren Naish.

  **Caption:** some more images from NFWP that highlight the unusual nature of the Giant otter. The flexible, muscular body, powerful limbs and dorsoventrally compressed, ‘wing-like’ tail (which explains its generic name, this being *Pteronura*) are all obvious here. NFWP bred Giant otters in 2023. Images: Darren Naish.

  **Caption:** lone European otter currently on show at NFWP. Little commented on is how remarkable the range of this species is: it extends from the European Atlantic fringes all the way east to Japan and the Korean Peninsula, and also includes southeast Asia and Sumatra, a good chunk of northern Africa, in addition to far southern India and Sri Lanka. Images: Darren Naish.

  **Caption:** the North American river otter can look pretty odd from a Eurasian perspective. It’s especially big and muscular, and animals in some populations (but not all) have a tremendously whiskery, grizzled face. NFWP had two individuals on show in 2016, both visible here. Images: Darren Naish.

What about those eponymous owls? Little owl Athene noctua, Tawny owl Strix aluco, Great grey owl S. nebulosus, Long-eared owl Asio otus, Eurasian eagle owl Bubo bubo, Snowy owl B. scandiacus and Common barn owl Tyto alba are present today, a collection clearly selected to reflect the mostly ‘native European’ theme of the park in its modern form. Other owls, some very much ‘less European’, were there as recently as 2016, including Great horned owl B. virginianus, Burrowing owl Athene cunicularia, Brown wood owl Strix leptogrammica, Ural owl S. uralensis, and Northern hawk owl Surnia ulula.

**Caption:** an owl montage depicting species kept at NFWP today and in the past. Clockwise from upper left: Great horned owl, Northern hawk-owl, Great grey owl, Snowy owl. Owls are afroavians (yes, that’s the vernacular term we should be using) with the latest studies showing that they are indeed close to hawks, eagles and Old World vultures (the accipitriforms). Images: Darren Naish.

Since 2010, the site has been known as the New Forest Wildlife Park and it very much emphasises its role in conservation, many of its animals being part of captive breeding programmes. I have actually written about the NFWP before – way back in 2007 – though on that occasion I used my visit as an excuse to write about peculiarities of the UK’s large mammal fauna in general.

The grounds. I’ve said before that you can and should judge the quality of a zoo or wildlife park by the quality, extent and landscaping of its grounds. NFWP scores well on this front, the spaces between and around enclosures mostly being forested and well vegetated, often featuring plants grown for their usefulness to birds and insects. Bird feeders*, bee hives and insect and hedgehog wintering sites are dotted about. The enclosures for many of the animals – including lynxes, wolves and deer – feel like they’re continuous with the surrounding woodland, generally contain the right amount of trees and understory, and have a natural-looking, complex topography.

We always thought it was a good thing to feed the birds. In recent years it’s been shown that certain species – finches in particular – have died off due to diseases transmitted by the sharing of plastic and metal bird feeders. Today, it may be deemed better to not feed the birds.

Caption: big hoofstock really should have access to large green spaces, and NFWP is able to provide this for its deer and bison. This photo from February 2012 (when light snow was on the ground, see foreground) shows Red deer and European bison using the field present at the park’s northern edge. Image: Darren Naish.

Caption: the grounds and surrounds of zoos, wildlife parks and so on are as important as the enclosures and their animals. You feel appropriately surrounded by trees at NFWP. The lynx enclosure, shown at right, includes a viewing window (note the bird stickers to prevent collisions) as well as an electrified fence. Images: Darren Naish.

Caption: tits photographed at NFWP feeders in 2016. Coal tit Periparus ater at left; Great tit Parus major and European blue tit Cyanistes caeruleus at right. The grounds attract a large number of locally occurring birds and other animals. Image: Darren Naish.

Some sections of the park are massively dominated by rhododendron. This creates what looks like a good, heavily vegetated habitat, but it’s one that’s actually devoid of things other than rhododendron: it’s a smothering, non-native plant that creates toxic leaf litter and has a dense root network that stops other plants from growing. On that note, a section in the middle of the park doesn’t feel as healthy and occupied by living things as it should, and this includes a swamp in the rhododendron-dominated area and two large ponds that are advertised as ‘frog ponds’. I checked and didn’t see a single amphibian (not even a tadpole), so I think that redesign and habitat management is needed for that section.

**Caption:** one of the two large ponds in the middle of the park. It looks pretty good in this shot but changes could be made to make it better suited for amphibians. Tadpoles (and spawning Common frogs *Rana temporaria*) really benefit from extensive shallow areas, so wildlife ponds should be constructed with this in mind. Image: Darren Naish.

Having mentioned amphibians, a single species is on show, namely Common toad Bufo bufo. Actually, I initially thought that the lone toad I saw was a European green toad Bufotes viridis (which is not a British native)… but no. Other than the aforementioned birds, there are no reptiles on show.

**Caption:** male Common toad at NFWP. Common toads are geographically widespread but the news about their persistence is not good, with massive population crashes being reported across Europe. One hypothesis is that the obliteration of young metamorphs on roads (and not adults so much) is a primary factor in this decline. Image: Darren Naish.

Smaller mammals. Located close to the entrance are a number of well-vegetated enclosures that house Red fox Vulpes vulpes, European badger Meles meles and both European polecat Mustela putorius and their domestic brethren the Ferret M. p. furo. Some are designed within the ethos of ‘Nature Quest’ discussed above, such that you peer through small openings and the animals don’t know you’re watching. It’s effective and I like this feature a lot. The badgers are rescue animals and the polecats are part of a successful breeding programme. Polecats were in severe decline in the UK but this has definitely been in reverse since the late 20th century.

**Caption:** I didn’t see any badgers on this trip, but here’s wooden art depicting one on the viewer’s side of their enclosure. The small viewing ports allow the animals to be observed without being aware of it. Image: Darren Naish.

While on mustelids, I should add that Pine martens Martes martes are present in a large caged enclosure near the deer and wildcats (discussed below). You get good views of them, especially when they climb about in the tunnel that allows them to leave the main part of the enclosure. The Pine marten, like the European polecat, was formerly in dire straits across the UK but it has increased in range and numbers in recent decades, and wild animals now occur in the New Forest again. NFWP has two martens and has bred four litters of kits between 2015 and 2023, all of which have been passed to other collections or released into the wild.

**Caption:** Pine marten, photographed (with rodent food object) inside wire tunnel. Martens of tradition seem not to be monophyletic, since both fishers (*Pekkania*) and the American marten *Martes americana* might be closer to wolverines or tayras (I guess this goes for the Pacific marten *M. caurina* as well). American and Pacific martens need a new generic name if this is right. Image: Darren Naish.

On rodents, rats, dormice, squirrels and others have been kept at the site in the past but I think that only Eurasian harvest mouse Micromys minutus are there today, which is good because they’re about the most visible and non-cryptic of European rodents in captivity.

**Caption:** despite their name, harvest mice aren’t necessarily animals of arable fields, since they also occur in rough grassland and reedbeds. The species familiar in Europe occurs across Asia as well, and a related one (the Indochinese harvest mouse *M. erythrotis*) occurs in China, Vietnam and presumably elsewhere in eastern Asia. Image: Darren Naish.

Hoofstock 1: deer. Moving now to hoofed mammals, three deer species are currently on show. Fallow deer Dama dama and Sika deer Cervus nippon are both contained within a large walk-through area where they’re only semi-enclosed. The collection used to include several Sika but I only saw one during my latest visit (a hind called Saffy), and it was lying down with a Fallow group. It was interesting to compare and contrast its appearance to that of the Fallow: both are cervine deer, but members of very distinct lineages.

**Caption:** a reclining deer group consisting mostly of Fallow, but with a lone Sika at lower left. The Fallow deer at NFWP include very dark individuals as well as standard white-spotted ones. Note how just about everything about the Sika is different once you start checking. Image: Darren Naish.

A small group of Red deer C. elaphus are present too, though in a separate fenced area. At this time of year (April), stags lack antlers but antler buds were prominent on Maverick, the big stag within the group. Sika and Fallow are both introduced to the UK (though now very much naturalised and clearly here to stay) whereas the Red is a true native, as is the smaller and highly abundant Western roe deer Capreolus capreolus, the only capreoline deer that survived in the UK into modern times (Moose Alces alces and Reindeer Rangifer tarandus were formerly present here, and reindeer have since been reintroduced).

**Caption:** reclining Red deer group at NFWP (clearly, I was looking at them during deer nap time). Maverick the stag is obvious at left. Red deer are big, a male generally weighing 240 kg and standing 1.2 m at the shoulder, but not in the same league as the east Asian/North American Wapiti *C. canadensis*, where males are not uncommonly over 400 kg and 1.5 m at the shoulder. Image: Darren Naish.

  **Caption:** the same Red deer antler at NFWP, photographed in 2016 (at left) and 2024. Note the good amount of rodent gnaw damage that’s now obvious, presumably caused by Grey squirrel *Sciurus carolinensis*. Images: Darren Naish.

Chinese muntjac Muntiacus reevesi used to be kept at the park but they’re not there today, despite still being on the signage. This tricks me every time: I love muntjac and always embark into the park expecting to see them. Muntjac are well known for having a major destructive impact on woodland understorey and are regarded as pests. I’m assuming that, for that reason, their keeping is now discouraged in British wildlife collections.

**Caption:** deer photographed at NFWP in the past, all female. Left to right: Chinese muntjac, Sika, Fallow. The Fallow deer at far right is giving a helpful illustration of how flexible the deer neck is: that neck is twisted and inverted such that the animal’s head is almost completely upside-down. I have a feeling that the photo was featured on SV-POW! for this reason. Images: Darren Naish.

Hoofstock 2: bovids. Moving now to bovids, NFWP also has a small flock of Mouflon Ovis gmelini. Old World wild sheep fall into three main species groups: mouflons, urials and argalis (we absolutely must avoid the species- and subspecies-level taxonomy for the purposes of this article). Mouflons are the most westerly of the three, their various populations occurring in western Asia, part of the Middle East and on some Mediterranean islands. There’s a lot to say about the diversity, taxonomy and history of wild sheep, and in fact I have written at length about these animals before. This is shorthand for saying that I’m not going to say more much about them now.

The NFWP mouflon are relatively recent additions, having joined the collection in 2014, and they’ve been successfully breeding since 2015 (6 lambs were born in 2023 alone). Their enclosure is big and spacious. I’ve learnt that most people have never heard the word ‘mouflon’ nor have any concept of what such an animal might be.

**Caption:** reclining male Mouflon. There are several ideas on what to do about Mouflon taxonomy but it all depends on how independent the population in question is from domestic sheep (*Ovis aries*). Those Mouflon that aren’t of domestic ancestry should be*Ovis gmelini* Blyth, 1841. Note that the horn tips of this male have been clipped so that they don’t keep growing and pierce his face: yes, this does happen! Image: Darren Naish.

Also on bovids, European bison or Wisent Bison bonasus are present too. European bison are nice and all but they’re virtually always smaller and less impressive than classic American ones (which I don’t think I’ve never seen, not even in captivity). In part, this is because European bison are the hybrid products of Steppe bison B. priscus and Aurochs Bos primigenius crossings, the result meaning that they’re more ‘cow x bison’ in appearance than their Pleistocene bison forebears (Barnett 2019). NFWP has three bison, all males, one of which was only born in 2020.

**Caption:** the three European bison currently on show at NFWP. The big male at right is Leszek (born at Fota Park in Ireland in 2008) and the male at the back is Heimdall (born in Barcelona Zoo in 2010). The young bull born in 2020 – Vlad – is at far left. I know all this biographical info thanks to the park’s excellent signage. Image: Darren Naish.

A constant area of discussion in UK rewilding circles concerns where and when bison will be released into the British countryside, and it won’t surprise you to know that we mostly don’t have sufficient space or ‘wildness’ to allow them to roam without encountering people. Then again, they’re not much more dangerous or reckless than cows and there are plenty of those roaming about the place (both standard modern breeds as well as archaic long-horned forms). Also part of the discussion is whether bison should be here anyway, since the evidence for their occurrence as wild-living natives in post-glacial UK is scant and controversial. Anyway, the ‘rewilding’ has started, since several were released into the wild of Kent in 2022 as part of the Wilder Blean Bison Project. One of these animals was pregnant on being released and hence ‘wild-born’ bison are in the UK already.

**Caption:** this photo is nothing whatsoever to do with the New Forest Wildlife Park, but is included here because it highlights the fact that cattle of various kinds – including the English longhorns shown here – occur in places across the British landscape. Most famously, we have the white Chillingham cattle (which I really must write about at some point). This photo was taken at Came Down (yes, real name), Dorset, in 2010. Image: Darren Naish.

The boars are back in town. At least we agree that Wild boar Sus scrofa is definitely a British native, and one that’s now back and very much present in the country due to releases both accidental and deliberate. A big, adult male boar is an impressive and formidable animal, so much so that people have long been inspired to depict these animals in sculpture and other forms of art; that’s an interesting topic that I’ve been planning to write about here ever since I first read about the Roman boar statue, dating to the 1550s, on display at the Ashmolean in Oxford.

**Caption:** the Italian boar statue today on display at the Ashmolean, where it’s been located since 1845 after being taken from a garden in Rome. Art historians and researchers have known about the statue since the 1550s but it’s clearly older than that. Its hooves and a few other sections have been repaired. Various bronze and marble statues based on this one are located elsewhere in Italy. I see this statue (and its copies) as celebrations of our appreciation of the wild boar, an amazing animal. Image: Darren Naish.

Boars living in the UK today – and those elsewhere in Europe – are a bit of a mess in genetic terms, typically (if not ubiquitously) incorporating genes from various domestic pig breeds. I don’t think that anyone has worked out, or cares all that much, what a ‘pure’ European or British wild boar might look like, but it’s notable that many of the boars we see in the UK today have black or near-black hair and a ‘dished’ (concave), short-snouted face. True, wild-type boar are browner and have a flat forehead and very long snout.

**Caption:** boars kept at NFWP in the past, a sow photographed in 2007 at left, and a boar (as in, a male boar, ha) photographed in 2009 at right. Note that the sow is extremely bulky, short-tailed and black, all features which indicate that she carries a lot more domestic pig genes than the other boars shown here. Images: Darren Naish.

  **Caption:** the big male boar – Duke – on show at NFWP in 2024. He (and a female called Duchess) arrived from Wildwood Trust in Kent in 2021. Note how different his long, tasselled tail is from at least one of the boars shown in the pictures above (an issue relevant to the discussion on the **Kleinmachnow lion case of July 2023**). Images: Darren Naish.

The key value of boar, whether they’re part domestic pig or not, is that they plough and overturn the ground while foraging, and thus act as ecosystem engineers. This role is crucial in regeneration and soil health, but it means that they quickly turn any small enclosed patch into a sodden quagmire, and such has happened with both boar enclosures at NFWP. Three boars are there at the moment, all new since 2021 and originating from Wildwood Trust in Kent. They bred last year but I didn’t see the piglets… or boarlets, if you prefer. Boar have been at NFWP park beforehand, as you can see from the photos above.

Cats of Europe. Two European cat species are on show, namely European wildcat Felis silvestris and Eurasian lynx Lynx lynx. The wildcats are in a large and recently constructed cage with numerous wooden platforms and ramps. This doesn’t strike me as the sort of enclosure they might prefer if they had a say in the matter (I would assume something heavily vegetated), but it does mean that they can be readily seen. UPDATE: I realise now that the enclosure is too new to be properly planted. It will become greener very soon.

**Caption:** the wildcat enclosure as of 2024, with a sleeping/resting cat visible as a black blob on a platform at upper right. It’s clearly a recently constructed enclosure, and note that the plants within it are new and young. Give it another few years and the space here will be substantially greener and more vegetated. Image: Darren Naish.

  **Caption:** a wildcat at NFWP, photographed in 2016 in a different enclosure from the one they’re present in now. The large amount of white on this animal’s face shows – *I think* – that it’s a different individual from the two on show today. Image: Darren Naish.

The signage has it that these are specifically Scottish wildcats, and that they’re thus F. s. grampia. However, it’s no longer thought that British wildcats are worthy of taxonomic distinction from those of mainland Europe (Kitchener 1991). I’ll say something else I often say when discussing British wildcats: the name ‘Scottish wildcat’ is misleading as it creates the impression that this animal is and was unique to the north, but it wasn’t, occurring as recently as 1800 in Wales and England too (Langley & Yalden 1977). It only became ‘Scottish’ because of persecution.

**Caption:** two wildcats are present in the enclosure as of 2024, and here’s one of them about to leap from one shelf to another. Scottish wildcats are always said to look much like domestic tabby cats but for their thicker, blunt-tipped tail, generally broader face, and general lack of white on the chin and chest. Image: Darren Naish.

On lynxes, the site was formerly home to a splendid adult male called Odin, and more recently a female called Munchkin. Today, two sisters – Tora and Inga, both originally from the Highland Wildlife Park in Scotland – occupy the big lynx enclosure and have done since 2019. You’ve probably heard how notoriously cryptic lynxes are, but I’ve seen them at NFWP every single time I’ve visited, sometimes very close to the viewing area.

**Caption:** lynxes present at NFWP in the past. The photo at left was taken in 2007, and I think it shows Odin, a male. The one at right is from 2016 and I think it shows Munchkin, a female. They’re definitely different animals if you look at the spots on the limbs. Images: Darren Naish.

  **Caption:** the two females lynxes of NFWP, photographed in 2024. Pretty frustrating that I couldn’t stop the lens from focusing on that rhododendron branch, but at least both animals are in the same shot. The Eurasian lynx is a British native and was here at least as recently as the 8th or 9th century. Dialogue continues on whether there should be an official reintroduction programme. Images: Darren Naish.

Macropods of the UK. Anyone familiar with the UK’s mammal fauna will know why wallabies are present at NFWP: it’s because the Red-necked or Bennett’s wallaby Notamacropus rufogriseus (specifically its Tasmanian subspecies N. r. rufogriseus) has been introduced and/or has escaped into several locations within the British countryside. These include Herm in the Channel Islands, Bedfordshire in eastern England, Staffordshire in the West Midlands, the Weald in south-east England, the Isle of Man, the Derbyshire Peak District in the north, and Loch Lomond in central Scotland (Lever 1977, 2009, Yalden 1988, English & Caravaggi 2020). Some of these colonies were doing well for a time but have since dwindled and disappeared, though occasional sightings continue to be reported and might mean that low numbers persist (English & Caravaggi 2020).

**Caption:** NFWP Bennett’s wallaby photographed in the act of stretching. The syndactyl 2nd and 3rd toes on the feet should be visible. The British populations mostly feed on heather, but grasses, pine, bracken shoots, oak, rowan and birch are eaten too. Image: Darren Naish.

Included in the same area of NFWP as the lynx (which are way over at one end) and wallabies (which are in the middle) are the wolves Canis lupus. These wolves are specifically Northwestern, Alaskan or Canadian timber wolf C. l. occidentalis, one of the largest and longest-limbed of wolves. They’re surprisingly big and thick-coated. NFWP currently has four, all siblings born at Colchester Zoo in 2010 and relocated to NFWP in 2011. A fifth member of the group, a female called Cedar, sadly died in 2024. This relatedness explains their similar looks, since some individuals there possess a distinctive sloping upper surface to the muzzle with a dark midline stripe.

**Caption:** one of the park’s Alaskan timber wolves. The thick coat is presumably a winter feature, but note also the substantial variation in colour across the animal’s head, body and limbs. It’s generally agreed that the wolf became extinct in the UK in about 1700 when the last Scottish individuals died or were killed, and the species was gone from England by the late 1400s. Skeletal remains show that British wolves were large and superficially like timber wolves. Image: Darren Naish.

As they’ve aged, the wolves have become paler: some were very dark just a few years ago. The park had wolves before this though, since some of the photos you see here are from 2012 yet show adults.

**Caption:** the NFWP wolf pack photographed in 2016, this meaning that it shows the same individuals as those living there today (with the exception of a female who died this year). Lots of interesting body language is visible here, including raised, tucked and horizontal tail postures, and it should be obvious who the leader is (and… we don’t use the term ‘alpha’ anymore). Image: Darren Naish.

  **Caption:** the NFWP wolf enclosure at it looked in February 2012 (that’s snow on the ground), showing a different set of wolves than the Colchester pack depicted above. Image: Darren Naish.

Final comments. And that ends my look at the New Forest Wildlife Park. As should be clear, it includes a good selection of animals that – with a few exceptions – very much feel part of the same, western European theme. Those exceptions include the Giant otters, but I so enjoy seeing them that I still regard their presence as a major draw. The landscaping and enclosures are good but I do think there’s room for involvement here and there: I really want to see those ponds improved, the rhododendron-spoilt area should be modified, and the fact that the boars have ruined their enclosures means that they should be given something better. Yes, I know that that’s easier said than done…

A real stand-out feature of the park is the quality of its signage. A massive amount of tidily-presented information is presented to the public, and there are small signs explaining the histories of the individual animals on show and what their names are.

**Caption**: a final reminder of the very special visage of the South American Giant otter, a truly remarkable mammal. Image: Darren Naish.

On other matters, the park has a decent café and shop, the parking is fine, admission is not expensive relative to similar attractions, and there’s a good mix of things for families with kids, and things for people who just want to go and look at animals. The New Forest Wildlife Park is definitely on my recommended list: go there if you can. Its website is here.

We finish with my wholly subjective scoring system…

Selection of species: 7 out of 10
Zoo nerd highlights: Giant otter, Scottish wildcat, Alaskan timber wolf
Quality of signage: 9 out of 10
Value for money: 9 out of 10
Overall worthiness: 9 out of 10

And for previous articles in my zoo reviews series, and articles relevant to some of the topics touched on here, see…

Britain: wildlife theme-park, December 2007
Tet Zoo Reviews Zoos: Edinburgh Zoo, August 2016
Tet Zoo Reviews Zoos: Colchester Zoo, October 2017
TetZoo Reviews Zoos: the Isle of Wight Zoo, February 2020
Ross Barnett’s 2019 The Missing Lynx: the Past and Future of Britain’s Lost Mammals, May 2021
TetZoo Reviews Zoos: ZSL Whipsnade Zoo, June 2021
TetZoo Reviews Zoos: Bristol Zoo, July 2021
TetZoo Reviews Zoos: Tripoli Zoo in Libya, January 2023

Refs - -

Barnett, R. 2019. The Missing Lynx: the Past and Future of Britain’s Lost Mammals. Bloomsbury Wildlife, London.

English, H. M. & Caravaggi, A. 2020. Where’s wallaby? Using public records and media reports to describe the status of red-necked wallabies in Britain. Ecology & Evolution 10, 12949-12959.

Kitchener, A. C. 1991. The Natural History of the Wild Cats. Christopher Helm, London.

Langley, P. J. W. & Yalden, D. W. 1977. The decline of the rarer carnivores in Great Britain during the nineteenth century. Mammal Review 7, 95-116.

Lever, C. 1977. The Naturalized Animals of the British Isles. Hutchinson & Co, London.

Lever, C. 2009. The Naturalized Animals of Britain and Ireland. New Holland Publishers, London.

Yalden, D. 1988. Feral wallabies in the Peak District, 1971-1985. Journal of Zoology 215, 369-374.

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It’s been said that the present is a Golden Age for cryptozoology books, cryptozoology being the ostensible study of creatures known from legend, account or anecdote but not accepted as valid by science...

But while it might be that a great many new cryptozoology-themed books have appeared in print over recent years, it doesn’t follow that their existence makes this a ‘Golden Age’… I mean, some might say exactly the opposite, since many of the books concerned are neither good nor useful. Matt Bille’s *Of Books and Beasts: A Cryptozoologist’s Library* (Bille 2022) is unusual, potentially valuable, and very much relevant to this increase in the subject’s literature since it lists, reviews and critiques it. Bille is no newcomer, but a stalwart who has been involved and contributing for several decades. He’s previously published two books devoted to the subject – *Rumors of Existence* (Bille 1995) and *Shadows of Existence* (Bille 2006) – and for some years edited and compiled the cryptozoology-themed newsletter *Exotic Zoology*.

**Caption:** Bille’s previous books on cryptozoology include *Rumors of Existence* (1995) and *Shadows of Existence* (2006). The thylacine at left was illustrated by Karen Whitman; the Okapi and Saola at right by William M. Rebsamen.

Of Books and Beasts is an annotated bibliography, covering works devoted to the topic of cryptozoology as well as others deemed relevant to the subject by the author. Here’s your reminder that an interest in cryptozoology does not necessarily denote any link between the author and a personal belief in alleged mystery creatures, since said author may well be addressing the subject’s connection to folklore, social or pop-culture phenomena, or the history of thought, discovery and exploration more generally. For the record, Bille works from a sceptical perspective and I can confirm from statements he's made elsewhere that his inclusion of, say, books on bigfoot does not reflect a personal opinion that bigfoot is an undiscovered species.

The bulk of mystery animal research (wherever the researcher sits on the ‘flesh and blood’ vs ‘sociocultural phenomenon’ spectrum) relies on the interpretation and evaluation of reports that exist in the published record. Partly for that reason, many (most?) people who write about the field are collectors of the literature, a literature that’s arcane and involves rare, long out of print items that are often hugely expensive ‘collector’s items’ today. For these reasons combined, a few efforts to compile a cryptozoological bibliography have been produced in the past. Karl Shuker and Stephen Shipp published one, in instalments, in the 1990s (Shuker & Shipp 1995a, b, 1996) (an online version is here). How complete is Bille’s review, and how does it shape up?

**Caption:** a miscellany of cryptozoological books. This subject is sufficiently arcane, yet at the same time popular, that many of its older books are much sought after and increasingly valuable. A few of the books visible here are now impossible to get at what I consider to be affordable prices. Image: Darren Naish.

  **Caption:** more cryptozoological books from my own collection, including a good number of works on sea and lake monsters. The books visible in the middle (on merfolk and dogmen) highlight the amorphous nature of the subject. Is cryptozoology only about ‘sensible’ alleged creatures, or does it also include those most often associated with myth and legend? Image: Darren Naish.

On completeness, an issue that has to be mentioned – Bille raises it in the Introduction – is that his coverage is limited to the English language. I’m no polyglot, but I’m firmly of the opinion that proper interest in a subject requires consultation of (or, at least, awareness of) works published in languages other than one’s own. This is especially obvious in cryptozoology given that its famous founder (Bernard Heuvelmans) wrote mostly in French, and that a number of his works only exist in that language. However, I appreciate that Bille had to draw the line somewhere. He also opted to exclude many books devoted to bigfoot and the Loch Ness Monster since their inclusion would “overwhelm this entire book” (p. xi). I suppose that that’s a necessity, but I admit to being disappointed.

**Caption:** the foundational impact of author and researcher Bernard Heuvelmans – whatever you think of his output – is increasingly appreciated, but less well known is that many of his works only exist in French. Certain books *about* Heuvelmans – like Barloy’s *Bernard Heuvelmans: Un Rebelle de la Science* – also only exist in French.

The book is divided into five sections (1: Cryptozoology Books, 2: Related Sciences, 3: Crypto-Fiction, 4: A Marvelous Miscellany, and Afterwords). Bille’s entries feature mini-reviews of the contents, approach and usefulness of the books he covers. My bias (speaking as a sceptical and scientifically qualified author) means that I mostly find Bille’s evaluations valuable. I dislike his use of the present tense when discussing the contents of a book: he argues in the Introduction that its use improves readability but, when reading a book, you’re seeing an author’s words from the past; they’re not speaking to you in the present (and, yes, you will appreciate the irony that I just wrote that sentence in the present tense).

To qualify my use of the word ‘mostly’ in the above paragraph, there are a few cases where Bille fails to explain a given author’s approach or argument with sufficient clarify, nor is he sufficiently critical of their scholarship. Andy McGrath’s Beasts of Britain, for example, is not a good or reliable guide to the UK’s cryptids, but a deeply idiosyncratic work where an ‘anything goes’ approach is applied to the British crypto-fauna. Think British bigfoot 1 km outside of downtown Bristol and colour-changing plesiosaurs off the coast of Devon. Meanwhile, Bille’s review of Michael Woodley’s In the Wake of Bernard Heuvelmans doesn’t mention the book’s raison d’etre: that Woodley analyzed sea monster reports anew and came up with a list of new taxonomic proposals that are even more bizarre than those proposed by Heuvelmans.

**Caption:** the ‘British bigfoot’ crowd sure are a special bunch of individuals. Yes, there are authors and researchers who seriously argue that bigfoot lives in the UK. Several interesting phenomena are at play here, but I don’t think that any of them are zoological in basis.

I hope I can be forgiven for being interested in what readers and reviewers say about my 2016 Hunting Monsters (Naish 2016). I very much appreciate Bille’s kind and fair words about the book (he regards it as an important contribution that does a good job of laying out its primary thesis), but I admit frustration in being criticized for not saying more about select topics, or for outright excluding the coverage of others. I have never worked for a single publisher that gave me free rein on wordcount: vast amounts are always excised during the editorial process or disallowed because of space and mandate. Incidentally, plans are afoot for an enlarged second edition of Hunting Monsters.

**Caption:** cryptozoology is a divisive subject. A group of people who *tend* to share a set of other ideas relating to politics, religion, human rights, and the societal role of science and scientific knowledge are highly aggressive, sometimes comically so, when it comes to whether cryptids are ‘real’ or not. This is obvious when you look at reviews of books like **my 2016 *Hunting Monsters***.

A problem one encounters in writing about cryptozoology is where the subject begins and ends. Are books on the scientific discovery of any and all animal species relevant? What about books on extinct animals that have been deemed (at one time or another) relevant to cryptozoology? What about books on folklore and indigenous knowledge that mention unusual animals, on belief systems and superstitions that involve the imagining of monsters and other weird creatures? It’s difficult. In deciding what to include and exclude, Bille has ultimately included any and all works that he regarded as consistent with or complimentary to the scientific, zoology-focused view of cryptozoology he endorses. His Section 2 therefore includes works on palaeontology, evolutionary theory, oceanography and exploration.

**Caption:** books on such matters as evolutionary history, zoological discovery, extinction and natural history are certainly relevant to cryptozoology at large, but where do you draw the line when including such books in an overview of mystery animal research?

I, personally, would have preferred it if the book didn’t include this section at all; I can appreciate that the featured works are relevant to cryptozoological research in its broadest sense, but their inclusion gives the volume a more random, potpourri flavour than the cryptozoology-focused one I would have preferred. That first section devoted to ‘pure’ cryptozoology, in fact, only occupies the first 123 pages of this 311 page book.

What about Section 3, on crypto-fiction? I’ve read a bit of crypto-fiction in my time and generally enjoyed it, and the weird netherworld that cryptozoology inhabits requires that investigators generally need to be aware of the coverage a given mystery beast has received in all genres. We’re all aware of cases where tales that originated as fiction later got retold or re-framed as if they were real, and fiction is also important in showing how a given cryptid is imagined, described or perceived in pop culture. And there’s now so much crypto-fiction out there that a compilation is welcome.

**Caption:** books classed as ‘crypto-fiction’ range from ‘serious’ thrillers, horror novels and adventure stories – some very much written for adults – to graphic novels and annotated picture books. I don’t own any of Hawthorne’s Kronos Rising books (nor do I ever intend to), but I’ve enjoyed the crypto-fiction works I’ve read. I may or may not have a pseudonymous connection to certain of these books.

Section 4 (A Marvelous Miscellany) is not as miscellaneous as I was expecting (which is a good thing), but mostly covers monster-themed books whose inclusion is totally appropriate for this work. In fact, I’m not sure why the entries here weren’t crowbarred into Section 1. The Afterwords section contains brief comments on Bille’s own books (mentioned above) and thoughts on how he selected the books he did. Of Books and Beasts ends with lists of all the books included, and their authors, but these don’t function as indices, unfortunately, since they’re not cross-referenced with page numbers. While there isn’t an index, I should add that the books included in each section are listed alphabetically.

**Caption:** another motley of cryptozoological works which can again be used to make the point that many of these books are today considered highly collectible. Works like Heuvelmans’s *On the Track* saw several editions, and dedicated researchers and collectors tend to have more than one. I own three at the moment. I don’t own a first edition of Gould’s *The Loch Ness Monster* of 1934, the first book ever published on the Loch Ness monster (copies of which are currently available at around £400). The 1969 reprint is easy to get and remains affordable.

Finally, some words on design. Of Books and Beasts is ergonomic and functional. It’s a book of text and that text is presented in no-nonsense fashion. I would have preferred a more attractive layout, with the book titles in a different font, a denser, less spaced-out text format, and some illustrations here and there (such as the covers of seminal or noteworthy books). The CG cover art (by Doug Hajicek of MonsterQuest fame) is a curious choice. I like the theme but not the execution. I get that the book’s look and layout were likely a consequence of budget, but it was released by an actual publisher and is not self-published. The lack of a publication date is also a bit irksome (though, as you’ll have guessed from the citation provided in this article, I found it online). I make these constructive criticisms in the hope that they might be useful for future editions.

Overall, Of Books and Beasts is good, and does what it was meant to. Veteran researchers will find it useful as a summary, including of works that they might not have seen or obtained themselves. Meanwhile, those who might not know the literature so well will leave with a clear idea of what’s considered valuable and worthy of consultation. The caveat is that the book is very much written for those who share Bille’s view that cryptozoology is a ‘flesh-and-blood’ field closely allied to conventional zoology. That might make it less appealing to some.

Bille, M. 2022. Of Books and Beasts: A Cryptozoologist’s Library. Hangar 1 Publishing. ISBN 9781955471275, softback, no illustrations, pp. 311. *Here at Amazon. £19.70/$23.99.*

For previous Tet Zoo articles on cryptozoology, see…

If Bigfoot Were Real, June 2016
Usborne’s All About Monsters, April 2019
Sea Monster Sightings and the ‘Plesiosaur Effect’, April 2019
Books on the Loch Ness Monster 3: The Man Who Filmed Nessie: Tim Dinsdale and the Enigma of Loch Ness, August 2019
The 1972 Loch Ness Monster Flipper Photos, August 2020
Monsters of the Deep, a Ground-Breaking Exhibition at the National Maritime Museum, Cornwall, October 2020
The Lake Dakataua ‘Migo’ Lake Monster Footage of 1994, February 2021
What Was the Montauk Monster? A Look Back to 2008, October 2021
Santa Cruz’s Duck-Billed Elephant Monster, Definitively Identified, November 2021
Legend of the Black Dog, August 2022
The Strange Case of the Minnesota Iceman, Part 1, August 2023
The Strange Case of the Minnesota Iceman, Part 2: A Review of Heuvelmans's Neanderthal, September 2023

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

Refs - -

Bille, M. 1995. Rumors of Existence: Newly Discovered, Supposedly Extinct, and Unconfirmed Inhabitants of the Animal Kingdom. Hancock House Publishers.

Bille, M. 2006. Shadows of Existence: Discoveries and Speculations in Zoology. Hancock House Publishers.

Bille, M. 2022. Of Books and Beasts: A Cryptozoologist’s Library. Hangar 1 Publishing.

Naish, D. 2016. Hunting Monsters: Cryptozoology and the Reality Behind the Myths. Arcturus, London.

Shuker, K. P. N. & Shipp, S. 1995a. A bibliography of cryptozoological and zoomythological books, part one. Animals & Men 6, 26-29.

Shuker, K. P. N. & Shipp, S. 1995b. A bibliography of cryptozoological and zoomythological books, part two. Animals & Men 7, 30-35.

Shuker, K. P. N. & Shipp, S. 1996. A bibliography of cryptozoological and zoomythological books (part three). Zoomythology. Animals & Men 8, 17-21, 30.

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Now would be a good time to publish a massive and comprehensive overview of where we’re at in our understanding of colubrid snake diversity and phylogenetic history. But that’s not happening today. Instead…

**Caption:** a garter snake in the water, on show at The Deep in Hull, UK. The Deep is a big aquarium located on the edge of the Humber, and it’s home to tropical amphibians and reptiles as well as numerous marine species. Image: Darren Naish.

…. I here provide thoughts on a colubrid I recently got to see in captivity: the Lake Zacapu garter snake Thamnophis eques insperatus, several specimens of which are currently on show at The Deep, Hull, UK. They’ve been there since 2016, have bred there, and The Deep is the only collection in the UK – and indeed the whole of Europe – to house and exhibit this snake. Here’s an article announcing this news.

**Caption:** a baby Zacapu garter snake, as featured in news articles released by The Deep. Image: (c) The Deep.

On natricines and taxonomy. Garter snakes are a group of (mostly) North American colubrids, deeply embedded within the colubrid sub-group Natricinae. Here I have to mention that some experts argue that ‘Colubridae’ of tradition should be split up into various family-level sub-groups (e.g., Vidal et al. 2007, Zaher et al. 2009, 2019), in which case Natricinae becomes Natricidae. This is my personal preference (and.. yes, I can justify that. It’s to do with the divergence times of the respective lineages and how they compare to family-level groups in other tetrapod clades). Colubridae, in this classification, becomes restricted to that group that contains the Eastern racer Coluber constrictor and its numerous close relatives, and Colubridae sensu lato then becomes Colubroidea… which is a bit confusing as this name is (in the traditional taxonomic system) already in use for the clade that contains viperids and elapids in addition to colubrids sensu lato. My sense from recent publications is that workers mostly want to stick with a tweaked version of the traditional nomenclature (e.g., Pyron et al. 2011, Figueroa et al. 2016, Deepak et al. 2021), in which case we still recognize Natricinae within Colubridae.

**Caption:** substantially simplified colubrid phylogeny, based on the results of Figueroa *et al*. (2016), and using the taxonomy where the clades concerned are regarded as colubrid ‘subfamilies', not as ‘families’. Other articles – coming soon – will discuss various of these other groups. Images: Sibynophiinae: **Thomas Brown**, CC BY 2.0 (**original here**); Natricinae: **Orchi**, CC BY-SA 3.0 (**original here**); Pseudoxenodontinae: **Umeshsrinivasan**, CC BY-SA 3.0 (**original here**); Dipsadinae: **Geoff Gallice**, CC BY 2.0 (**original here**); Grayinae: Kate Jackson, used with permission; Calamariinae: in public domain; Ahaetuliinae: **Rushenb**, CC BY-SA 4.0 (**original here**); Colubrinae: **Dawson**, CC BY-SA 2.5 (**original here**).

Natricinae is a large group (containing about 40 extant genera) mostly associated with North America and Eurasia, though it does have representation in Africa and one outlier (the Common or Mair’s keelback Tropidonophis mairii) has made it to Australasia. Natricines are often associated with wetlands and some regularly eat fishes and amphibians. Crayfish, other crustaceans, and worms are eaten by some species.

Oviparity (egg-laying) is the norm for natricines outside of North America (just two Asian species are viviparous), but all American members of the clade are viviparous. Does this have any association with the aquatic proclivities of these snakes? That was tested in a recent study; read on. From the provincial west European perspective, natricines are among the most familiar snakes of all, since the species we encounter most frequently – as a British person I’m thinking of the Barred grass snake Natrix helvetica of western and central Europe – are members of this group. And if you haven’t heard the news, the ‘Grass snake Natrix natrix’ of tradition has proven to be three species, and the one present here in the west of Europe is N. helvetica (Kindler et al. 2017).

**Caption:** beautiful photo of a Barred grass snake. This species mostly occurs in the southern half of the UK, the Netherlands, France, Switzlerland and Italy, and is the most familiar snake species to those of us who live in western Europe. It’s diverse enough across this range that five subspecies are currently recognized. Image: **Benny Trapp**, CC BY-SA 4.0 (**original here**).

A brief rant on old Tetrapod Zoology material. One more thing on colubrids in general: I’ve published a fair amount about these snakes – their diversity, natural history, phylogeny and taxonomy – in past Tet Zoo articles. But those articles – the ones hosted at Scientific American (that’s ver 3) in particular – are now completely ruined. They no longer have their images (which were key), are paywalled, and I can’t find intact versions at wayback machine. That’s very dispiriting given that my intention in covering obscure material of this sort is, and always has been, to release it into the wild. There’s therefore no point in linking to it anymore.

**Caption:** the vast majority of old Tetrapod Zoology articles are ruined. I tried to access this (at left) 2012 Tet Zoo article on colubrid snakes, but all the images have been removed… aaaand you can only look at it for a second anyway before (at right) the note about subscription pops up and makes it invisible. Yes, I have archived versions of these articles (many thanks to those who obtained them and sent them on to me) and will republish them here in time. And I already have so much other stuff to do….

Lake Zacapu’s garter snakes, a brief history. Anyway…. as you might guess from its name, the Lake Zacapu garter snake is unique to Laguna de Zacapu, an upland lake (2000 m above sea level) in Michoacán, western Mexico. This is large for a lake (1 km wide and 400 m wide) but small as the entire geographical range of a subspecies, and it’s the only known place where this snake occurs, making it of conservation concern. So far as I can tell, I don’t think we can say more than that, since not enough is known to properly evaluate its conservation status. Lake Zacapu is spring-fed and feeds numerous irrigation canals that sustain crops. It was apparently not shown on maps until the mid-1960s (Conant 2003).

**Caption:** Conant’s 2003 map showing the location of Mexican lakes and drainage areas relevant to garter snake discovery, the caveat being that this depicts their appearance prior to European invasion, in c 1500. Image: Conant (2003).

I have to mention in passing that the lake is home to an endemic ambystomatid or mole salamander, Ambystoma andersoni, named in 1984. In fact, the history of the discovery and recognition of this salamander – a neotenous species similar to the better known Axolotl A. mexicanum* – is tied to that of the region’s garter snakes since the herpetologists who made these snake and salamander discoveries were co-operating and even on the same trips together (Conant 2003). I wrote about endemic Mexican mole salamanders in 2012, but it’s another article now rendered inaccessible.

As ever, terms like ‘discovery’ and ‘recognition’ refer to the formal scientific canon, not to collective human knowledge. These animals were certainly known to the people who lived alongside them, as is usual.

Caption: Anderson’s salamander, one of several ambystomatid salamanders endemic to Mexico’s lakes. Most of these species are geologically young and among the most recently evolved of species known to science. Image: Arican, CC BY 3.0 (original here).

Also of interest is that T. e. insperatus is, like Anderson’s salamander, a recently discovered, recently described taxon, having been named in 2003 for a specimen collected as roadkill in 1961 (Conant 2003). This was found 6 km away from any suitable lake-side habitat, and this explains why Roger Conant gave it the subspecific name insperatus, meaning ‘unhoped for; unexpected’ (Conant 2003, p. 26). The specimen was an unsightly mess, and Conant didn’t figure the whole animal for that reason, instead only figuring the diagnostic scalation and pigmentation present on its body. Following this initial description, no further specimens were reported, leading to concern that the animal might not be extant.

It would appear, however, that this was due to insufficient knowledge of the location, since visits to the lake by garter snake specialist Steven Bol found the snakes relatively easy to find (Bol 2012). Bol reported that the Zacapu garter snake is variable in pigmentation, some individuals being light and prominently marked with yellow, others being darker and lacking yellow markings entirely.

**Caption:** I haven’t seen the published version of Bol’s 2012 article announcing the rediscovery of the Zacapu garter snake, but a well-illustrated online version **is here**; above is a screengrab. Image: (c) Steven Bol.

The Zacapu population is currently regarded as a subspecies of the Mexican garter snake, a large and heavy built garter snake (second in size only to the Giant garter snake T. gigas of California) that can reach 1.3 m in total length. I mostly think of garter snakes as slender animals less than 40 cm long, so the fact that there are hefty, thick-bodied species over a metre in length is surprising. Within natricines, garter snakes are especially close to the Nerodia water snakes, a wholly North American group (Figueroa et al. 2016).

Phylogenetic studies indicate that natricines originated in Asia around 40 million years ago (during the Late Eocene), and among the several dispersals that occurred within the history of the group was an eastwards one that involved the colonizing of North and Central America (Deepak et al. 2021). It appears that ancestral natricines were mostly terrestrial but there are indications that anuran-based diets and ‘aquatic generalist’ diets were important in the group’s early history. Vermivory – which is widespread in garter snakes and may be the main way they make a living in suburban environments – appears to be a specialized habit. Viviparity evolved independently at least three times, but not in correlation with aquatic habits (Deepak et al. 2021). Yes, viviparity is certainly useful for aquatic snakes (and other reptiles too), but you don’t need to be aquatic for viviparity to evolve, as is demonstrated by the numerous times it occurred in thoroughly terrestrial squamates.

**Caption:** a garter snake montage, depicting some of the variation in size and pigmentation present across the 35 or so species. Clockwise from left: Eastern garter snake *T. sirtalis sirtalis*, Coast garter snake *T. elegans terrestris*; Mexican garter snake *T. eques*. Images: **Wilson44691**, public domain (**original here**); **Steve Jurvetson**, CC BY 2.0 (**original here**); **NS777**, CC BY-SA 4.0 (**original here**).

A diversion on scale microornamentation. While looking at the captive Zacapu garter snake shown here, I noticed that the meniscus was bending around the snake to such a degree that the snake’s surface looked hydrophobic. Is this the case? Would a snake (or any squamate or reptile) benefit from having a hydrophobic surface? Animals known to be hydrophobic – like aquatic insects – are like this because they ‘want’ to remain on the water surface, not be immersed in or swim through it. However, a hydrophobic surface is also helpful because it prevents the attachment of air bubbles that otherwise interfere with buoyancy (e.g., Ishii et al. 2016).

**Caption:** the bending of the meniscus about the scales of a garter snake. This might not mean anything. It might just be normal for an object part-submerged in water. Buuut… does it also mean that this snake has water-repellent scales? Image: Darren Naish.

I’m not sure that anyone imagines squamate scales to simply be flat sheets of featureless keratin, but they’re anything but once you learn about them at the microscopic level. Indeed, anatomists have known since at least the 1870s that squamate scales possess microornamentation or nanotexture (take your pick) that perhaps has a function in epidermal integrity, in the frictional properties of the skin, or in anti-fouling and self-cleaning. Does this microornamentation also provide water-repellent properties and thus make life easier for species that swim? Exploring this topic properly would require an article all its own and I might have to come back to it. Some particularly fascinating research concerns the hydrophobic properties of rattlesnake scales (where the scales are used to harvest rainwater and even sleet and snow; the snakes then drink the collected water from their own skin; Phadnis et al. 2019) and those of Gaboon vipers Bitis rhinoceros (Spinner et al. 2014). In the latter, nanoridges and other nanostructures make the scales superhydrophobic, this being most prominent on the black scales. These have a velvety texture.

**Caption:** rainwater droplets clinging to rattlesnake scales (at left), with a close-up image of a scale at right. As discussed by **Phadnis *et al*. (2019)**, the scale microornamentation appears adapted for this role.

  **Caption:** at left, *Bitis* has an extremely complex scale surface microornamentation, which is hierarchical on some scales (like the black ones shown in B): there are leaf-shaped protrusions marked with micro-ridges. As the diagram shows, microornamentation varies according to which region of the body the scales come from. At right, we see how the black, velvety scales have superb water-shedding qualities. Images: **Spinner *et al*. (2014)**.

More recent work has also shown that lipid coatings on snake scales provide lubrication and wear protection (Baio et al. 2015). It also appears that these lipids make the scales hydrophobic, and work done on the microornamentation of natricine scales – including that of garter snakes – indicates that these snakes have a surface pore pattern where cup-like chambers may “assist in the retention of the exudate on the surface of the scale. Such a coating of fatty material on the surface might form a water barrier on the skin especially advantageous to aquatic taxa” (Chiasson & Lowe 1989, p. 109). So… are garter snake scales water-repellent and, if they are, is this an adaptation for amphibious life? Yes, it would appear so.

Finally… yeah, I’m so angry about the situation with the old colubrid-themed articles from Tet Zoo ver 3 that I’ll be republishing them here, with updates.

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

For previous Tet Zoo articles on squamates, see… [UPDATE: as per the comments above, it now seems that at least some of these articles have been made inaccessible by Sci Am]

The enormous liolaemine radiation: paradoxical herbivory, viviparity, evolutionary cul-de-sacs and the impending mass extinction, June 2013
It’s high time you were told about Psammodromus, July 2013
Tale of the Takydromus, September 2013
Racerunner Lizards of the World Unite, October 2014
What's With All These New Chameleon Names?, Part 1, February 2016
By the Horns of Trioceros, the Casque of Calumma, the Brood of Bradypodion--Chameleons, Part 2, February 2016
Palleon, Archaius, Kinyongia, Nadzikambia--The Last Chameleons, Part 3, March 2016
The Tropidurine Treerunners, December 2017
The Tet Zoo Guide to Mastigures, August 2018
The Remarkable Basilisks, May 2023
Do Lizards Really Have ‘Mite Pockets’?, March 2024

Refs - -

Baio, J. E., Spinner, M., Jaye, C., Fischer, D. A., Gorb, S. N. & Weidner, T. 2015. Evidence of a molecular boundary lubricant at snakeskin surfaces. The Journal of the Royal Society Interface 12, 20150817.

Bol, S. 2012. The rediscovery of Thamnophis eques insperatus (the Zacapu Mexican Garter Snake) in Zacapu, Michoácan, Mexico. The Garter Snake 17 (1).

Chiasson, R. B. & Lowe, C. H. 1989. Ultrastructural scale patterns in Nerodia and Thamnophis. Journal of Herpetolology 23, 109-118.

Conant, R. 2003. Observations on Garter Snakes of the Thamnophis eques complex in the Lakes of Mexico’s Transvolcanic Belt, with descriptions of new taxa. American Museum Novitates 3406: 1-64.

Deepak, V., Cooper, N., Poyarkov, N. A., Kraus, F., Burin, G., Das, A., Narayanan, S., Streicher, J. W., Smith, S.-J. & Gower, D. J. 2021. Multilocus phylogeny, natural history traits and classification of natricine snakes (Serpentes: Natricinae). Zoological Journal of the Linnean Society 195, 279-298.

Ishii, D., Yamasaki, H., Uozumi, R. & Hirose, E. 2016. Does the kinorhynch have a hydrophobic body surface? Measurement of the wettability of a meiobenthic metazoan. Royal Society Open Science 3, 160512.

Kindler, C., Chèvre, M., Ursenbacher, S., Böhme, W., Hille, A., Jablonski, D., Vamberger, M. & Fritz, U. 2017. Hybridization patterns in two contact zones of grass snakes reveal a new Central European snake species. Scientific Reports 7, 7378.

Phadnis, A., Manning, K. C., Schuett, G. W. & Rykaczewski, K. 2019. Role of scale wettability on rain-harvesting behavior in a desert-dwelling rattlesnake. ACS Omega 4, 21141-21147.

Spinner, M., Gorb, S. N., Balmert, A., Bleckmann, H. & Westhoff, G. 2014. Non-contaminating camouflage: multifunctional skin microornamentation in the West African Gaboon viper (Bitis rhinoceros). PLoS One 9, e91087.

Vidal, N., Delmas, A.-S., David, P. Cruaud, C., Couloux, A. & Hedges, S. B. 2007. The phylogeny and classiﬁcation of caenophidian snakes inferred from seven nuclear protein-coding genes. C. R. Biologies 330, 182-187.

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Over 200 modern lizard species possess slits, apertures and folds in the skin that lead to epidermal structures termed mite pockets…

**Caption:** a mite pocket montage, showing a mite pocket in a scrub lizard (at left), the mite-filled nuchal pockets of a tropidurid (at upper right), and (at lower right) all the locations where pockets can be present. Images: McCoy *et al*. (2012); Carvalho *et al*. (2018); Arnold (1986).

Variously located on the sides of the neck, armpit region, and in front of and behind the top of the thigh, these are often inhabited by parasitic mites, specifically chigger mites of the families Trombiculidae and Leeuwenhoekiidae. Because the pockets seemingly don’t have a function other than their use by mites, it was proposed back in the 1980s that lizards and mites have co-evolved such that the pockets are specialized mite aggregation sites.

And because the lizards with these pockets belong to far-flung branches of the squamate family tree (they’re present in various geckos, phrynosomatids, tropidurids, oplurids, chameleons, lacertids and skinks), the pockets must have evolved at least seven times.

**Caption:** a highly simplified squamate phylogeny (based on recent molecular results) serving to emphasize how pockets (P) have evolved several times in distantly related groups. Even within Iguania (represented here as a single lineage), pockets are present in more than four groups. Image: Darren Naish, produced for my **in-prep textbook**.

Supporting this co-evolutionary idea is data indicating that the pockets are most prevalent, and largest, in lizards that occupy the same sort of humid, terrestrial habitats as those favoured by mites, and that lizards in arid or arboreal environments generally don’t have them (Arnold 1986, Curtis & Baird 2008, Reed 2014). Also worth noting is that males tend to have higher mite loads than females. I should also add at this point that the mites we’re talking about here are definitely parasitic: none are phoretic mites (those that use larger animals as transport to get from A to B and don’t exact much cost on said larger animal). Scale mites and ticks also use the pockets in some species.

**Caption:** at left, mites clustered in the right nuchal pocket of a Florida scrub lizard *Sceloporus woodi*. At right: perhaps casting doubt on the idea that the pockets have co-evolved with mites is the fact that mites cling opportunistically to the skin folds and flaps present in lizards without pockets, like this *Agama cristata* from Guinea. Images: McCoy *et al*. (2012); Wagner *et al.* (2009).

To scientists familiar with lizards (and the literature devoted to them), the existence and claimed function of mite pockets is well known, and indeed the structures have received high-profile coverage in the scientific literature (Benton 1987)… at least, as ‘high profile’ as you can expect for something like this. A memorable back-and-forth in the technical literature also brought the topic to wider attention than it might otherwise have had (Arnold 1986, 1993, Bauer et al. 1990, 1993). And, today, I get the impression from articles online and comments in the literature that the hypothesis that these structures are (1) adaptive and (2) evolved in-step with mite parasitism appears essentially accepted.

**Caption:** mite pocket distribution as portrayed on an archetypal lizard. Going from head to tail, they are: n = nuchal; a = axillary; pa = post-axillary; i = inguinal; pf = postfemoral. Image: Arnold (1986).

Prior to the appearance of the ‘mite pocket’ idea, these structures were usually given descriptive names related to their anatomical position, such that they were axillary and postfemoral pockets, axillary and inguinal pits, or postfemoral dermal pockets (Arnold 1986, pp. 1-2). The term ‘mite pockets’ was first used by Loveridge (1925) who (after observing them in Asian bent-toed geckos) also suggested that the term acarodomata (singular: acarodomatum) might be used for them*. That technical name wasn’t original to him but came from the botanical literature.

Because, however, calling them ‘mite pockets’ implies that this is certainly what they are, it seems more appropriate to me to be less committal, and for that reason I’m just going to call them ‘pockets’ from hereon.

The spellings acarodomatia and acarodomatium have also been used (Bertrand & Modrý 2004). Mite-housing structures in hymenopteran insects (yup, they too are a thing) are termed acarinaria, and you might argue that this would be a better term than one devised for plants.

Caption: other organisms have mite-housing structures that have been seen as precedents to the mite pockets of lizards, including plants and insects. These images show the acarinaria of the small Eurasian wasp Allodynerus delphinalis. The wasps transport the mites to their nests where they help protect Allodynerus from attacks by parasitoids. This is therefore a mutualistic relationship. Image: Okabe & Makino (2008).

In memory of Dr Nick Arnold. The seminal work on these structures was published by herpetologist and lizard specialist Nick Arnold in 1986, and here I must make a brief diversion, since Nick died last year and I thought very highly of him, and still do. Dr E. N. Arnold, formally Edwin Nicholas Arnold and based at London’s Natural History Museum where he was Curator of Herpetology, was a brilliant scientist, and he leaves behind an impressive legacy of interesting and compelling studies of lacertid and agamid lizards, recently extinct island-dwelling tortoises and much else. An aspect of Nick’s writing that often stands out is that he was especially good at interpreting or framing data within historical or phylogenetic hypotheses.

**Caption:** Nick Arnold (1940-2023) was honoured with an obituary in *The Times* (**original is here**, but paywalled). Few of us will be considered worthy of being memorialized in this way. Image: (c) *The Times*.

Many naturalists unfamiliar with the technical literature were also exposed to Nick’s writings, since (with J. A. Burton) he authored the highly successful and very good Collins book A Field Guide to the Reptiles and Amphibians of Britain and Europe, first published in 1978 and illustrated by Denys Ovenden. This was the standard go-to guide to the European herpetofauna for over three decades, and in part still is. Finally: as is true of all the best scientists, Nick was also a phenomenally nice person and was appropriately generous with time and resources when opportunity allowed.

**Caption:** Nick Arnold’s Collins field guide to the reptiles and amphibians of Europe saw several editions. I don’t have all of them… but I’m working on it. Burton is not credited as a coauthor on the later editions; I don’t know why.

Anyway… back to the pockets. After analysing the anatomy of pockets in geckos, phrynosomatids, chameleons and lacertids, Arnold (1986) concluded that the skin within the pockets was essentially adapted for mite use: the pockets aren’t just epidermal invaginations, they’re invaginations where the skin is distinctly thickened, and where the pocket interior has a modified skin surface, such that scalation is reduced or even absent, and blood supply is rich (Arnold 1986, p. 19). In some pocket-bearing lizards, there are indications that lymphoid cells are densely clustered where mites occur (suggesting a sort of enhanced local immune response to the mites’ mouthparts), but Arnold (1986, p. 17) expressed uncertainty as to whether this was adaptive on the part of the lizards or a consequence of mite activity.

**Caption:** Arnold (1986) featured several histological investigations of pocket anatomy and what this meant for mite presence. At left, this section through a nuchal pocket on a fence lizard shows how the collagen (c’) within the pocket is thinner than that elsewhere across the body, and how the epidermis around the mite feeding site (the stylostome, marked s) consisted mostly of lymphoid cells (l). At right, this section through a mite and the post-axillary pocket of a *Pristurus* gecko from Oman shows how the skin in the vicinity of the stylostome (s) is densely packed with lymphoid cells (l). Images: Arnold (1986).

The main take home from Arnold (1986) is that the pockets “reduce the deleterious effects” (p. 18) of parasitic mites by providing mite-friendly areas that cause the mites to cluster in select areas rather than spread across the body, this minimizing tissue damage and irritation more broadly. Arnold’s argument was termed the ‘damage limitation’ or ‘damage-amelioration’ hypothesis by later authors (Bauer et al. 1990, Reed 2014).

**Caption:** mites clustered in an axillary pocket of a chameleon (*Rieppeleon brevicaudatus*, though included in *Rhampholeon* when Arnold was writing). The mites are obviously distinct from the “sculpted surfaces of the scales lining the pockets” and are actually feeding from “the solf folded interstitial skin between the scales” (Arnold 1986, p. 8).

Some quick and dirty speculations. Incidentally, Arnold’s hypothesis that lizards have repeatedly evolved special mite-nurturing pockets raises all sorts of speculative possibilities for lizard-parasite evolution.

If these really are ‘mite pockets’, should we – for example – expect the evolution of dedicated pocket-dwelling parasites of lizards? Think sessile, limbless mites or even pocket-dwelling parasites belonging to other groups that have moved in to take advantage. What about the development of ‘pocket-raiding’ habits by conspecifics or even by other animals? Might it even be that the supposed mite-nurturing anatomy of the pockets could be co-opted to something else, like the feeding of conspecifics? If the pockets really provide enhanced blood flow and have specialized internal skin that’s co-evolved with the feeding on body fluids… well, the parallels with the evolution of lactation in mammals are unavoidable.

**Caption:** I’m old-fashioned and still rely extensively on physical printed literature. This isn’t necessarily because I want to, but because so much of the material I need to access is *still* not easily findable in digital form. Here are physical reprints of various of the sacred texts discussed in this article. Image: Darren Naish.

But are the pockets really ‘mite pockets’? You should have noticed so far a note of scepticism about Arnold’s ‘damage limitation hypothesis’. A follow-up to Arnold (1986) was published by Aaron Bauer, Anthony Russell and Norman Dollahon in 1990 following their analysis of hindlimb pockets in the New Caledonian gecko Rhacodactylus (Bauer et al. 1990).

**Caption:** a left, Gargoyle gecko *Rhacodactylus auriculatus* in captivity, the species studied by Bauer *et al*. (1990) in their test of Arnold’s hypothesis. At right, the ventral surface of the hindlimb of *R. auriculatus*, the popliteal fold and its entrance marked with an arrow. Images: **Generish**, CC BY-SA 4.0 (**original here**); Bauer *et al*. (1990).

And they concluded that the pockets weren’t anything to do with co-evolution and parasites, but a byproduct of the way lizard skin grows, folds or has evolved, the pockets – in cases – being related to use of the skin in crypsis and/or locomotion (Bauer et al. 1990). The mites, according to this view, are therefore taking advantage of sites of vulnerability on the lizards, and indeed damage caused to lizards by mites carries the same costs whether it occurs within the pockets or elsewhere on the body; it isn’t ‘limited’ as it should be according to Arnold’s hypothesis (Bauer et al. 1990).

An exchange between both sets of authors followed (Arnold 1993, Bauer et al. 1993), the ultimate conclusion (as seen by an outsider) being that the pockets are “neutral or mildly maladaptive “phylogenetic baggage”” (Bauer et al. 1993, p. 868). This was always my take on the phenomenon all along: that the pockets originated for reasons unrelated to co-evolution with a parasite, and that demonstrating lizard/mite co-evolution was a big ask, and a task that hadn’t been fulfilled. The similarity that the pockets have with various prominently marked or pigmented skin folds and flaps suggests, I think, that at least some of them are elaborations of structures that have roles in signalling or crypsis, and thus any use of them by parasites is incidental.

**Caption:** this is Flame the Central bearded dragon *Pogona vitticeps*, and like many lizards she has skin folds on the neck and shoulder region that are associated with distinctive ornamentation and markings. This suggests that the evolution of pockets (in the nuchal region at least) is an exaptation, and that the structures started out as being associated with signalling and display. Image: Darren Naish.

Mite pockets in the 21st century. 1993 is a long time along now. As I said at the start of this article, the general impression you might get from the literature today is that Arnold’s hypothesis has been accepted. I don’t know if this is because researchers really have been convinced by his proposal, or whether they’re aware of his initial argument (Arnold 1986, Benton 1987) but not the follow-up. Publications on lizards that have reason to refer to the pockets tend to just term them ‘mite pockets’ without equivocation (e.g., Rocha et al. 2008, Mockett 2017, Carvalho et al. 2018), though not all authors have done this (Fajfer 2012).

**Caption:** male specimen of the Bolivian-Brazilian tropidurid *Tropidurus chromatops* showing the deep and complex nuchal pockets of this species, in this case full of mites. Note how the pockets are also associated with distinctive pigmentation. Image: Carvalho *et al*. (2018).

Has any actual work been done on the pockets since the Arnold vs Bauer exchange of the 80s and 90s? Yes, yes it has, though let’s start with an observational report more than targeted experimental work. Bertrand & Modrý (2004) reported the discovery of numerous scale mites packed under a skin fold on the neck of the endemic Kenyan Elmenteita rock agama Agama caudospinosa. This lizard lacks proper pockets, and yet here was a case where clustering scale mites were causing a fold to become an incipient pocket.

Though the authors didn’t say it, this discovery potentially weakens the hypothesis that pockets evolved as mite aggregation sites: it looks instead like mite clustering is opportunistic, and that any suitable flap or skin fold is exploited.

**Caption:** at left, Elmenteita rock agama *Agama caudospinosa* in the field. At right, a specimen reported by Bertrand & Modrý (2004) in which numerous scale mites were packed under one of the nuchal skin folds. Images: rjq, CC BY-NC 4.0 DEED (**original here**); Bertrand & Modrý (2004).

Now for the big 21st century news on mite pockets: yes it’s true, Jay Clark Reed completed an entire 2014 doctoral thesis at the University of Michigan on the pockets and their possible role and evolution, albeit only in phrynosomatids (fence lizards and kin). So far as I can tell, a technical publication reporting the results of this work has yet to appear, so I feel a bit guilty discussing its results and hope I’m ok to do this. Reed’s (2014) primary conclusion was that the pockets probably did co-evolve with parasitic mites, since the included phylogenetic analyses showed that phrynosomatids evolved larger and more complex pockets on moving into mite-rich habitats. Meanwhile, pockets were reduced and lost in groups that – over the course of their evolutionary history – moved away from mite-heavy areas (Reed 2014). Yeah yeah, correlation isn’t causation, but this is at least suggestive, and consistent with Arnold’s hypothesis.

Reed’s mate selection hypothesis. Recall the mention early on that males carry higher tick loads than females? It’s well known that animals vetting others as potential mates partly base their evaluations on parasite load. Reed (2014) hypothesized that the pockets might have enlarged such that the deleterious presence of mites could be concealed, and thus make a parasite-loaded male more appealing to a female. This would mean that the pockets evolved as dishonest signals…. the theoretical problem here being that cheating about your parasite load (if that load is deleterious and hence worth concealing) would be bad in the long run, since it would result in offspring that are less fit than those of your competitors: those offspring would then lose out via natural selection. Err, unless everyone is lying about their parasite load I guess.

**Caption:** Yarrow’s spiny lizard *Sceloporus jarrovii*, the species used in Reed’s studies on mate selection and mite occurrence. *S. jarrovii* is a phrynosomatid iguania endemic to Mexico and the southern USA. Image: **Greg Schechter**, CC BY 2.0 (**original here**).

Anyway, via a series of ingenious tests (some involving the addition of painted-on pretend mites), it was shown that females did not make their mate selections on the basis of visible parasite load (Reed 2014). Ironically, females in some tests preferred males with the highest mite loads, but this is likely because big males – who are the most desirable – have the highest loads simply as a consequence of size. The idea that mate selection might have driven pocket evolution does not, therefore, seem supported (Reed 2014).

And that basically brings us to where we are. While further research is needed, mite pockets might really be mite pockets after all, and I find it exciting that a number of high-level evolutionary phenomena – exaptation, convergence, co-evolution, sexual selection and dishonest signalling among them – have all been suggested to have a bearing on this surprisingly odd and complex bit of anatomy. In addition, the fact that lizards have evolved such structures in the first place activates the speculative part of my brain…

**Caption:** certain species within the world of the Squamozoic have evolved elaborate pockets that have a very specialized function. In the giant subterranean amphisbaenian *Graboidus*, deep slits along the posterior part of the body lead to partitioned internal cavities inhabited by juveniles. Here, they feed on nutritious fluid expressed from the walls of the cavities. Image: Darren Naish.

I’ve been thinking about writing an article on mite pockets for years, but it was the sharing of a picture and its associated comments by digital artist and creature designer Kate Pfeilshiefter that inspired me to write this entire overly long article. Thank you, Kate.

For previous Tet Zoo articles on squamates that have some connection to this one, see…

The enormous liolaemine radiation: paradoxical herbivory, viviparity, evolutionary cul-de-sacs and the impending mass extinction, June 2013
It’s high time you were told about Psammodromus, July 2013
Tale of the Takydromus, September 2013
Racerunner Lizards of the World Unite, October 2014
What's With All These New Chameleon Names?, Part 1, February 2016
By the Horns of Trioceros, the Casque of Calumma, the Brood of Bradypodion--Chameleons, Part 2, February 2016
Palleon, Archaius, Kinyongia, Nadzikambia--The Last Chameleons, Part 3, March 2016

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

Refs - -

Arnold, E. N. 1986. Mite pockets of lizards, a possible means of reducing damage by ectoparasites. Biological Journal of the Linnean Society 29, 1-21.

Arnold, E. N. 1993. Comment – function of the mite pockets of lizards: an assessment of a recent attempted test. Canadian Journal of Zoology 71, 862-864

Bauer, A. M., Russell, A. P. & Dollahon, N. R. 1990. Skin folds in the gekkonid lizard genus Rhacodactylus: a natural test of the damage limitation hypothesis of mite pocket function. Canadian Journal of Zoology 68, 1196-1201.

Bauer, A. M., Russell, A. P. & Dollahon, N. R. 1993. Function of the mite pockets of lizards: a reply to E.N. Arnold. Canadian Journal of Zoology 71, 865-868.

Benton, M. J. 1987. The mite pockets of lizards. Nature 325, 391-392.

Bertrand, M. & Modrý, D. 2004. The role of mite pocket-like structures on Agama caudospinosa (Agamidae) infested by Pterygosoma livingstonei sp. n. (Acari: Prostigmata: Pterygosomatidae). Folia Parasitologica 51, 61-66.

Carvalho, A. L. G., Rivas, L. R., Céspedes, R. & Rodrigues, M. T. 2018. A new collared lizard (Tropidurus: Tropiduridae) endemic to the Western Bolivian Andes and its implications for seasonally dry tropical forests. American Museum Novitates 3896, 1-54.

Curtis J. L. & Baird, T. A. 2008. Within-population variation in free-living adult and ectoparasitic larval trombiculid mites on collared lizards. Herpetologica 64, 189-199

Fajfer, M. 2012. Acari (Chelicerata) – parasites of reptiles. Acarina 20, 108-129.

Loveridge, A. 1925. A mite pocket in the gecko, Gymnodactylus lawderanus Stoliczka. Proceedings of the Zoological Society of London 1925, 1431.

McCoy, E. D., Styga, J. M., Rizkalla, C. E. & Mushinsky, H. R. 2012. Time since fire affects ectoparasite prevalence on lizards in the Florida scrub ecosystem. Fire Ecology 8, 32-40.

Mockett, S. 2017. A review of the parasitic mites of New Zealand skinks and geckos with new host records. New Zealand Journal of Zoology 44, 39-48.

Okabe, K. & Makino, S. 2008. Parasitic mites as part-time bodyguards of a host wasp. Proceedings of the Royal Society B 275, 2293-2297.

Reed, J. C. 2014. Analysis of the Function and Evolution of Mite Pockets in Lizards. University of Michigan Dissertation.

Rocha, C. F. D., Cunha-Barros, M., Menezes, V. A., Fontes, A. F., Vrcibradic, D. & Van Sluys, M. 2008. Patterns of infestation by the trombiculid mite Eutrombicula alfreddugesi in four sympatric lizard species (genus Tropidurus) in northeastern Brazil. Parasite 15, 131-136.

Wagner, P., Ineich, I., Leaché, A. D., Wilms, T. M., Trape, S., Böhme, W. & Schmitz, A. 2009. Studies on African Agama VI. Taxonomic status of the West African Agama (Sauria: Agamidae) with prominent tail crests: Agama boulengeri Lataste 1886, Agama insularis Chabanaud, 1918 and Agama cristata Mocquard, 1905. Bonner Zoologische Beiträge 56, 239-253.

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Among the proudest of my achievements is the publication of Dinosaurs: How They Lived and Evolved, DHTLE for short, co-authored with Professor Paul Barrett and published by the Natural History Museum, London. I think it’s fair to say that it’s the flagship ‘dinosaur book’ of the museum. It’s also one of only a handful of dinosaur-themed books written at ‘adult level’. “Finally, a modern, intelligent, trade book on dinosaurs for thoughtful readers”, to quote a reviewer at Quarterly Journal of Biology.

**Caption:** first edition at left, second edition at right, third edition in the middle. Image: Darren Naish.

The book went to second edition in late 2018; I wrote about its publication here. It got a new cover (this time featuring a fuzzy Tianyulong by Bob Nicholls), but the number of changes made to the text was actually low, predominantly because Paul and I were both chronically overworked at the time and unable to do a thorough job of updating it.

The book has gone to third edition (Naish & Barrett 2023) because, simply put, it continues to sell well. But rather than simply release the book with a few small corrections, we opted to use this as an excuse to correct errors present in the second edition, and to properly change numerous sections of text in view of recent discoveries and proposals. In the rest of this article, I want to note the changes we made: metacommentary is fun (and where else can I do it but here), but to get the full picture you will, of course, want to check the book itself. Buy it here!

**Caption:** I think often about the ‘ancestors’ of *Dinosaurs: How They Lived and Evolved*, the most important of which are Tim Gardom and Angela Milner’s *The Natural History Museum Book of Dinosaurs* (first published 1993), and Alan Charig’s *A New Look at the Dinosaurs* (two editions of which are shown here, first published 1979). Both works have their strengths and weaknesses. Image: Darren Naish.

Without further ado…

Dinosaur origins and dinosaur cousins. Our text and diagrams reflect the recent discovery that lagerpetids – a group of small, long-legged Triassic archosaurs with vaguely theropod-like proportions – are closer to pterosaurs than to dinosaurs, and we’ve also accounted for new discoveries of silesaurids, a near-dinosaur group that have proved diverse in body size and feeding biology. I never did like the John Sibbick reconstruction of Marasuchus included in the previous editions, and this has been replaced by a new one by Gabriel Ugueto.

**Caption:** massively simplified depiction of the Ornithoscelida phylogenetic hypothesis, as first published by Baron *et al*. (2017). The image was used in the Tet Zoo article **here**. Other phylogenetic models have been outed since, some of which even nest Ornithischia within Theropoda. Image: Darren Naish.

Moving to dinosaurs themselves, a decision made early on is that we’d retain our section on the Ornithoscelida hypothesis, this being the idea that theropods and ornithischians are more closely related to one another than either is to sauropodomorphs. This is because at least some of the studies published since the hypothesis was first outed in 2017 have found that those competing views that now exist are equally well supported by the data. It’s not true that the original authors of the first Ornithoscelida paper (of which Paul is one) have abandoned the hypothesis, by the way.

Dinosaur diversity. Numerous small changes have been made to the theropod section, some of which take account of new discoveries pertaining to Dilophosaurus, noasaurid ceratosaurs and baryonychine spinosaurs. We modified the text on Spinosaurus to better reflect continuing disagreement on what it was like, and we also replaced my old skeletal reconstruction of the tyrannosauroid Eotyrannus with a new one provided by Dan Folkes (based on the new data described in my 2022 monograph on this taxon, co-authored with Andrea Cau; Naish & Cau 2022).

**Caption:** assorted theropod illustrations that appear in the third edition of *DHTLE*, nearly all of which are new for this edition. (A) *Spinosaurus*, now with modified tail; (B) *Eotyrannus* skeletal reconstruction by Dan Folkes; (C) the woodpecker *Campethera*; (D) *Deinonychus* skeletal reconstruction by Scott Hartman; (E) the pheasant *Phasianus*, and (F) the penguin *Spheniscus*. The birds all feature in a neornithine cladogram. Images: (c) Dan Folkes, (c) Scott Hartman, Darren Naish.

Megaraptorans get slightly more mention than they did in previous editions, the conflict over whether they belong within allosauroids or tyrannosauroids requiring that they get a mention in both sections (I currently prefer a tyrannosauroid position). I made the mistake of implying that megaraptorans are wholly Gondwanan, something that’s not true given the inclusion of Japan’s Fukuiraptor and Thailand’s Phuwiangvenator within the group (a few other possible Laurasian megaraptorans are known are well).

Our ornithischian text has received a reasonable overhaul. A new cladogram (more in keeping with the look of the other cladograms in the book) is featured, and the text on heterodontosaurids has been changed to reflect competition between the view that they’re ‘archaic’ ornithischians outside the clade that includes thyreophorans and cerapodans versus one where they’re within Cerapoda and close to marginocephalians.

**Caption:** another new illustration included in the third edition of *DHTLE*, albeit only at small size and in a cladogram. It’s a fuzzy heterodontosaurid. Image: Darren Naish.

Moving to thyreophorans (armoured ornithischians), the text on Scelidosaurus has been updated in view of David Norman’s 2020-2021 monographs, and we also take account of Jakapil, a bipedal ornithischian from the Late Cretaceous of Argentina that, if correctly identified, shows that Scelidosaurus-like thyreophorans persisted for about 100 million years than previously thought (we are of course aware that other identifications for Jakapil have been proposed).

In the ankylosaur section, new text has been added that takes account of the Canadian nodosaurid Borealopelta and the details it preserves on its colour scheme (the immaculately preserved specimen retains the melanosomes it possessed in life). We also take account of the proposed existence of Parankylosauria, a Gondwanan clade named in 2021. We weren’t able to change the ankylosaur text to take account of Raven et al.’s 2023 reclassification of ankylosaurs (this study resurrected Polacanthidae, Struthiosauridae and Panoplosauridae, and did not recognise a formal Nodosauridae) (Raven et al. 2023), so that’ll be something to remember for next time. The stegosaur section has been changed in view of new information on the anatomy of the African Kentrosaurus. It seems that those ‘parascapular’ spines belong on the hips after all.

**Caption:** the incredible holotype of the Canadian ankylosaur *Borealopelta*, published in 2017, has to be regarded as one of the best 20 or so Mesozoic dinosaur specimens reported this century. We didn’t have space to show images of it in *DHTLE*, but we do at least discuss its significance. Here it is on show at the Royal Tyrrell Museum in Alberta. Image: ケラトプスユウタ, CC BY-SA 4.0 (**original here**).

Biology, physiology, behaviour. Moving now to the chapters on palaeobiology, the most noteworthy changes are those concerning physiology, eggs and nesting behaviour.

Over time I came to quite dislike the illustration of a ground-feeding Diplodocus I created for the previous editions, both because it’s having to indulge in an unsightly over-extension of its head-neck joint (equivalent to you looking straight up into the sky for an long stretch of time) and because I gave it rows of scales that don’t much resemble the scales known from any sauropod fossil. So a new illustration has been produced. I’m not sure I like that one either.

**Caption:** a good artist can draw something with a very complex texture (like the scaly body of a giant dinosaur) and use clever artistic shortcuts in order to avoid doing the ridiculous… like, say, drawing every single scale. I lack that skill, and thus am destined to perform the unthinkable. This close-up shows part of my illustration of a diplodocid, the final version of which appears in the third edition of *DHTLE*.

Recently published studies encouraged us to modify our section on dinosaur physiology, partly to take account of Jasmina Wiemann et al.’s 2022 histology-based study. Therein, non-bird dinosaurs were mostly found to be endothermic, though some ornithischian lineages were posited as secondarily ectothermic (Wiemann et al. 2022). I think that the argument for ‘mesothermic’ dinosaurs – popular at about the time that we produced the first edition – is looking weak today and a few tweaks to the text of the third edition reflect this.

On eggs and nests, the most obvious change is incorporation of the discovery that soft-shelled eggs (‘leathery-shelled’ is a better descriptor) were present in at least some dinosaur groups. Recent discoveries pertaining to the colour and patterning of Mesozoic maniraptoran eggs are also alluded to.

The great extinction, and birds! Our section on the end-Cretaceous extinction always did, a critical reader might argue, somewhat miss the mark. This is because we opted to endorse an ‘integrated scenario’ where the effects of volcanism were combined with the Chicxulub impact in a terrible dual event. I now think that that was a mistake and that it was the bolide impact alone that’s sufficient to explain the extinction. Recent research backs this up, and in fact one 2020 study found that the atmospheric release of volcanic ash may have mediated the impact by lowering temperatures that would otherwise have skyrocketed (Chiarenza et al. 2020).

**Caption:** new cladograms appear in the third edition of *DHTLE*. Images: Darren Naish.

One of the points we emphasize throughout DHTLE is that birds are dinosaurs, and that we’re now well past the point at which we can ignore this. Palaeontologists who work on the dinosaurs of the Mesozoic often pretend that birds don’t exist, as if the word ‘dinosaur’ is synonymous with ‘non-bird dinosaur’. Those paying attention will know that much has happened in neornithine bird phylogeny this century, a consequence being that there’s an approximate consensus on the shape of the tree. Some of the details depicted in the tree used previously doesn’t reflect that consensus, so a new one appears in the third edition. I owe thanks to palaeornithologist Albert Chen for discussion of this issue.

**Caption:** your reminder that ‘dinosaur’ is not synonymous with ‘non-bird dinosaur’. Stop pretending that it is. Also, it’s just not true that all workers specializing on Mesozoic dinosaurs are unfamiliar with modern ones, and for much more on that subject see my **October 2023 article on Feduccia’s bizarre *Romancing the Birds and Dinosaurs* book**. Images: Darren Naish.

We also added a new illustration of Gastornis. We previously featured one by John Sibbick where the animal is shown as an arch-predator, grabbing a small horse. A big herbivorous bird might still have grabbed small horses on occasion, but that image is looking dated now so it’s gone.

And that’s where we’ll end things. This is a non-trivial amount of change, enough to make the third edition worth owning even if you own the second or first ones. And if you don’t own the book at all, now is definitely the right time to obtain a copy that’s more up to date with respect to our present state of knowledge.

**Caption:** Bob Nicholls is an absolutely joy to work with, and is an incredibly hard worker to boot. Massive thanks are owed to him for the brilliant cover art he produced for *Dinosaurs: How They Lived and Evolved*. **His website is here.** Images: (c) Bob Nicholls.

Oh – I mentioned earlier than the second edition featured a wonderful and entirely novel reconstruction of Tianyulong on its cover, produced by British palaeoartist Bob Nicholls. I’m very pleased to say that Bob was once again able to generate novel cover art, this time depicting the English baryonychine Ceratosuchops. That animal – named in 2021 by a mostly British team (Barker et al. 2021) – is fitting for the cover of a book published by the UK’s premiere palaeontological institution, associated as it is with the naming and study of the baryonychine exemplar Baryonyx itself by the museum’s Alan Charig and Angela Milner.

**Caption:** *Baryonyx* has a special place in the history of British dinosaurs, in our understanding of spinosaurid theropods, and in the palaeontological research of London’s Natural History Museum. These two images are fairly iconic: at left, the late Bill Walker holds the holotype claw; at right, a life reconstruction by John Holmes, produced in 1987. Images: (c) Natural History Museum, London.

Thanks to everyone who’s bought Dinosaurs: How They Lived and Evolved – whatever edition that might be – and special thanks to those who’ve said positive things about it in reviews, have recommended it to others, and have had cause to use it as a way of introducing other people to our current understanding of dinosaurs and their world.

Finally, it’s worth saying that the third edition is, so far, difficult to get in some parts of the world, and I know that some people have made an online purchase of what they thought was the third edition, only to then receive the second edition in the post. In view of this you have to be ultra-careful if attempting to buy the third edition online. For those in the UK, the edition on sale via the NHM shop is definitely the third edition, and in fact purchase from that site is probably the best way to ensure that you’re getting it (other than by picking it up by hand in a shop).

**Caption:** proof that the third edition is on sale in the Natural History Museum shop, a photo from January 2024. Image: Darren Naish.

For previous Tet Zoo articles relevant to the issues covered here, see…

Naish and Barrett's Dinosaurs: How They Lived and Evolved, November 2016
Ornithoscelida Rises: A New Family Tree for Dinosaurs, March 2017
The Second Edition of Naish and Barrett’s Dinosaurs: How They Lived and Evolved, November 2018
Dr Angela Milner and the Discovery of Baryonyx, August 2021
Two New Spinosaurid Dinosaurs from the English Cretaceous, September 2021
The Eotyrannus Monograph is Finally Published, July 2022

My research and writing (including the material that appears here) is supported by the contributions I receive via patreon. If you value what I do, please consider supporting it here.

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Barker, C. T., Hone, D. W. E., Naish, D., Cau, A., Lockwood, J. A. F., Forster, B., Clarkin, C. E., Schneider, P. & Gostling, N. J. 2021. New spinosaurids from the Wessex Formation (Early Cretaceous, UK) and the European origins of Spinosauridae. Scientific Reports 11: 19340.

Baron, M. G., Norman, D. B. & Barrett, P. M. 2017. A new hypothesis of dinosaur relationships and early dinosaur evolution. Nature 543, 501-506.

Chiarenza, A. A., Farnsworth, A., Mannion, P. D. & Allison, P. A. 2020. Asteroid impact, not volcanism, caused the end-Cretaceous dinosaur extinction. Proceedings of the National Academy of Sciences 117, 17084-17093.

Naish, D. & Barrett, P. M. 2023. Dinosaurs: How They Lived and Evolved (Third Edition). The Natural History Museum, London.

Naish, D. & Cau, A. 2022. The osteology and affinities of Eotyrannus lengi, a tyrannosauroid theropod from the Wealden Supergroup of southern England. PeerJ 10:e12727.

Raven, T. J., Barrett, P. M., Joyce, C. B. & Maidment, S. C. R. 2023. The phylogenetic relationships and evolutionary history of the armoured dinosaurs (Ornithischia: Thyreophora). Journal of Systematic Palaeontology 21, 2205433.

Wiemann, J., Menéndez, I., Crawford, J. M., Fabbri, M., Gauthier, J. A., Hull, P. M., Norell, M. A. & Briggs, D. E. G. 2022. Fossil biomolecules reveal an avian metabolism in the ancestral dinosaur. Nature 606, 522-526.

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Regular readers here, and those who follow me on social media (@TetZoo on Twitter/X and Instagram; I’m on Facebook too), will know that I’m heavily invested in the Common frog Rana temporaria population that lives in and around the scruffy ‘garden’ areas that surround my house….

**Caption:** the shallow end of pond 2 as of February 5th. When spawn is fresh, it’s tight, compact and rubbery, but as it matures its jelly absorbs water, expands and becomes much less firm. It’s possible to age the spawn by days and even hours (approximately) on the basis of its look. Image: Darren Naish.

Well, RIGHT NOW – early February – is the most exciting time of the year if you’re interested in explosively breeding frogs like R. temporaria, since this is the time when they suddenly appear in and around ponds, and then go about their competing and calling, and ultimately their spawning. Here’s a 2020 article on the Common frog and my efforts to boost their numbers and help them out.

I keep close tabs on all of this and report it on social media, and such is interest in this whole thing – I call it spawnwatch (with a hashtag where appropriate for social media) – that I’ve just decided to add things here at the blog too. But I’m going to do this in a slightly unusual fashion: I’m going to keep updating this article as long as there’s stuff worth talking about. Come back later to see what I mean!

**Caption:** these photos give you some idea of how packed things get, of how many frogs there are here. The image at right (taken from distance from an upstairs window) basically shows a massive brawling scrum of 30 or more frogs. Images: Darren Naish, Toni Naish.

  **Caption:** it’s the 29th January and frogs are gathering for a weeks-long event. No spawn had appeared at this point. The shallow part of the pond at far right is where the frogs (mostly) spawn. No females are visible in this photo. Image: Darren Naish.

I’ll start by adding that we’re already about a week into spawnwatch. Frogs began gathering in late January, with the first spawn appearing on January 30th, this being the first time we’ve had a spawning in that month. Right now (5th February), there are at least 15 clutches. At least 24 appeared in 2023, so my assumption is that that number will be exceeded in 2024. There are likely more adult frogs in pond 2 (the main breeding pond here) here than last year: over 60, as opposed to around 50 in 2023. An issue I now see with pond 2 is that the shallow part of the pond (which is where frogs prefer to spawn) is not big enough for the amount of spawn we might get, meaning that some of it is being kicked into deeper water (where it won’t develop: it really needs to be at the surface to receive direct sunlight). I’m therefore removing some spawn into an adjacent large container that’s serving as a makeshift pond. I might modify the pond in future in view of this.

**Caption:** heavy frog activity on February 3rd. You’ll notice from these photos that individual frogs can be distinguished on the basis of their markings. I’m making an effort to identify individuals and monitor them year to year. Other such projects are underway elsewhere and very much ahead of me in terms of the data collected. Image: Darren Naish.

OK, enough for now… come back soon for updates added below. Oh, I’ve just returned from the Herpetofauna Workers Meeting 2024 (this year held in Whiteley, Hampshire), so have just spent the last few days immersed in the latest news on British and Irish herpetological science news, truly a fantastic meeting. Here are a few relevant scenes…

**Caption:** Chris Packham welcomes delegates to the Herpetofauna Workers Meeting 2024, this year held at Solent Hotel and Spa, Whiteley, Hampshire. Two days of talks, workshops and events on where we’re at with respect to work on newts, anurans and squamates of Britain and Ireland. Image: Darren Naish.

  **Caption:** select images from the Herpetofauna Workers Meeting 2024. Many books were on sale, including from NHBS and Pelagic. I did something that I’m still trying to process and purchased the massive and substantial *Salamanders & Newts of the World* by Jean Raffaëlli. The card newt head was made during the quiz. I think you can agree that we done good (my team came third overall). Images: Darren Naish, Steve Allain.

UPDATE 1: we’re now up to 17 spawn clutches. But what’s interesting is that both new clutches are in unusual places relative to what the frogs seem to prefer: one is in open water in the middle of the pond (albeit on floating vegetation), and the other is at the heavily vegetated end of the pond (which is shallow, but not ideal as the water is always shaded, not in direct sunlight). I wonder if these clutches were laid where they were because the otherwise ideal shallow end is too packed full of busy, competing frogs?

On the subject of the shallow, busy end., I just watched and photographed a pair in amplexus move into the shallow end with seeming intent. I wondered if I might get to watch actual spawning happen (something I’ve never seen in person). The ‘elevated head’ posture maintained by the male is interesting: was he acting like this to intimidate other males and/or vocalize? Note also that the male is surprisingly plump. Come back for more updates soon…

**UPDATE 2:** the pond has been extremely active today (February 6th), and an additional five spawn clutches have appeared overnight, bringing us up to 22 clutches so far. 58 adult frogs were counted in the pond this morning, with at least some out of sight because they were submerged or in pond 1 when I was doing the count. I’ve seen two frog pairs still in amplexus, so there’s definitely more to come. I am also increasingly concerned that the shallow end of the pond – the part where the frogs want to spawn – is just not big enough for the number of frogs we have, and indeed one of the new spawn clutches is a second middle-of-pond one. There might still be a week or more of this yet to go! I’ll end with an updated version of the graph…

**UPDATE 3:** things have quieted down now (February 9th) and no new clumps have appeared over the past 24 hours, but this puts us at 24 clutches (the photo below was taken on Feb 8th). Part of the reason for the slow down is that it’s now rained continually for two straight days, and this has dampened the frogs’ enthusiasm for calling and competing at the water’s edge. The substantial amount of rain has also raised the level of the pond by about 10 cm, which is not great because the shallow edges they prefer for spawning are no longer as shallow. So that might be about it…but stay tuned, I’ll report what happens next.

**UPDATE 4:** things have essentially wound down, with no new spawn having appeared for the past two days. The final count is 27 clutches as of Feb 13th, and below you can see the resulting graph. Given that several frogs are still hanging around in the pond, it’s not impossible that more spawn clutches will appear, but croaking is no longer occurring and there are never more than five or six frogs visible above the surface. In years past, spawning usually occurred on or after Valentine’s Day (Feb 14th), yet this year the main event occurred a week before this, all of which is consistent with national trends: frogs across the UK now spawn at least one (and often two) weeks earlier in the year than they did approximately five years ago. This appears correlated with increasing average temperatures.

I think that this is likely to be the last update; to those who did so, many thanks for coming back to check things as they got posted. UPDATE! Another clutch has appeared as of 14th February, bringing the total to 28. The graph below is a new one that takes account of this. UPDATE!! More clutches have appeared since Feb 14th, the newest (at the time of writing) being on February 22nd. This has taken us to 31 clutches. And thus another new graph has been created…

For previous Tet Zoo articles on frogs, see…

Green-boned glass frogs, monkey frogs, toothless toads, November 2007
In pursuit of Romanian frogs (part III: brown frogs), August 2011
The toads series comes to SciAm: because Africa has toads too, September 2011
20-chromosome toads, September 2011
Glassfrogs: translucent skin, green bones, arm spines, January 2013
Everybody loves glassfrogs, February 2013
African tree toads, smalltongue toads, four-digit toads, red-backed toads: yes, a whole load of obscure African toads, December 2014
Gladiatorial glassfrogs, redux, January 2015
Frogs you may not have heard of: Brazil’s Cycloramphus ‘button frogs’, January 2015
It's the Helmeted water toad… this time, with information!, January 2015
The Terrible Leaf Walker Frog, March 2017
A Love Letter to the Common Frog, August 2020
What, If Anything, Is A Strabomantid? January 2024
Small Obscure African Toads; Toads Part 14, January 2024

Tetrapod Zoology is dependent on funds raised at patreon. Please help support this blog if you consider it worthwhile and want to see me continue doing it, thanks!

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Giant tortoises are among the most remarkable of reptiles to have ever evolved….

**Caption:** I’m not lucky enough to have ever seen any giant tortoises in the wild. In fact, I’ve never seen any tortoises, of any sort, in the wild, ever. But here are some captive and museum giant tortoises I have seen. Images: Darren Naish.

I don’t know how I’m going to justify that statement but I’ll leave it there as it is, so let’s just live with it. I like giant tortoises enough that I’ve had reason to write about them a few times here in the past (see links below). But while the living species are very nice and all, they do sort of pale in comparison to their extinct giant relative: Megalochelys, a geologically long-lived Eurasian taxon that was alive during the Miocene, Pliocene and Pleistocene and lived from central Asia (and perhaps eastern Europe) in the west to Indonesia in the east, including Borneo and Sulawesi. Recent finds indicate that it also occurred in the Philippines.

**Caption:** the very familiar mounted *Megalochelys* (AMNH 6332) on show at the American Museum of Natural History in New York. This specimen was collected by Barnum Brown in the Siwalik Hills of India in 1922 and was restored from thousands of fragments likened to the pieces of a jigsaw puzzle. Some of the features here are not especially realistic (look at those hands) but others are close to reality: note the forked epiplastral structure at the front of the plastron, the tall, ‘sawn off’, steep-fronted nose, and the wide, oval anterior opening to the shell. Image: Claire Houck, CC BY-SA 2.0 (original **here**).

You might know this tortoise better as Colossochelys, the name that was in use for much of the 20th century and the one still prevalent in books that discuss or mention it. But Megalochelys has nomenclatural priority, having been published by Hugh Falconer and Thomas Cautley in 1837. Colossochelys was published by the same authors in 1844. The reason that Colossochelys won out (for a while) was not exactly empirical: Murchison (1868, p. 359), in notes attached to Falconer’s never-finished c 1837 memoir on the animal, noted that Megalochelys* was abandoned because it “was thought not to convey a sufficiently expressive idea of the size”!

Falconer died in 1865 and left behind a substantial volume of notes on fossils he had been studying from the Indian Siwalik Hills. This work was tidied up and published in two volumes by Charles Murchison, both published in 1868.

Caption: back in 2021, I published two articles on the occasional carrion-eating proclivities of tortoises. Hodari Nundu was sufficiently inspired to produce this excellent illustration, in which a Megalochelys disputes carcass ownership with the hyaenid Pachycrocuta. The venue is Pleistocene south Asia. Image: Hodari Nundu, used with permission.

An additional issue affecting the nomenclature of these tortoises is that they were – like virtually all big tortoises living and extinct – subsumed at times into the super-inclusive catch-all versions of Testudo and Geochelone that were once in fashion. Those names are, today, restricted to specific lineages within the tortoise family (the type species for Testudo is the Spur-thighed tortoise T. graeca, and that for Geochelone is the Indian star tortoise G. elegans) and we now use a system where multiple genus-level names are in use.

There are several Megalochelys species, and the biggest individuals of the biggest species (M. atlas) have a curved carapace length (CCL) of over 2 m and an estimated mass of over 900 kg. There are suggestions of even bigger individuals: Ren Hirayama and colleagues reported one from Myanmar with a carapace length of 2.7 m (Hirayama et al. 2015), though it's difficult to determine there if they were reporting straight-line carapace length or CCL, which makes a big difference. Whatever, Megalochelys was very big, and for an illustration produced in 2020 for my still in-prep giant textbook project… stops to sigh and peer out of the window for a few moments… I did a bit of research into working out what it might have looked like when alive. The text you’re reading here was previously published at the Tet Zoo Patreon but I’m recycling it here. Just think, you could have read this already, four years ago, if you supported me at patreon.

**Caption:** a familiar reconstruction of *Megalochelys* if you share a reading history at all similar to mine. This is Robert Bakker’s reconstruction from his 1986 *The Dinosaur Heresies*. The silhouette depicting *Megalochelys* might be somewhat oversized relative to the human and the modern giant tortoise… but then again, it might not given the size variation present in the three species depicted here. Image: (c) Robert Bakker.

Massive, forked epiplastra, scutes and thorn-like scales. An especially interesting feature of Megalochelys is that it has a massive, thick, robust, bilobed structure at the anterior end of the plaston, formed by the paired epiplastra (Falconer & Cautley 1837, Murchison 1868, Setiyabudi 2009, Srivastava & Schleich 2018). It’s about 15 cm thick (as in: deep in the vertical plane) in big adults, and we know from fossils of juveniles, subadults and young adults that it increased massively as maturity was achieved (Srivastava & Schleich 2018). When covered in its scutes (the gulars), this structure would have been even bigger. It’s narrow from side to side, allowing lots of room for forelimb movement, but is so similar to the forked epiplastra of some living tortoises (like Centrochelys) that it was surely used in similar fashion. That is, as a weapon during intraspecific combat.

**Caption:** the bifurcated, gently curved epiplastral fork or ploughshare of *Megalochelys*. At left, the reconstructed epiplastral fork of *M. sivalensis*, figured by Srivastava & Schleich (2018) and based on the holotype figured by Falconer & Cautley (1837). At right (the three photos labelled a, b, c), the same structure in (a) dorsal, (b) ventral and (c) right lateral views, this time belonging to a *Megalochelys* cf. *sivalensis* specimen from Java described by Setiyabudi (2009). The ploughshare is highly variable in precise size and shape, as is typical for structures that have a role in sexual combat.

The idea of two of these giants pushing and shoving with their ploughshare-like plastron edges is a compelling one that really needs depiction in palaeoart: I've seen one drawing which hints that it might be going on, but nothing really exciting or dynamic. So here’s a challenge to palaeoartists… can we get some thrilling, scary art depicting this behaviour birthed into existence, perhaps making the most of the size and determinedness of these tortoises and their lack of regard for the environment, adjacent foliage and maybe even unlucky nearby animals.

**Caption:** a long, bifurcated epipastral ‘ploughshare’ is present in various tortoises in addition to *Megalochelys*, and is used by them in shoving and fighting. African spurred tortoise or Sulcata tortoises *Centrochelys* sulcata – like the two shown here – do this, as do Madagascan Angonoka or Ploughshare tortoises *Astrochelys yniphora*. Image: a screengrab from Fred’s Reviews58 YouTube video **here**.

Some shells of these tortoises are well enough preserved that we can determine the position of their scutes (the scutes – some experts insist that they really should be called scales – do not align with the underlying bones of the shell), so I added those to my reconstruction after getting the form of the shell right. I then used the known limb bones to reconstruct the lengths of the limbs, and I deliberately gave the animal a relatively erect limb carriage where the legs are held in column-like fashion, this being based on the way the humeral heads fit into the scapulacoroid glenoid. At least some of the several Megalochelys species were continental animals, not island-dwellers... which led me to wonder if at least some of them might be especially well protected on the limbs, as are some big, continental tortoises of today (like Centrochelys). I added thorn-like scales and large, polygonal scales, and I think they look pretty good. The Megalochelys species might have been variable with respect to these details.

**Caption:** at left, a reconstructed *Megalochelys* carapace and plastron. Features to note include relatively large openings for the limbs and a highly domed shape. If you do plan to reconstruct this tortoise, be sure to view the shell from all angles, since it’s wide relative to its height. The scale bar is 10 cm. This specific specimen is from Flores and was reported by Setiyabudi (2016). Yes, these tortoises were on Flores during the Pleistocene, and hence lived alongside Komodo dragons, small stegodonts, giant storks and so on. At right, a life reconstruction that incorporates information from this specific shell. Images: Setiyabudi (2016); Darren Naish.

What else do we know? We have skull material for Megalochelys. Not only does this allow us to work out how large (or small) the head was relative to the rest of the animal (it was about similar in length to the humerus, so over 30 cm long in a very large individual), we also know that this was a tortoise with a short, deep snout and deep nasal region. In other words, the head was shaped much like that of the living Aldabran giant tortoises, so I used these animals as models for the look of the head shown in my reconstruction.

**Caption:** a *Megalochelys* specimen with a CCL (curved carapace length) of 2 m would look stupendous. Here I’m going to follow the long tradition of showing this animal to scale with a modern human: the one used here has a standing height (without hat) of 1.6 m. Image: Darren Naish.

There is, of course, a vast amount of additional stuff that could be said about this fascinating group of tortoises. And, yes, we should get into the habit of referring to them as a group rather than as a singular entity (given that there were several species). The good news is that Jura did a pretty good job of condensing information on them over at Reptilis.net back in 2016. That’s where we’ll end things for now, but I promise to return to turtles at some point in the near future.

For previous Tetrapod Zoology articles on tortoises and other turtles, see…

Gilbert White’s pet tortoise, and what is ‘grey literature’ anyway?, October 2008
Giant African softshells – wow!, Jan 2010
Giraffe-necked giant tortoises, May 2010
Giant fossil matamata turtles (matamatas part V), April 2011
In case you forget, softshell turtles are insanely weird, December 2011
Turtles that eat bone, rocks and soil, and turtles that mine, April 2014
Turtles I Have Recently Seen, June 2015
Letters from the World of Turtle Evolution, August 2017
Tortoises as Consumers of Carrion, March 2021
Tortoises as Consumers of Carrion, Part 2, June 2021
Leopard Tortoises, January 2022

If you enjoy Tetrapod Zoology, please consider supporting its persistence at patreon. Thanks to those who keep things going already, you are the finest of humans.

Refs - -

Falconer, H. & Cautley, P. T. 1837. On additional fossil species of the Order Quadrumana from the Siwalik Hills. Journal of the Asiatic Society of Bengal 6, 354-360.

Hirayama, R., Sonoda, T., Takai, M., Htike, T., Maung Thein, Z. M. & Takahashi, A. 2015. Megalochelys: gigantic tortoise from the Neogene of Myanmar. PeerJ PrePrints 3: e961v1.

Murchison, C. D. 1868. Palaeontological Memoirs and Notes of the late Hugh Falconer: With a Biographical Sketch of the Author Compiled and Edited by Charles Murchison. Robert Hardwicke, London.

Setiyabudi, E. 2009. An Early Pleistocene giant tortoise (Reptilia; Testudines; Testudinidae) from the Bumiayu Area, Central Java, Indonesia. Journal of Fossil Research 42, 1-11.

Setiyabudi, E. 2016. Pleistocene reptiles of the Soa Basin (Flores, Indonesia): adaptation and implication for environment. Jurnal Geologi dan Sumberdaya Mineral 17, 107-124.

Srivastava, R. & Schleich, H. H. 2018. Fossil Turtles and Crocodiles from the Caenozoic of S Asia. Society for Amphibian and Reptile Conservation of Nepal, Munich.

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Once again, it’s January 21st, meaning that Tetrapod Zoology the blog – initiated one dark night in the long-ago age of 2006 – has reached another birthday. It’s 18th, no less. And thus it’s once more time to look back at the previous year from the very biased, wholly whimsical and personal perspective of Tet Zoo- themed events…

**Caption:** a montage of things relevant to Tetrapod Zoology’s 2023. From left to right: a cassowary in Florida, a basilisk (top), a muscovy duck, and some 3D-printed dinosauroid models. Images: Darren Naish.

2023 was another manic year in the history of things Tet Zoo. Some indication of this is given by the fact that the 17th birthday article wasn’t published on time, in part because I was busy returning home from fieldwork in Florida. And said fieldwork was the grand cassowary relocation event, covered at length some months later during the year (go here).

Ancient Sea Reptiles and Monsters of the Deep. Among the first personally significant events of the year happened during February: I mean the widespread release of Ancient Sea Reptiles: Plesiosaurs, Ichthyosaurs, Mosasaurs and More by the Natural History Museum in the UK and Smithsonian Books in the US (Naish 2022). Ancient Sea Reptiles saw print in October 2022 but it wasn’t really out and available in shops until early 2023. I have many thoughts on this book and the general message it aims to convey, and for more on those things I direct you to this February article. Another article promoting the book and its message was published in Evolve, the magazine of the Natural History Museum (Naish 2023a). Ancient Sea Reptiles has done well in sales, sufficiently so that a second impression (impression, not edition) was put into production only a few months later in the year. This allowed me to correct a few typos and broken sections of text that somehow made it into the first printing.

**Caption:** ancient sea reptiles get covered in the Natural History Museum’s *Evolve* magazine (message me if you want a pdf of the article), the same issue that announces the museum’s hosting of Titanosaur, an exhibit devoted to the biology and anatomy of the giant sauropod *Patagotitan*. Images: (c) Natural History Museum, London.

Regular readers of Tetrapod Zoology will know that I was guest curator for Monsters of the Deep (MOTD), a museum exhibition hosted at the National Maritime Museum, Cornwall in Falmouth. Alas, MOTD came to the end of its run at Falmouth in January 2023, which is a shame because I sure loved using it as an excuse to visit that part of the world. So… what next?

The good news is that MOTD had and has life beyond Falmouth, since in April 2023 it re-opened at Chatham Historic Dockyard in Essex. Museum spaces being what they are (no two museums are alike), the exhibition had, of course, to be re-arranged and re-designed, and the final result was very different from the Cornish version. But I’m saying all of this from photos, since I was unfortunately never able to get to Essex to see it myself. And, indeed, I never will see it, since it ended its run in Essex in November (herpetologist Steve Allain wrote an article about the exhibition here). What next? Well, news is due to be announced, so stay tuned.

**Caption:** Monsters of the Deep as it looked at Chatham Historic Dockyard, Essex, during November 2023. **See Steve’s article** for more images taken during his visit. Image: Steve Allain.

2023, year of frogs. I’ve been paying some reasonable amount of attention to our local frogs for as long as I’ve lived here, and regular readers might recall that we started out with essentially no frogs in the area before seeing a general uptick that was fully in swing by 2020. I’m talking here about the lone amphibian species that occurs in my area: the Common frog Rana temporaria. We have no toads here, no newts. Depauperate herpetofauna donchaknow.

**Caption:** the shallow end of pond 2 during February 2023. Common frogs need very shallow areas in which to spawn, since their eggs will only develop if in sunlight. Something I hadn’t appreciated before 2023 is that large numbers of frogs have the ability to modify the shallow parts of ponds through their activity, since their competing, fighting and constant moving removes leaf litter and sediment. Images: Darren Naish.

  **Caption:** fresh spawn is tight, compact and rubbery, and individual clutches are easy to distinguish. But as the spawn absorbs water, it stretches out and individual clutches become hard to distinguish. We definitely ended up with at least 24 masses of spawn, but that number is conservative. Image: Darren Naish.

A concern I have about our newer pond – it’s called pond 2 – is that it’s deep, often in shade due to tall hedges, and with a substantial amount of duckweed (and other floating plants) on its surface. This is bad for frogspawn and tadpoles, since they develop fastest when ponds are shallow and sunlit, and relatively warm overall. My plan for 2023 was to reduce the height of the hedges, and also remove the duckweed. On the first point, I wasn't much successful. I just never found the time. On the second, I largely succeeded. Anyway, the breeding season for 2023 was exceptional, with more than 50 adult frogs present and at least 24 spawn clutches. What a massive success! How did this happen? (1) the creation of ponds, and (2) the maintenance of deliberately ‘wild’ areas around the ponds, with substantial understorey, leaf litter and brush piles.

**Caption:** a depiction of spawn clutch number present in the ponds here at Tet Zoo Towers, recorded over the time that we’ve had ponds. A long period of almost nothing followed by a rapid, exponential rise. Image: Darren Naish.

I look forward to seeing how things go in 2024. Have we reached peak frog, or are numbers set to increase even more? I’ll report events on social media (I’m still active on Twitter/X as @TetZoo).

Baryonychines part 1. As per most years, my work on Mesozoic dinosaurs – in particular those of the Lower Cretaceous Wealden Supergroup of southern England – continued in the background. A few papers that have been on the backburner for years now, err, remained on the backburner, and those that made it into print were led by my colleague Chris Barker and represent parts of his PhD (which is devoted to baryonychine spinosaurs). The first of these for 2023 appeared in Journal of Anatomy in February. It concentrates on baryonychine brain anatomy as inferred from CT scans (Barker et al. 2023a) and I wrote it about here.

At some point during March I appeared as a guest on the Plastic Plesiosaur Podcast run by Miles Greb and Trey the Explainer. Each episode of the Plastic Plesiosaur Podcast is based around the theme of a specific episode of the much-loved TV series MonsterQuest, and the episode I appeared in was devoted to an episode (number 19 of season 2) titled Jaws in Illinois, the main focus of which is the alleged presence of bull sharks in fresh- and brackish-water locations in the US. It’s a bit weird that Miles and Trey chose me to contribute seeing as I’m not exactly a shark expert, but I think I was deemed topical because of the recent publication of Ancient Sea Reptiles. The episode was released in late March and is here. Incidentally, I starred in an episode of MonsterQuest back in the day, I think devoted to the British big cat phenomenon.

**Caption:** I’m lucky enough to live close to the New Forest National Park and get to visit it often. Here’s a scene from February 2023. It’s an interesting place for lots of reasons, including its massive bracken meadows and ‘wild’ ponies. A very problematic thing about the forest – remember, it’s labelled a ‘National Park’ – is that it’s generally seen by people as a dog playground, meaning that literally thousands of dogs run about the forest every single day. So, good luck conserving ground-nesting birds, the wildlife of ponds and streams, ground water quality and so on. I say all of this as a dog owner myself. Image: Darren Naish.

  **Caption:** having mentioned New Forest ponies, here’s a montage. These ponies were all photographed on the same day (23rd May 2023). Image: Darren Naish.

Dorling Kindersley’s Ask An ‘Ologist. March was also when the first of the ‘Ask An ‘Ologist’ Dorling Kindersley events happened. This was an interactive event in which school kids from across the UK would submit questions on a given subject and have them answered by a relevant ‘ologist’. I might not be a professional palaeontologist but I play one on TV, and I’ve worked for DK on numerous occasions since the early 2000s. Hence it was appropriate that I get roped in to answer dinosaur-themed questions. Other experts dealt with questions on Egyptology, modern zoology, and science in general.

**Caption:** my relationship with Dorling Kindersley has been long and fruitful. As I said in **this article from 2022**, the 2001 *Encyclopedia of Dinosaurs & Prehistoric Life* (which I co-authored with David Lambert and Liz Wyse) should probably be considered the most significant book I’ve been involved in. Images: Darren Naish.

The school that asked ‘the best’ question (as decided by we ‘ologists) won a ton of books, and also got a special visit from the relevant ‘ologist. I visited St Richards Reynolds Catholic College in Twickenham, London, in June, where I spoke to a class of Year 4 children (ages 8-9). There were lots of questions, and all were brilliant (I think the most memorable was "Would dinosaurs have howled at the moon?"). Thankfully, no-one asked the dreaded "How do you become a palaeontologist?". A few of the children hung around even as they were supposed to have left, and a girl asked me "Are there any girl palaeontologists?". I said that the subject is for everyone and that there are many great women in palaeontology. Another girl wanted to tell me that she was from Mexico since one of the issues that came up in the Q&A was the Chicxulub impact site. This was a hugely positive experience and I’m really grateful to have been involved in it. Everything at the school was well organised, DK did an excellent job in pulling it all together, and the children and staff were great and really fun to talk to.

**Caption:** in a London school during June 2023, I tell a group of Year 4 children what it’s like to get to work on books. Image: Hattie Hansford/Dorling Kindersley.

It was considered such a success that plans were soon made for a US version, and this happened in early November. Sadly, DK weren’t prepared to fly us out to the US to visit the school that gave us the ‘best’ question, oh well.

Also during late April, I attended the Lyme Regis Fossil Fair (again), this time as a guest of the Lyme Regis Museum where I gave a talk (and participated in a book signing) on Mesozoic marine reptiles, this of course again being a tie-in with Ancient Sea Reptiles. The only room available for the event was small and most people who turned up – including members of my own family – had to be turned away!

**Caption:** scenes from the Lyme Regis Fossil Festival of 2023. At right, a replica miniature *Neovenator* skeleton owned by a company that specializes in the digitizing and printing of fossils. I’ve shown **images of this replica before**. Images: Darren Naish.

Prehistoric Planet Season 2. One of the biggest personal events of the year was of course the May 2023 release of season 2 of the Apple TV/BBC Studios series Prehistoric Planet. I’m chief scientific consultant for the series and being involved is an incredible experience; involvement now marks a significant chunk of my adult life and it’s been an absolutely joy to work at the BBC Natural History Unit and to collaborate with the many amazing and talented people involved in the making of the series.

We didn’t have publicity events in Los Angeles this time round, but events associated with the release of the series did happen at several venues during May and June, including Queen Elizabeth Hall, the Everyman Cinema at Broadgate, London Comic Con and the Festival of Nature in Bristol. Our official day of dealing with journalists and abusing local bars happened in early May, and here’s where I got hold of the big Prehistoric Planet season 2 poster board I now own.

**Caption:** a Prehistoric Planet season 2 promotional event at Queen Elizabeth Hall on London’s South Bank. At left, Liz Bonnin hosts. At right, (left to right) executive producer Mike Gunton, series producer Tim Walker and consultant Darren Naish field question. Behind and around us are members of the BBC National Orchestra of Wales. Image: Amber Cherry Eames.

  **Caption:** at another Prehistoric Planet season 2 event, series producer Tim Walker and I discuss the making of the series from the main stage of London’s MCM Comic Con during May 2023. Images: Toni Naish, Darren Naish.

A novel experience for 2023 was the recording of a podcast dedicated to Prehistoric Planet, some of which happened at my house, some of which happened in special studios. The several different episodes were released throughout May. They’re available here.

**Caption:** images from the Prehistoric Planet season 2 media event held at the offices of MPC in Soho (London) in early May 2023. Dinosaur-shaped biscuits at left; my buddy Prehistoric Planet producer Dom Walter at right, outside The Ship. Images: Darren Naish.

Some readers here will know that I’ve been tweeting long threads on the scientific decisions behind each of the Prehistoric Planet episodes. For season 2, time has just not allowed me to get through these at all quickly, and even now (January 2024) I’ve only gone through two of them. The first – dedicated to Islands (episode 1 of season 2) – was released in June (it’s here at Threadreader), the second – dedicated to Swamps (episode 3 of season 2, but wrongly thought by me to be episode 2) – appeared in November. The others are coming, I promise.

**Caption:** a selection of Prehistoric Planet objects I own. The poster board might be familiar. As for the models: sadly, Apple aren’t interested in producing Prehistoric Planet merchandise, but fans of the series have made their own incredibly good versions, some of which are available for sale online. Image: Darren Naish.

  **Caption:** a close-up of those amazing (but unofficial) Prehistoric Planet models. The large swimming *Tyrannosaurus* at left is by **Ancestors Models** (sadly not available for sale at the moment). The remainder are by Marco Makes whose products are **here on Etsy**. Image: Darren Naish.

  **Caption:** more detailed images of the amazing Prehistoric Planet-inspired swimming *Tyrannosaurus* model made by **Ancestors Models** (sadly not available for sale at the moment). It’s built to float (though I admit that I haven’t tested this) and is beautifully painted. The company concerned is set to produce more models of this quality. Images: Darren Naish.

Also on Prehistoric Planet, I was interviewed for the All Creatures podcast in June and for the I Know Dino podcast in July.

Moving now to something quite different… thanks to a tip-off from Dan Garrick, I made an emergency trip to Marwell Zoo in late May to watch the Crocodile monitor Varanus salvadorii there consume an entire rabbit. It was a rewarding experience and I took tens of photos. The lizard used the trunks of nearby trees to help position and reposition the carcass within its mouth. We also got to see the zoo’s two young Mainland clouded leopards Neofelis nebulosa. I’ve seen clouded leopards before but not those two.

**Caption:** a captive (and not yet fully grown) Crocodile monitor consumes a furry vertebrate object. How predators process, manipulate and swallow prey is a hugely important component of their natural history: for many animals we still have comparatively little information on this aspect of their evolution and biology, so observation and study of it is always welcome. Images: Darren Naish.

  **Caption:** the clouded leopards are among the most beautiful and anatomically remarkable of all cats. **I wrote about them briefly in 2007**, back when the Sunda clouded leopard *Neofelis diardi* was resurrected from synonymy. It’s fitting that the two Marwell Zoo individuals shown here were photographed while partly concealed by foliage. Images: Darren Naish.

Baryonychines part 2. At the end of May our second spinosaurid paper appeared, this one being devoted to our analysis of a single tooth from a relatively old part of the Wealden Supergroup. It was published in PeerJ and thus is open access. As discussed in the article (Barker et al. 2023b) and the resulting Tet Zoo article, here was an excuse to test the hypothesis that all isolated baryonychine teeth from the Wealden should be referred to Baryonyx. No, they shouldn’t.

**Caption:** it’s pretty scary how much science can be done on isolated teeth. They are the very opposite of useless and disposable bits of evidence. The paper being depicted here is, again, open access. Images: **Barker *et al*. (2023b)**.

Work on additional English baryonychine specimens continues in the background. On that note, I think it goes without saying that… at any one time, five or more technical papers are being compiled, finished or reviewed ‘in the background’. I tend to hold off talking about them until they see print, but – as of the time of writing (January 2024) – I have several papers on dinosaur palaeobiology, anatomy and diversity in various stages of completion. And they’re not all on baryonychines, ha ha.

Not going to Brazil. An amazing trip was due to happen in June. I was an invited speaker for the 9th Pterosaur Symposium in Crato, Brazil. I’ve visited Brazil once before (in 2013, and that was for the 6th Pterosaur Symposium: I wrote about it here), but that was in Rio, on the coast. This Crato trip would surely be a very different experience. I couldn’t wait.

But the problem with flying is that it involves airports, planes, schedules, weather, and all other kinds of incompatible things. And thus it was that an intense storm over Central Europe delayed my first plane by about an hour, meaning that I (and about a hundred other people) missed our long-haul flight to São Paolo. After an overnight stay involving an ‘emergency field bed’ in the airport (my financially bereft backstory requires that I’m not able to consider paying for airport hotels), I went to arrange a new flight the next day, but nothing could be sorted that would get me there with sufficient time to make it worthwhile. Ultimately, I had to give up and return home. How dispiriting. I later sent apologies to Taissa Rodrigues and the other organisers. Because I’d booked time off work, I made the most of it and went on a much-needed family holiday, albeit only in the UK.

**Caption:** your connecting flight was late, but you rush to the gate anyway… and find that the plane you were hoping to get is still there! Maybe they’ll see you and let you on. Nope, you can do nothing but watch it leave. How depressing. At right: a nighttime view from a walk about the town of Ilfracombe in coastal north Devon. Images: Darren Naish.

  **Caption:** there are several good reasons to go to Ilfracombe, and one of them is Verity, the 20-metre-tall statue designed by Damien Hirst, put in place in 2012 and cast by Pangolin Editions in 40 separate sections. On her right (seaward) side, she is flensed, such that her skull, musculature, internal breast tissue and foetus are visible. She is intended as an allegory for truth and justice, and stands atop a pile of books and holds a sword aloft. Images: Darren Naish.

  **Caption:** as a seaside town, Ilfracombe is of course home to gulls, and I love gulls. This lucky Herring gull *Larus argentatus* is taking advantage (or trying to take advantage) of a gurnard discarded at the dockside by a fisherman. Images: Darren Naish.

Fish and false German lions. A terrible secret was revealed in the June of 2023 when Jeff Liston and I released a co-authored study (albeit only an abstract and conference presentation) for that year’s EAVP (European Association of Vertebrate Palaeontology) meeting at Sabadell in Spain (I didn’t attend the meeting). The catch is that it was on fishes, specifically the giant Jurassic pachycormid Leedsichthys, and specifically on the plausibility of it sometimes engaging in breaching behaviour (Liston & Naish 2023). The fact is that I’ve actually written a truly vast amount about fishes – all of them – and will… eventually… at some point… have published more words and more pages of text on fishes than most other living humans. I refer of course to the still unpublished contents of The Vertebrate Fossil Record, but the less said about that the better. It will see print eventually.

A newsworthy event that saw widespread coverage in online news sources was the alleged observing of a live lion in a suburb of Berlin, Germany, during July. I was confident immediately that the footage did not show a lion but either a calf or a boar (I was eventually convinced by the boar identification), and I said so at Tet Zoo. This was reported in various German news outlets, was ultimately supported by other people with relevant experience, and the story then died a death as people realized that there never was reason to believe in a lion in the first place. This was a classic ‘flap’ (as defined in the Fortean literature) and a person with good knowledge of the history on alleged ‘animal escape’ stories could have written a very accurate script on how things would play out.

**Caption:** at left, the Tetrapod Zoology evaluation of the Kleichmachnow lion-that-wasn’t. At right, a German news item (here, translated into English; **original here**) that covers this interpretation. It turned out that the version of the video shared by the media was selectively cropped in order to remove the unambiguous boar footage present both before and after the ‘lion’ section, and that the person who filmed the footage never said it was a lion anyway.

  **Caption:** a Prehistoric Planet event at the Festival of Nature in late June 2023. The silhouette depicts the tyrannosaurid *Qianzhousaurus* (which appeared in Prehistoric Planet season 1); the skull is that of a *Mosasaurus*. Image: Darren Naish.

Things to do in a garden in June. I must have mentioned here and there at Tetrapod Zoology that one of my main concerns these days is the constant, never-ending erosion and destruction of the natural world that I witness about me all the time. I’m sure that this is more immediately visible here in tiny little overbuilt southern England than it is in many other parts of the world, but it’s a global phenomenon and one that we all have to be aware of. It often seems that individual people can do little about this, since the real issues are ‘big’ things beyond our control.

But we as individuals can at least do a little to help: we can be more tolerant in letting ‘weeds’ grow, can create ponds and pools, and can stop being so insanely keen to remove leaf litter and decomposing wood. Writing entirely from a provincial, ‘little England’ perspective (those qualifiers are important), I tweeted a thread during June 2023 on things that individual people can do to help (the thread is here). The good news is that this went soft-viral and was shared about the web here and there (here’s a version at IFL Science). Obviously, I think my advice is good and hope that it makes a tiny bit of difference.

**Caption:** part of the space around our house is paved, which is not ideal if we want to help nature. But we can improve things by having planted areas and plants growing in pots, as here. Since this photo was taken in June 2023 I’ve acquired another water butt. Image: Darren Naish

What I’d neglected to appreciate is that some of my advice – like, that advocating the creation of new sources of standing water – is considered bad, if not illegal, in some tropical places. I apologise for that and re-emphasize that I was only thinking about things from the British perspective.

**Caption:** proof that Slow worms *Anguis fragilis* still exist in southern England. This one was encountered at Highcliffe, Dorset, during July 2023. Slow worms used to occur in the part of the UK where I live but they’ve mostly disappeared from the places where I used to be able to find them. This is an animal whose survival is contingent on the things mentioned above: they need green spaces with natural cover, arthropods and other small animals, and so on. Image: Darren Naish.

  **Caption:** frogs again, this time encountered foraging in the garden during July 2023. Frogs have white eyeshine and incredible low-light eyesight. Images: Darren Naish.

A summer whale watching adventure. During July, I and a team of buddies once more boarded the Pont-Aven to travel across the Bay of Biscay in search of whales and other marine wildlife. These trips, which involve official monitoring and recording, are organized by the marine conservation charity ORCA and I intend to be involved in them as often as possible. They didn’t happen over the course of the pandemic, and in fact this was the first one to be arranged since 2019. That 2019 trip was a massive success in terms of which whale species we saw (an article about it is here).

**Caption:** I’m naturally very lazy and hence don’t ordinarily see sunrise. But I do when I go whale watching, and here is one. It was shortly after 7am on a July morning. Image: Darren Naish.

  **Caption:** that grey blurry mass on the right is a cloud dumping some quantity of rain, and we’re about to go through it. It rained a lot on our 2023 trip and was extremely windy at times, so much so that we had to stay off deck for part of the trip. Image: Darren Naish.

The 2023 trip wasn’t quite as memorable, but was still great. I had to make emergency measures immediately beforehand to get a new lens for my half-decent camera (a Canon EFS 500D); I succeeded, and it served me reasonably well. We saw dolphins of a few species, a substantial number of Fin whales Balaenoptera physalus (sometimes two and three together) and some gannets and petrels. Numbers of certain of these animals (seabirds in particular) were low, perhaps because of the high sea surface temperatures that were a concern at the time, die-off caused by bird flu, or both.

**Caption:** these are Sowerby’s beaked whales *Mesoplodon bidens*, a new species for me. This is the only beaked whale in the region with a long, slender rostrum, a low melon, and a greyish overall colour. It has small, paired lower jaw teeth located halfway along the lower jaw and was the first species of the highly speciose genus *Mesoplodon* to be scientifically described. Images: Alex Srdic.

  **Caption:** we saw an incredible number (for me!) of Fin whales on our 2023 trip, certainly over 20, mostly distant blows. Fin whales roll quickly, such that the dorsal fin is soon directly beneath the remnant of the still-hanging blow. Images: Alex Srdic.

No Bay of Biscay trip would be complete without sightings of at least some beaked whales, and we were blessed with observations of two species: Cuvier’s beaked whale Ziphius cavirostris and Sowerby’s beaked whale. My associate Alex Srdic has once again kindly allowed me to use his photos. The ship docked in Plymouth in the UK, and we used that as an opportunity to visit both the National Marine Aquarium and The Book Cupboard, one of the UK’s most impressive second-hand book shops.

**Caption:** Plymouth’s National Marine Aquarium is great, and I’ve been there several times. I like life-sized animal models, and here’s a leatherback sea turtle, humpback whale and killer whale in the large hall that’s also home to the giant Eddystone Reef exhibit. Image: Darren Naish.

  **Caption:** when in Plymouth, we visited the Barbican Prawn (officially The Leviathan), and here it is (at left) in less than ideal weather conditions. At right, my book haul from The Book Cupboard, an impressive lot if you share my interests. Images: Darren Naish.

  **Caption:** I live in, and am unable to escape from, the suburban environment. Wildlife is mostly in chronic decline in such places. But it remains surprising how adaptable and tenacious some species are. Look here for the cryptic Roe deer *Capreolus capreolus*, photographed in August 2023 only a few metres across the road from my house. Image: Darren Naish.

Exotic Norfolk; oh, and Tajikistan. I went to north coastal Norfolk in late July, partly to see wildlife there while visiting my friend and collaborator Steve White. I saw Harbour seals Phoca vitulina and numerous gulls, and on that front the impact of bird flu was obvious, since numerous dead and dying gulls were present at the colonies we visited.

**Caption:** more hot Herring gull action, this time from Hunstanton, Norfolk. This picture is part of a whole sequence in which the two birds challenged each other and eventually fought. Image: Darren Naish.

  **Caption:** part of a gull colony at Wells-next-the-sea, Norfolk, photographed during July 2023. You can see the live, normal-looking gulls, I’m sure, but look harder and there are a worrying number of sick, dying and dead ones here too (there’s plastic waste as well, but of course). Image: Darren Naish.

  **Caption:** the seals I mostly see around the shores of the UK are Grey seals *Halichoerus grypus*, but these are Common or Harbour seals (photographed at Wells-next-the-sea, Norfolk). A nice, tranquil scene. All was well and good until a guy on a paddleboard, with his two kids, insisted on coming right up to the seals, this causing them to flee. Image: Darren Naish.

But the big event of this part of the year – one of the big events of the year for me – was my trip to Tajikistan. I’ve always wanted to go to Central Asia, both for the incredible landscape and for the wildlife, and this didn’t disappoint. I didn’t see any wild mammals (though we did frequently observe bear scat, and we at least hear altercations occurring between local dogs and wild boar) but numerous birds were observed and badly photographed: for more on those see my articles on birdwatching in Tajikistan, part 1 and part 2.

**Caption:** I was pleased that we flew across Asia during daylight, meaning that I got views from the plane like this. I’m not quite sure what country or region this spectacular mountainous scenery belongs to; let me know if you can work it out. Images: Darren Naish.

The main reason for the trip was cryptozoological, or ethnozoological if you prefer, since the region is associated with observations and tales concerning both a wildman-type creature termed the gul as well as tigers (this, in a part of Asia where tigers are supposed to be extinct). We collected copious relevant data and are still in the process of getting it properly written up. The results were sufficiently encouraging that we plan to return.

**Caption:** the Tajik countryside was truly spectacular. I’ve never seen so many amazing mountains. Unfortunately, the regions we visit were far from ‘remote’, however, and signs of human presence and use were everywhere. Image: Darren Naish.

  **Caption:** Tajikistan is a land of massive, raging rivers, and numerous wooden bridges have been built to allow their crossing, some safer than others. A massive series of storms passed across the region where we were staying during the latter part of the trip and several bridges were lost. In the photos here, Richard Freeman is visible at left and Jon Hare at right. Images: Darren Naish.

During September I was supposed to be in Lyme Regis for a special fossil-themed event, but that didn’t happen for reasons. I did, however, go to the Fossil and Mineral Fair at Lyndhurst in the New Forest, to the coast, to Bristol, and to London to see the titanosaur exhibition (a tie-in with Prehistoric Planet). We were also finally ready – months of preparation were involved – to release tickets for TetZooCon 2023, and off on sale they went.

**Caption:** at the 2023 Mineral and Fossil Fair (in Lyndhurst, Hampshire), Kate Acheson-Dumbravă is in charge of the ZOIC PalaeoTech stall. She has plush dinocarididans (and more)! Kate and I go way back and did fieldwork in Romania together, back in the day. Image: Darren Naish.

  **Caption:** I visited a massive number of local places where you can look at wildlife during 2023, but I mostly haven’t discussed them here as this article is already way too long. Here’s an interesting shot showing Sturt Pond, Milford-on-Sea, during October 2023, where a Herring gull is pretending that it’s a wader, perhaps because it was aiming to steal from the Little egret *Egretta garzetta* just ahead of it. Image: Darren Naish.

Loch Ness again, part 1. I mentioned in the previous birthday article that I did filmed interview pieces for two Loch Ness Monster documentaries during 2022, and the results of both appeared on screen in 2023. The first – titled Loch Ness: Hunting the Monster – appeared on the UK’s Channel 5 in late September.

**Caption:** title card for Channel 5’s *Loch Ness: Hunting the Monster*. I have no idea how accessible it is outside of the UK, but to those within it **it’s available here on the My5 app**.

Hunting the Monster was 2 hours long and focused on the way belief in the monster was compiled through the supposed accruing of new evidence, and the soundbites from experts (including myself) mostly explained how people interpreted that evidence at the time. It also mostly did a fair job of saying what happened to that ‘evidence’ in the end, with their treatment of the Rines flipper photos (the Tet Zoo treatment of which is here) being especially good. They never said that the MacNab 1955 photo was a boat wake though, nor that Dinsdale’s 1961 film shows a boat. I featured throughout. My performance was a bit stilted but maybe that’s because I was trying to sound authoritative.

**Caption:** a less than ideal image of that Darren Naish guy appearing in yet another documentary on the Loch Ness Monster. I just know that the two or three Nessie truthers that still exist out there *love* it when they see me appear on these shows.

The Tetrapod Zoology Animal Figure Collection on film. At some point during August (while I was in Tajikistan actually), conservationist, author and TV presenter Chris Packham asked if I’d be interested in having a film made about my animal figure collection. I said yes, of course, and in October I was visited by Chris, camera operator Joe Cooper and junior presenter Aneeshwar Kunchala.

**Caption:** Darren, young Aneeshwar, and Chris (at right) talk dinosaur models and toys at Tet Zoo Towers. Aneeshwar is holding the giant Beasts of the Mesozoic *T. rex* figure; I’m holding the much-sought-after Dinosaurs in the Wild one. Image: Anil Kunchala.

The relevant segment was for the online TV series 8 out of 10 Bats and the relevant episode, released in early November, is here: the segment with me starts at 21:41. This was a fun and very rewarding experience, and I hope that my message about the educational value of animal toys and figures made it through to those interested. It was also very gracious of Chris to give plugs for Ancient Sea Reptiles, Mesozoic Art and TetZooCon as well! Many thanks to Chris, Megan McCubbin, and Aneeshwar and his family for making it happen.

In case it’s not obvious (I actually have no idea whether it is or isn’t), I really struggle to make time for blog-writing these days, so it’s quite a big deal when I complete and publish an especially long, complex article. That in mind, October’s publication of my very long review of Alan Feduccia’s book Romancing the Birds and Dinosaurs was notable. Because it was written in co-operation with the German AG EvoBio site that aims to combat creationist claims, a German language version also appeared.

**Caption:** at left, the cover of Ethan Kocak’s amazing *Moistly Harmless* of 2023, with associated merch and artwork. At right, some of the beautiful art (some by guest artists) that appears within.

Ethan Kocak’s Moistly Harmless (Kocak 2023), a graphic novel written in celebration of salamanders, arrived in October too. I did the foreword and hence feel special connection to it. And my article on the making of Prehistoric Planet – I think the only article I’ve published on this subject – appeared in the Natural History Museum’s Evolve magazine (Naish 2023b), sadly only available to the museum’s members.

Dinosaurs: How They Lived and Evolved, third edition. During late 2022, Paul Barrett and I received news that our Dinosaurs: How They Lived and Evolved was going to get a third edition. The field of dinosaur science moves fast, so here was an excuse to modify numerous sections of the book in keeping with new data and hypotheses, and even to swap out certain illustrations. We made a start on this in December 2022, worked on it through the early months of 2023, and were mostly done by April. Bob Nicholls agreed to produce new cover art and we opted to replace the fuzzy Tianyulong of the second edition with a beautiful rendition of the baryonychine Ceratosuchops.

**Caption:** goodbye second edition, hello third. Bob Nicholls has done the cover art for both of these editions, and let’s see if we can keep this tradition going into the future. Images: Natural History Museum Publishing/Darren Naish.

My copies arrived in early November. I’m pleased to get the book (Naish & Barrett 2023) to a third edition: my ultimate hope is that it gets at least as many new editions (I think four) as one of its predecessors, Alan Charig’s A New Look at the Dinosaurs. I’ve been planning to explain how this third edition is different from the second and first but, again, time has not allowed. I obtained copies for sale at TetZooCon 2023 and also for the 200 Years of Dinosaurs conference at the Natural History Museum (on which, more below).

Regular readers here will know that I’m not only interested in new editions of my books, but also in foreign-language versions. I feel it’s quite the win when you learn that your book is being translated, especially if the country or region is one that isn’t exactly blessed with an abundance of relevant books written in the relevant language. So it’s good news that a Dutch language edition of Dinosaurs: How They Lived & Evolved appeared in March 2023. It has a totally different design and look from the existing editions. The full title is Dinosauriërs: Hoe Spectaculaire Fossielen en Nieuwe Technieken ons Beeld Steeds Completer Maken, meaning something like 'Dinosaurs: How Spectacular Fossils and New Techniques Are Increasingly Completing Our Picture'. It's published by VeenMedia, in association with New Scientist. And it's interesting to see John Sibbicks's spectacular ceratosaur vs brachiosaur painting make the cover, an image that so many of us associate with an iconic dinosaur book from the 1990s.

**Caption:** at left, cover of the extremely well designed *Dinosauriërs: Hoe Spectaculaire Fossielen en Nieuwe Technieken ons Beeld Steeds Completer Maken*. I like the late-1980s John Sibbick cover art, even though it does seem an odd choice for a book published in 2023. Many of us of course best know that piece from the cover of the 1990 *Dinosaurs: A Global View* by Sylvia and Stephen Czerkas. Images: Darren Naish.

Loch Ness again, part 2. Episode 92 of the podcast was released, and it was another marine reptile special, this time focusing on mosasaurs. And November also saw the release of the second of those Loch Ness Monster documentaries, this one titled They Created a Monster and currently available (in the UK) via BBC iplayer. The film mostly focuses on notorious Loch Ness Monster faker and huckster Frank Searle, one of many characters involved in the hunt for Nessie, and one I’ve had reason to mention in previous articles on the Loch Ness Monster.

**Caption:** the BBC iplayer card for *They Created a Monster*. If you know your Loch Ness Monster history, you might recognize the artistic depictions of various of the players.

Also on monsters, I featured on a Halloween ‘scary science’ segment on the BBC World Service Unexpected Elements show. I think you need a BBC account to listen to it, but it’s here (starting at 14:17). I inadvertently said that mothman is associated with New Jersey, not West Virginia, during the interview, oops.

TetZooCon, Godzilla, and In Conversation With David Lindon. December. And here we come to another massive personal event of 2023: the 10th TetZooCon. Given that this was THE TENTH ONE, it follows that we’d make it something big and extra special, and I think we succeeded. Two-and-a-bit days of talks, events, workshops, merchandise sales and more, with bonus electrical faults, air con and heating failures, and concomitant freezing temperatures. I won’t say any more here since a very long review of the event appeared in December. TetZooCon 2023 marks another watershed in the history of the event, and there’s no going back now. Stay tuned for news on 2024’s meeting.

**Caption:** Jennifer Campbell-Smith very kindly produced this image of a Lammergeier *Gypaetus barbatus*, one of my favourite animals., as a sort of personal thanks for her involvement in TetZooCon 2023 (she spoke on corvids). It was fantastic to have her there. Image: Jennifer Campbell-Smith.

Of other relevant events in December, I saw Godzilla Minus One at the cinema. Godzilla (and other movie monsters) is part of the Tet Zoo remit since thoughts on creature-building and the speculative biology devised to explain them is relevant to our interests, as demonstrated by various articles that have been published at Tet Zoo over the years (and see Godzilla vs. Kong: A scientist decides who would really win).

A zoom conversation I had with David Lindo was released as part of his In Conservation With series during late December (it’s here on YouTube). David is best known as a birder but it turns out that he has a serious, secret interest in cryptozoology, and that’s the subject we discussed. I’ve previously met David in real life (he spoke at TetZooCon 2015, hosted at the London Wetland Centre) and in the digital realm (when he spoke at TetZooMCon 2020, this time on the possible persistence of supposedly extinct birds).

**Caption:** your humble author, at left, with the always excellent David Lindo (and a particularly good background. I really must get a greenscreen). Images: The Urban Birder (**original video here**).

The inevitable introspection. And so finally we come to January, and the run-up to today. For reasons, I published on frogs at Tet Zoo (here on strabomantids, here on obscure African toads), but this of course is the part of the year when I reflect hard on the events of the previous 12 months (more so than I do in December), all in preparation for birthday articles such as the one you’re reading now.

I didn’t just sit around through the whole of January though, since I’ve only recently returned from 200 Years of Dinosaurs: New Perspectives on an Ancient World at the Natural History Museum. This was a phenomenal event, featuring about 30 talks on cutting-edge dinosaur research worldwide. I got to visit the titanosaur exhibition again, this being just a few days before it was due to be closed down.

**Caption:** scenes from the excellent 200 Years of Dinosaurs conference, held at the Natural History Museum, London. Prof Paul Barrett opens the meeting at upper right. The original *Megalosaurus* lower jaw was present and had been specially brought along by arrangement with Oxford University Museum of Natural History. I include (middle image, lower row) an image from Matt Lamanna’s talk on Antarctic dinosaurs since it shows the Prehistoric Planet season 2 depiction of *Imperobator*… will that reconstruction (which I and others designed) stand the test of time? Images: Darren Naish, Michael Pittman (lower right).

And thus here we are, the 18th birthday of Tetrapod Zoology. As per tradition, let’s review the blog’s subject coverage during the year…

Miscellaneous

The 17th Year of Tetrapod Zoology: 2022 in Review, January 2023
Five Years of Tetrapod Zoology ver 4, July 2023
Announcing the 10th Tetrapod Zoology Convention, October 2023
New Species Round-up for 2023, Part 1, November 2023
New Species Round-up for 2023, Part 2, November 2023
At TetZooCon 2023, November 2023
TetZooCon 2023 the report, December 2023

Amphibians

What, If Anything, Is A Strabomantid?, January 2024
Small Obscure African Toads; Toads Part 14, January 2024

Mammals

Piltdown Man and the Dualist Contention, March 2023
The Discovery of the Okapi, Part 1, April 2023
The Discovery of the Okapi, Part 2, April 2023
The Slightly Surprising Diversity of Zebras, Part 2, June 2023
The Kleinmachnow Lion is Definitely Not a Lion, July 2023

Squamates

The Remarkable Basilisks, May 2023

Permian and Mesozoic swimming reptiles

Ancient Sea Reptiles Is Out Now, February 2023
News from the World of Ancient Sea Reptiles, May 2023

Non-bird dinosaurs

A brain for Baryonyx: using CT-scanning to examine British spinosaurid brains, February 2023
Discovering 'Hidden' Diversity Within Wealden Spinosaurid Dinosaurs, May 2023
Alan Feduccia’s Romancing the Birds and Dinosaurs: Forays in Postmodern Paleontology, October 2023

Birds

A History of Larks: Twists and Turns and Overlooked Convergence, February 2023
Larks Part 2: of Subspecies and Brickbats for Carl, March 2023
Birdwatching in Oregon, June 2023
Controversies in Ratite and Tinamou Evolution (Part I), July 2023
The Great Florida Cassowary Relocation Event of 2023, July 2023
Laysan Ducks: Not as Degenerate as People Used to Think, August 2023
The Remarkable Muscovy Duck, September 2023
Birdwatching in Tajikistan, Part 1, October 2023
Birdwatching in Tajikistan, Part 2, October 2023

Cryptozoology

The Strange Case of the Minnesota Iceman, Part 1, August 2023
The Strange Case of the Minnesota Iceman, Part 2: A Review of Heuvelmans's Neanderthal, September 2023

SpecBio

My Own Personal Dinosauroid, May 2023

As ever, there’s nothing like fair taxonomic balance here. Evidently, I never found time to cover stem-mammals, croc-line archosaurs, turtles and pterosaurs during 2023, and amphibian and squamate articles for 2023 are rare. Birds and mammals win out again in terms of coverage, I think because I saw, or was involved in, interesting things on those groups that warranted coverage. My plans to write about turtles and various other groups during the year just didn’t come to fruition as I couldn’t find time to generate new content, and I’m also a bit surprised that I never wrote more about fossil marine reptiles. I did plan to, what with the Ancient Sea Reptiles book.

As per usual, I’m deeply unhappy with the spread of subject coverage and do wonder why I even pay attention to this issue at this point. And, similarly, I remain frustrated that so many things never got finished during the year, assorted books, papers and articles among them. I am, by now, resigned to the fact that these things take literally years when you can’t devote proper time to them, which I can’t. Those who support me at the Tet Zoo Patreon will know that some progress was made throughout 2023 on the Cryptids of Bernard Heuvelmans project, but not nearly enough to get the thing anywhere close to being finished. I’m hoping to have it done by late 2024.

**Caption:** Flame the bearded dragon is alive and well, and here she is foraging in the garden during May 2023. On the same day, she swam in the pond (hence the bits of duckweed stuck to her arm). Her bright orange tones indicate that this was all a positive thing from her point of view. Image: Darren Naish.

Anyway…. you will all, of course, agree with me that it’s more important to focus on the positive things more than the negative. Obviously, 2023 was a significant year in the history of Tetrapod Zoology both as a place where content is published, and as a brand: things of note include the wide release of Ancient Sea Reptiles, the third edition of Dinosaurs: How They Lived and Evolved, the publication of two technical papers (Barker et al. 2023a, b) and two magazine articles (Naish 2023a, b), fieldwork in Tajikistan, whale watching, numerous TV and podcast appearances, my involvement in Prehistoric Planet* season 2 and the TENTH – and by far the biggest and busiest – TetZooCon.

I’m not going to discuss Caspar et al. (2023) since it’s only a preprint. But here’s your heads-up that it exists.

Caption: here’s proof, of a sort, that the amazing people who support me at patreon do get bespoke gifts on occasion. Images: Darren Naish.

It just remains for me to say, as ever, a massive thanks to everyone who visits this blog and leaves comments, and to those who support or assist me. Here’s hoping for a happy and healthy 2024!

For previous TetZoo articles on birthdays and other landmarks, see…

Happy first birthday Tetrapod Zoology (part I), January 2007
Happy first birthday Tetrapod Zoology (part II), January 2007
Happy second birthday Tetrapod Zoology (part I), January 2008
Tetrapods of 2007 (happy birthday Tet Zoo part II), January 2008
Happy THIRD birthday Tet Zoo, January 2009
Tet Zoo = 4 years old today, January 2010
2009, a year of Tet Zooery, January 2010
Four years of Tet Zoo: to infinity... and beyond!, April 2010
It is with some dismay that I announce Tet Zoo's first hemi-decade, January 2011
Tet Zoo 5th birthday extravaganza, part II, January 2011
Happy Birthday Tetrapod Zoology: SIX YEARS of blogging, January 2012
Happy 6th Birthday, Tetrapod Zoology (part II), January 2012
Tetrapod Zoology enters its 8th year of operation, January 2013 (I cannot find any intact versions of this article, thanks SciAm)
Today marks NINE YEARS of Tetrapod Zoology, January 2015
Tetrapod Zoology 10th-Birthday Extravaganza, Part 1: 2015 in Review, January 2016
Tetrapod Zoology 10th Birthday Extravaganza, Part II: the Rest of 2015 Reviewed, January 2016
Tetrapod Zoology 10th-Birthday Extravaganza, Part 3: Tet Zoo's Tetrapod Treatment in 2015, January 2016
Today Is Tet Zoo's 11th Birthday, January 2017
The 12th Year of Tet Zoo, January 2018
The Tet Zoo 12th-Birthday Event, Part 2, January 2018
The Much Belated Final Part of the Tetrapod Zoology 12th Birthday Event, December 2018
Tetrapod Zoology Is A Teenager Now, January 2019
Tetrapod Zoology's 14th Year of Operation, 2019 in Review, January 2020
On Tetrapod Zoology’s 15th Birthday, the Year in Review, January 2021
Happy 16th Birthday, Tetrapod Zoology, February 2022
Four Years of Tetrapod Zoology ver 4, July 2022
The 17th Year of Tetrapod Zoology: 2022 in Review, January 2023
Five Years of Tetrapod Zoology ver 4, July 2023

Refs - -

Barker, C. T., Naish, D. & Gostling, N. J. 2023b. Isolated tooth reveals hidden spinosaurid dinosaur diversity in the British Wealden Supergroup (Lower Cretaceous). PeerJ 11: e15453.

Barker, C. T., Naish, D., Trend, J., Michels, L. V., Witmer, L., Ridgley, R., Rankin, K., Clarkin, C. E., Schneider, L. & Gostling, N. J. 2023a. Modified skulls but conservative brains? The palaeoneurology and endocranial anatomy of baryonychine dinosaurs (Theropoda: Spinosauridae). Journal of Anatomy doi: 10.1111/joa.13837

Caspar, K. R., Gutiérrez-Ibáñez, C., Bertrand. O. C., Carr, T., Colbourne, J., Erb, A., George, H., Holtz, T. R., Naish, D., Wylie, D. R. & Hurlburt, G. R. 2024. How smart was T. rex? Testing claims of exceptional cognition in dinosaurs and the application of neuron count estimates in palaeontological research. bioRxiv 2024.01.10.575006.

Kocak, E. 2023. Moistly Harmless: How to Appreciate News and Salamanders. Endangered Art Books.

Liston, J. J. & Naish, D. W. 2023. “Once more into the breach, dead friends”: dynamic behaviour in the giant suspension-feeding pachycormid Leedsichthys. In Alba, D. M., Marigó, J., Nacarino-Meneses, C. & Villa, A. (eds). Book of Abstracts of the 20th Annual Conference of the European Association of Vertebrate Palaeontologists, 26th June – 1st July 2023. Palaeovertebrata, Special Volume 1-2023: 22.

Naish, D. 2022. Ancient Sea Reptiles. Natural History Museum, London.

Naish, D. 2023a. Ancient sea reptiles. Evolve 52, 32-37.

Naish, D. 2023b. Step back in time. Evolve 53, 30-37.

Naish, D. & Barrett, P. M. 2023. Dinosaurs: How They Lived and Evolved (Third Edition). Natural History Museum Publishing, London.

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Long-time readers of Tetrapod Zoology will know of my long-term plans to complete a series of articles that cover the TOADS OF THE WORLD, the first part of which was published in 2009….

**Caption:** a montage of animals relevant to this article. (A) a typical *Bufo* toad, purely for scale-related reasons; (B) Merten's smalltongue toad *Werneria mertensiana*, one of the species discussed in this article; (C) tadpole of a red-back toad (*Schismaderma*); (D) even stranger tadpole of *Werneria*. Images: Darren Naish.

Various complications require that this series has never been finished, and that long gaps in its publication exist. None of this is helped by the fact that Tetrapod Zoology has (by necessity, not design) moved sites four times now, a consequence being that any long-running series is now spread across three of those four different versions of the blog, this in turn meaning that some of its constituent parts can only be observed intact at wayback machine*.

For those who don’t know, the collapse of ScienceBlogs (where Tet Zoo ver 2 is hosted) and the lack of interest at Scientific American Blogs (where Tet Zoo ver 3 is hosted) resulted in the Tet Zoo articles hosted there being reformatted and ruined, mostly through the removal of images (and comments). This presumably happened to all blogs hosted at those sites over the same timeframe.
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  **Caption:** the montages above show – just in case you don’t know – how much toad-related progress has been made at Tet Zoo over the years (specifically, between 2009 and 2014). See the bottom of this article for links to these articles and others.
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Every so often, I aim to pick up where the series last left off and continue, and that’s something I aim to do this year. But in an effort to work out where we left off, and to migrate material here to ver 4 where things can be salvaged, I’m going to start by republishing the most recently published article in the series… which isn’t ‘recent’ at all, since it was published in the long-ago time of 2014. Here it is, with updates and new citations added where necessary…

In the most recent articles in the series – they were published way back in September and November 2011 (see below for links) – we looked at toads that have been identified by some as belonging to an endemic African clade. Several other lineages discussed in the same part of the series aren’t necessarily part of the same clade, but do belong to the same general region of the cladogram (Van Bocxlaer et al. 2010, Pyron & Wiens 2011).

**Caption:** highly simplified phylogeny for Bufonidae with some of the main evolutionary and biogeographical events marked at appropriate places. Based predominantly on Van Bocxlaer *et al*. (2010). *Atelopus* by Giovanni Alberto Chaves Portilla CC BY-SA-2.5; *Rhaebo* by Brian Gratwicke CC-BY-SA-2.0; *Anaxyrus* by LA Dawson CC BY-SA 2.5; *Rhinella* by Froggydarb CC BY-SA-3.0; *Mertensophryne* by Vladimir Dinets, used with permission; *Bufo* s. s. by Kruczy89 CC BY-SA 3.0; *Bufotes* by Richard Bartz CC BY-SA 3.0; *Ansonia* by Thomas Brown CC BY-SA 2.0; *Duttaphrynus* by L. Shyamal CC BY-SA 2.5.

Nectophryne, Werneria and *Wolterstorffina. The African tree toads of the genus Nectophryne (the name ‘African tree toads’ is also used for species in the genus Nectophrynoides as well) are small, poorly known rainforest toads. Hardly any information is available on them bar the fact that they lay their eggs in little shallow pools, and that the males then guard the eggs. Molecular data suggests that Nectophryne forms a clade with the smalltongue toads (Werneria) and the Wolterstorff toads (Wolterstorffina) (Frost et al. 2006, Van Bocxlaer et al*. 2010, Pyron & Wiens 2011), all of which mostly occur in western Africa. If these small toads really do form a clade, they apparently represent an endemic African radiation, and the fact that they seem to have diverged ‘early on’ within Bufonidae lends additional support to the idea of a Gondwanan origin for toads as a whole.

**Caption:** Bate’s tree toad *Nectophryne batesii*, one of the two species in the genus. It has been reported across a relatively large part of the African tropics, from Nigeria and Cameroon in the west to Central African Republic in the east and Democratic Republic of the Congo and Gabon in the south. This distribution might say something about geological antiquity, buuut might not. Image: Bernard Dupont, CC BY-SA 4.0 (**original here**).

  **Caption:** Merten's smalltongue toad *Werneria mertensiana*. These are mid-sized anurans, with SVLs of 30-50 mm. They hide in leaf litter as well as being associated with streams. Drawing by Darren Naish, based on photo by Mary E. Gartshore.

The smalltongue toads or torrent toads (Werneria) of Gabon, Cameroon and Equatorial Guinea are small (30-40 mm SVL), nocturnal toads of fast-flowing streams. Six species are currently recognised, two of which – W. iboundji and W. submontana – were named in 2004 (Rödel et al. 2004). They’re highly dependent on running water and take refuge it in when disturbed. They eat beetles and have “a dorsolateral ridge that extends from the groin to the snout” (Graybeal & Cannatella 1995, p. 122), though this is very subtle. Parotoid glands and tympani are absent and the snout protrudes relative to the position of the mouth. Smalltongue toad tadpoles are specialised stream-dwelling animals. They’re dorsoventrally compressed with a broad head, dorsally positioned eyes and nostrils and sucker-like mouth disc.

Wolterstorff toads (also known as forest tree toads) were found by Graybeal & Cannatella (1995) to be paraphyletic with respect to a Nectophryne + Nimbaphrynoides clade, and none of the characters used to diagnose the genus appear unique. However, in at least some species the males have particularly large bony crests on their humeri, and it’s plausible that this is a synapomorphy at some level. In at least some Wolterstorff toad species (three are recognized), the tadpoles have a small ventral sucker.

**Caption:** *Werneria* tadpole in lateral view. The sucker-like mouth is fringed with soft, flap-like extensions. Illustration by Darren Naish, based on a drawing from Rödel *et al*. (2004). These tadpoles are around 17-24 mm long in total.

Four-digit toads. Didynamipus sjostedti, sometimes called the Four-digit toad, is a small bufonid restricted to Cameroon, Equatorial Guinea and Nigeria. It also lacks a tympanum and columella (what is it with all these toads losing their ears?) and have a reduced phalangeal and digital formula (as suggested by the name, only four toes are present). Two competing phylogenetic views have been proposed for this taxon. One idea – based on an analysis of morphological characters – is that it's part of an endemic African clade that also includes Nimbaphrynoides (the Nimba toads of Liberia, Ivory Coast and Guinea), Parker’s tree toad Laurentophryne parkeri and the Nectophryne African tree toads (Graybeal & Cannatella 1995). The other idea is that, together with the Nectophrynoides tree toads, it's close to the Asian stream toads Ansonia (Pyron & Wiens 2011). There are suggestions that this toad might exhibit direct development (that is, there’s no tadpole phase) but I don’t know if this has yet been confirmed, or otherwise.

**Caption:** Four-digit toad *Didynamipus sjostedti*, photographed in the wild by Václav Gvoždík. It’s tiny, females having an SVP of 19 mm, males 16 mm. Image CC BY-SA 2.5 (**original here**).

Having mentioned Parker’s tree toad: it also lacks a tympanum and columella, and in fact is like the four-digit toads in possessing only seven presacral vertebrae.

**Caption:** excellent illustration of Red-backed toad by Willems van der Merwe. *Schismaderma* toads is a bit different from the other toads discussed in this article: it’s relatively large size, granular dorsal skin texture, toxicity and large egg clutches make it part of the ‘range-expansion phenotype’ identified in toads by Van Bocxlaer *et al*. (2010) (for more on that concept, see the 2011 article **The toads series comes to SciAm: because Africa has toads too**). Image: Willems van der Merwe, used with permission.

The Red-backed toad… I mean, toads! Finally, for now, we look at the Red toad or Red-backed toad Schismaderma carens. This is a highly distinctive African toad, conventionally included in Bufo and regarded as representing the ‘Bufo carens group’. Its inclusion within Bufo sensu lato was always peculiar, as it lacks parotoid glands. The larvae are distinctive in possessing a U-shaped fold on the dorsal surface (Graybeal & Cannatella 1995) and adults are also easily identified thanks to their reddish backs (the underside is creamy or light grey), prominent lateral skin fold, and twin dark spots on the back.

Until very recently, Schismaderma was monotypic and thus ‘Red-backed toad’ was synonymous with S. carens. However, a second species was named in 2021: the Angolan red toad S. branchi, named for late herpetologist Bill Branch (1946-2018). It’s smaller than S. carens, with less prominent, less numerous palmar tubercles and bolder ventral patterning. Some individuals are fully as red as some individuals of S. carens, but S. branchi always lacks the prominent dark spots on the back and the black lateral stripes typical of the former species (Baptista et al. 2021).

**Caption:** Red-backed toad tadpole in lateral view. These tadpoles are relatively large, reaching 35 mm. They’re social and move about in tightly packed ball-shaped shoals. Illustration by Darren Naish, based on illustration in Channing (2001).

The affinities of Schismaderma have been contentious. Pramuk (2006) found it to belong to a clade that also included the African 20-chromosome toads (Amietophrynus) and to be the sister-taxon to the Asian ‘Bufo melanostictus group’ (named Duttaphrynus by Frost et al. (2006)). Frost et al. (2006) also recovered Schismaderma and Duttaphrynus as close relatives, but found Bufotes (Old World green toads) and Peltophryne to form a clade with Duttaphrynus and Schismaderma too. Van Bocxlaer et al. (2009, 2010) found Schismaderma to be the sister-taxon to a Churamiti + Nectophrynoides clade while Pyron & Wiens (2011) found it to be the sister-taxon to the Old World green toads.

**Caption:** holotype male specimen of *Schismaderma branchi* Baptista *et al*., 2021. The species is so far unique to the Luando River sub-basin, bordering the Cuanza-Congo watershed, of Angola, but there are suspicions that its distribution might be broader. Image: Baptista *et al*. (2021).

And with that... we’re done with another segment of the toad family tree. Well, I couldn't finish the section on the Ethiopian toads because I have yet to find any goddam pictures than I can use. On that note, more on toads at some point in the future, oh yes.

For previous articles in the Tet Zoo toads series see...

Toadtastic - the invasion begins!, October 2009
Bidder's organ and the holy quest for synapomorphies, October 2009
Our sex lives in words and pictures (or, On the reproductive biology of the Bufonidae), October 2009
Skulls, crests, snouts and giant poison glands: the heads of toads, October 2009
Toads of the world: first, (some) toads of the north, October 2009
The Natterjack, its life and times, October 2009
The resurrection of Anaxyrus, November 2009
South America, land of toads, part I: harlequins, redbellies and plump toads, January 2010
South America, land of toads part II: tree toads, Truebella, Frostius... oh, and did I mention the COMMUNAL NESTS?, August 2010
Blomberg’s toad and its omosternum-bearing buddies, August 2010
The toads series comes to SciAm: because Africa has toads too, September 2011
20-chromosome toads, September 2011

And for articles on other hyloid anurans, see…

Green-boned glass frogs, monkey frogs, toothless toads, November 2007
Glassfrogs: translucent skin, green bones, arm spines, January 2013
Everybody loves glassfrogs, February 2013
Gladiatorial glassfrogs, redux, January 2015
Frogs you may not have heard of: Brazil’s Cycloramphus ‘button frogs’, January 2015
It's the Helmeted water toad… this time, with information!, January 2015
The Terrible Leaf Walker Frog, March 2017
What, If Anything, Is A Strabomantid?, January 2024

Tetrapod Zoology is dependent on funds raised at patreon. Please help support this blog if you consider it worthwhile and want to see me continue doing it, thanks!

Refs - -

Baptista, N. L., Pinto, P. V., Keates, C., Edwards, S., Rödel, M.-O. & Conradie, W. 2021. A new species of red toad, Schismaderma Smith, 1849 (Anura: Bufonidae), from central Angola. Zootaxa 5081, 301-332.

Channing, A. 2001. Amphibians of Central and Southern Africa. Cornell University Press, Ithaca & London.

Graybeal, A. & Cannatella, D. C. 1995. A new taxon of Bufonidae from Peru, with descriptions of two new species and a review of the phylogenetic status of supraspecific bufonid taxa. Herpetologica 51, 105-131.

Pramuk, J. B. 2006. Phylogeny of South American Bufo (Anura: Bufonidae) inferred from combined evidence. Zoological Journal of the Linnean Society 146, 407-452.

Pyron, R. A. & Wiens, J. J. 2011. A large-scale phylogeny of Amphibia including over 2,800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution 61, 543-583.

Rödel, M.-O., Schmitz, A., Pauwels, O. S. G. & Böhme, W. 2004. Revision of the genus Werneria Poche, 1903, including the descriptions of two new species from Cameroon and Gabon (Amphibia: Anura: Bufonidae). Zootaxa 720, 1-28.

Van Bocxlaer, I., Loader, S. P., Roelants, K., Biju, S. D., Menegon, M. & Bossuyt, F. 2010. Gradual adaptation toward a range-expansion phenotype initiated the global radiation of toads. Science 327, 679-662.

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Welcome to 2024! And we kick things off with frogs. What, I hear you ask, is a strabomantid?

**Caption:** a hyloid montage. What is that small creature in the red circle? It’s a terraranan, and it’s here to shake things up…

Strabomantids are a group of terrestrial South and Central American frogs, sometimes termed terrestrial-breeding frogs, landfrogs or cloud forest landfrogs. Just about all are small (20-50 mm SVL), direct-developing animals associated with forests. They’re mostly animals of the forest floor, but some are arboreal. Some strabomantids (like the pristimantine Serranobatrachus) are cloud-forest animals that hide in leaf litter whereas others (like Yunganastes and Atopophrynus*) have some association with moss-covered rocks. The term ‘direct-developing’ refers to the reproductive strategy where no tadpole phase exists, and where fully-formed miniature froglets emerge directly from eggs. Oh, before I continue, let me say that I thought I’d written about the group on Tet Zoo before, but I can’t find anything in the archives, ho hum.

SVL = snout to vent length. Body length, basically.

Strabomantid anatomy. As is typical for animal groups whose members are mostly regarded as allies on the basis of molecular data (read on), it’s difficult at this point in history to point to anatomical traits that might be regarded as diagnostic for this group. We can at least make some generalisations about them though.

**Caption:** two representatives of Strabomantidae, demonstrating typical morphology for the group. At left, Two-lined robber frog *Bahius bilineatus*, a holoadenine that occurs across the Brazilian state of Bahia and is predicted to occur in Minas Gerais as well. At right, Savage’s goias frog *Barycholos ternetzi* of central Brazil. *Barycholos* is also a holoadenine. Images: Rafael O. Abreu, CC BY-SA 4.0 (**original here**); Lucas Grandinetti, CC BY-SA 2.5 (**original here**).

Slightly expanded digital pads are typical for strabomantids, and these are expanded into proper discs in arboreal species (including those included in Pristimantis and Strabomantis). These discs are supported internally by hook-like lateral processes on the terminal phalanges, and T-shaped and knob-bearing phalanges are also present in the group (Hedges et al. 2008). The fingers are unwebbed, and the toes usually are too, though the toes do sometimes have webbing at their bases. The fourth finger is reduced and even absent in some taxa. A single, medially located vocal sac is generally present but some (like Holoaden) lack it.

External ears are generally present in these frogs, but some (like the mostly Andean Bryophryne species and the possibly extinct Sonson frog Atopophrynus syntomopus of Colombia) lack them. Atopophrynus, incidentally, was originally described as a poison-dart frog (Lynch & Ruiz-Carranza 1982). I say that it’s possibly extinct because no specimens have been discovered since the type specimens were collected in 1981, and this is despite repeated searching at the type locality.

**Caption:** images of the Sonson frog *Atopophrynus syntomopus* from the original description. In the head, note the lack of features around the ear region. In the figures of the hands (A and B) and feet (C and D), note the strongly reduced first toe (C and D) and the T-shaped distal phalanges. This frog is tiny: the scale bars are 2 mm. Images: Lynch & Ruiz-Carranza (1982).

Pristimantis, final boss of tetrapod genera. Strabomantids might not be familiar, but they really should be, since one of the most noteworthy features of the group is that it includes the most speciose tetrapod genus of all time, this being Pristimantis, named by Spanish herpetologist Marcos Jiménez de la Espada back in 1870. As of the time of writing (January 2024), 602 species in this genus are recognized. As you might guess for such a speciose group, new species are named on a regular basis: 12 new Pristimantis species were named in 2023, for example. Pristimantis species tend to be large-eyed, slim-limbed little frogs marked with small spots or fine striping on their dorsal surfaces. There’s a fair amount of variation in snout shape and skin texture, some being smooth-skinned, others being bumpy or granular. Some are named rainfrogs (also written rain frogs), rubber frogs and robber frogs. These names are apparently all onomatopoeic references to their calls and nothing to do with anatomy or behaviour.

**Caption:** a very useful figure from Hedges *et al*. (2008), highlighting the continual rise over time in our knowledge of strabomantids and their kin, the terraranan frogs. This figure also makes the point that a small number of experts have driven this trend. It’s important to remember that this sort of thing (which is common across taxonomic groups) reflects actual discovery, not fashion or novel approaches to taxonomy (as is sometimes argued). Image: Hedges *et al*. (2008).

A good many Pristimantis species were originally included in Eleutherodactylus, a genus whose affinities actually lie elsewhere within Hyloidea. That is, it’s not a strabomantid. There always was a suspicion that the traditional version of Eleutherodactylus was what we in the trade term a taxonomic wastebasket – it was a dumping ground for small, anatomically nondescript Neotropical hyloids that couldn’t be tidily allocated to any of the familiar hyloid groups – and molecular data accrued during the 21st century has demonstrated that this is indeed the case (Frost et al. 2006, Hedges et al. 2008). There’s a lot to say on the dissolution of Eleutherodactylus and what’s happened to its constituent parts but this isn’t the place for that.

**Caption:** a montage of *Pristimantis* species. So pretty! The species shown here are (clockwise from upper left) *P. orcesi*, *P. erythros*, *P. loujosti* and *P. pycnodermis*. These species were figured together in the 2018 description of *P. erythros* of the Ecuadorian Andes, named therein the Blood rain frog! Image: **Sánchez-Nivicela *et al*. (2018)**.

Some select strabomantids with unusual names. Including the Sonson frog, 20 currently recognized genera are included within Strabomantidae. In addition to those I’ve already mentioned, it’s worth making comments on a few of the others… though not all of them, or I’ll never get this article finished. I’ve opted here to discuss those taxa whose names catch my eye.

**Caption:** representative strabomantids showing some of the variation in head size and shape. At left, *Noblella pygmaea*, a tiny Peruvian species named in 2009. At right, the Common big-headed frog *Oreobates quixensis*, a relatively large and short-snouted member of its genus. Images: Alessandro Catenazzi, CC BY-SA 2.5 (**original here**); Pavel Kirillov, CC BY-SA 2.0 (**original here**).

Euparkerella was named in 1959 in honour of British herpetologist H. W. Parker and contains small, narrow-headed, short-fingered frogs of the Brazilian Atlantic forest. Molecular studies indicate that substantial cryptic diversity exists within this genus and that much micro-endemism is present (Fusinatto et al. 2013), so more species are set to be named in coming years. Holoaden is another endemic Brazilian genus (named in 1920 by Brazilian herpetologist Alìpio de Miranda-Ribeiro), its species notable for their association with highland, mountainous environments and for a highly glandular dorsal surface.

**Caption:** *Euparkerella brasiliensis*, photographed in Rio de Janeiro State. Image: **Fusinatto *et al*. (2013)**, CC BY 4.0 (**original here**).

Niceforonia, a close relative of Holoaden according to molecular data (Padial et al. 2014), is named for Colombian herpetologist Nicéforo María and includes 15 species endemic to northern South America. This is one of those taxa where the terminal phalanges (the bones at the tips of the digits) are T-shaped. Lynchius is another relative of Holoaden (according to some studies; see Pyron & Wiens 2011), and again its name commemorates a herpetologist noted for his work on tropical American amphibians, this time John D. Lynch. At least 11 frog and salamander taxa have been named in Lynch’s honour so far.

Finally, Qosqophryne was named in 2020 for species previously included in Bryophryne and is endemic to the Department of Cusco in Peru. Its generic name uses the Quechua spelling – Qosqo – of Cusco (combined with phryne, Greek for ‘frog’) (Catenazzi et al. 2020).

Where do strabomantids fit in the anuran family tree? There’s no doubt that strabomantids are hyloids: that is, part of the major clade that includes glassfrogs, treefrogs, true toads, poison-dart frogs and many others. Within that clade, molecular studies indicate that strabomantids are part of the clade that also includes the craugastorid fleshbelly frogs and kin, the eleutherodactylid New World rain frogs and the sometimes toxic brachycephalid toadlets and robber frogs. This massive clade – containing over 1000 species – is termed either Terrarana or Brachycephaloidea and a substantial amount of work has been published on its phylogeny, taxonomy and diversity since about 2008 (e.g., Hedges et al. 2008, Pyron & Wiens 2011, Canedo & Haddad 2012, Padial et al. 2014, Heinicke et al. 2018, Motta et al. 2021).

**Caption:** I’ve published hyloid cladograms several times here at Tetrapod Zoology, and this is the most recent version, dating to 2017 and produced to accompany **an article on poison-dart frogs**. It depicts a sort of consensus based on the cladograms of Frost *et al*. (2006), **Grant *et al*. (2006)** and Pyron & Wiens (2011), with the taxonomy proposed by Frost *et al*. (2006) superimposed on to the branches. Note that ceratophryids (horned frogs) are depicted as being close to bufonids (true toads) and dendrobatids (poison-dart frogs). And where are terraranans? I excluded them entirely. Image: Darren Naish, **images produced for the textbook**.

  **Caption:** several new hyloid cladograms have been published since I produced the version shown above. They differ sufficiently in details that it’s hard to produce a consensus: this is a simplified version of the consensus **featured at AmphibiaWeb**. Note the unresolved polytomy that includes terraranans and kin. The topology shown here differs sufficiently from that of Frost *et al*. (2006) that the names they proposed (including Leptodactyliformes and Agastorophrynia) have, potentially, very different memberships from the ones they preferred, and for that reason I haven’t bothered applying them here. Image: Darren Naish, **images produced for the textbook**.

The taxonomic history of terraranans is hopelessly complicated and the hypothesis that they form a clade composed of multiple ‘family’-level groups is recent, effectively starting with a study published by S. Blair Hedges and colleagues in 2008. That 2008 study is also the one in which the group name Strabomantidae was first coined. Hedges et al. (2008) didn’t provide anything like a phylogenetic definition for Strabomantidae but did list 49 character states that they used to help define this group. Many of these are references to the absence of unusual features present in other terraranan groups (like the vertebral shields and fusion between the skull bones and overlying skin present in some brachycephalids), and working out which are truly diagnostic for strabomantids specifically would be a lot of work.

**Caption:** brachycephalid frogs – sometimes called three-toed toadlets – have been mentioned a few times in this article, and this is what they look like. Fairly hilarious. Also quite extreme in anatomical terms. The species shown here are (left) *Brachycephalus izecksohni* and *B. olivaceus*, both from Brazil and both named anew in 2017. Images: **Ribeiro *et al*. (2017)**, CC BY-SA 4.0.

Within Terrarana/Brachycephaloidea, some variation in results means that authors have disagreed on which taxonomy best reflects phylogeny. Things will become needlessly complicated if I recount all the to and fro that’s happened in the literature, but I’ll make my life easier by saying that I’m following those authors (e.g., Hedges et al. 2008, Blackburn & Wake 2011, Heinicke et al. 2018, Jetz & Pyron 2018, AmphibiaWeb 2024) who find a topology where Strabomantidae and the closely related Craugastoridae are separate groups. The key problem is that Strabomantis itself hops around, ha ha, within phylogeny, such that it (and kin) are sometimes outside of a clade that includes Craugastor and kin, and are sometimes within it. Those competing results lead to different taxonomic outcomes (Blackburn & Wake 2011, Pyron & Wiens 2011, Motta et al*. 2021).

Yeah, it’s irritating that both names are in current use, and both have effectively the same meaning. I can see it being useful that one is defined as more inclusive than the other, but I don’t think that anuran workers have done anything like this yet. An additional complication is that some authors have opted to go with the emended spelling Terraranae because this is, apparently, etymologically more correct. The latter point was made by Dubois (2009), and later by Duellman et al. (2016) in their naming of Arboranae, a hyloid clade that includes hylid treefrogs and kin.

Caption: several different topologies have been recovered for Terrarana, though the majority find eleutherodactylids and brachycephalids to be outside a craugastorid + strabomantid clade, with brachycephalids closest. This topology is from the graphical abstract of Heinicke et al. (2018).

The diversity within Strabomantidae has resulted in the naming of four subfamily-level divisions within the group: Strabomantinae, Pristimantinae, Holoadeninae and Hypodactylinae. Which of those is actually worth recognizing in view of the group’s phylogenetic structure obviously depends on which study you consult. Again, this article is not the place for a discussion of that complexity.

And that’s where we’ll end things. As is hopefully clear at this point, the name Strabomantidae is mostly (albeit not universally) applied to a clade of tropical American terraranan or brachycephaloid hyloids that are allied to craugastorids. An alternative view posits that the group’s constituent taxa are better placed within Craugastoridae, in which case a strabomantid clade might still exist but only as a ‘subfamily’. Either way, the lineages concerned are firmly placed within Terrarana/Terraranae/Brachycephaloidea.

It’s also important to note that many frogs relevant to this discussion are endangered or potentially extinct, since they’ve proved hard or even impossible to find since being first named. Degradation and loss of habitat, the impact of fungal infection, and climate change are all connected to strabomantid decline. Also worth noting is that quite a few species do not have agreed-upon conservation status, since data is currently deficient.

**Caption:** all frogs are great, but hyloids include some of my favourite groups. Clockwise from upper left: the poison-dart frog *Phyllobates*, the marsupial treefrog *Hemiphractus*, the white-lipped frog *Leptodactylus*, and the helmeted water frog *Calyptocephalella*! Check the links below for articles on most of these animals. Images (clockwise from upper left): Darren Naish; Santiago Ron, CC BY-ND 2.0 (**original here**); Darren Naish; José Grau de Puerto Montt, CC BY-SA 3.0 (**original here**).

I like all the anurans, but hyloids are among my favourites and I’ve covered them quite a few times on Tet Zoo before. For previous articles, see…

Green-boned glass frogs, monkey frogs, toothless toads, November 2007
The toads series comes to SciAm: because Africa has toads too, September 2011
20-chromosome toads, September 2011
Glassfrogs: translucent skin, green bones, arm spines, January 2013
Everybody loves glassfrogs, February 2013
African tree toads, smalltongue toads, four-digit toads, red-backed toads: yes, a whole load of obscure African toads, December 2014
Gladiatorial glassfrogs, redux, January 2015
Frogs you may not have heard of: Brazil’s Cycloramphus ‘button frogs’, January 2015
It's the Helmeted water toad… this time, with information!, January 2015
The Terrible Leaf Walker Frog, March 2017

Tetrapod Zoology is dependent on funds raised at patreon. Please help support this blog if you consider it worthwhile and want to see me continue doing it, thanks!

Refs - -

AmphibiaWeb: Information on amphibian biology and conservation. [web application]. 2024. Berkeley, California: AmphibiaWeb. Available: https://amphibiaweb.org/. (Accessed: 2024).

Blackburn, D. C. & Wake, D. B. 2011. Class Amphibia Gray, 1825. In Zhang, Z.-Q. (ed) Animal biodiversity: an outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148, 39-55.

Canedo, C. & Haddad, C. F. B. 2012. Phylogenetic relationships within anuran clade Terrarana, with emphasis on the placement of Brazilian Atlantic rainforest frogs genus Ischnocnema (Anura: Brachycephalidae). Molecular Phylogenetics and Evolution 65, 610-620.

Catenazzi, A., Mamani, L., Lehr, E. & von May, R. 2020. A new genus of terrestrial-breeding frogs (Holoadeninae, Strabomantidae, Terrarana) from southern Peru. Diversity 12, 184.

Dubois, A. 2009. Miscellanea nomenclatorica batrachologica. Class-series nomina are nouns in the nominative plural: Terrarana Hedges, Duellman & Heinicke, 2008 must be emended. Alytes 26, 165-175.

Duellman, W., Marion, A. B. & Hedges, S. B. 2016. Phylogenetics, classification, and biogeography of the treefrogs (Amphibia: Anura: Arboranae). Zootaxa 4104, 1-109.

Fusinatto, L. A., Alexandrino, J., Haddad, C. F. B., Brunes, T. O., Rocha, C. F. D. & Sequeira, F. 2013. Cryptic genetic diversity is paramount in small-bodied amphibians of the genus Euparkerella (Anura: Craugastoridae) endemic to the Brazilian Atlantic Forest. PLoS ONE 8 (11), e79504.

Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F. B., Kok, P. J. R., Means, D. B., Noonan, B. P., Schargel, W. E. & Wheeler, W. C. 2006. Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History 299, 1-262.

Hedges, S. B., Duellman, W. E. & Heinicke, M. P. 2008. New World direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa 1737, 1-182.

Heinicke, M. P., Lemmon, A. R., Lemmon, E. M., McGrath, K., & Hedges, S. B. 2018. Phylogenomic support for evolutionary relationships of New World direct-developing frogs (Anura: Terraranae). Molecular Phylogenetics and Evolution 118, 145-155.

Jetz, W. & Pyron, R. A. 2018. The interplay of past diversification and evolutionary isolation with present imperilment across the amphibian tree of life. Nature Ecology and Evolution 2, 850-858.

Lynch, J. D. & Ruiz-Carranza, P. M. 1982. A new genus and species of poison-dart frog (Amphibia: Dendrobatidae) from the Andes of northern Colombia. Proceedings of the Biological Society of Washington 95, 557-562.

Motta, A. P., Taucce, P. P. G., Haddad, C. F. B. & Canedo, C. 2021. A new terraranan genus from the Brazilian Atlantic Forest with comments on the systematics of Brachycephaloidea (Amphibia: Anura). Journal of Zoological Systematics and Evolutionary Research 59, 663-679.

Padial, J. M., Grant, T. & Frost, D. R. 2014. Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria. Zootaxa 3825, 1-132.

Sánchez-Nivicela, J. C., Celi-Piedra, E., Posse-Sarmiento, V., Urgiles, V. L., Yánez-Muñoz, M. & Cisneros-Heredia, D. F. 2018. A new species of Pristimantis (Anura, Craugastoridae) from the Cajas Massif, southern Ecuador. ZooKeys 751, 113-128.

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The tenth TetZooCon – that’s the Tetrapod Zoology Convention – happened between Friday December 1st and Monday December 4th, and I think I’ve just about recovered. Hosted once again at Bush House, King’s College London (KCL), it was the biggest, busiest and most successful of our events so far, as is fitting for the tenth one…

**Caption:** the TetZooCon 2023 banner, composed of Mesozoic marine reptiles but a good number of additional animals too. Image: George Lathouris, used with permission.

Getting Bush House as a venue is big deal and we owe massive thanks to Chris Manias of KCL for helping to set it all up. It means that we have access to a massive auditorium, two or three upstairs rooms for workshops and breakout events, and a large foyer where we have stalls, a reception area that can be used for drinks, and an exhibition space. Chris’s involvement also means that we’re running in co-operation with the Popularizing Palaeontology network that he organises, and a consequence is that we now host a greater diversity of events than we might without him. This year, these included the panel event ‘Engaging with Extinctions – Past, Present and Future’ on the evening of the Friday, and a lunchtime screening of 1925’s The Lost World on the Sunday, introduced by Dave Hone.

**Caption:** TetZooCon 2023 merch! **Buy it here.**

Before I launch into more specific thoughts, let me say a massive thank you to our speakers, participants, vendors and assistants, without which none of this would happen. Thanks also to those who took photos and shared them, or allowed use of their photos here. Thanks in particular to professional photographer Alfred Barwick.

**Caption:** a montage of animals… all archosaurs… specially relevant to TetZooCon 2023. Left to right: Northern cassowary *Casuarius unappendiculatus*, American crow *Corvus brachyrhynchos*, the recently extinct Madagascan croc *Voay robustus* (art by Alex Lovegrove), and an assortment of captive Nile crocodiles *Crocodylus niloticus* (I think with a single Sacred crocodile *C. suchus* in there somewhere). Images: Darren Naish; (c) Alex Lovegrove.

The extinction panel event. And thus it was that, after spending the better part of Friday at the Natural History Museum with my daughter Hel and my friend Aidan Williams Dale (visiting from Canada, specifically for TetZooCon), we made the trek to Bush House. There were drinks, there was the setting up of stalls and such ahead of Saturday, and then there was the panel event.

**Caption:** a short pre-TetZooCon trip to the Natural History Museum involved us looking at Hope the Blue whale (at left) and posing with the Darwin statue. Darren Naish at left, Aidan Williams Dale at right.

  **Caption:** some Natural History Museum trip highlights. Clockwise from upper left: giant salamander model and skeleton, Giant armadillo *Priodontes maximus* taxiderm specimen, *T. rex* animatronic in festive attire, Black caiman *Melanosuchus niger* in the Reptiles and Amphibians gallery. Images: Darren Naish.

The panelists (Aidan and myself, memory studies researcher Clara De Massol De Rebetz and artist and cultural geographer Amy Cutler) introduced ourselves before discussing the various extinction-related points introduced by Chris. I used my introduction to cover two of my favourite points relating to extinction: (1) that all extinctions start out as local extinctions, and that the tree removal, plastic pollution, degradation of local green space via over-use from people and their pet dogs, roadkill of animals large and small and so on is all part of the bigger, global problem. Extinction is not some remote thing happening only in Amazonia, Indonesia or Madagascar. (2) Long-extinct animals – looking mostly at you, non-bird Dinosauria – are essential tools for science literacy, and we shouldn’t be ashamed to use them in getting people interested in topics like ecology, evolution and extinction. Readers with good memories might recall that that second point was the main focus of my United National Science Summit presentation of September 2022.

**Caption:** I think the contents of this slide speak for themselves. All about me I see plastic pollution, road-killed animals large and small, and the constant removal of trees. Images: Darren Naish.

In the discussion that followed our opening pitches, we covered various points relating to how extinction is reflected in our work, projects and outlook. I’m strangely proud of the fact that I managed to incorporate a quote by the great Stewart Lee, make of that what you will. Amy Cutler turned up very late – I don’t want to embarrass her too much by saying why (though I will say that she went to the wrong venue) – but she delivered, her time culminating with the showing of a short film generated by an AI.

**Caption:** another scene from the Friday evening extinction panel, here showing Amy Cutler talking about visual representations of dead animals in film and how this relates to our understanding of extinction. Image: George Lathouris, used with permission.

At this point I have to mention the fact that the main auditorium wasn’t functioning properly. Last year, this same room overheated when kept in use for the better part of the day, and our timetable for 2023 was built around us leaving it empty for hours at a time in order to counter this. But fate is a cruel mistress, and this year a very opposite problem existed. The main auditorium was perpetually freezing cold, with no heating and cold air blasting in from the ventilation. I went on stage without a coat and succeeded, via violent thermogenesis, in burning off a fair amount of the spare calories I carry around.

We did what we could to get this fixed – numerous visits were made to KCL staff and numerous phonecalls happened – but all to no avail. A similar problem affected the foyer. This situation was such a farce that few people in attendance will forget it. But I hope that the quality and number of our talks and events and the good social vibe means that they might.

A marine reptile symposium. TetZooCon 2023 was partly based around Mesozoic marine reptiles, and this is because 2023 was the year in which I published Ancient Sea Reptiles (Naish 2022) [UPDATE: actually, the book saw print in 2022. But 2023 was the year in which it went on sale in shops and other outlets]. I spoke about fossil marine reptiles in general and was joined in the same session by Judyth Sassoon on plesiosaur semiotics (semiotics = nothing to do with partly formed ear regions but instead the topic of how meaning is conveyed via imagery) and Emily Swaby on temnodontosaurs, one of my favourite ichthyosaur groups.

**Caption:** Judyth Sassoon in the marine reptiles session, her opening slide visible in the background. Image: Alfred Barwick, used with permission.

Later in the day, Dean Lomax discussed the story behind the Rutland Sea Dragon, a spectacular British ichthyosaur – a temnodontosaur no less – still under study (one paper on the specimen has recently appeared: Larkin et al. 2023). Dean has also been involved in the recently completed movie Why Dinosaurs? which just had its Hollywood premiere, so we were also treated to a showing of the trailer. This was followed by a marine reptile panel event in which I was joined on stage by Emily, Dean and Luke Muscutt (the other marine reptile speakers had other engagements). Topics discussed included our inspirations and the people who’d aided and assisted us early on (I slipped a tenner to Luke, who then explained how his primary scientific role model was that Darren Naish guy), our thoughts on what other researchers might have gotten wrong, and what we’re doing next on the subject.

**Caption:** Dean Lomax talks at TetZooCon 2023 on the Rutland ichthyosaur, quite fairly bigged up as the most spectacular ichthyosaur specimen yet discovered in the UK. Infamously dubbed the ‘sea dragon dinosaur dolphin’ by a British journalist, it represents an animal that would have been around 10 m long. Image: Darren Naish.

  **Caption:** the marine reptile discussion panel, showing (l to r) Darren Naish, Luke Muscutt, Emily Swaby and Dean Lomax. Yes, I appreciate the irony of the backdrop being some fish-themed montages. Image: Alfred Barwick, used with permission.

A planned proper book signing for Ancient Sea Reptiles never happened as I couldn’t make time for it, but I sold my entire stock anyway and now need more. I also sold all the copies I brought of Dinosaurs: How They Lived and Evolved (the new third edition) and nearly all my stock of Dinopedia too.

In a separate marine reptile session on Sunday, Luke spoke about Flip, the world’s first realistic plesiosaur robot. Flip was in attendance and those present were allowed to see it in action (and even, in special cases, operate it themselves). This event was filmed and some of this material will appear in a documentary. UPDATE: part of the TetZooCon 2023 outing of Flip has been released on YouTube.

**Caption:** Luke Muscutt (standing at left) discusses his work on Flip, the robot plesiosaur that had its first public outing at TetZooCon. Flip stars in the BBC documentary *Attenborough and the Giant Sea Monster*, to be screened on BBC One on New Years’ Day 2024. So does Luke! Image: Alfred Barwick, used with permission.

In the same session, Richard Forrest spoke about his experiences with the excavation of plesiosaur fossils. I missed both of these talks due to the modern archosaur session happening at the same time. Ah yes, that…

The modern archosaur session. Said aforementioned modern archosaur session involved Jennifer Campbell-Smith on corvids and fieldwork on them, Todd Green on cassowaries, a cassowary discussion panel, and Evon Hekkala on crocodiles. Quite the lineup. But while we were setting things up the entire electrical box at the lectern – supplying power to the computers, microphones and everything else there – died. We did everything we could to see if this could be fixed (it was most likely a blown fuse) and sought help from staff… but this too was all for nothing. And thus we had to resort to jumping from room to room, which was partly a relief given the low temperatures caused by the heating/ventilation problem. I won’t explain the fully sorry saga as it’s long-winded and embarrassing, but it was a total shambles and we ended up moving three times. Having said all that, the talks still happened, and they were great.

**Caption:** Todd Green talks cassowaries. I worked with Todd and his team in Florida to relocate several adult cassowaries in early 2023: **here’s the Tetrapod Zoology article on that event**. To learn more about the cassowary-themed movie Todd and colleagues are currently putting together, **go here**. Image: Georgia Witton-Maclean, used with permission.

Jenn’s corvid talk was a highlight for many people. She discussed crow diversity in general, and her own fieldwork on American crows and how they transfer and learn information. Jenn is an amazing artist and (like so many who attend TetZooCon) an outstanding natural historian, and it was an honour to have her join us. Next up was Todd who gave a rundown on cassowary research and where we’re at with respect to our knowledge today. Todd and colleagues (namely Kerrie and AJ Dodd, both present at TetZooCon 2023 as well) are making a film (you can help support it here) and his talk was followed by a panel event in which I, Todd and AJ – chaired by Jenn – discussed cassowaries some more. Todd, AJ and Kerrie also had a stall where they sold and auctioned cassowary-themed objects including a model head and 3D-printed foot.

**Caption:** Jennifer Campbell-Smith talks corvids. I’m going to steal some of Jenn’s own words on TetZooCon 2023: “It was an utterly delightful mix of professional and enthusiast zoology nerds who could all come together to discuss and enjoy our passions in a welcoming, safe, fun environment. The sincere interest in my talk and both the compliments and excellent conversation I received afterward were humbling, heart-warming, and genuinely had a deep impact on me”. Image: Todd Green.

The final talk of the session was Evon Hekkala’s on ‘Maneaters, Mummies and Madagascar; Sacred and Secret Tales from Cryptic Crocodiles’. This was another highlight; it covered Evon’s research on the crocodiles of both mainland Africa and Madagascar, and discussed this work within the broader context of her research and career.

**Caption:** Evon Hekkala on her crocodile research. This has involved the resurrection of *Crocodylus suchus* as distinct from *C. niloticus*, the recovery of new specimens of the recently extinct *Voay* and its phylogenetic position, and more. An amazing talk. Image: Georgia Witton-Maclean, used with permission.

Additional talks that weren't part of these ‘themed’ sessions happened too. Herpetologist Steve Allain spoke about his research on the non-native midwife toads living in England (e.g., Allain et al. 2019a, b, Allain & Shimbov 2021), Hanna Ayoob spoke about her work in running zoology-themed outreach and scicomm events, and Amber Coste discussed her studies of extinct dolphins from New Zealand, notable for their remarkably odd and sometimes tusk-like protruding teeth (Coste et al. 2023a, b). Amber was a PhD student of fossil marine mammal specialist Ewan Fordyce who, sadly, died this year, so it’s fitting that her talk was dedicated to him.

**Caption:** Steve Allain (at right) is introduced by James Pascoe. One of several talks I really wanted to see but couldn’t get to, alas. Image: Georgia Witton-Maclean, used with permission.

  **Caption:** Amber Coste talks weird fossil dolphins, but at this point of the talk is discussing the contributions of her PhD supervisor, the late Ewan Fordyce. Image: Alfred Barwick, used with permission.

  **Caption:** more Amber. Her talk featured a good amount of interesting, and sometimes necessarily speculative, palaeoart. Image: Georgia Witton-Maclean, used with permission.

An eternal winter of palaeoart. Palaeoart is a mainstay of TetZooCon and we’ve always had a palaeoart workshop of some sort, usually accompanied by one or more exhibitions. This year we decided to ramp things up and basically have palaeoart events running for most of the wekend. John Conway engaged Luis Rey in a discussion on Luis and his work, and Joschua Knüppe led a show-and-tell (whereby numerous artists spoke about their projects).

**Caption:** Luis Rey (at left) discusses his palaeoart work with John Conway. Those numerous small boxes in the background contain small, pliosaur-shaped objects used in a workshop event that occurred later on the same day. Image: Alfred Barwick, used with permission.

  **Caption:** Joschua Knüppe explains the background to his palaeo-map project, here showing efforts that have gone beforehand (specifically, maps produced by Ray Troll). Image: George Lathouris, used with permission.

James Pascoe led a paint-a-pliosaur event in which a pre-booked number of people decorated a TetZooCon exclusive pliosaur model. I wasn’t able to attend any of this due to my presence in other sessions and was especially worried about not getting to claim ownership of a patented Pascoe pliosaur. But I needn’t have worried, because….

**Caption:** a very large Pascoe pliosaur (it’s 46 cm long) has joined the Tet Zoo Towers collection, and I am pleased. Image: Darren Naish.

James also brought along several wonderful dioramas depicting Mesozoic dinosaurs in miniature landscapes, though quite how he transported these without them getting smashed to bits still amazes me. Also within the palaeoart event, I led a Prehistoric Planet discussion, which again was only available to a limited number of attendees due to room size.

**Caption:** some of (or parts of some of) the very impressive Jurassic and Cretaceous dioramas created by James Pascoe, available on show at TetZooCon 2023. James notes Paul Glynn as among his inspirations, and Paul has showcased his own dioramas at TetZooCon in the past. Image: George Lathouris, used with permission.

Sales and stalls. The main foyer of the venue was fully occupied by stalls and tables. We’ve been hosting more and more of these year on year as TetZooCon has grown, and this year were at maximum capacity, with as many tables as we could get. On that note, I never had time to look properly at all the stalls, let alone buy anything from them. Everything was amazing and exciting, from Rebecca Groom’s palaeoplushies to Steve White’s art prints, to the range of models and figures made by Sam St Leger, Paul Glynn and others, and to the stickers, cards and more offered by others. I sold books and various animal figures, as well as a range of TetZooCon merchandise, including cups, t-shirts and tote bags (all of which, and more, are available here). Dougal Dixon was in attendance again and also shifted a good number of books.

**Caption:** a palaeoplushie-themed table, hosted by Leo Chilvers (at left) and Rebecca Groom. Note the size of some of the pieces for sale here (visible at far right). Also worth looking for is the *Prehistoric Planet*-style *Mononykus* (sitting in the cardboard popcorn box) next to Leo. Image: Georgia Witton-Maclean, used with permission.

  **Caption:** I’ve mentioned a few times here the range of *All Yesterdays* figures produced by Sam St Leger of **Splendid Editions**. See below for close-ups of the figures Sam has produced so far (more are on the way). When taking this photo I inadvertently photographed Natalia Jagielska presenting her super-floofy *Sinosauropteryx* plushie as if it might be the second coming of Christ. Image: Darren Naish.

  **Caption:** more detailed views of Sam St Leger’s *All Yesterday*’s figures, with other figures visible too (including *Moschops* and a Tarasque). These are available from the **Splendid Edition shop on Etsy**. Image: Darren Naish.

Several things were on sale at TetZooCon 2023 as exclusives, or for the first time ever. Perhaps most notable among these were the scale Velociraptor statues made by Jed Taylor and Ruadhrí Brennan. These are models of superb and frightening craftmanship, and – as is appropriate – they aren’t cheap. In view of this, I had a mild concern that they might not sell. They sold out within the first 20 minutes of the Saturday.

**Caption:** scenes from the Jed Taylor and Ruadhrí Brennan stall… argh, such treasures! I sincerely planned to at least buy some fridge magnets (those numerous skull models on the dark plate) but didn’t even get round to that. Hopefully Jed and Ru will be selling wares at future events, and if that’s not a draw I don’t know what is. Images: Ruadhrí Brennan.

  **Caption:** each of those little orange slips next to a Taylor-Brennan *Velociraptor* figure says SOLD. In the end, even the giant one at far left was purchased as well. I planned to get the whole set for my own collection but…. yeah, that didn’t exactly work out as planned. Image: Jed Taylor.

As might be obvious to those who were there, let me say again that we’re basically at maximum capacity in terms of what can fit in the foyer. That’s a bit of a problem, since our plan is to expand and host more stalls. I’m not yet sure how we’re going to get round that for 2024 and beyond – stay tuned.

**Caption:** one of so many excellent art-themed stalls we had at TetZooCon 2023. This is Lee Brown’s; Lee runs DailyDinoSketch and sells art, stickers, rock art and more **here at Etsy**. Image: George Lathouris, used with permission.

  **Caption:** another stall, this time run by Emily Higgs. Stickers, prints and tons more! The **Emily Higgs Shop is here on Redbubble**. Image: Georgia Witton-Maclean, used with permission.

  **Caption:** I had to deliberately avoid looking at this stall for too long or my finances would look bleaker than they already do. This is Paul Glynn’s range of figures and models, and oh my god. I love those Wealden dinosaur models, but the sharks and pike very much have my full attention as well. Paul’s work is available on **Etsy at TheYoungerEarth**. Image: George Lathouris, used with permission.

Cosplay! If you remember the article on TetZooCon 2022, you might recall the mention of the impromptu cosplay event that happened there, entirely thanks to the good work of palaeontology Phd student Meghan Jenkinson. I’ve been thinking for years of hosting cosplay at TetZooCon. After all, we’re always joined by an incredibly talented, creative number of individuals whose skills and interests are very much relevant to cosplay. And thus TetZooCon 2023 hosted a cosplay competition, led by Meghan and Hel Naish.

**Caption:** the cosplay event, with Meghan (in hazmat suit) and Hel (with giant tail) present at far left. Image: Alfred Barwick, used with permission.

A good number of attendees made an effort, the three winners being Connor Temple from Primeval (played by Armin Reindl), Mary Anning (played by Mélissa Delteil) and Nigel Marven (played by James Appleby). Prizes included those kindly provided by our friends at Everything Dinosaur and Palaeoplushies.

The Marven Event, and the quiz. I’ve been trying for a while to book film-maker, writer, TV presenter and natural historian Nigel Marven for TetZooCon, and I’m pleased to say that Nigel’s busy schedule finally allowed this to be fulfilled this year. The broken nature of the main auditorium meant that we had to run his talk in an upstairs room not nearly as big, so it was totally overcrowded. Nigel spoke about his involvement in those TV shows depicting prehistoric animals – Prehistoric Park, Primeval and the Walking With specials – as well as his various adventures with living reptiles, his recent film-making projects in Slovakia and elsewhere, and more. He was good enough to hang around for signings, photos with the fans and so on and here’s where it became obvious that a good number of our attendees – I believe they call themselves the palaeostream team – were all wearing specially made Prehistoric Park t-shirts.

**Caption:** we were very pleased to have Nigel Marven speaking at TetZooCon 2023, and here he is with his cover slide in the background. Image: Alfred Barwick, used with permission.

  **Caption:** a packed-out lecture room during Nigel’s talk (too packed-out; but we were meant to be in the giant main auditorium, alas). Image: Alfred Barwick, used with permission.

  **Caption:** Nigel Marven (in the middle) surrounded by a happy throng of Palaeostream friends, plus various Mesozoic-themed props. Image: Alfred Barwick, used with permission.

Occurring in parallel to the Marven Event – apologies to those who wanted to do both – was our quiz. Richard Hing was the winner this year, with Kelvin Britton and Albert Chen being runner-ups. Prizes were kind donations from various of our vendors and also Everything Dinosaur: thank you all so very, very much. And thus things drew to a close…

Fieldtrip! Except they didn’t, because a good number of us – around 40 – went to London Zoo on the Monday. Despite the potential for bad weather, it turned out all right and was a success. We looked at interesting aspects of the zoo’s historical architecture – I always enjoy telling people about the stories behind the Elephant and Rhino House (now the Casson Pavilion), the Goat Hills of the Mappin Terraces and more – and we were lucky with the animals.

**Caption:** the TetZooCon 2023 fieldtrip contingent in the walk-through aviary. Numerous Scarlet ibis *Eudocimus ruber* are visible, as is a lone Abdim’s stork *Ciconia abdimii* on the roof at upper right. As discussed below, we’ll be making the fieldtrips (which have been a constant TetZooCon feature since the first one) a more integrated thing from 2024. Image: Darren Naish.

Highlights included Western gorillas Gorilla gorilla having sex, a vicious attack on my good self by a juvenile Scarlet ibis, a very close-up view of one of the Linne’s two-toed sloths Choloepus didactyla, eye contact with a Chinese muntjac Muntiacus reevesi, and brief viewings of an aye-aye Daubentonia madagascariensis. We also had excellent views of two young Sumatran tigers Panthera tigris sondaica playing and even carrying sticks and small logs around.

**Caption:** a London Zoo montage, showing (clockwise from upper left) co-operative Chinese muntjac, a slightly blurry Mohol bushbaby *Galago moholi*, and a detail of the stonework around the entrance of the (now closed) Reptile House, showing a parent Shingleback skink *Tiliqua rugosa* with its babies (these lizards have extended parental bonds, and it’s interesting to see that this was known during the 1920s, the time when the Reptile House was built). Images: Darren Naish.

I’ll finish here by saying that the ad hoc, impromptu nature of the trip meant that we weren’t able to take advantage of anything intelligent like a group discount. That’s changing from 2024, on which read on.

To the future. TetZooCon 2023 was hard work. I, personally, have never been so rushed and overworked at any event, and it was bad enough that I was often unable to get time to eat or even drink. We were also let down by a cascade of failures at the venue that were beyond the control of any of us running TetZooCon, all of which makes it clear how bad universities are at catering to external events outside of normal hours. Nevertheless, it was enough of a success that we’re already thinking about plans for 2024. I can say to begin with that we will be in the same venue, will almost certainly be running things across a Friday evening and a weekend, and will again be hosting a fieldtrip of some sort (hold that thought).

**Caption:** I think it can be agreed that we have a pretty impressive selection of stalls at TetZooCon 2023. We only hope that this will continue into the future. The montage here shows (clockwise from upper left) images from the **Neo Jurassica** stall (the skeleton is that of the Brazilian noasaurid theropod *Berthasaura*); prints from **Mark Witton**; and replica skulls and models manufactured by **Paul Glynn**. Images: George Lathouris, used with permission.

A few things have to change though. We’ll be undergoing an upscaling of sorts, perhaps with a dedicated committee that betters handles and manages the event’s different components. We also have to become more expensive. I hope that people appreciate how cheap TetZooCon is compared to similar events (conventions and conferences), and the fact is that we’re not generating enough income to cover our outgoings. We’re currently considering making TetZooCon something bigger than a weekend-long event, on which stay tuned, and are also looking into turning the fieldtrip into a larger and more formalized thing. I’ve worked in the past as a tour guide and, yes, am thinking quite seriously about getting coach trips and maybe even overnight stays part of our remit. None of this is arranged yet but at least now you’ve been warned.

**Caption:** going to London Zoo and looking at Bactrian camels and excellent artwork is all well and good, but what about going further afield and looking at stuff way beyond the outskirts of the nation’s capital? *We shall see.* Images: Darren Naish.

At least part of TetZooCon 2024 will be devoted to cryptozoology, mostly because I hope to have another book out and published by then… yikes, no pressure. Other stuff will be happening as well though, and we’re also looking at booking special speakers on other subjects. For news watch the TetZooCon page at Facebook and #TetZooCon on Twitter/X and elsewhere.

It feels to me like we’ve reached another milestone in the history of this event, this tenth one being appropriately massive and well-attended. Indeed, the continuing success of TetZooCon demands that we have to change and upscale once again. It only remains for me to say a massive thanks to everyone who attended and made it what it was (big shout in particular to those who joined us from overseas, it was such a thrill to see you), to those who worked with me in running the whole thing, and to our speakers, stall holders and event-runners. See you again, I hope, in 2024.

**Caption:** the primary TetZooCon organizers, Darren Naish (left) and John Conway, with a Rebor American alligator in the middle. I really should have taken a holiday right away. But I couldn’t. Image: Hel Naish.

Other articles on TetZooCon 2023 will be listed here as and when they’re published. Steve Allain’s review is here.

For previous articles on TetZooMCon and TetZooCon, see…

The events of TetZooCon 2014, July 2014
TetZooCon 2015 Is On, July 2015
The Events of TetZooCon 2015, November 2015
Coming Soon: TetZooCon 2016, September 2016
The Day After TetZooCon, October 2016
The Fourth TetZooCon, September 2017
The TetZooCon of 2017, October 2017
Reasons to Attend TetZooCon 2018, September 2018
TetZooCon 2018: Best TetZooCon So Far, October 2018
Announcing TetZooCon 2019 – the Biggest Yet, August 2019
Final Call For TetZooCon 2019, October 2019
The Sixth TetZooCon, October 2019
TetZooCon 2020 + Zoom = TETZOOMCON 2020, December 2020
TetZooMCon 2020 an Unbridled Success, December 2020
Cronch Cats, Beasts of Gévaudan, Dinosauroids, Mesozoic Art and Much More: TetZooMCon 2021 in Review, September 2021
The Ninth and Largest of the Tetrapod Zoology Conventions, December 2022
Announcing the 10th Tetrapod Zoology Convention, October 2023
At TetZooCon 2023, November 2023

You can support the persistence of this blog and my research and writing in general by throwing money at my partreon. For as little as $1 a month, you can help make a difference and see unpublished and in-prep stuff.

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Allain, S., Gandola, R., Tighe, A. & Wilkinson, J. W. 2019a. An investigation into the provenance of Bedford’s midwife toads. Bedfordshire Naturalist 74 (1), 48-52.

Allain, S., Gandola, R. & Wilkinson, J. W. 2019b. One or multiple origins of Midwife toads in the UK? FrogLog 27 (1), 23.

Allain, S. & Shimbov, M. I. 2021. The mystery of the Thorney midwife toads (Alytes obstetricans). Nature in Cambridgeshire 63, 26-27.

Coste, A., Fordyce, R. E. & Loch, C. 2023a. A new dolphin with tusk-like teeth from the late Oligocene of New Zealand indicates evolution of novel feeding strategies. Proceedings of the Royal Society B: Biological Sciences 290 doi 10.1098/rspb.2023.0873

Coste, A., Fordyce, R. E. & Loch, C. 2023b. A new fossil dolphin with tusk-like teeth from New Zealand and an analysis of procumbent teeth in fossil cetaceans. Journal of the Royal Society of New Zealand doi 10.1080/03036758.2023.2267456

Larkin, N. R., Lomax, D. R., Evans, M., Nicholls, E., Dey, S., Boomer, I., Copestake, P., Bown, P., Riding, J. B., Withers, D. & Davis, J. 2023. Excavating the ‘Rutland Sea Dragon’: the largest ichthyosaur skeleton ever found in the UK (Whitby Mudstone Formation, Toarcian, Lower Jurassic). Proceedings of the Geologists’ Association 134, 627-640.

Naish, D. 2022. Ancient Sea Reptiles: Plesiosaurs, Ichthyosaurs, Mosasaurs & More. Natural History Museum Publishing, London.

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Yes, TetZooCon 2023 is happening right now… more or less (I’m writing this on Thursday 30th November)… and that explains the lack of action here lately. A summary of what will be happening at TetZooCon 2023 has already been published here, but here are random thoughts before I leave.

**Caption:** I don’t think it’s a secret that TetZooCon 2023 is, in part, based around Mesozoic marine reptiles, mostly because it’s the year in which I published *Ancient Sea Reptiles*. The montage, designed and compiled by John Conway, features many illustrations that appear in the book.

TetZooCon, the vision. My plan for TetZooCon – right from the first event of 2014 – was to get it to the point where there might be several talk sessions happening in parallel, where on-stage panel or roundtable events would be happening here and there on the schedule, and that it would be more ‘con’ (as in: convention) and less ‘conf’ (as in: conference). That last point means tables, stalls, things on display, books for sale and for signing, art exhibitions and so on. Incrementally, we’ve gotten closer and closer to that end. And here in 2023, we’re there, basically. This year, we have three parallel sessions, a foyer full of stalls with merch, books and more on sale, and several roundtable and Q&A sessions. This year’s TetZooCon is also the biggest in terms of attendance, though I don’t have a final count as we’re in the throes of getting things sorted as I write.

**Caption:** the first TetZooCon happened in 2014 (**you can read about it here**). It should be obvious from this montage that I’ve relied extensively on pulling favours from friends and colleagues: the speakers here (l to r) are Mark Witton, Paulo Viscardi and Mike Taylor.

TetZooCon is a big enough event on the calendar that part of the year now revolves around it, and it isn’t going away. Due to excellent co-operation we have with Dr Chris Manias at King’s College, we also have an excellent venue, and indeed the biggest challenge for running large events seems to be that very thing. The prohibitive cost of venues has killed other, seemingly successful events in the recent past.

A public event. I really hope that TetZooCon is understood as the outreach event designed for non-specialists that it is. It is not a technical conference where specialists talk to other specialists, but a publicly-facing event, open to anyone interested, where specialists do give talks, but present them to a general audience, often for the first time. We would like to see that point emphasized more. Here I should mention the fact that even technical conferences are, in theory, ‘open to the public’, but they typically have a structure and promotional model that very much relies on the attendance of active and working researchers (mostly PhD students, post-docs and teaching staff).

**Caption:** field outings have been a regular feature of TetZooCon right from the start, though the nature of our operation means that these are generally informal, ad hoc, and arranged at the last minute. We’ve been to Crystal Palace a few times. I love this photo, showing Mark Witton with one of the *Iguanodon*s… though it’s actually from a non-TetZooCon meeting, details details. Image: (c) Mark Witton.

Another point on being public-facing: this year, more young people (children and teenagers) are attending than ever. I welcome this and seek to expand on it.

Going abroad, or not. Discussions on social media often highlight how much interest there would be in holding TetZooCon outside the UK, most obviously in the USA. Those who attended our zoom-based events – ha ha, we called them TetZooMCon – generally enjoyed them and liked being part of the collective. While we would love to be more international, it ain’t gonna happen. This is a shoestring thing run by two people (myself and John Conway), there are no appreciable profits, and just flying across an ocean would cost enough to put us into negative equity. In future – if special funding of some sort were obtained – I would love to run a non-UK thing. But, clearly, there are serious limitations preventing it.

**Caption:** another point of success for TetZooCon (err, in my opinion) is that we now serve as an important venue for the sale, and sometimes even the launch, of relevant books. These ones – among others – were available for sale and signing at the 2022 event, and will be on sale again this year (though I regret that only *one copy* of *Mesozoic Art* is up for grabs!).

As for running the in-person TetZooCon meeting as a pay-to-view streaming event (something else that gets suggested a lot), we just can’t do it. Too few of us are running this, and the profits are nowhere near grand enough to cover the costs. So: nice idea, but we don’t have the time, the staff, or the finances for this to work, not yet.

**Caption:** that’s the 2023 banner at top, but this montage gives some idea of what happened at TetZooCon 2022. It was the biggest TetZooCon ever in the history of the universe (**you can read about it here**), but 2023 has it beat by some margin. Yes I am a little scared.

That’s where I’ll end things. I look forward to seeing many of you there and hope things go well, despite the worsening situation with British transport, the bad weather, the financial cost of doing anything in central London, the looming threat of continuing issues with public health, and the general circling the drain of all we hold dear. Come back soon for a post-TetZooCon report!

Much is due to be published here soon: there is constantly a long list of articles I’m hoping to get finished and published when workload allows. Topics receiving coverage soon including Australian feral mega-cats again, wolves, a tale from the forests of Madagascar, an overview of new(ish) cryptozoology books, zebras part 3 and more.

For previous articles on TetZooMCon and TetZooCon, see…

The events of TetZooCon 2014, July 2014
TetZooCon 2015 Is On, July 2015
The Events of TetZooCon 2015, November 2015
Coming Soon: TetZooCon 2016, September 2016
The Day After TetZooCon, October 2016
The Fourth TetZooCon, September 2017
The TetZooCon of 2017, October 2017
Reasons to Attend TetZooCon 2018, September 2018
TetZooCon 2018: Best TetZooCon So Far, October 2018
Announcing TetZooCon 2019 – the Biggest Yet, August 2019
Final Call For TetZooCon 2019, October 2019
The Sixth TetZooCon, October 2019
TetZooCon 2020 + Zoom = TETZOOMCON 2020, December 2020
TetZooMCon 2020 an Unbridled Success, December 2020
Cronch Cats, Beasts of Gévaudan, Dinosauroids, Mesozoic Art and Much More: TetZooMCon 2021 in Review, September 2021
The Ninth and Largest of the Tetrapod Zoology Conventions, December 2022
Announcing the 10th Tetrapod Zoology Convention, October 2023

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In the previous article we looked briefly at those new amphibian and mammal species named during 2023. This time we skip ahead to reptiles… including birds because – yes – birds are reptiles in the phylogenetic sense….

**Caption:** a ‘new’ species montage for 2023…. err, sorta. Clockwise from upper left: Chamba pitviper *Gloydius chambensis* Kuttalam *et al*., 2023, Khammouan karst dragon *Laodracon carsticola* Brakels, Sitthivong, Wang, Nguyen & Poyarkov, 2023 in karst habitat, goshawk montage (see below). Images: Kuttalam *et al*. (2023); Sitthivong *et al*. (2023); Louis Agassiz Fuertes, in public domain; at right, Norbert Kenntner, CC BY-SA 3.0 (**original here**).

Some snakes and some lizards. Yeah yeah, snakes are lizards, but you know what I mean. One thing is clear when you look at new squamate species named in any one year, and that’s geckos geckos geckos. 2023’s list includes new species of Paroedura from Madagascar, at least seven new Lygodactylus dwarf geckos, several new Australian Amalosia geckos, and at least three new bent-toed geckos (Cyrtodactylus). A revision that saw much shuffling of the geckos formerly included in Ptychozoon saw print (some are now included within Gekko), this occurring hand-in-hand with the naming of some new Gekko species (Lalremsanga et al. 2023). That’s a sufficiently complex issue that I’ll have to put it to one side for now. New anoles, chameleons, lacertids, teiids and more also saw print.

**Caption:** Mizoram parachute gecko *Gekko mizoramensis* Lalremsanga *et al*., 2023. There was a time when geckos of this sort were classified together in *Ptychozoon*, but at least some are now included within *Gekko*. Image: (c) Lal Muansanga.

New snakes for 2023 include Harrison Ford’s slender snake Tachymenoides harrisonfordi Lehr et al., 2023, a dipsadine colubroid from Peru and only the second species of its genus to be named. That genus was itself only recognised in 2022, having been previously included within Tachymenis. Yes, it’s named after Harrison Ford and is one of several species that honour him (the others are all arthropods, I think); the authors note that this is “in recognition of his work for Conservation International and his voice for nature” (Lehr et al. 2023, p. 203).

A study of dipsadines from Colombia, Ecuador and Panama resulted in the recognition of four new Sibon species and one new Dipsas, and one of these made the newswires: DiCaprio’s snail-eating snake Sibon irmelindicaprioae Arteaga & Batista, 2023, a Near Threatened species from Panama. It’s named in honour of “Irmelin DiCaprio (1945–present), mother of Leonardo DiCaprio, long-time advocate and supporter of biodiversity conservation around the world” (Arteaga & Batista 2023). Ah, eponyms… hold that thought.

**Caption:** holotype specimen of DiCaprio’s snail-eating snake *Sibon irmelindicaprioae* Arteaga & Batista, 2023. It’s an adult male, collected at Cerro Bailarín, Panama, in 2011. Image: **Arteaga & Batista (2023)**.

  **Caption:** at left, images of the holotype of Harrison Ford’s slender snake *Tachymenoides harrisonfordi* Lehr *et al*., 2023 in life. The red arrows point to the three longitudinal skin folds present in this species. The snake is 40 cm long. At right, the holotype (when alive) of the Chamba pitviper *Gloydius chambensis* Kuttalam *et al*., 2023. It’s 42.6 cm long. Images: Lehr *et al*. (2023), Kuttalam *et al*. (2023).

Four new species of the odd-scaled xenodermid genus Achalinus, all from China, were named during the year, as was the karst-dwelling pitviper Trimeresurus ciliaris Idiiatullina et al., 2023 from Thailand and the Chamba pitviper Gloydius chambensis Kuttalam et al., 2023 from India.

**Caption:** a montage featuring new (for 2023) species of the Asian draconine agamid *Diploderma*. At left, the *Diploderma* *jiulongense* holotype, from Yandai, Jiulong County in Sichuan Province, China. At right, the *Diploderma tachengense* holotype, from Tacheng Town in Yunnan Province, China. SVL in these lizards is around 55 mm in males. Images: **Liu *et al*. (2023)**.

The Khammouan karst dragon. New lizard species belonging to already known genera are named on regular basis, but new genera come along less frequently. The Khammouan karst dragon Laodracon carsticola Brakels, Sitthivong, Wang, Nguyen & Poyarkov, 2023 of the karstic massifs of central Laos is among this select number. It’s an agamid, and specifically a draconine: a member of the large clade that includes the flying Draco species, the pricklenape dragons (Acanthosaura), the forest or dragon lizards (Calotes) and so many others. Within the group, molecular data shows that Laodracon is especially close to the Diploderma mountain dragons (a genus that saw the naming of four new species in 2023, all from China and newly collected in 2022: Liu et al. 2023). Laodracon is black overall with white cross-bands, blotches and spots distributed across the body, and a grey reticulated pattern and blue spot on the throat (Sitthivong et al*., 2023). Key features include a swollen tail base decorated with enlarged, keeled scales on all of its surfaces.

This is another of these taxa where the authorship for the name is not the same as the authorship of the paper. I wish that this wasn’t a thing: it means you can’t look at a paper and assume that you know the name’s authorship from this information alone. Instead, you have to obtain and dig into the paper to find the naming section. Also, the authorship list was only given for the genus, not the species. I assume that both are the same.

Caption: a Laodracon montage, from Sitthivong et al. (2023). At left: the holotype male specimen in different views, key traits (relating to the form of the tail base) being visible in G. At right: the new taxon’s position in phylogeny, showing its closeness to the large Diploderma radiation. Images: Sitthivong et al. (2023).

At present, it’s very poorly known. Individuals have been seen perched and climbing on steep karst pinnacles 50-70 m above ground (Sitthivong et al. 2023). The first individual made known to science was observed during August 2022 by S. Xayyasith while birdwatching. A second specimen was observed (and collected) during October 2022 by a local guide who was taking tourists on a zipline trip. This one became the holotype (Sitthivong et al. 2023). Laodracon is poorly known to local people and is presently known only from two male specimens. Suspicions are that it specializes on eating ants. Its discovery emphasizes the need for conservation of the area’s biologically important karst habitat.

Birds of 2023. And we finish with birds, where there isn’t big 2023 news. Or is there? A few species listed as newly published for 2023 – among them the Principe scops owl Otus bikegila and Wangi-wangi white-eye Zosterops paruhbesar – were actually published in 2022, though here we come to the irksome issue of whether digital publication counts ahead of physical paper publication. UPDATE: as Albertonykus notes in the comments, a few new species were named in 2023 as a consequence of molecular analysis and the splitting of previously recognized species. They include Cracraft’s schiffornis Schiffornis cracrafti Lima et al., 2023 of southwestern Amazonia and Darién nightingale-thrush Catharus arcanus Halley et al., 2023 from Panama.

And having mentioned splitting… birders pay a lot more attention to ‘splittings’ and ‘lumpings’ than do other people interested in animals, so it’s worth saying that numerous taxonomic decisions of this sort saw print in 2023. Among the more interesting ones involve the splitting of the American goshawk Accipiter atrocapillus from the Eurasian A. gentilis and the distinction of the Western cattle egret Bubulcus ibis from the Eastern B. coromandus. As ever, those claiming that these splits are only done so that birders can compile longer life lists will be harshly admonished. Those claims are neither fair nor true.

**Caption:** a goshawk montage. The animals here were formerly lumped together as the Northern goshawk but new decisions formerly recognize a split between the two, such that we have the American goshawk *Accipiter atrocapillus* (Wilson, 1812) (at left) and the Eurasian goshawk *A. gentilis* (Linnaeus, 1758). Images: at left, Louis Agassiz Fuertes, in public domain; at right, Norbert Kenntner, CC BY-SA 3.0 (**original here**).

While I’m here, it’s worth making a brief point on the issue of bird taxonomy that everyone’s talking about right now. Namely, the American Ornithological Society’s stated aim to remove eponyms (or patronyms) from North American vernacular names for birds and have them replaced with descriptive ones. There are, of course, appropriate sociopolitical reasons for this move, and you can rail about it (no pun, ha ha) representing ‘woke’ culture if you want. But an argument can be made that eponyms are not good names, since they neither ‘honour’ the animal, nor connect the name with the animal.Yellow-headed blackbird is a good name. I previously commented on this exact issue in my 2019 article on Lyall Watson’s 1981 book Whales of the World and my argument there was that eponyms should go so long as there’s sufficient momentum. Well, now there is.

However… a complication is that some eponyms were published to honour scientists and even politicians and community leaders who did something truly notable or important with respect to the history of the organism concerned. They might have provided funding that resulted in its discovery or recognition, or provided vital stewardship for the species and its habitat. They might be beloved mentors who were the sole builders of conservation programmes, research groups or scientific projects in an otherwise neglected or beleaguered region. Ergo, this is not a black and white ‘eponyms are bad’ issue.

**Caption:** a classic tweet, from the pen of **Kristie**.

That’s where we’ll end things. Be sure to check out part 1 if you haven’t already done so. And for previous Tetrapod Zoology articles that cover certain of the subjects touched on here, see…

Ptychozoon: the geckos that glide with flaps and fringes (gekkotans part VIII), June 2010
Australia, land of dragons (by which I mean: agamids) (part I), January 2014
Australia, land of dragons (part II), February 2014
Remembering Lyall Watson’s Whales of the World, June 2019
New Species Round-up for 2023, Part 1, November 2023

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

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Arteaga, A. & Batista, A. 2023. A consolidated phylogeny of snail-eating snakes (Serpentes, Dipsadini), with the description of five new species from Colombia, Ecuador, and Panama. Zookeys 1143, 1-49.

Kuttalam, S., Santra, V., Owens, J. B., Selvan, M., Mukherjee, N., Graham, S., Togridou, A., Bharti, O. K., Shi, J., Shanker, K. & Malhotra, A. 2023. Phylogenetic and morphological analysis of Gloydius himalayanus (Serpentes, Viperidae, Crotalinae), with the description of a new species. European Journal of Taxonomy 852, 1-30.

Lalremsanga, H. T., Muansanga, L., Vabeiryureilai, M. & Mirza, Z. A. 2023. A new species of parachute gecko of the subgenus Ptychozoon (Sauria: Gekkonidae: Gekko) from the Indo-Burma Region. Salamandra 59, 125-135.

Lehr, E., Cusi, J. C., Fernandez, M. I., Vera, R. J. & Catenazzi, A. 2023. A new species of Tachymenoides (Serpentes: Dipsadidae: Tachymenini) from the puna of the Otishi National Park in Peru. Salamandra 59, 199-206.

Liu, S., Hou, M., Ananjeva, N. B. & Rao, D. 2023. Four new species of the genus Diploderma Hallowell, 1861 (Squamata, Agamidae) from China. ZooKeys 1148, 167-207.

Sitthivong, S., Brakels. P., Xayyasith, S., Maury, N., Idiiatullina, S., Pawangkhanant, P., Wang. K., Nguyen. T. V. & Poyarkov, N. A. 2023. Hiding on jagged karst pinnacles: A new microendemic genus and species of a limestone-dwelling agamid lizard (Squamata: Agamidae: Draconinae) from Khammouan Province, Laos. Zoological Research 44, 1039-1051.

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This is a time of ecological crisis and massive loss of animal diversity, make no mistake about it. But there’s still a vast amount of new stuff left to discover, and every year we see a significant influx of newly recognized species, even among tetrapods. In this and the next article, we take a whistle-stop tour of those tetrapod species new to science as of 2023. As ever, remember that new to science is not synonymous with new to humanity…

**Caption:** a recently-named-species montage, featuring a pangolin cladogram, the new gymnure *Podogymnura intermedia* Balete *et al*., 2023 and the Seepage siren. Images: Gu *et al*. (2023), Balete *et al*. (2023), Fedler *et al*. (2023).

A quick amphibian review. If, over the years, you’ve followed the Tetrapod Zoology articles on amphibians (oooor, if you’re well informed on amphibians in general) you’ll know that substantial numbers of new living amphibian species are named every year: over 100 each year, typically over 150. If you’re familiar with the idea that a handful of new bird and mammal species are named each year, that’s a lot. Non-specialists tend not to hear about these many species because they mostly don’t make the newswires.

**Caption:** a montage of just a few of the megophryid and strabomantid frogs named in 2023. Clockwise from upper left…. *Xenophrys pangdaensis* is a Tibetan megophryid; its authors suggested the common name Pangda horned toad (Shu *et al*. 2023); a montage showing the holotype female (a-c, e-f) and male (d) of the strabomantid *Pristimantis kopinangae* Means *et al*., 2023 from Mount Kopinang, Guyana; and the holotype specimen of *Pristimantis clarae* from Peru in (A) dorsal and (B) ventral views. Images: **Shu *et al*. (2023)**, **Means *et al*. (2023)****,** **Venegas *et al*. (2023)**.

At the time of writing, 106 new living amphibian species have been named in 2023. Most are frogs, and it’s obvious from the list that we’re in a very active phase of research on the study and naming of megophryid spadefoots from China, strabomantids from such nations as Ecuador, Guyana, Brazil and Peru, and Madagascan mantellids. Five new caecilians have been named so far in 2023, all Colombian and belonging to the genus Caecilia.

**Caption:** at least five new species of *Caecilia* caecilians were named in 2023. This is one of them: *C. macrodonta* Fernández-Roldán *et al*., 2023, a species noted for its large, recurved dentary teeth. It’s from the Cordillera Oriental of Colombia and the holotype was collected in 1980. Image: Fernández-Roldán *et al*. (2023).

Salamanders: new sirens and giants. About 14 new salamanders have been named so far during 2023, stand-outs for me including the Seepage siren Siren sphagnicola Fedler et al., 2023 from Florida and the Qimen giant salamander Andrias cheni Xu et al., 2023 of Anhui Province, China. The Seepage siren follows on the proverbial heels of the Reticulated or Leopard siren S. reticulata of Alabama, named in 2018, and is one of several taxa historically (but incorrectly) lumped together within S. intermedia, the Lesser siren (Fedler et al. 2023)

**Caption:** a male Seepage siren specimen, in this case an individual with a partly regenerated tail. This is the smallest living siren species, the longest specimen reported by Fedler *et al*. (2023) having a total length of about 19 cm. Image: Fedler *et al*. (2023).

We now move to mammals…

Two new gymnures. Gymnures or moonrats are poorly known east and south-east Asian lipotyphlans that belong to Erinaceidae, the hedgehog family. They look superficially like giant, woolly-coated shrews. Of the five extant gymnure genera, I think it’s fair to say that the least familiar is the endemic Philippine genus Podogymnura. Podogymnura was named in 1905 for the Mindanao gymnure P. truei but the genus was regarded as monospecific until the 1982 naming of the Dinagat gymnure P. aureospinula. The Philippines are an area where both small mammal diversity is high, and where numerous recent discoveries have been made, so perhaps it was always likely that additional Podogymnura species would be discovered, perhaps on various of the islands around Mindanao. Dinagat and Bucas Grande are among those islands, and both are inhabited by P. aureospinula*.

This year has seen the naming of a third species: the East Mindanao gymnure Podogymnura intermedia Balete et al., 2023. The specific name is a reference to the fact that it’s intermediate in size between P. auroespinula, the largest Podogymnura species, and the remaining members of the genus. Balete et al. (2023) also evaluated P. truei and found that the supposed subspecies P. truei minima Sanborn, 1953 differed sufficiently from the nominate form (in pelage and skin pigmentation, skull and tooth size and proportions and overall size) to warrant recognition as another species, so hello Podogymnura minima.

**Caption:** the holotype specimen of *Podogymnura intermedia* Balete *et al*., 2023, photographed in 2005 by D. S. Balete. The fact that the species was recorded in 2005 but not published until 2023 is another reminder of the fact that ‘publication date’ is very much not the same thing as ‘discovery date’. Image: Balete *et al*. (2023).

Quit it with ‘Eulipotyphla’. You don’t need it. Lipotyphla still functions just fine, even though its composition has changed over recent decades. This is basically the same argument as the one Prothero et al. (2021) made about the similarly redundant ‘Cetartiodactyla’.

The mysterious pangolin. Pangolins are, I think, globally better known today than they were, but it’s sad that this is only the case because the extent of the absurd and disgusting trade in their meat and scales has reached epidemic proportions. A consequence of this overexploitation is that materials intercepted during shipment are now routinely examined via genetic and morphological analysis.

Pangolin scales obtained in Hong Kong in 2012 and 2013 possess genetic traits different from those of any recognised Asian pangolin species (Hu et al. 2016), this suggesting the presence of an otherwise unknown species. However, the extracted gene sequences were fragmentary and mitochondrial only, and the suggestion was made that their unusual nature could be explained by them representing variation within one of the known species, or being exclusively mitochondrial. The news for 2023 is that scales sharing genetic markers with the Hong Kong samples – confiscated from Yunnan, China in 2015 and 2019 – have now been analyzed. Gu et al. (2023) performed appropriately thorough comparisons between the DNA of this putative new species and 138 whole-genome sequences from other pangolins, and also examined the morphology too.

**Caption:** a Philippine pangolin mother and baby, a species almost certainly close, in both phylogeny and appearance, to the Asian cryptic species new for 2023. Image: Shukran888, CC BY-SA 3.0 (**original here**).

The conclusion: the scales do indeed belong to a new species, one presently known only from illegally transported scales. It’s been named the Asian mysterious pangolin Manis mysteria Gu et al., 2023 and it’s, hopefully, only a matter of time before the relevant molecular and scale characters are identified in individuals known from entire bodies, preferably live ones [UPDATE: technically, the name has been published without the allocation of a type specimen, meaning that M. mysteria is a nomen nudum]. M. mysteria scales are small, so this species will likely be a diminutive, climbing species similar to its close relatives. A time-calibrated molecular analysis shows that M. mysteria is part of the East Asian Manis clade and diverged from the Philippine M. culionensis and Sunda/Malayan M. javanica pangolins during the Pliocene, around 5 million years ago (Gu et al. 2023).

**Caption:** phylogenetic trees generated from examination and comparison of pangolin DNA, showing how the new cryptic species is part of the Asian *Manis* clade and is likely closest to the Malayan and Philippine pangolins. Image: Gu *et al*. (2023).

Other mammals, briefly. Other new mammals named during 2023 include the Argentinian tuco-tuco Ctenomys pulcer Verzi et al., 2023, two new talpid moles from Iran (Talpa hakkariensis Gündüz et al., 2023 and T. streetorum Gündüz et al., 2023, both previously included within Père David’s mole T. davidiana), and Príncipe’s pipistrelle Pseudoromicia principis Juste et al., 2023. UPDATE: yes, I forgot the Red tigrina Leopardus narinensis Ruiz-García et al., 2023, a cat from Colombia named for a skin collected in 1989 and initially misidentified as that of an Ocelot L. pardalis. That’s ironic given that I tweeted about its publication back when it was new!

Ok, we’ll stop there. Reptiles next…

Several previous Tetrapod Zoology articles discuss various of the animals mentioned here. See…

THE AMAZING WORLD OF SALAMANDERS, October 2013
The Biology of Sirens, June 2016
The Amazing Caecilians, October 2022

If you enjoyed this article and want to assist me in what I do — and see behind-the-scenes stuff in development — please support me at patreon. Click anywhere on this text in bold.

Refs - -

Balete, D. S., Heaney,L. R., Rickart, E. A., Quidlat, R. S., Rowsey, D. M. & Olson, L. E. 2023. A re-assessment of diversity among Philippine gymnures (Mammalia: Erinaceidae: Podogymnura), with a new species from eastern Mindanao. Zootaxa 5228, 244-266.

Fedler, M. T., Enge, K. M. & Moler, P. E. 2023. Unraveling Siren (Caudata: Sirenidae) systematics and description of a small, seepage specialist. Zootaxa 5258, 351-378.

Fernández-Roldán, J. D., Lynch, J. D. & Medina-Rangel, G. F. 2023. On the identities of Caecilia degenerata Dunn, 1942 and of C. corpulenta Taylor, 1968 (Amphibia: Gymnophiona: Caeciliidae) with descriptions of three new species of Caecilia Linnaeus, 1758 from the Cordillera Oriental of Colombia. Zootaxa 5227, 205-228.

Gu, T.-T., Wu, H., Yang, F. & Yu, L. 2023. Genomic analysis reveals a cryptic pangolin species. Proceedings of the National Academy of Sciences 120, e2304096120.

Hu, J., Roos, C., Lv, X., Kuang, W. & Yu, L. 2020. Molecular genetics supports a potential fifth Asian pangolin species (Mammalia, Pholidota, Manis). Zoological Science 37, 538-543.

Means, D. B., Heinicke, M. P., Hedges, S. B., Macculloch, R. D. & Lathrop, A. 2023. Exceptional diversity of Pristimantis landfrogs (Anura: Terraranae) on the Wokomung Massif, Guyana, with descriptions of three new species. Journal of Vertebrate Biology 72, 1-26.

Shu, G., Li, K., Wu, Y., Liu, Q., He, Z., Li, L., Zhang, H. & Guo, P. 2023. A new species of Xenophrys (Amphibia, Anura, Megophryidae) from southern Tibet, China. ZooKeys 1182, 307-329.

Venegas, P. J., García-Ayachi, L. A., Marchelie, A., Ormeño, J. R. & Catenazzi, A. 2023. A new species of terrestrial-breeding frog, genus Pristimantis (Anura: Strabomantidae), from the Peruvian Yungas of Central Peru. Taxonomy 3, 331-345.

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On December 1st, 2nd and 3rd 2023, the 10th Tetrapod Zoology Convention – TetZooCon – happens at Bush House, King’s College, The Strand, London. With just over a month to go (yikes), now is time to buy a ticket and consider joining us. It’s going to be the biggest TetZooCon so far, and hopefully the best. Here’s a rundown of what’s due to happen…

**Caption:** a montage of things relevant to TetZooCon 2023. The books *Ancient Sea Reptiles*, *Mesozoic Art* and *Locked In Time* - and others - will be on sale, and talks on cassowaries will be happening. At far right: Bush House, our venue. Images: cassowary from Todd Green; others Darren Naish, *Locked In Time* via Columbia University Press.

Schedule and timetable. TetZooCon 2023 kicks off on the evening of Friday 1st December with the panel event Engaging with Extinctions - Past, Present & Future. A drinks reception and registration also happens on that Friday. From Saturday morning onwards, stalls are present throughout the meeting whereby people sell merch, books and more, and show art. I, personally, have a select number of animal figures and natural history books on sale. I expect most sales to occur via card machines as is typical for the modern age.

TetZooCon has reached the point in its life where it now has to involve parallel sessions throughout. Alas, gone are the days when we can all sit together and see *all* the presentations. This means that you’ll need to pay attention to the timetable if there are specific talks or events you want to attend. Saturday and Sunday are the main ‘event days’, involving a series of talks, on-stage panel discussions and at least two workshops. This year, we welcome cosplay (something that had a soft launch last year; honorary mention again of Rebecca Groom and Meghan Jenkinson, the instigators). We recommend that cosplay is limited to Saturday afternoon (post-lunch), and an on-stage judging – with prizes – happens at around 6pm on Saturday. It’s followed by an art exhibition and drinks reception.

Marine reptile extravaganza. Due to the publication of my Natural History Museum/Smithsonian Books book Ancient Sea Reptiles, TetZooCon 2023 is partly focused on marine reptiles. I will be selling and signing copies, and giving a talk about Mesozoic marine reptiles too. And I will be joined by a list of colleagues: we have Judyth Sassoon, Richard Forrest and Luke Muscutt talking plesiosaurs, and Emily Swaby and Dean Lomax talking ichthyosaurs.

         Luke is bringing the robotic plesiosaur he’s been working on, a world exclusive, and Dean will be selling and signing copies of his book *Locked in Time*. All marine reptile speakers will engage in the panel event *Marine reptile science: problem areas, and… where next?*

**Caption:** Luke Muscutt with robot plesiosaur. Image provided by Luke Muscutt.

Other talks on Saturday involve fossil dolphins (Amber Coste: Daunting Dentitions - Inside the Mouths of Dolphins That Bit the Dust), sci-comm and outreach (Hana Ayoob: Audiences and Other Animals) and herpetology (Steve Allain: On the Trail of Midwife Toads in Great Britain).

      **Palaeoart and media events.** In contrast to previous years, our palaeoart session isn’t so much of a session, but a constantly running series of events happening in parallel to the rest of the meeting. Luis Rey will be leading a discussion – *A Discussion on the Past and Future of Palaeoart* – and James Pascoe is hosting a *Paint-a-Pliosaur Event* on the Saturday. Joschua Knüppe is also running a grand show-and-tell event on Sunday.

   **Caption:** pliosaur models by James Pascoe. A good number of them will be in attendance at TetZooCon 2023. Images: James Pascoe.

Sunday lunchtime also sees a screening of 1925’s The Lost World, introduced and chaired by Dave Hone, who’ll also be providing commentary. The movie is 106 minutes long, and shouldn’t overlap with the events happening post-lunch on the day. They include my Prehistoric Planet-themed event (which is happening in the same room as the palaeoart meetings) and Rebecca Wragg Sykes’s talk Things We See In The Dark: From Shadows on Cave Walls to (Homo) Naledi In the Sky.

**Cassowaries, crows, crocodiles**. In other events, we’re joined on the Sunday by Todd Green, who’ll be talking about his research on cassowaries and the cassowary research renaissance. This is followed by a cassowary-themed panel event in which Todd and I are joined by film-makers Kerrie and AJ Dodd. Also appearing on the Sunday are Jennifer Campbell-Smith on *Cautious Crows: How to Trick Corvids Into Letting You Study Them* and Evon Hekkala on *Maneaters, Mummies and Madagascar; Sacred and Secret Tales from Cryptic Crocodiles*.

      Sunday ends with our special guest, Nigel Marven, who’ll be in attendance for part of the afternoon and also giving the talk *Filming Adventures With Dinosaurs and Other Reptiles*. As usual, we end with the TetZooCon quiz, again with prizes.

It should be obvious that we’re talking here about *a lot* of stuff, and that this is going to be a packed meeting. Tickets have already been on sale for a few weeks and the number we’ve sold indicates that this is set to be the biggest TetZooCon so far (which is in keeping with the long-term trend). At this point, masking is down to personal preference. I would say that masking might be wise given current concerns about new covid strains, but a mask mandate is not in operation.

Ok. For more information (including a map, the timetable, and reviews of previous years) visit the TetZooCon page here, and that’s where tickets are available too. Things are coming together as planned, but even now there are things yet to be confirmed or finalized. For breaking news and developments, keep an eye on the TetZooCon facebook page. This will probably be the last update I provide here at Tet Zoo, so I look forward to seeing you in London in December!

For previous articles on TetZooMCon and TetZooCon, see…

The events of TetZooCon 2014, July 2014
TetZooCon 2015 Is On, July 2015
The Events of TetZooCon 2015, November 2015
Coming Soon: TetZooCon 2016, September 2016
The Day After TetZooCon, October 2016
The Fourth TetZooCon, September 2017
The TetZooCon of 2017, October 2017
Reasons to Attend TetZooCon 2018, September 2018
TetZooCon 2018: Best TetZooCon So Far, October 2018
Announcing TetZooCon 2019 – the Biggest Yet, August 2019
Final Call For TetZooCon 2019, October 2019
The Sixth TetZooCon, October 2019
TetZooCon 2020 + Zoom = TETZOOMCON 2020, December 2020
TetZooMCon 2020 an Unbridled Success, December 2020
Cronch Cats, Beasts of Gévaudan, Dinosauroids, Mesozoic Art and Much More: TetZooMCon 2021 in Review, September 2021
The Ninth and Largest of the Tetrapod Zoology Conventions, December 2022

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Welcome to the second of my articles on birdwatching in Tajkistan in Central Asia….

**Caption:** a montage of passerines discussed in this article….

As you’ll know if you read the first article, I wasn’t in Tajikistan for bird-related reasons and in fact I only got to engage in opportunistic ‘fly-by’ birding. Nevertheless, I was thrilled with what I saw. In the previous article I covered the non-passerine species: various gallinaceous birds, waders, pigeons and raptors as well as the odd woodpecker, bee-eater and hoopoe. This time we get to the group that contains the bulk of extant avian diversity, the passerines or perching birds. And boy were there a lot of ‘difficult’ birds here, many of which have remained a mystery to me (in terms of identification).

**Caption:** an indication of what the scenery can be like in Tajikistan. This is the landscape around Lake Iskanderkul, a massive glacial lake on the northern side of the Fann Mountains. Image: Darren Naish.

  **Caption:** features of the Romit Valley in Tajikistan, a location where many of the birds discussed in these articles were seen. The valley featured wide, raging rivers (like the Kafinigan River shown here), and tall, jagged mountain peaks. Image: Darren Naish.

  **Caption:** another very scenic shot, this time in Saratag/Sarataq (both spellings seem to be in use), about 140 km north of Dushanbe, the capital. Note the mostly coniferous woodland across the slopes. Bears were here (as evidenced by fresh dung and conversations with locals). Image: Darren Naish.

For preamble and a short introduction on what Tajikistan is like and on what it’s like to look for birds there, be sure to check out the previous article. Alright, let’s get to it…

Shrikes, corvids and some surprising kin. We’ll start with corvoids. Several shrike species occur in Tajikistan, some of which only occur in the east and not here in Europe. I had good views of Long-tailed shrikes Lanius schach [UPDATE: this article previously had the species identified as Red-backed shrike L. collurio, a species I’ve seen before in eastern Europe], and what’s either Isabelline shrike L. isabellinus or Turkestan shrike L. phoenicuroides. The Long-tailed shrike is (in males, anyway) a very attractive, boldly marked bird with a reddish-brown back and light grey head. The other shrike is about impossible to pin down since both female and juvenile male Isabelline and Turkestan shrikes are sandy-brown with a less distinct mask than those of most other shrikes. Big thanks to Tim Worfolk (see comments) for his help here. Shrikes are one of my favourite bird groups, but so far I’ve only seen a handful of species. I spoke previously about them in this article on birdwatching in China.

**Caption:** I had bad luck in my efforts to photograph shrikes, buuuut then… I had bad luck in photographing just about all of the birds. Two Long-tailed shrikes are visible in this photo, I hope. Long-tailed shrikes occur widely across Asia and also occur in Australasia. Numerous subspecies have been named. Image: Darren Naish.

  **Caption:** more shrikes. The Long-tailed shrike photo at left isn’t great, but at least you can clearly see the grey head, black mask and pinkish sides. At right, an Isabelline or Turkestan shrike seen in shrubby vegetation close to a river. Images: Darren Naish.

Corvids are not especially diverse in Tajikistan. There are no exotic jays or magpies, nor any of those unusual crows with white head markings or massively overbuilt bills. Carrion crows Corvus corone were seen on numerous occasions at many rural (and not urban) places, but it was obvious immediately that they were distinct from those I know from Europe.

**Caption:** Carrion crows were seen in flocks of 10-15 on more than one occasion, implying that these Central Asian populations might be more social than the European ones. It should also be obvious that these crows are more slender and lightly built than western forms of this species. Image: Darren Naish.

  **Caption:** Carrion crow seen flying overhead. I like the asymmetrical nature of the primary feather damage. It may or may not mean something (like handedness). Image: Darren Naish.

I had the impression that they were slimmer, shallower in the neck and with a proportionally longer, pointier bill than the Carrion crows I know here in the UK. Perhaps this is consistent with the view that these Asian birds represent the subspecies C. c. orientalis, the Eastern carrion crow. This occurs from Iran in the west to Japan in the east and is larger on average than the nominate form. That’s interesting, because my first (incorrect) impression on seeing these Tajik birds was that they were ravens. On that note, I was surprised to never once see ravens of any sort.

Magpies were seen on regular occasion, mostly in rural locations but also at roadside restaurants and parking areas. Recent studies show that the magpies previously lumped together in Pica pica have a phylogenetic structure that warrants the recognition of several species within Eurasia, in part because the American species (which have always been regarded as distinct species) are surrounded by Eurasian lineages. Magpies in Tajikistan belong to the bactriana subspecies, which groups with European and west Asian magpies in molecular studies (Song et al. 2018). This means that they should stay within P. pica, which makes sense as the Tajik birds I saw looked like European ones.

**Caption:** Tajik magpies. The bird at left – photographed in a carpark in the Fann Mountains – was missing some head feathers and hence not great in appearance. At right, a different bird, this time in Saratok. Images: Darren Naish.

A big surprise was the Indian paradise flycatcher Terpsiphone paradisi, a bird you might associate more with tropical south-east Asia than the very different habitats of Central Asia. Certainly I did. I have to mention upfront that the species I’m dealing with is one of three formerly lumped together as the ‘Asian paradise flycatcher’ (the others are the Chinese or Amur paradise flycatcher T. incei and the Oriental or Blyth’s paradise flycatcher T. affnis), and this – obviously – is the name used for the species in Ayé et al. (2012) and most other sources on Tajik birds.

**Caption:** Indian paradise flycatcher, a most handsome bird. Males and females are alike in possessing head crests, but males also have white primaries and very long central tail feathers. This is a female, and she belongs to the rufous form: there’s a white one where all the feathers are white excepting those on the head, neck and wing tips. Images: Darren Naish.

Also worth mentioning upfront is that Terpsiphone isn’t a ‘flycatcher’ in the way that most of us imagine this term: it’s not a member of the muscicapoid clade that includes conventional Old World flycatchers, but is a monarchid and thus part of Corvoidea. Within Corvoidea, monarchids are close to shrikes and corvids (Jønsson et al. 2016). They’re a widespread group with a complex phylogeographical history that involves tropical Asia, Africa, eastern Asia and the islands surrounding all of these places (Fabre et al. 2012).

**Caption:** action take-off shots of the same bird shown above. What these photos reinforce, partly at least, is that birds leap or fall into flight when launching from perches. They don’t start flapping and then take off. Images: Darren Naish.

A big surprise about this species is how agile it is in the arboreal environment. It doesn’t just perch on horizontal branches and sally back and forth, as you might predict for a bird with ‘flycatcher’ in its name. It clings to vertical trunks while probing the bark and will also move along a branch with its long axis parallel to that of the branch. A dark-headed, brown bird behaving in this way was identified by a member of our team as a woodpecker – meaning that we had a mystery bird on our hands – until we worked this out.

**Caption:** not a great photo, but an interesting one because it shows how paradise flycatchers are good at clinging and climbing about on trunks in woodpecker-like fashion. Image: Darren Naish.

Finally on corvoids, I saw a golden oriole in the same orchard at Hakimi where I got the woodpecker photos (see the previous article). As is typical for golden orioles, it was elusive and flew very quickly when eventually emerging from the foliage, so no photos. Two golden oriole species – the Indian Oriolus kundoo and European O. oriolus – occur across Central Asia but it looks like the latter is absent from Tajikistan. That matches what I saw, since the Tajik bird (a male) was very bright yellow with comparatively little black on its face.

We now move to the remaining passerines, the enormous, complex group technically termed Passerides.

Tits, or titmice if you’re that way inclined. Tits (Paridae) are an unusual group that were formerly regarded as not belonging to any of the major clades within Passerides. The most recent molecular studies show that they’re most likely an early-diverging group within Sylviida. A large tit, seen at close range in an orchard, was superficially similar to the Eurasian Great tit Parus major but grey on its shoulders and mantle and white on its sides and belly, rather than olive greenish dorsally and yellow ventrally. It can only have been one of ‘bokharensis group’ within the Great tit: a group of central and east Asian subspecies that were previously grouped together as the ‘Turkestan tit’. Some authors do still regard this as a distinct species (P. bokharensis).

**Caption:** a Tajik great tit, photographed foraging on a tree in an orchard, sometimes reaching under bark with its bill. This is obviously one of the grey ‘Turkestan tit’ subspecies, very distinct from the great tits of the west. Images: Darren Naish.

A few other species were far less familiar. A small tit with a boldly marked, black and white head, tall feather crest, yellowish nape and reddish vent area was seen on a few occasions in the Romit Valley. This is the Rufous-naped tit Periparus rufonuchalis, a widespread species that occurs from India north and east to China. The Periparus species – the best known is the Coal tit P. ater – are exclusive to Eurasia and northern Africa and a feature I now associate with species in the genus is a relatively long, slightly downcurved bill. That’s different from the shorter, more conical bill of tits like the Cyanistes species. And having mentioned the Cyanistes species….

**Caption:** a comparatively long-billed tit with a tall head crest, a reddish vent area and a lack of pale spots on its coverts. It can only be a Rufous-naped tit. Image: Darren Naish.

These very odd, mostly white and grey, yellow-breasted tits are Azure tits C. cyanus, a variable and polytypic, mostly Asian species that looks like a washed-out Blue tit. The two hybridize in western Russia and the resulting animal is known as Pleske’s tit. It was formerly regarded as a distinct species. Actually, the yellow-breasted form of the Azure tit I saw in Tajikistan is a distinct, Central Asian population that’s regarded as a distinct subspecies C. c. flavipectus by some authors, and as a species (the Yellow-breasted tit C. flavipectus) by others.

**Caption:** all my initial sightings of Azure tits (seen at Saratag in the Fann Mountains) involved the birds being seen in near-total silhouette, a classic example of the sort of bad luck that happens when you try and photograph birds. The only photos I got that don’t involve the birds being silhouetted are instead out of focus, like the example on the right. Images: Darren Naish.

Swallows and martins. Hirundines – swallows and martins – are a distinct lineage within Sylviida too. Individuals belonging to a few species were seen in the Romit Valley, but I never got any good looks nor any good photos. A smallish, light brown species seen flying over rivers was, I presume, Sand martin Riparia riparia, but Pale martin R. diluta, is also possible. The reddish throat, dark dorsal colour and long tail streamers of a second species made me think that it was Barn swallow Hirundo rustica.

**Caption:** terrible photos of small, fast-moving, brown hirundines. I’ve assumed that these are sand martins, but what’s with that white area on the tail? I remain confused. Image: Darren Naish.

  **Caption:** hirundines of, I think, two different species. The bird in flight at left is a Sand martin like those shown above, but the perched one at right is dark dorsally and has a dark, V-shaped area on its breast. These features mean that it has to be Barn swallow. Image: Darren Naish.

Warblers. Ok, I need to talk about warblers. Because boy did I see a lot of warblers. They fell into three groups: leaf warblers, acro warblers, and sylviid warblers. As covered on Tet Zoo before (see this article from February 2022), Old World warblers are part of Sylviida (formerly Sylvioidea) – the great passerine group that includes babblers, bulbuls, white-eyes, cisticolas and so on – and those groups we call ‘warblers’ aren’t close relatives (e.g., Oliveros et al. 2019).

Because I know you all love being reminded about the basic structure of the passerine family tree, here’s a reminder of the basic structure of the passerine family tree (albeit sticking only with Passerides), modified since the last time it appeared here…

**Caption:** a more elaborate version of the warbler-flycatcher-sparrow section of passerine phylogeny than we’ve seen here before, using the ‘up-ranked’ taxonomy used by Cracraft *et al*. (2004) and then modified by Oliveros *et al*. (2019). The clade we’re seeing here is now termed Passerides (whereas before it was Passerida), which means that the clades previously termed Sylvioidea, Muscicapoidea and Passeroidea are now up-ranked to Sylviida, Muscicapida and Passerida. These illustrations and the cladogram itself are part of my in-prep textbook project, which will one day be finished, I promise. **Support it – and other projects – here.** Image: Darren Naish.

Leaf warblers, or phylloscopids, are a mostly greenish, slender-billed group, often bearing a prominent supercilium (a pale eyebrow stripe). The numerous species are often difficult to tell apart and experts use song and specifics of wing feather length to differentiate them, features that can’t be applied to fleeting observations of live birds seen at distance. The leaf warbler you see here was perching on boulders and branches adjacent to the Kafinigan River in the Romit Valley. It has a yellowish supercilium, olive green mantle, a pale ‘covert bar’, light green secondaries, darker primaries, and legs that have pinkish and yellowish tones. It’s not long-billed and looks (on the basis of eye size and proportions) ‘mid-sized’ within the group. The species that seems to be the best match is the Greenish warbler Phylloscopus trochiloides, and that would make sense as this is one of the most widespread and common leaf warblers of the region. Other identifications are possible; let me know what you think if you know leaf warblers!

**Caption:** a mid-sized, fairly green leaf warbler, seen in the Romit Valley, that was moderately co-operative. I *think* it’s a Greenish warbler. Image: Darren Naish.

Here's another warbler, and this time it’s not a leaf warbler. The profile of its head and tendency to adopt a ‘skulking’ posture made me think that it might be an acro: an Acrocephalus warbler, the group that includes reed warblers. It’s brown, greyish ventrally, has a poorly defined supercilium (with some yellow just above the eye), a dark stripe through the middle of the eye, and a pale area adjacent to the wrist. The yellowish tint to the throat and belly is due to reflection from the adjacent leaves, I think. It’s also long-billed and has greyish legs.

**Caption:** a possible acro warbler that’s possibly a Blyth’s reed warbler. In this image, the bird is in a low, ‘skulking’ pose, with the feathers on its head depressed close to the skull. However, look at the other photo below to appreciate the different profile the bird has when not doing this. Image: Darren Naish.

  **Caption:** the exact same bird shown in the photo above, this time with a more erect pose and raised head feathers. Small birds often reflect the colour of nearby foliage on their pale parts meaning that they can look darker or lighter than they do in the hand, or in the field guide. Image: Darren Naish.

This description is most consistent with two species that occur in the region – Paddyfield warbler A. agricola and Blyth’s reed warbler A. dumetorum – and, of the two, Paddyfield warbler is my preferred identification because it’s a confirmed denizen of Tajikistan (Blyth’s reed warbler presence in Tajikistan is marked with question marks by Ayé et al. (2012)) and because it has a deeper supercilium than Blyth’s reed warbler, as does the bird I saw. Again, I could be wrong and I welcome alternative suggestions. UPDATE: following comments (see below), I’ve changed this and am now preferring the Blyth’s reed warbler suggestion.

A second, bigger acro was seen at Saratok, in the Fann Mountains north of Dushanbe. My only photos are fuzzy. This bird was warm brown dorsally, greyish ventrally, had a tall, dark forehead, a thin yellowish supercilium that extended a short distance behind the eye, short primary projection, and pinkish legs. Several possible identifications exist for this one, with Blyth’s reed warbler again on the list in addition to – outside bet – the poorly known Large-billed reed warbler A. orinus, which is present in Tajikistan but not well recorded there. I keep coming back to the possibility that it might be a Blunt-winged warbler A. concinens, since it looks so right. According to eBird, this species is confirmed for western Tajikistan, so this isn’t impossible. But maybe it’s just a Eurasian reed warbler A. scirpaceus in especially good light (they normally look duller brown and darker at the wingtips). Thoughts appreciated.

**Caption:** blurry photos of a warm brown warbler that has a tall forehead, yellowish supercilium and short wings. Am I right in thinking that this is an *Acrocephalus* warbler, and – le shock – is it a Blunt-winged warbler? Images: Darren Naish.

Finally, what about sylviids? I saw these on numerous occasions, both as singletons and in small groups of 2-3, and assumed I was seeing Ménétriés’s warbler Sylvia mystacea or a close relative. But no: the birds I photographed were relatively large, entirely white ventrally, and with a sharp line demarcating the white submoustachial area from the dark grey ear coverts. These features mean that they’re most likely Eastern orphean warbler Curruca crassirostris, a species that occurs from the Balkans east to eastern Kazakhstan and eastern India. I saw these low in the Karatag Valley as well as near Lake Timur Dara (higher in the valley).

**Caption:** a montage showing what I think are Eastern orphean warblers. The ‘orphean’ in the name (there’s a Western orphean warbler *C. hortensis* too) is a reference to Orpheus of Greek mythology, the musician, poet and singer. I presume the name is a reference to the singing of these birds. Images: Darren Naish.

  **Caption:** more Eastern orphean warblers. The individuals with light grey heads shown here must be female, but the bird on the right – with the very dark head – would appear to be a male. Images: Darren Naish.

Thrushes, Old World flycatchers, and starlings. I saw two thrushes: Mistle thrush Turdus viscivorus (no photos) and Eurasian blackbirds T. merula. The latter were seen on numerous occasions, though the only bird I got useable photos of was in a shabby state. I also had the best views I’ve ever had of Common nightingale Luscinia megarhynchos. This bird was foraging on ground adjacent to an orchard and often cocked its tail.

**Caption:** female Eurasian blackbird, photographed foraging in a vegetable garden. The feather loss on the head might be due to disease. Image: Darren Naish.

  **Caption:** I’ve never had especially good views of nightingales before. This bird, photographed adjacent to a gardened shrine in the village of Hakimi, was especially co-operative in its posing. Images: Darren Naish.

Muscicapidae – the so-called Old World flycatcher family – is a large group that includes wheatears, chats, robins proper, nightingales, flycatchers proper (Muscicapa, of course, and kin) and birds with the word ‘thrush’ in their name, like rock thrushes. They are of course the core group within Muscicapoidea/Muscicapida, a clade that also includes waxwings and kin, kinglets, and creepers, wrens and kin (Certhioidea).

**Caption:** more Spotted flycatchers (**others were shown in part 1**). It might just be my bad colour vision (or the settings on my PC), but the one on the left has a pinkish hue to my eyes. *Muscicapa striata* was scientifically named by Simon Pallas in 1764 but included by him in *Motacilla*. The content of *Muscicapa* expanded after it was named by Mathurin Brisson in 1760 but recent studies have shown that its traditional version is polyphyletic, a consequence being that some species have been placed in other genera or given their own generic names. Images: Darren Naish.

As mentioned in the previous article, Spotted flycatchers Muscicapa striata were seen in some abundance, and the countryside was obviously ideal for their needs and preferences. I photographed as many individuals as I could, since it’s sometimes interesting to see variation (or the converse) within populations. I was aware that a second, similar flycatcher species – the Rusty-tailed flycatcher Fidecula ruficauda – occurs in the region, and was hoping that I might see it. And as you can see from this photo: I did! Ayé et al. (2012) of course have this as a species of Muscicapa, but molecular studies published more recently show it to instead be part of Ficedula (Hooper et al. 2016). For a time there were suggestions that it should get its own genus (Ripleyia, later changed to Ripleyornis due to preoccupation by a fossil mollusc).

**Caption:** Rusty-tailed flycatcher in the Romit Valley, very helpfully displaying its key field sign. It’s partly migratory, the Central Asian populations moving to India for the winter. Images: Darren Naish.

Wheatears of several species live in Tajikistan. One species was encountered in rock-strewn fields in heavily grazed parts of the Romit Valley. Its long primary projection, white rump and white sides to the proximal two-thirds (or so) of the tail show that it was likely a Northern wheatear Oenanthe oenanthe, and not a male as they have light grey upperparts and a black mask. Incidentally, the word ‘wheatear’ originated as a corruption of ‘white-arse’, this being a reference to the white rump of this species. UPDATE: thanks to a suggestion from Simon Woolley, I’m changing this from Northern wheatear to Pied wheatear O. pleschanka on account of its higher contrast, darker plumage.

**Caption:** at left, female Pied wheatear in a boulder-strewn field. I don’t think that this is an especially controversial suggested identification. At right, what might be a Variable wheatear. If that identification is correct, this has to be the black-belled *opistholeuca* subspecies. Images: Darren Naish.

What I think was a second wheatear species – the mostly black Variable wheatear O. picata – was seen on boulders near the Kafinigan River. This species is really, err, variable and there are white-capped, white-bellied and white-rumped forms as well as populations where all of those things are absent. As you can see, the one I saw (assuming it was a Variable wheatear) was black apart from a white base and sides to the tail. This makes it the ‘black-bellied’ O. picata opistholeuca, and the good news is that this form is said to belong to northern Afghanistan and at least part of Tajikistan (Ayé et al. 2012). However, read on…

What might have been a third wheatear – though I’m even less confident about this one – was seen in an area of small fruit trees in the Karatag Valley. The bird was chat-like in form, with an off-white breast, buff flanks, belly, undertail and rump, dark wings and tail, and grey-brown dorsal plumage. I thought at the time that this might be a female Pied wheatear O. pleschanka, though most populations of this bird have a dark throat. However, the ‘vittata’ form – which might occur in Tajikistan and is of unresolved taxonomic status – has a white throat. I don’t think that this identification is right though. Any better ideas?

**Caption:** at left, a chat-like bird from the Karatag Valley that might be a female Pied wheatear. If so, it must be the white-throated ‘*vittata*’ form. At right, a female redstart, perhaps a Black redstart. Images: Darren Naish.

A scruffy looking, probably female redstart was seen in the Romit Valley. I think it’s a Black redstart Phoenicurus ochruros in view of the pale streaking on the head. Eversmann’s redstart P. erythronotus occurs in Tajikistan but only as a winter visitor, and it has obvious pale borders to its coverts.

Finally on chats and kin, what I think was Plumbeous water redstart P. fuliginosus was seen perched on rocks in the Kafinigan River. Suspiciously, this bird was only several metres away from the ‘wheatears’ suggested above to be Variable wheatear, so now I wonder if my possible ‘Variable wheatears’ were additional Plumbeous water redstarts. Sitting on boulders next to a river is not a very wheatear-like thing to do, after all.

**Caption:** possible Plumbeous water redstart. If correct, it must be a female since males are blue with a red tail. The species was formerly placed in its own genus (*Rhyacornis*) but post-2010 molecular studies have found it to belong to the redstart genus *Phoenicurus*. Image: Darren Naish.

I mentioned in the previous article that Common or Indian mynas Acridotheres tristis were among the most frequently seen of Tajikistan’s birds. This species occurs from Iran all the way to peninsula south-east Asia, but it’s been introduced to numerous other places where it’s doing quite well, including Japan, Australia, New Zealand, Fiji, various islands in the Atlantic and Indian Ocean, and parts of Canada (!) and of course Florida. Given how adaptable the species is, it certainly has the potential to out-compete natives, so this spread is a source of concern.

Acridotheres is a starling (Sturnidae). Various different affinities have been suggested for starlings in the past, among the most popular being that they’re corvoids or close to icterids and kin. Molecular data shows that they’re part of Muscicapida, and in fact close to thrushes and Old World flycatchers.

**Caption:** Common mynas were seen regularly in gardens, in fields with livestock, and in villages. White patches on the dorsal surfaces of the wings are very prominent in flight. Image: Darren Naish.

Dippers. Unsurprisingly for a mountainous nation with abundant, fast-flowing streams and rivers, Tajikistan has dippers, a muscicapoid group also close to thrushes and Old World flycatchers.

As you’ll know if you keep up with technical publications on avian evolution and morphology, that old claim about dippers lacking any and all specializations for aquatic life is not at all accurate. Smith et al. (2022) looked at all aspects of dipper anatomy in the interests of comparing them to other wing-propelled diving birds and reported diving adaptations in the nostrils, plumage, musculature and much else. Dippers do have diving adaptations, but these are weakly expressed relative to those of other diving birds, perhaps because they’re relative newcomers to the aquatic realm (Smith et al. 2022), or perhaps because they’re ecologically distinct in doing a form of shallow-water, near-surface foraging different from that of other diving birds. It should also be added that the ‘no aquatic specializations’ claim mostly related to skeletal anatomy anyway and – as Steve Ormerod has reminded me – it’s long been known that dippers have numerous soft-tissue adaptations related to aquatic life, including in their uropygial gland, feathers, eyes, nostrils and overall proportions.

**Caption:** I saw White-throated dippers at close range on several occasions, typically as they were flying quickly up or down the Karatag River. The only co-operative ones, like this one, were observed at great distance. Image: Darren Naish.

Two species can be found in Tajikistan: the White-throated dipper Cinclus cinclus (which I know well from Europe) and the Brown dipper C. pallasii, and I was most excited about potentially seeing the latter. I’m pleased to say that I saw (and photographed) both, though a complication I was aware of is that there’s a dark brown form of the White-throated dipper (C. c. baicalensis) that lacks the white throat! However, it has a light brown neck and breast, whereas the Brown dipper is dark brown overall.

**Caption:** a Brown dipper foraging at the edge of a river. These birds are happy divers and bottom-walkers, but they also wade in fast-flowing water, as is obvious here. In the image at bottom left, the white nictitating membrane is obvious. Image: Darren Naish.

Wagtails. We now move to the final major clade within Passerides: Passeroidea/Passerida, the group that includes sparrows, finches, buntings, icterids and so on. And somewhere within this lot (surprisingly, close to sparrows and finches) are the wagtails and pipits, or motacillids.

Wagtails were seen frequently at streams and rivers during my trip, specifically Grey wagtail Motacilla cinerea and White wagtail M. alba. The white wagtails were of the ‘Masked’ subspecies M. a. personata, which is unusual in that the black throat and breast are continuous behind the eye with the black nape and cap. These two zones are separated by white in most other white wagtail forms… but not all: M. a. subpersonata of Morocco has this too, as does the very black M. a. alboides of China and the Himalayas (Alström et al. 2003).

**Caption:** the Grey wagtails of Tajikistan look essentially the same as those of Europe. It’s an especially long-tailed wagtail. Images: Darren Naish.

  **Caption:** masked, *personata*-type White wagtails, seen at different Tajik locations (but always close to rivers or streams). Images: Darren Naish.

Finches. One of the birds that surprised me the most, of all the Tajik birds I saw, actually belonged to a species – or does it? – that I already know from home, namely the European goldfinch Carduelis carduelis. I knew that goldfinches were present in the Karatag Valley before I got to look at them properly, since I heard their distinctive ‘tinkling’ calls and saw them, at distance, flying to, and feeding from, thistle heads. But when I got to see them properly, up close, I was surprised, since they were really odd-looking, being pale, lacking the black crown and nape typical of European goldfinches, and with a long, extremely pointed bill.

**Caption:** the strangest goldfinches. These photos show the same individual, but the other goldfinches I saw were like this one in bill form: it’s normal for this population. Images: Darren Naish.

Ayé et al. (2012) informally term this the ‘Eastern goldfinch’, but the taxonomy is a bit confused as more than one subspecies might be involved here (C. c. paropanisi and C. c. subulata among them). Indeed, goldfinches are variable across their extensive range and around 14 subspecies are recognized by some authors, these falling into the mostly western carduelis group and the mostly eastern caniceps group. I wonder if the long, especially pointy bill on the birds I saw is an adaptation for plants specific to the region.

A group of finches seen foraging on the ground in an orchard confused me until I realized that I was seeing a strongly dimorphic species where females are grey-brown (and whiteish ventrally) but males have a bright red head, breast and rump. I think that this is Common rosefinch Carpodacus erythrinus, a widespread Eurasian cardueline finch that, in the west, appears to be increasing its range across Europe. In the east, some populations winter in the Asian tropics.

**Caption:** on a partly bare slope at the edge of an orchard, a group of what I think were rosefinches were watched foraging on the ground. The red heads, necks and breasts of the males are obvious. Image: Darren Naish.

Mystery ‘finches’. I also saw a few mystery finches that I haven’t been able to identify. In ‘finch 1’, the plumage is nondescript and a key feature is a deep, conical bill. Reddish legs were visible in ‘finch 2’, a Karatag Valley bird. Are these more rosefinches, or are they something else, like Trumpeter finch Bucanetes githagineus? That species is not confirmed for Tajikistan, its nearest population being in Uzbekistan [UPDATE: several suggested identifications have been made for this bird in the comments; I most like Alan’s suggestion that it might be Plain mountain finch Leucosticte nemoricola]. Perched above Snake Lake (which, yes, was named for the former abundance of snakes there) was ‘finch 3’, a reddish-and-black bird with a pink, slightly down-curved bill and pinkish legs, streaked breast, and black wing and tail feathers bordered by white. It was cardueline-like in shape and stance. I’m totally stumped by this one – any suggestions?

**Caption:** a miscellany of mystery finches, or finch-like passerines. I look forward to seeing suggested identifications. Images: Darren Naish.

A finch-like bird – I think it might be a bunting – was seen on an acacia-like tree in the Karatag Valley. The tail, secondaries, coverts and tertials are black lined prominently with yellowish-brown, a pale eye-ring is connected to a stripe that separates the crown from the ear coverts, and the legs are pale. I reckon this is a female Red-headed bunting Emberiza bruniceps, a common species that’s present throughout Tajikistan and the surrounding countries. Other suggestions welcome!

**Caption:** a bunting (or similar passeroid/passeridan) with brownish, streaked plumage across the head and neck and where the wing feathers are black and lined with yellow-white. I’m suggesting that these are female Red-headed bunting, a familiar and common species in Tajikistan and adjacent nations. But I could be wrong. Images: Darren Naish.

And that – finally – is where things come to an end. You’ll recall from the first article my table of non-passerines I saw. Here’s a table showing the passerines. The count continues from the non-passerines (where 16 species were observed)…

So that’s at least 49 species in total (32 of which are passerines). Not bad for casual, fly-by birdwatching, I think. It includes a few specialities of the region, some enigmas that are tough to identify at distance, and a huge amount that was entirely new to me. Of interest incidental here, perhaps, is the less than perfect nature of various of my photos. It should be obvious that I struggle to photograph fast-moving, small animals, for various reasons. I presume that everyone who photographs live animals has the same problem, it’s just that we don’t usually have any reason to see their bad or very bad photos. Aware of the substandard nature of many of my images, I provided the following poll, and look at the results….

So, you only have yourselves to blame.

And that is that. More discussion of what happened in Tajikistan is due to be covered here in time. For now, we move away to other topics. For previous articles on birdwatching in far-flung locations, see…

To the Sahara in quest of dinosaurs (living and extinct), December 2008
From Morocco, with larks, babblers, gazelles, owls and GIANT DINOSAUR BONES, December 2008
Birding in Brazil: a view from suburban Rio de Janeiro, June 2013
Birdwatching in Suburban China, May 2019
Suburban Birdwatching in Queensland, Australia, November 2019
Birdwatching in Oregon, June 2023
Birdwatching in Tajikistan, Part 1, October 2023

Refs - -

Alström, P., Mild, K. & Zetterström, B. 2003. Pipits and Wagtails of Europe, Asia and North America. Christopher Helm, London.

Ayé, R., Schweizer, M. & Roth, T. 2012. Birds of Central Asia. Christopher Helm, London.

Cracraft, J., Barker, F. K., Braun, M., Harshman, J., Dyke, G. J., Feinstein, J., Stanley, S., Cibois, A., Schikler, P., Beresford, P., García-Moreno, J., Sorenson, M. D., Yuri, T. & Mindell, D. P. 2004. Phylogenetic relationships among modern birds (Neornithes): towards an avian tree of life. In Cracraft, J. & Donoghue, M. (eds), Assembling the Tree of Life. Oxford University Press (Oxford), pp. 468-489.

Fabre, P.-H., Irestedt, M., Fjeldså, J., Bristol, R., Groombridge, J. J., Irham, M. & Jønsson, K. A. 2012. Dynamic colonization exchanges between continents and islands drive diversification in paradise-flycatchers (Terpsiphone, Monarchidae). Journal of Biogeography 39, 1900-1918.

Hooper, D. M., Olsson, U. & Alström, P. 2016. The Rusty-tailed flycatcher (Muscicapa ruficauda; Aves: Muscicapidae) is a member of the genus Ficedula. Molecular Phylogenetics and Evolution 102, 56-61.

Jønsson, K. A., Fabre, P.-H., Kennedy, J. D., Holt, B. G., Borregaard, M. K. Rahbek, C. & Fjeldså, J. 2016. A supermatrix phylogeny of corvoid passerine birds (Aves: Corvides). Molecular Phylogenetics and Evolution 94, 87-94.

Oliveros, C. H., Field, D. J. , Ksepka, D. T., Barker, F. K., Aleixo, A., Andersen, M. J., Alström, P., Benz, B. W., Braun, E. L., Braun, M. J., Bravo, G. A., Brumfield, R. T., Chesser, R. T., Claramunt, S., Cracraft, J., Cuervo, A. M., Derryberry, E. P., Glenn, T. C., Harvey, M. G., Hosner, P. A., Joseph, L., Kimball, R. T., Mack, A. L., Miskelly, C. M., Peterson, A. T., Robbins, M. B., Sheldon, F. H., Silveira, L. F., Smith, B. T., White, N. D., Moyle, R. G. & Faircloth, B. C. 2019. Earth history and the passerine superradiation. Proceedings of the National Academy of Sciences 116, 7916-7925.

Smith, N. A., Koeller, K. L., Clarke, J. A., Ksepka, D. T., Mitchell, J. S., Nabavizadeh, A., Ridgley, R. C. & Witmer, L. M. 2022. Convergent evolution in dippers (Aves, Cinclidae): the only wing‐propelled diving songbirds. The Anatomical Record 305, 1563-1591.

Song, G., Zhang, R., Alström, P., Irestedt, M., Cai, T., Qu, Y., Ericson, P. G. P., Fjeldså, J. & Lei, F. 2018. Complete taxon sampling of the avian genus Pica (magpies) reveals ancient relictual populations and synchronous Late-Pleistocene demographic expansion across the Northern Hemisphere. Journal of Avian Biology 49, e01612.

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I’ve recently returned from the most extraordinary trip to Tajikistan in Central Asia, an expedition that I and a team of colleagues made for… well, I can’t say precisely what for, but I will say that it involved field research on enigmatic, alleged large mammal species.

**Caption:** a Tajik (non-passerine) bird montage, and a map showing Tajikistan’s location (**go here if you need a larger version**). East of Uzbekistan, south of Kyrgyzstan, west of China, north of Afghanistan. Images: birds by Darren Naish; map (c) Google maps.

I’ve longed to go to Central Asia for years but haven’t had the funds or opportunity, so this really was a dream come true. While there, I looked for wildlife, as is my want. And, naturally, I saw a good many birds, and that’s what I’m here to talk about now. A disclaimer is that I only ever have the opportunity to engage in what I call ‘fly-by’ birdwatching, by which I mean that everything is opportunistic, not ‘targeted’. No trips out to special birding places, no use of an accompanying guide, and no accruing of lists of hundred of species. Just the viewing (and photographing) of species that happen to be in the right place at the right time. One day I’ll do things right, but that day hasn’t arrived yet. It might, nevertheless, still be impressive how many species I get to see: an indication of how rich and diverse bird life still is, and how even non-specialized, non-dedicated visits can result in great luck if you’re paying attention.

Some necessary preamble. What to do when preparing to go see birds in Tajikistan? These days, I have a camera and lens (a Canon EFS 500D, with a 55-250mm lens) that’s reasonably good at photographing objects tens of metres away. Alas, it’s not great at landscape shots, meaning that I’m also using a Fujifilm FinePix S4200 (a good bridge camera) and my phone (Galaxy Samsung S9), which has an excellent camera. A consequence of constantly juggling three cameras is that I can’t also use binoculars since I only have two hands.

**Caption:** key tools… a moderately decent camera, a field guide… and my note books too. Image: Darren Naish.

A key item needed should you wish to identify birds is a field guide. The good news is that the region is well-served by one in particular: Raffael Ayé et al.’s 2012 Birds of Central Asia (Ayé et al. 2012). I had it on me at all times, and I fully recommend it.

What is Tajikistan like as a place to look for birds? Tajikistan is landlocked, mountainous and elevated (it’s mostly over 3000 m above sea level), highly rural, and with a low human population of less than 10 million. Arable and livestock farming dominate the land: orchards were common in the places we visited, and sheep and goat farming and bee-keeping were everywhere. A human impact on the landscape was present everywhere. Livestock were everywhere (don’t forget that this includes domestic honeybees) and trees across many places had been removed.

**Caption:** an indication of what the landscape is like in the Karatag Valley of western Tajikistan, the region where we did much of our fieldwork. Image: Darren Naish.

  **Caption:** more mountainous terrain in western Tajikistan, this time the infamous Anzob Pass in the Fann Mountains. Image: Darren Naish.

Having said that, Tajikistan was everything I’d hoped for, this being my first ever trip to any of the Asian ‘stan’ nations. It was a place of ridiculous mountain vistas, high and jagged, sometimes snowy, peaks, abundant raging rivers and forested slopes. The region we were in (the north-west) is temperate at altitude and subtropical in general, with mixed deciduous woodland across most of the countryside and coniferous woodland above 1500 m. Add all of this together, and it sounds like a place where you’d expect gallinaceous birds like partridges, abundant hawks, eagles and vultures, arid-adapted birds like sandgrouse and bustards, and open-country and waterside passerines like shrikes, larks, chats, dippers, pipits and finches.

**Caption:** more mountain porn. I took thousands of photos. These peaks are above the Romit Valley. Image: Darren Naish.

Familiar birds, birds of the water. Ok, let’s get to it. Two bird species were seen repeatedly at most of the locations we went to, namely White wagtail Motacilla alba and Spotted flycatcher Muscicapa striata. The area’s partly arable, well-watered, partly wooded habitats seem ideal for them. On commensal species, House sparrows Passer domesticus and feral Rock pigeons Columba livia were seen in Dashunbe, the capital and largest city in the west. Oh, I must avoid discussing the status of ‘Rock pigeons’/’Rock doves’ in Central Asia…. boy are things complicated (Stringham et al. 2012).

**Caption:** I think that these are *all* Spotted flycatchers; three of among many I saw. Images: Darren Naish.

Eurasian collared doves Streptopelia decaocto and Laughing dove S. senegalensis are additional pigeon species frequently seen in and around towns and villages in the region. I saw the latter but not the former. Common mynas Acridotheres tristis were also seen in towns and villages just about everywhere we went. I haven’t spent enough time in Asia for this to be a familiar bird and hence went to some trouble to photograph the birds I saw.

**Caption:** an urban Laughing dove that was comfortable wandering about a restaurant in search of food. I initially identified this bird as a Collared dove, in which case the black ‘collar’ that the species is named for would have to be concealed. Thanks to reader comments, I now think that the shape and reddish hue demonstrates that it’s a Laughing dove. Image: Darren Naish.

Tajikistan is landlocked, and the lakes we visited didn’t have notable mudflats or adjacent muddy shores. Partly for these reasons, we didn’t see much in the way of seabirds, wildfowl or waders, though a single wader was seen close to streams and pools in the Karatag Valley: the Common sandpiper Actitis hypoleucos. Two were seen, both walking around pools and also in flight. A ‘mystery wader’ seen flying past our Karatag Valley campsite, always at great speed, was likely the same species. It was very white ventrally (including on the tail) and with very dark wings, and my thoughts when photographing it was it was too white and too dark to be Common sandpiper. UPDATE: nope, this wasn’t another Common sandpiper, but a Green sandpiper Tringa ochropus, with the solid black wings being the main giveaway. Thanks to Tom Worfolk for this identification. We didn’t see any grebes, pelicans, herons or cormorants.

**Caption:** sandpipers of the Karatag Valley. The bird on the right is certainly a Common sandpiper. The blurry one on the left probably is too… but is it? UPDATE: it’s a Green sandpiper! Images: Darren Naish.

Birds of gallinaceous and columbiform sort. On gallinaceous birds, I saw a group of female See-see partridges Ammoperdix griseogularis running near the road in the barren, rocky surrounds of the Fann Mountains on our way to Lake Iskanderkul. A captive Chukar Alectoris chukar was encountered at Labidjay, near the Karatag River.

**Caption:** the Chukar is the only *Alectoris* species that occurs in Central Asia, though it’s polytypic, with around six subspecies recognized across the region. I don’t know why this one was being kept in a cage, and it obviously wasn’t being kept in a cage that’s at all adequate for its wellbeing. Image: Darren Naish.

Having mentioned pigeons above, I should add that Wood pigeons Columba palumbus were seen on several occasions when away from towns. They sometimes looked much like European wood pigeons, but a wood pigeon I photographed in flight in the Karatag Valley looked different, with olive-brown on the upper surface of the wing. I thought for a while that it might therefore be the east and south Asian subspecies C. p. casiotis… but no. That animal has a buff patch on the side of the neck, not the white one typical of C. p. palumbus. While poorly known here in the west and mostly reported in low numbers, C. p. casiotis has recently been shown to be relatively abundant in some Asian countries (Dangaura et al. 2023).

**Caption:** a montage of bad pigeon photos (I mean that the photos are bad; I’m sure that the pigeons were quite amicable). The two on the left are Rock pigeons/doves, though the little crest on the bird at upper left is interesting, and I presume a fluke caused by ruffled feathers. The big pigeon at right looks more brightly coloured than I’d expect for a Wood pigeon but does appear to be typical of the species. Images: Darren Naish.

I didn’t see any owls but did hear one while out at night in the Romit Valley. I didn’t record it and don’t remember the sound well enough to work out which species it might have been.

Raptors! I had high hopes of seeing some of the numerous accipiters, Buteo hawks and eagles that inhabit Tajikistan, and in the end I was slightly lucky, but… only slightly. While sat at camp in the Karatag Valley, a brownish, mid-sized accipiter flew directly in front of me while in pursuit of a Carrion crow. It had heavily barred ventral plumage, dark primaries and at least four narrow transverse bands on the tail. These features show that it was a juvenile Shikra Accipiter badius, a first for me. Incidentally, I would have thought that a Carrion crow was outside the range of prey for this species, but the behaviour I saw sure looked like hunting, and the crow was turning and diving as if its life depended on it.

**Caption:** a fast-flying accipiter hawk – a juvenile Shikra – in pursuit of an Eastern carrion crow *Corvus corone orientalis*. Image: Darren Naish.

  **Caption:** at left, the best image I was able to get of the Shikra as it flew past our camp. At right, a bad photo, but one that shows the evasive manoeuvres the crow was undertaking and the fact that the Shikra was actively pursuing it. Images: Darren Naish.

Tajikistan is home to at least seven eagle species. While close to the jagged peaks above the Romit Valley, we watched a large eagle soar and wheel about among the clouds. I assumed that it was a Golden eagle Aquila chrysaetos – which does occur across Tajikistan as well as in the surrounding nations – but could it have been an Eastern imperial eagle A. heliaca, another big species that occurs across the region too? The latter has a proportionally shorter tail, a straighter trailing margin to the wing, and holds its wings in a flatter pose than the Golden. Looking through my photos, I see a longish tail that makes me think Golden eagle.

**Caption:** there’s something inescapably melancholy about the sight of an immense lone eagle, soaring in the cold, cloudy, grey skies at massive altitude. I doubt the eagle feels that way though. Images: Darren Naish.

Finally, what about vultures? I really hoped to see Lammergeier Gypaetus barbatus – one of my favourite birds, and a species I never have seen in life – but no luck there. A distant griffon was seen flying over some rocky hills as we were driving to the capital near the end of the trip. I hoped it would be a Himalayan griffon Gyps himalayensis but the brown (rather than near-white) underwing showed that it was instead a Eurasian griffon G. fulvus, a species I’ve seen at pretty close range in Spain. Whatever, I was very pleased to see it.

**Caption:** Old World vultures are among my favourite birds. The Eurasian griffon is, to date, the only species I’ve seen in the wild. Image: Darren Naish.

Woodpeckers, bee-eaters, hoopoes. Woodpeckers and kin (the piciforms) and the rollers, kingfishers, bee-eaters and their kin (previously lumped together as coraciiforms) are united within the Picocoraciae…. wow I hate words with too many vowels. What’s Tajikistan like for birds of that sort?

A single woodpecker occurs in the region, the White-winged woodpecker Dendrocopos leucopterus. This is very similar to the Great spotted woodpecker D. major, a species I see on regular basis at home in the UK, but differs in having so much more white on the closed wing. I saw (and heard) individuals on many occasions at several locations but was mostly unsuccessful in photographing them: their flight is too fast, and when observed in trees they were usually too veiled by twigs and leaves to allow direct line of sight. Some passable photos of a foraging individual were taken in an orchard at Hakimi, a village to the east of Dushanbe.

**Caption:** I saw White-winged woodpeckers often, but this bird was the only one for which I had unobstructed views. The black (rather than red) nape shows that this is a female. Image: Darren Naish.

  **Caption:** more woodpeckerage, showing an out-of-focus bill and eye on the out-of-focus head. I deliberately included the apples in the third photo so we can be sure that the bird was foraging in an apple tree. Images: Darren Naish.

European bee-eaters Merops apiaster were abundant across the region and I saw them on numerous occasions, often hunting and catching bees. I wonder what local bee-keepers think of them. Despite talking to many local people I never thought to ask. The only ones seen at close range – typically perched on electrical wires – were witnessed while I was in a vehicle, so all my photos are of birds seen across distance. Very frustrating. I also saw Eurasian hoopoe Upupa epops, both on the ground and in flight. No good photos, only hilariously bad ones.

**Caption:** bee-eaters have an unmistakeable profile in flight. And the Common bee-eater (and others species too, I’m sure) has a colour scheme that’s extremely recognisable even at distance. The bird at left (from the Karatag Valley) is living up to its name and has a bee in its bill. Images: Darren Naish.

  **Caption:** this photo is bad, but at least you can tell that it’s a hoopoe. Well, I *think* you can tell. Image: Darren Naish.

Falcons. Moving now to that group of predatory birds that are not closely allied to other predatory birds – I’m speaking of falcons – I had reasonable luck. Tajikistan has a diverse falcon assemblage: at least two of the larger species (Peregrine Falco peregrinus, Barbary F. pelegrinoides) occur there, as do Common F. tinnunculus and Lesser F. naumanni kestrels, Merlin F. columbarius and Eurasian hobby F. subbuteo.

A somewhat surprising, unusual falcon was seen perching on the stone-covered bank of the Kafinigan River in the Romit Valley. It was big, stocky, pale, finely barred on its light grey ventral surface and seemed to lack a dark mask. My thinking at the time (and still) is that it was a Saker F. cherrug, which is interesting as this is a species whose range in Tajikistan is marked with question marks in the field guide (Ayé et al. 2012). It definitely was present in the country in the early 20th century and migrant birds were reportedly still being captured for falconry as recently as the 1990s (Dixon 2009). The camera resolutely refused to focus on it (it was lightly raining at the time, which can mess with the autofocus function), so no photos.

Kestrels of undetermined species were seen while on the road. A pair of distant falcons seen high above us in the Romit Valley, sometimes landing on a rugged cliff-face, were thought (on account of a dark mask) to be peregrines. However, the photos I got revealed that they were a bit smaller than you’d expect for peregrines, and red underneath the tail. This means that they can only have been hobby, which is said to be common across the region (Ayé et al. 2012).

**Caption:** poor photos of Eurasian hobby seen in the Romit Valley of western Tajikistan. Image: Darren Naish.

That brings an end to this discussion of the non-passerines I saw in Tajikistan. 16 species: not bad, but not great… though let me remind you again about fly-by birding. Here’s a table…

Ok, we’ll end things there for now, since next we come to the elephant bird in the room: the passerines or perching birds. I saw many, and sometimes got useable photos of them… though mostly I did not. Tune in next time! For previous articles on birdwatching in far-flung locations, see…

To the Sahara in quest of dinosaurs (living and extinct), December 2008
From Morocco, with larks, babblers, gazelles, owls and GIANT DINOSAUR BONES, December 2008
Birding in Brazil: a view from suburban Rio de Janeiro, June 2013
Birdwatching in Suburban China, May 2019
Suburban Birdwatching in Queensland, Australia, November 2019
Birdwatching in Oregon, June 2023

Refs - -

Ayé, R., Schweizer, M. & Roth, T. 2012. Birds of Central Asia. Christopher Helm, London.

Dangaura, H., Tiwari, V., Chaudhary, S., Dangaura, K. D., & Chaudhary, A. 2023. Record numbers of Common Wood Pigeon (Columba palumbus casiotis) observed in western Nepal during December 2022. Nepalese Journal of Zoology 7, 60-64.

Dixon, A. 2009. Saker Falcon breeding population estimates. Part 2: Asia. Falco 33, 4-10.

Stringham, S. A., Mulroy, E. E., Xing, J., Record, D., Guernsey, M. W., Aldenhoven, J. T., Osborne, E. J. & Shapiro, M. D. 2012. Divergence, convergence, and the ancestry of feral populations in the domestic rock pigeon. Current Biology 22, 302-308.

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For something like four decades, Dr Alan Feduccia of the University of North Carolina has been arguing that everyone is wrong about dinosaurs….

Feduccia’s primary contention – expressed in technical papers and opinion pieces but mostly in his books *The Origin and Evolution of Birds* (Feduccia 1996) and *Riddle of the Feathered Dragons* (Feduccia 2012) – has been, and remains, that birds cannot have evolved from theropod dinosaurs but instead have their origins in non-dinosaurian, climbing quadrupedal reptiles of the Triassic. He holds this view because he regards theropods as too specialized for terrestrial, cursorial life and because he contends that birds, bird flight and feathers can only have evolved in an arboreal, ‘trees down’ setting.

**Caption:** the Feduccian oeuvre, these books being (left to right) from 1996, 2012 and 2020. The more recent works have a fringe, contrarian taint.

Unfortunately for Feduccia, the ‘birds are theropods’ paradigm is well supported and firmly embedded within the modern view of what dinosaurs are. Not only is the origination of birds regarded as a significant event within dinosaurian history, worse is that aspects of avian anatomy, behaviour and biology have been identified in, and extrapolated for, dinosaur groups of all sorts. Put another way, the view that non-bird dinosaurs were near-avian is very much part of mainstream thinking.

Romancing the Birds and Dinosaurs: Forays in Postmodern Paleontology (RTBAD from hereon) is not an instruction manual for palaeozoophiles, nor does it include homage or reference to the 1984 movie Romancing the Stone. Rather, it’s composed of 23 essays on the state of dinosaur science as Feduccia sees it today. The title is inspired by Feduccia’s idea that he’s responding to John Ostrom’s 1987 ‘Romancing the dinosaurs’, an odd choice given that Ostrom’s article can’t be considered familiar or influential. Discussions on bird origins are in the book but this might not be obvious, since about a third of its content (chapters 1-7) is devoted to criticism of work on dinosaurian physiology, on what dinosaurs are, and on the popularization of dinosaurs that has occurred since the Dinosaur Renaissance of the 1960s through to the 90s. Chapters 8-11 are mostly devoted to contentious issues in bird evolution, like palaeognath biogeography and paedomorphism, and avian digit homology.

**Caption:** alas, *Romancing the Birds and Dinosaurs* does not include dating advice, no matter how alluring those sexy dinosaurs might be. Image: **Lyn Joyce**, used with permission.

Finally, chapters 12-21 reinterpret the feathered theropods of the Jurassic and Cretaceous. Those with hair-like filaments (like little Sinosauropteryx) do not really have a filamentous pelt, Feduccia argues, but are covered in misidentified collagen fibres. Those regarded as bird-like non-birds (the turkey-like Caudipteryx, four-winged Microraptor and so on) are, Feduccia contends, misidentified members of the avian radiation, or (in the case of the small scansoriopterygids) not theropods at all. The last two chapters include thoughts on the end-Cretaceous extinction event, and a summary and call to arms.

**Caption:** a few of the main theropod taxa that get extensive mention in *RTBAD*. All belong to the theropod group Coelurosauria. *Caudipteryx* and *Microraptor* belong to the coelurosaurian group Maniraptora. Image: Darren Naish.

What is RTBAD all about, exactly? RTBAD can best be summarized as an effort to show that the view of dinosaurs promoted by specialists is chaotic, shallow and unthinking, and driven by a quest for popularity and an adherence to cult cladism.

Most modern books on fossil animals celebrate the current golden age of palaeontological discovery. RTBAD, in contrast, is infused with negativity and devoted to the idea that palaeontologists are mostly wrong about everything. Said palaeontologists are part of a cabal where essentially all work being done on bird-like dinosaurs and dinosaur-like birds is driven by an extreme form of confirmation bias: Feduccia accuses palaeontologists of practising a ‘theory-laden’, ‘monolithic’ science, and states repeatedly that conclusions are made to “accommodate the cladogram”, to fit predetermined narratives that are based on phylogenetic hypotheses. Feduccia misses the irony that his own research is transparently driven by an even more targeted form of confirmation bias than the one he erroneously identifies in others. In his efforts to show that the avian hand is not homologous with the theropod one, that integumentary filaments on theropods must be collagen fibres, and that Caudipteryx and other feathered dinosaurs are members of the avian radiation, Feduccia’s research programme has been devoted to verifying his ‘birds are not dinosaurs’ position. To this end, he has been unsuccessful, as he admits himself, at one point lamenting that his previous book “never gained popularity among paleontologists” (p. 311).

**Caption:** I will say this for Alan Feduccia… his books do feature some nice artwork (thinking in particular of the pieces by John P. O’Neill). The cover of *RTBAD* features this excellent image by Qiuyang Zheng, showing a fruit-eating jeholornithid bird while a terrestrial dromaeosaurid forages nearby. Image: (c) Qiuyang Zheng.

Because science and scientists welcome and celebrate scepticism, error-correction and out-of-the-box thinking, an argument can be made that Feduccia’s role is valuable, perhaps necessary. Throughout the book, Feduccia opines that he and a select band of colleagues are the true scientists here, the ones with a proper methodology, the ones practising true, honest scepticism, and the ones who work independently of trends in popular culture. They alone can guide the reader through the fickle, shallow, chaotic mess of claims made about dinosaurs and archaic birds by the scientifically naïve, power-hungry, juvenile popularists of our age. Why, some of these people “have Twitter accounts with large followers [sic], dealing with everything from paleontological discoveries to sports and politics!” (p. 21). Take that, Steve Brusatte!

The preface and early chapters of RTBAD are devoted to the admonishing of modern palaeontology and palaeontologists (Chapter 1: ‘Burning dim: the new theory-laden study of fossils’; Chapter 2: ‘The road to paleontological postmodernism’; Chapter 3: ‘Make it new! The dinosaur renaissance’). I’m among the terrible people that Feduccia has in mind here and receive appropriate excoriation. I especially enjoy Feduccia’s inability to apply due diligence to an article I published on April 1st (‘Mass Survival of Multitudinous Dinosaur Lineages across the K–Pg Boundary’).

**Caption:** there are several reasons for the cessation of my April 1st articles, and one of them is that some people were and are unable to recognise them as parody. **Tricking mokele-mbembe truthers** is one thing, but **catching a Distinguished Professor Emeritus hook, line and sinker is another**. I feel so guilty.

Alan Feduccia, j’accuse. But try as I might to think of the author as a wise sage and friend of the reader, the fact is that RTBAD is cartoonishly overloaded with bias and dirty tricks. Whataboutism, intellectual dishonesty, erroneous repetition, strawmanning and naïve falsification appear throughout as the author aims to diminish positions he dislikes, the most vehement of his arguments representing nothing more than personal incredulity.

‘Whataboutism’ is the arguing tactic whereby facts and issues tangential to the matter at hand are mentioned in an effort to weaken an opponent’s position. The composite ‘Archaeoraptor’ and the misidentified Oculudentavis (pp. xii-xv) represent crass errors it’s true, and it’s also true that some theropod fossils have been reinterpreted, sometimes many times (pp. 5-6). Feduccia’s point in discussing these is to show that the scientists behind them are trying to install themselves as “new authority figures” (p. xii), and that a sort of cladistic fundamentalism has taken over. All I see is scientists trying to do science, and sometimes making mistakes. It’s not fair to say that Feduccia behaves like a creationist, but it’s difficult to avoid this comparison given that those repeat references to ‘Archaeoraptor’ look awfully similar to that eternal creationist trick of never failing to mention Piltdown man.

**Caption:** the ‘Archaeoraptor’ specimen – outed in the popular media (*National Geographic* magazine) as important in our developing ideas on bird-dinosaur evolution – proved to be a composite, or a hoax if you want, as demonstrated by Rowe *et al*. (2001). What specific relevance does this embarrassing mistake have for studies on archaic birds and allied dinosaurs at large? Well, none, and the only people who might return to it as some sort of faux dog whistle are creationists. Oh, and Alan Feduccia. Image: Rowe *et al*. (2001).

The repetition that Feduccia uses is especially telling. If you want someone to accept something that’s (at best) arguable and (at worst) wrong, keep repeating it in the hopes that they’ll accept it. Feduccia desperately wants readers to accept that the ‘plumed’ Triassic Longisquama had aerial capabilities and is relevant to avian origins, so every mention of it in RTBAD works in the idea that it was a ‘parachutist’. We even get “… the Triassic arboreal parachutist Longisquama, a small arboreal archosaur parachutist…” (p. 87). The hypothesis that Longisquama might have used its plumes in gliding is interesting, but that’s not the same as saying that it’s well supported… by which I mean that it’s not supported at all.

**Caption:** *Longisquama* (at left) and the drepanosaurids (like the two animals shown at right) are small, quadrupedal, climbing Triassic diapsid reptiles. They aren’t archosaurs or close kin of archosaurs, and there’s no good reason to think that they’re relevant to the ancestry of birds. Feduccia and his colleagues have repeatedly emphasized the possibility that these animals are closer to birds than are (non-bird) theropod dinosaurs. Image: Darren Naish.

Feduccia and the sauropod neck pose debate. Arguably the most insightful chapter in the book is that on neck posture in sauropods, and by ‘insightful’ I mean with respect to Feduccia’s vision of vertebrate palaeontology, not the debate in question. Two teams of specialists disagree on sauropod neck pose. Team 1 used digital modelling to argue that sauropods were constrained to semi-horizontal neck poses and that neck mobility was relatively restricted due to the nature of the overlapping facets between the vertebrae. Team 2 argued that x-ray data from living animals indicates that elevated neck poses should be assumed, that experimental evidence demonstrates that the overlapping facets allowed more mobility than was assumed by Team 1’s digital models, and that Team 1’s digital modelling has created an impression of certitude about neck anatomy that’s not supported by the specimens (all of which are deformed). Disclosure: I’m a member of Team 2.

**Caption:** **Taylor *et al*. (2009)** argued that data from living tetrapods – including amphibians, mammals, squamates, crocodylians and birds – show how an elevated neck posture should be assumed as the default for extinct members of the group, in the absence of compelling evidence to the contrary.

Feduccia expresses strong preference for Team 1’s position and is dismissive of Team 2’s, labelling the authors “antagonists” (p. 37). Is this because he knows of methodological reasons why Team 2’s arguments should be dismissed? No, it’s because he argues that Team 2’s conclusions are based on emotional attachment to artwork in which sauropods were depicted as swan-necked (pp. 32-33). In discussing Team 2’s position, Feduccia only cites one of the relevant papers (Taylor et al. 2009) and incorrectly dismisses the data it contains as if it comes only from domestic mammals. He also points out that sauropods wouldn’t have been swan-like in neck flexibility, but no-one promotes that anyway. It’s strawmanning.

Why is this example ‘insightful’? Feduccia appears to regard the argument for semi-horizontal neck poses as the conservative, biologically more sensible view. But arguments over matters like sauropod neck pose are complex, and numerous authors have applied different approaches to this issue. To imply that a given view on the subject can be framed as reactionary, promoted for aesthetic preference, and favoured because it’s radical is dishonest and Feduccia is, again, guilty of using strawmanning and personal incredulity as his primary weapons.

**Caption:** Feduccia’s discussion of sauropods would have the reader believe that the ‘erect neck’ hypothesis is poorly founded, and promoted by radicals aiming to upset a more conventional view. It’s interesting that Feduccia does such a bad job of discussing the work that’s actually been published on this topic. The image at left shows postulated neck mobility in *Spinophorosaurus*, from **Vidal *et al*. (2020a)**. The image at right shows *Diplodocus* in the habitual neck pose inferred by **Taylor *et al*. (2009)** on the basis of data from living tetrapods.

Of interest is that a lot has happened since Team 1 and Team 2 published their opening volleys. Studies of stress dissipation (Christian & Dzemski 2011), orientation of the vertebral column as a whole (Vidal et al. 2020a) and comparative studies between giraffe and sauropod ranges of vertebral motion (Vidal et al. 2020b) have been published. They conclude that sauropods held their necks in elevated poses. Feduccia doesn’t cite or mention any of this work of course, which is in line with his habit of ignoring evidence that contradicts the positions he prefers.

Feduccia and phylogenetics. Intellectual dishonesty or naïve falsification (or both) are at play where Feduccia discusses archosaur phylogeny, his aim being to show that the entire field is chaotic and uncertain, and that groups are artificial and poorly defined. Chapter 7 (‘Dinosaur: What’s That?’) opens with an anecdote whereby Feduccia asked an unnamed colleague to define Theropoda. Said unnamed colleague stated that “It’s whatever they say it is!” (p. 93); Feduccia goes on to note that Dinosauria is diagnosed by “hardly any morphological features other than large size and gait” (p. 94) and that those characters used to define Theropoda might be the consequence of convergent evolution to bipedality. After all, he states, we know that certain non-dinosaurian archosaurs of the Triassic were superficially theropod-like. He notes in passing that certain theropod characters are not present in all theropods, the implication being that this is a problem. This is either naïve or another dirty trick, since the phenomenon termed evolution means that not all members of a group have features present in the group’s early members.

**Caption:** a highly simplified depiction of archosaur phylogeny as we currently understand it. A number of superficially dinosaur-like members of the ‘crocodile line’ within Archosauria are known (like the poposauroid *Effigia*, shown here at the top of the diagram) and a substantial amount of anatomical data shows that they are indeed part of the croc-line clade, not the bird-line clade. The hypothesis shown here is based on a substantial number of studies that cite and discuss an enormous number of anatomical observations and discoveries. Image: Darren Naish.

A philosophical problem that Feduccia keeps bumping into here is his insistence that group membership within phylogenetics should be demonstrated by a few obvious anatomical traits. But animals are complicated objects, and if phylogenetics has taught us anything it’s that substantial amounts of data need to be analyzed if we want to recover something approaching ‘true’ phylogeny. The quest for two or three ‘key characters’ is a forlorn prospect, and pointing to a superficially dinosaur-like archosaur as if its possession of a deep snout and pointy teeth are enough to make it a dinosaur is woefully naïve given the tens of anatomical details that place it elsewhere in the archosaur tree.

Feduccia and neornithine birds. We move now to Feduccia’s content on the evolution of neornithine birds, an area that those critical of Feduccia have tended to avoid. Feduccia tries to score points in RTBAD by arguing that modern findings justify his proposal of a post-Cretaceous ‘big bang’ in neornithine evolution, the implication being that this model was, and remains, smart and innovative. I don’t understand. Feduccia may well be right that neornithines did not originate as deep in the Cretaceous as some workers have argued, but that view is hardly novel to him. Furthermore, evidence does show that the bulk of the neornithine radiation occurred just after the end-Cretaceous event: again, this is a consensus view, not one that we only accept because of his work.

**Caption:** today, both the fossil record and time-calibrated molecular phylogenies converge on a similar answer…. neornithine birds originated during the Late Cretaceous but not right at its end. This cladogram from Field *et al*. (2020) shows how the main neornithine clades had perhaps diverged by 100 million years ago. It’s still the case that the vast bulk of neornithine lineages originated after the Cretaceous, however. Image: Field *et al*. (2020).

However, evidence (example: the recently described galloanserine Asteriornis) shows that this radiation was well underway long prior to the very end of the Cretaceous, the latest work showing that the main neornithine clades (palaeognaths, galloanserines and neoavians) had diverged somewhere between 100 and 75 million years ago (e.g., Field et al. 2020). Feduccia’s own diagrams show a Late Cretaceous origination for neornithines (Feduccia 2020, p. 297), yet he contests the existence of Late Cretaceous neornithines throughout his text. It’s a confusing picture and I find it dishonest that Feduccia claims to have introduced something valuable.

**Caption:** the version of the neornithine ‘big bang’ portrayed in *RTBAD*. This version differs from ones that Feduccia has published before in that they show the neornithine radiation originating in the Late Cretaceous. Image: Feduccia (2020).

In an additional example of strawmanning, Feduccia attempts to diminish efforts to use phylogenetic systematics by saying that the recovery of a grebe-loon clade demonstrates how such efforts are doomed to failure. Yet again: yes, people make mistakes in science, and the claimed recovery of a grebe-loon clade can be put down to the inclusion of insufficient data. It’s unfair to use this as some kind of excuse to throw up your hands and stop doing science. And, hey, it’s not as if more qualitative efforts to determine the relationships between organisms are somehow superior, as is demonstrated by the wrong phylogenetic hypotheses endorsed aplenty in Feduccia’s own research (Feduccia 1977a, b, 1978, Feduccia & Olson 1980a, b).

**Caption:** it’s true, Joel Cracraft once argued that divers/loons (upper left), grebes (upper right) and hesperornithines (below) were close kin within a group termed Gaviomorphae. Other authors have found a diver + grebe clade more recently. Is this proof that efforts to determine avian phylogeny via cladistics are a waste of time, doomed to failure? No, it’s proof that scientists sometimes get things wrong. Images: diver by John Picken, CC BY 2.0 (**original here**); others by Darren Naish.

In similar vein, Feduccia attempts to mislead the reader by reeling off a list of phylogenetic hypotheses no longer regarded as correct. The examples he uses include the once-popular linking of whales with mesonychians, the suggested placing of dromornithids within ratites, and a proposed affinity between stiff-tailed ducks and the big, unusual, musk ducks. Feduccia’s point here is that phylogenetics as practiced by cladists is fatally flawed. But – again – that’s not what these examples show. They show that phylogeneticists make mistakes by failing to incorporate relevant data or, even better, data that was unknown when the hypothesis in question was proposed. This is hardly a fatal flaw but an inevitable consequence of progress.

Feduccia also includes here a claim that pterosaurs were once regarded as a sister-group of theropods. No: the claim was that pterosaurs are close kin of dinosaurs, something that still appears to be the case. Feduccia mixes this up with the claim that pterosaurs were regarded as “obligatorily bipedal” and took to flight from the ground up. While connected to the argument that pterosaurs are close kin of dinosaurs, this was presented as an argument from functional anatomy, and it’s out of place in a list of contested phylogenetic hypotheses.

**Caption:** an iconic image of the Jurassic pterosaur *Dimorphodon*, by J. Kevin Ramos, used in several of Kevin Padian’s papers and articles published during the 1980s. Feduccia is fixated on this work, since he’s frequently drawn attention to this specific view of pterosaurs (which is not endorsed by anyone other than Padian). Image: J. Kevin Ramos.

Yet another strawman is set up in Feduccia’s claim that ornithologists widely and uncritically accept the old-fashioned view that ratites owe their distribution to vicariance. This view was promoted during the 1980s by Joel Cracraft, and – yes – it’s been promoted by science writers (Feduccia quotes Richard Dawkins). But is it accepted today? A substantial amount of work has replaced it, and it’s misleading to argue that it has sway now.

Also on ratites and kin, Feduccia once again promotes the view that these birds owe their anatomy to paedomorphosis. This is a historically popular hypothesis, to be sure, but it has a lot counting against it and may be wrong: see the Tetrapod Zoology article Controversies in Ratite and Tinamou Evolution (Part I). You wouldn’t know this from Feduccia’s perspective, since he presents an entrenched position that a paedomorphic explanation for ratite morphology is so self-evident that it shouldn’t be challenged. Time and time again, Feduccia’s coverage of an issue comes across as misleading and misrepresentative of the position favoured by those working in the field.

**Caption:** the hypothesis that ratites are paedomorphic has been popular. We don’t know that it’s true; it’s a hypothesis, and **not one that matches the data all that well**. Images: Darren Naish; public domain.

That never-ending debate on avian digit homology. Over recent years, Feduccia has devoted time to the homology of the avian hand, his argument being that avian and theropod digital complements are different, and – ergo – birds can’t be theropods. He argues that his work was done “to shed light” (p. 138) on the issue. I’m cynical. I think that it was specifically pursued in an effort to erode the bird-dinosaur link (you’ll recall the comment about targeted confirmation bias). Claims that identification of avian hand digits as I-II-III “accommodate the cladogram” (p. 142) are not fair; the suggested pattern is based on homology observed across phylogeny, and this exists whether one uses cladograms or not.

**Caption:** at left, ostrich embryo right hand in dorsal view. At right, the same hand (A) compared to that of (B-F) non-bird dinosaurs. Integral to Feduccia’s identification of digits in the avian hand is the argument that ostrich embryos are pentadactyl, with those little spurs at the base being digits I and V. But is that so? Vargas & Fallon (2005, pp. 241-215) noted that these “condensations of the wing appear at a much later stage than the digital condensations of the functional digits, are very small sized, and only transiently detectable. Therefore, these mesenchymal condensations cannot be compared with much certainty to specific digits that were lost in the adult”. Feduccia (2020, p. 151) quotes the following from Henry Gee: “I can’t make much of this image; it reminds me of a Rorschach Test”. Images: left, Feduccia (2020); right, Feduccia (2002).

Any effort to understand what consensus might exist on the development of the avian hand will reveal that different, distinct models have recently been considered, and – importantly, given Feduccia’s bias – that even those finding a II-III-IV configuration (e.g., Čapek et al. 2013, de Bakker et al. 2021) are not deemed inconsistent by their authors with theropod ancestry. Some studies on gene expression show Hox genes supporting the I-II-III pattern (Vargas & Fallon 2005, Salinas-Saavedra et al. 2014), though not all do (Stewart et al. 2019, de Bakker et al. 2021). It is, nevertheless, amusing to see Feduccia bend over backwards to explain the genetic studies away, noting that “[just] because something can be shown experimentally does not mean it happened in nature” (p. 144).

In an effort to reconcile an avian II-III-IV pattern with the I-II-III generally thought correct for tridactyl theropods, some authors have proposed a ‘frame shift’: an event where the pattern of digit formation ‘shifted’ to a different location in the hand (embryonic hand formation is supposed to start with digit IV). Feduccia is dismissive, asking “what’s the point?” (p. 144), later arguing that a frame shift is nonsensical since it wouldn’t serve any function. It should be obvious why the frame shift hypothesis is worth considering, since it provides a potential explanation for otherwise surprising results. While Feduccia might be more interested in birds than other theropods, the fact is that non-bird theropods have remarkable hands too, the presence of didactyl and even monodactyl lineages meaning that we might well be talking about frame shifts within theropod manual evolution even if birds didn’t exist.

**Caption:** based on the pattern of hand anatomy as seen across evolutionary history, it looks obvious that the bird hand is formed of digits I, II and III. However, looks can be deceiving… what does molecular data say? By looking at the expression of HoxD genes in the developing hand, some studies report molecular support for the I-II-III configuration. Image: **Salinas-Saavedra *et al*. (2014)**.

As for ‘function’, we don’t know why these events occur. The fact that they’ve been reported for the hands of certain salamanders and skinks (Feduccia implies that they’re unknown outside the proposed theropod/bird example) is no more significant for the function and structure of these animals than it theoretically is for birds and other theropods.

The ‘collagen model’ for dinofuzz: bad science, not good. Another debate in which Feduccia seemingly holds a fixed, contrarian position is that concerning the integumentary filaments known from various dinosaur and pterosaur fossils. Feduccia tries hard to make the reader believe that these have been debunked as collagen fibres. This hypothesis was proposed early in the modern wave of (mostly Chinese) dinosaur and pterosaur finds and it’s hard to escape the inference that Feduccia wants these filaments to be collagenous because, again, it helps derail the idea that dinosaurs and pterosaurs might have been bird-like.

**Caption:** an opinion promoted by some in the discussion on bird origins is that being highly sceptical of dinosaur ‘fuzz’ represents a good, honest stance. And scepticism on this issue is, obviously, not ‘bad’. But immediately casting aspersions (which is what Feduccia did: Morell 1997, p. 38) was and is a sign of massive bias. These images (from Currie & Chen 2001) show two of the *Sinosauropteryx* specimens that exhibit integumentary filaments ((a) NIGP 127586 and (b) NIGP 127587), plus close-ups of the filaments preserved across their bodies (and not just on their dorsal midlines, as Feduccia and colleagues have stated). Images: Currie & Chen (2001).

Feduccia’s inference in RTBAD is that good, solid science has demonstrated a collagenous identification for the filaments. Even the most generous evaluation reveals that this is inaccurate. At best, it’s a hypothesis. “Why have paleontologists refused to consider the possibility that these filaments represent collagen fibres?” (p. 184), Feduccia asks. Because the two don’t look alike, because dinosaur filaments possess features showing that they’re keratinous, not collagenous, and because those aiming to convince others that the filaments are collagenous have either screwed up or pushed really hard – cheated, you might say – in their efforts to make collagen fibres resemble dinosaur filaments. I say ‘cheated’ because Feduccia figures anatomical preparations where collagen fibres have been teased out of bone in an effort to create objects that look like filaments. You’d be justified in asking what relevance a laboratory preparation has for fossils of fuzzy-coated dinosaurs and pterosaurs. After all, “[just] because something can be shown experimentally does not mean it happened in nature” (p. 144).

**Caption:** integral to Feduccia’s argument that filaments on *Sinosauropteryx* are collagen fibres is the interpretation of said filaments as “beneath the skin outline” (p. 185), internal to the “clearly demarcated body outline” (p. 165). In the *Sinosauropteryx* specimen (IVPP V12415) you see here (this image shows part of its tail), said ‘skin outline’ or ‘body outline’ is marked with red arrows. It’s no ‘outline’, but the demarcation between an area where the matrix has been broken off by preparation tools versus that where the matrix remains. This is the case for all the specimens concerned. Most observations used to support the collagen fibre hypothesis are of this calibre. Image: Darren Naish.

Feduccia also figures slender, multi-branched ligaments carefully extracted from modern birds and tidily photographed to show how similar they are to dinosaurian and pterosaurian filaments. Superficially similar in external appearance they might be, but where do ligaments occur? Do they project in pelage-like orientation from the outside of the body, as they do in the relevant dinosaurs and pterosaurs? No, they are unique to specific regions, like the digits and neural spines.

The only worker beside Feduccia who has properly argued for the collagenous interpretation is the late Theagarten Lingham-Soliar. Lingham-Soliar is discussed on numerous occasions throughout RTBAD, mostly because he was Feduccia’s only supporter on the issues at hand. A brief digression on Lingham-Soliar – Solly to his friends and correspondents (of which I was one) – is worthwhile.

**Caption:** at left, one of Lingham-Soliar’s ichthyosaur skin fibre images, showing the fibres very much internal to the external body outline and thus deeply embedded within the skin. At right, a close-up of ichthyosaur skin fibres as provided by Lingham-Soliar & Wesley-Smith (2008). Note that the numerous tiny fibrils do not taper toward their ends and are united in thick, rope-like structures. They don’t resemble dinosaur filaments in detail. Images: left, from Lingham-Soliar (2001), scale bar = 10 cm; right, Lingham-Soliar & Wesley-Smith (2008), scale bars = 2 microns, 1 micron in insets.

During the late 1990s and early 2000s, Lingham-Soliar published papers on the skin of ichthyosaurs, his primary interest being their complex, multi-layered networks of collagen. These are embedded within the skin and consist of overlapping, grid-like fibre arrays. They don’t look like the integumentary filaments of dinosaurs and pterosaurs. Alas, Lingham-Soliar did not hold this view himself, and between 2007 and 2015 published papers and books in which, like Feduccia, he aimed to show that the filaments on dinosaurs are misidentified collagen fibres. I’ve often wondered if Lingham-Soliar only began considering ichthyosaur collagen fibres relevant to the dinosaurian and pterosaurian data after being approached on this matter by Feduccia.

Outside of Feduccia’s writings, how were Lingham-Soliar’s arguments received? The primary response was rejection, since better and more thorough tests on the dinosaur filaments found them to be non-collagenous, external, and possessing a microscopic anatomy unique to integumentary filaments (Mayr 2010, Zhang et al. 2010, Godefroit et al. 2014, 2020, Mayr et al. 2016, Smithwick et al. 2017). Some of the features integral to Lingham-Soliar’s argument – that is, that the filaments in dinosaurs like Sinosauropteryx are misinterpreted collagen fibres – are mistakes caused by the low magnification he was using, or are misidentified preparation marks or holes in the matrix. Feduccia avoids mentioning any of this, but in discussing Smithwick et al.’s (2017) rejection of Lingham-Soliar’s proposals, he write that their study is “not based on confirmable evidence” (p. 186). As per the Hox gene example noted above, that sure sounds like a way of dodging a conclusion you don’t like.

**Caption:** Lingham-Soliar argued that several long, straight, non-tapering ‘fibres’ surrounding the fossils of the theropod *Sinosauropteryx* can be identified as collagen fibres. The features he had in mind were preparation marks made by tools, like those shown here (from **Smithwick *et al*. 2017**). Scale bars = 10 mm. Images: **Smithwick *et al*. (2017)**.

The bottom line on this whole issue is that the ‘collagen camp’ that Feduccia promotes is based on superficial similarity and assertion, and the taphonomic and microstructural work published so far does not support Feduccia’s claim that there is “collagen, collagen, everywhere!” (p. 190).

An immobile position on the flight origins debate. Feduccia’s chapter on WAIR (the ‘wing-assisted incline running’ model of flight origins) is yet another section packed with personal incredulity and strawmanning. Another of Feduccia’s immobile positions is that flight (and feathers, and birds) originated in the trees, ergo any model positing a ground-up origin can only be dismissed. WAIR – published by Ken Dial and colleagues in and around 2003 – enjoyed time in the limelight in providing a seemingly good explanation for how maniraptorans perhaps made their first forays into flight. Things began to become undone once it became clear how specialised for flight a bird has to be in order for WAIR to work, and today WAIR is not popular as an explanation for the beginnings of avian flight. Feduccia knows us better than we know ourselves though, and insists that WAIR is alive and well.

Feduccia also contends that models positing wings as providing propulsion from the ground “have come and gone” (p. 224), but that isn’t accurate either as such views have been incorporated into modern hypotheses of flight origins (Dececchi et al. 2016, Naish & Barrett 2018, Larsson et al. 2020). As for Feduccia’s statement that “no vertebrate has [evolved flight] from the ground up” (p. 225; his use of italics), maybe this explains why birds are so different from other volant vertebrate groups, all of which have hindlimbs incorporated into a wing membrane.

**Caption:** **Dececchi *et al*. (2016)** showed that at least some non-bird maniraptorans do not have the right combination of anatomical features to benefit from WAIR as originally envisioned. Image: **Dececchi *et al*. (2016)**.

A position encountered throughout the book is that a given area of contention has been settled thanks to a new discovery or study: that science there is finished, always in the direction of Feduccia's preferred perspective. What Feduccia fails to note, or perhaps is unable to perceive, is that he’s guilty of promoting specific arguments and conclusions consistent with his preferred view as if they represent the ‘last word’ on the specific issue. This is tone-deaf given the view promoted elsewhere in the book (namely, that the relationships between archaic birds and bird-like maniraptorans are difficult to disentangle). One example comes from his criticism of work on the Romanian Balaur. Feduccia doesn’t like studies that regarded it as a dromaeosaurid but is happy with one (Cau et al. 2014) that shows it to be “little more than a secondarily flightless bird”. But the study concerned – I’m one of its authors – can’t be considered a ‘last word’ on this matter given the labile nature of Balaur in phylogenetic studies.

**Caption:** interpreted as a jeholornithid-grade member of Avialae, as per **Cau *et al*. (2015)**, *Balaur* might have looked less ‘*Velociraptor*-like’ than depicted in some recontructions. But would it be fair to then describe it as “little more than a secondarily flightless bird”, as Feduccia suggests? Image: Jaime Headden, used with permission.

Physiology. Feduccia’s arguments about dinosaur physiology are also dishonest. The majority of scientists working on non-bird dinosaurs think that these animals had an ‘elevated metabolism’ of some sort. It’s likely that dinosaurs of most or all sorts were endothermic, but at least some work promotes mesothermy or heterothermy, and it could be that dinosaurs used diverse strategies. This is more in step with our understanding of living animals than a simple dichotomous view, since we know that heterothermy and even ectothermy is present in some mammals, that true endothermy has evolved in cartilaginous and ray-finned fishes, that partial endothermy is present in some lizards, and so on. It’s shades of grey, as are so many things in the world of biology.

**Caption:** modern big reptiles, like pythons and alligators, are not dinosaur-like at all. But a point made several times in Feduccia’s writings is that the existence of such animals is really very much relevant to the fact that there are (or were) non-bird dinosaurs. Images: Darren Naish.

Feduccia argues that non-bird dinosaurs were assuredly ‘cold-blooded’, and he mentions several reasons for favouring this view. One is that pythons and alligators are capable of fantastic feats of behaviour despite their physiology, therefore dinosaurs. Another is that the Mesozoic was a perpetual global hothouse. He states this with lamentable regularity, describing the Cretaceous as “monotonously hot” numerous times. It’s true that long sections of the Mesozoic were very warm. But this was not true for the entirety of Mesozoic time: dinosaurs and their contemporaries were alive when conditions were temperate, cool or even cold (Cavalheiro et al. 2021, Wang et al. 2023). The Lower Cretaceous deposits of Liaoning Province – famous for their filamentous small theropods and pterosaurs, and numerous feathered non-bird maniraptorans and archaic birds – appear to have been deposited in cool, Alpine habitats, to take one example. He mentions ‘ectothermic growth rings’ as if they’re a slam-dunk for a non-endothermic physiology, despite the fact that the old black-white view on these structures is known to be wrong. In short, Feduccia’s entire chapter on physiology is rambling and chaotic, ending with what reads like a stream of consciousness on zoological miscellanea.

**Caption:** these global temperature maps for the Valanginian in the Early Cretaceous, from **Cavalheiro *et al*. (2021)**, show projected temperatures of less than 12 degrees C across large areas of North America and Eurasia. Numerous dinosaur fossils are known from this timeframe, from the cool and cold areas. Note the sea ice projected for the polar regions too. Yes, there were long warm spells across the Mesozoic but we’re long past the time where the whole of the Mesozoic (or even whole of the Cretaceous) can be framed as a perpetual hothouse. Image: **Cavalheiro *et al*. (2021)**.

For the love of Greg Paul. Some reasonable section of RTBAD is devoted to the idea that maniraptoran theropods are misidentified birds, and that these animals are not related to other theropods, instead representing a lineage that emerged from among ‘avimorph thecodonts’ of the Triassic. After arguing for decades that animals like Deinonychus, Velociraptor and Oviraptor do not have any important similarity to archaic birds, Feduccia is now of the opinion that these dinosaurs and their kin are part of the bird clade… but are somehow not dinosaurs.

The notion that maniraptorans like Velociraptor might be the flightless descendants of Archaeopteryx-like forms is the brainchild of Greg Paul, who explained it most cogently in his 2002 Dinosaurs of the Air: the Evolution and Loss of Flight in Dinosaurs and Birds. Combine the indisputable evidence for the existence of feathers on maniraptorans with Feduccia’s hardline insistence that feathers maketh the bird, and we have Feduccia cosying up to Paul and adopting a broken version of Paul’s ‘all maniraptorans are birds’ hypothesis (broken, because Paul, unlike Feduccia, still regards maniraptorans as coelurosaurian theropods). That’s amusing given the utter disdain Feduccia has expressed for Paul in the past.

**Caption:** the late Larry Martin was first to use the term ‘Paulian School of Bird Origins’ to describe endorsement of Paul’s ‘neoflightless’ hypothesis. *RTBAD* confirms Feduccia’s endorsement of Paul, and *Dinosaurs of the Air* is the sacred text.

In order to divorce maniraptorans from other coelurosaurs and theropods, you must explain away the existence of maniraptoran-like non-maniraptorans. Feduccia attempts this by claiming that all non-maniraptoran theropods are an utterly different class of animal life from maniraptorans seeing as they have “short, stubby hands” (p. 132, p. 163); they had, he opines, so reduced their forelimbs that evolving long, maniraptoran-like ones would be in contradiction of Dollo’s Law. He makes this argument with a straight face, devoting an entire chapter (Chapter 10: ‘You Can’t Go Home Again; Dollo’s Law’, pp. 129-135) to this claim.

In reality, those coelurosaurs that are not maniraptorans – they include tyrannosauroids, compsognathids and ornithomimosaurs – have elongate, slender hands and limb proportions indicative of a trend in forelimb elongation within the group. They appear, if you’re prepared to admit it, very much like the animals you’d expect to be ancestral to maniraptorans. We know that Feduccia is aware of these animals since he has reason to mention them here and there in his writings (including in RTBAD), so the argumentation here can’t represent naivety; it can only be intellectual dishonesty.

**Caption:** contra Feduccia, it simply isn’t true that there’s a trend of forelimb reduction across those theropods surrounding maniraptorans in the family tree. As implied by the simplified phylogenetic hypothesis shown here, tyrannosauroids and ornithomimosaurs are both long-armed, like maniraptorans, and indeed are proportionally long-armed relative to theropods less close to birds. I’ve included two icons for Maniraptora because archaic maniraptorans (like therizinosaurs) are proportionally shorter-armed than the more bird-like ones. Images: (c) **Scott Hartman**, used with permission.

Among the newest of Mesozoic maniraptoran groups to be recognised are the unusual scansoriopterygids, and Feduccia devotes some attention to this group, often (but not always) wrongly referring to them as ‘scansoriopterids’ [sic].

Scansoriopterygids are fascinating animals, combining small size with a short-snouted skull, probable climbing adaptations in the hindlimb and long-fingered forelimbs equipped (in some taxa, not all) with flight membranes. They look like bird ancestors of a sort if you want (note those words) birds to have evolved from small tree-climbers, and it’s not surprising that they’re of interest to Feduccia. The problem for Feduccia is that scansoriopterygids are – like other maniraptorans – part of Theropoda, allied either to oviraptorosaurs or located within the dromaeosaurid-bird clade. Feduccia’s schtick – it’s not novel to RTBAD but was published previously by Stephen Czerkas and Chongxi Yuan, and by Czerkas and Feduccia – is that scansoriopterygids “lack salient dinosaurian features” (p. 243) and “lack definite theropod characters” (p. 247).

**Caption:** scansoriopterygids are really fascinating animals. All are small, ranging from sparrow-sized to pigeon-sized. At left, we see the *Scansoriopteryx* holotype as figured by Stephen Czerkas and Feduccia. The scale bar is 10 mm! At right, a life reconstruction of *Yi qi*, a member of the group with membranous wings supported by a long, spur-like additional forelimb element. Images: Czerkas & Feduccia; John Conway (**original here**).

The hypothesis here, clearly, is that scansoriopterygids are relevant to bird ancestry but are not theropods or dinosaurs. Again, we’re seeing what Feduccia would like us to think is supported, rather than what is supported. The claim that scansoriopterygids are so different from theropods (and other dinosaurs) that they might not belong to this group is false, since theropod features are present throughout their skeletons. Furthermore, the scansoriopterygid features that Feduccia regards as making them ‘different’ from other theropods (like an especially long third finger, a propubic pelvis, partially closed acetabulum and lack of a supra-acetabular crest on the ilium) are not a problem for their theropod identity: it’s just that scansoriopterygids are weird, and most of these features evolved elsewhere within theropods anyway.

Consilience, not consensus. RTBAD’s final chapter claims to represent ‘A search for consilience, not consensus’. Feduccia uses it as a reminder of the fixed nature of his position, all while admonishing those who support a dinosaurian ancestry for birds as making extraordinary claims – predictably, he cites Sagan; p. 313 – that haven’t been subjected to appropriate scepticism. Waitaminute. The feathered maniraptorans are clearly close kin of compsognathids, tyrannosauroids and so on, which are themselves clearly close kin of allosauroids and so on. To argue at this point that maniraptorans descend from quadrupedal, arboreal non-dinosaurian reptiles is an extraordinary claim that should be subjected to appropriate scepticism. It has, and has been found not just wanting, but inconsistent with all the data we have.

**Caption:** maniraptorans (the theropod dinosaur group that includes birds) do not exist within a phylogenetic vacuum. That is, they are not somehow unlike all other theropods, and share numerous anatomical features with ornithomimosaurs, compsognathids and tyrannosauroids: so many, that any phylogenetic position favoured for maniraptorans has to ‘pull’ those other groups along too. This image shows a highly simplified phylogenetic hypothesis for these groups. Image: Darren Naish.

Further irony is evident from Feduccia’s reiteration of his mantra that “if it has feathers and avian flight wings, it’s a bird” (p. 312). Given the fact of evolution, what about an animal with ‘prototype’ or incipient versions of avian flight wings, and what about animals with integumentary structures that look like incipient feathers? Such animals are predicted to exist – even better, we’ve found them in the form of non-maniraptoran coelurosaurian dinosaurs – but Feduccia is here reminding us that they cannot exist within the paradigm he wants us to accept. That’s not just unscientific, it’s anti-scientific. You’re supposed to favour hypotheses and theories that are built on data, not the opposite!

All of which leads me to my final point. As might be obvious, I did not enjoy this book. I was perpetually frustrated by the author’s cherry-picking of data points and citations, his dismissal and attempting rubbishing of contemporary views of the Mesozoic world, and his persistent promotion of the idea that scientists working on dinosaurs are desperate self-popularists and megalomaniacs operating within a cult. As discussed above, his use of whataboutism, strawmanning, naïve falsification, repetition and more are obvious throughout and make RTBAD frustrating, biased and intellectually problematic.

**Caption:** for decades, Alan Feduccia has promoted the view that he and his colleagues “know birds”, and that those who promote the ‘birds are dinosaurs’ hypothesis do not. This has never been true, and it’s becoming increasingly less true over time. Whatever, we can turn it around and say that Feduccia certainly doesn’t “know” non-bird dinosaurs. Images: Darren Naish.

But for all this, I say from the point of view of someone fascinated by contrarian and unorthodox views in science that I cannot help but be glad that this book exists. Yes, it is a great irony that I enjoy the fact that this book has been published, and that Alan Feduccia continues to pen his contrarian, intellectually problematic, scientifically dishonest and technically misleading, wayward thoughts on the evolution and biology of dinosaurs and their kin. Do we benefit from the existence of Feduccia’s efforts to invalidate and rebuff scientific consensus, as some argue? I think the opposite is true, but I admit a guilty pleasure in enjoying the weird.

Alan Feduccia, 2020. Romancing the Birds and Dinosaurs: Forays in Postmodern Paleontology. BrownWalker Press, Irvine, Boca Raton. ISBN 978-1-59942-606-8. Softback.

Acknowledgements

This review happened thanks to the generosity of Martin Neukamm of AG EvoBio. Martin has recently written a long article on Feduccia and the use of his writings by creationists: How do we know that birds are living dinosaurs? A critical analysis of creationist argumentation. I also thank colleagues who provided commentary and checked sections of the text, namely Albert Chen, John Harshman, Gerald Mayr, Mike P. Taylor and Mathew Wedel.

For previous Tetrapod Zoology article relevant to the topics covered here, see…

A quick history of tree-climbing dinosaurs, July 2008
The skin of ichthyosaurs, September 2008
Epidexipteryx: bizarre little strap-feathered maniraptoran, October 2008
Yi qi Is Neat But Might Not Have Been the Black Screaming Dino-Dragon of Death, May 2015
The Climbing, Flying Babies of Deinonychus, June 2015
The Romanian Dinosaur Balaur Seems to Be a Flightless Bird, June 2015
The Integrated Maniraptoran, Part 1, May 2016
The Integrated Maniraptoran, Part 2: Meet the Maniraptorans, May 2016
The Integrated Maniraptoran, Part 3: Feathers Did Not Evolve in an Aerodynamic Context, May 2016
The “Birds Are Not Dinosaurs” Movement, November 2017
Controversies in Ratite and Tinamou Evolution (Part I), July 2023

Refs - -

de Bakker, M. A. G., van der Vos, W., de Jager, K., Chung, W. Y., Fowler, D. A., Dondorp, E., Spiekman, S. N. F., Chew, K. Y., Xie, B., Jiménez, R., Bickelmann, C., Kuratani, S., Blazek, R., Kondrashov, P., Renfree, M. B., Richardson, M. K. 2021. Selection on phalanx development in the evolution of the bird wing. Molecular Biology and Evolution 38, 4222-4237.

Čapek, D., Metscher, B. D. & Müller, G. B. 2013. Thumbs down: a molecular‐morphogenetic approach to avian digit homology. Journal of Experimental Zoology (Molecular and Developmental Evolution) 9999B, 1-12.

Cavalheiro, L., Wagner, T., Steinig, S., Bottini, C., Dummann, W., Esegbue, O., Gambacorta, G., Giraldo-Gómez, V., Farnsworth, A., Flögel, S., Hofmann, P., Lunt, D. J., Rethemeyer, J., Torricelli, S. & Erba, E. 2021. Impact of global cooling on Early Cretaceous high pCO2 world during the Weissert Event. Nature Communications 12: 5411.

Christian, A. & Dzemski, G. 2011. Neck posture in sauropods. In Klein, N., Remes, K., Gee, C. T. & Sander. P. M. (eds) Biology of the Sauropod Dinosaurs. Indiana University Press, Bloomington and Indianapolis, pp. 251-260.

Currie, P. J. & Chen, P.-j. 2001. Anatomy of Sinosauropteryx prima from Liaoning, northeastern China. Canadian Journal of Earth Sciences 38, 1705-1727.

Dececchi, T. A., Larsson, H. C. E. & Habib, M. B. 2016. The wings before the bird: an evaluation of flapping-based locomotory hypotheses in bird antecedents. PeerJ 4: e2159.

Feduccia, A. 1977a. The whalebill is a stork. Nature 266, 719-720.

Feduccia, A. 1977b. Hypothetical stages in the evolution of modern ducks and flamingos. Journal of Theoretical Biology 67, 715-721.

Feduccia, A. 1978. Presbyornis and the evolution of ducks and flamingos. American Scientist 66, 298-304.

Feduccia, A. 1996. The Origin and Evolution of Birds. Yale University Press, New Haven & London.

Feduccia, A. 2002. Birds are dinosaurs: simple answer to a complex problem. The Auk 119, 1187-1201.

Feduccia, A. 2012. Riddle of the Feathered Dragons: Hidden Birds of China. Yale University Press, New Haven & London.

Feduccia, F. 2020. Romancing the Birds and Dinosaurs: Forays in Postmodern Paleontology. BrownWalker Press, Irvine, Boca Raton.

Field, D. J., Benito, J., Chen, A., Jagt, J. W. M. & Ksepka, D. T. 2020. Late Cretaceous neornithine from Europe illuminates the origins of crown birds. Nature 579, 397–401.

Godefroit, P., Sinitsa, S. M., Cincotta, A., McNamara, M. E., Reshetova, S. A. & Dhouailly, D. 2020. Integumentary structures in Kulindadromeus zabaikalicus, a basal neornithischian dinosaur from the Jurassic of Siberia. In Foth, C. & Rauhut, O. W. M. (eds) The Evolution of Feathers. Springer Nature, Berlin, pp. 48-65.

Godefroit, P., Sinitsa, S. M., Dhouailly, D., Bolotsjy, Y. L., Sizov, A. V., McNamara, M. E., Benton, M. J. & Spagna, P. 2014. Response to Comment on ‘A Jurassic ornithischian dinosaur from Siberia with both feathers and scales’. Science 346, 434.

Larsson, H. C. E., Dececchi, T. A. & Habib, M. B. 2020. Navigating functional landscapes: a bird’s eye view of the evolution of avialan flight. In Pittman, M. & Xu, X. (eds) Pennaraptoran Theropod Dinosaurs: Past Progress and New Frontiers. Bulletin of the American Museum of Natural History 440, American Museum of Natural History, New York, pp. 321-332.

Lingham-Soliar, T. 2001. The ichthyosaur integument: skin fibers, a means for a strong, flexible and smooth skin. Lethaia 34, 287-302.

Lingham-Soliar, T. & Wesley-Smith, J. 2008. First investigation of the collagen D-band ultrastructure in fossilized vertebrate integument. Proceedings of the Royal Society B 275, 2207-2212,

Mayr, G. 2010. Response to Lingham-Soliar: dinosaur protofeathers: pushing back the origin of feathers into the Middle Triassic? Journal of Ornithology 151, 523-524.

Mayr, G., Pittman, M., Saitta, E. T., Kaye, T. G. & Vinther, J. 2016. Structure and homology of Psittacosaurus tail bristles. Palaeontology 59, 793-802.

Morell, V. 1997. The origin of birds: the dinosaur debate. Audubon 99 (2), 36-45.

Naish, D. & Barrett, P. M. 2018. Dinosaurs: How They Lived and Evolved. The Natural History Museum, London.

Olson, S. L. & Feduccia, A. 1980a. Relationships and evolution of flamingos (Aves: Phoenicopteridae). Smithsonian Contributions to Zoology 316, 1-73.

Olson, S. L. & Feduccia, A. 1980b. Presbyornis and the origin of the Anseriformes (Aves: Charadriomorphae). Smithsonian Contributions to Paleobiology 323, 1-24.

Rowe, T., Ketcham, R. A., Denison, C., Colbert, M., Xu, X. & Currie, P. J. 2001. Forensic palaeontology: the Archaeoraptor forgery. Nature 410, 539-540.

Salinas-Saavedra, M., Gonzalez-Cabrera, C., Ossa-Fuentes, L., Botelho, J. F., Ruiz-Flores, M. & Vargas, A. O. 2014. New developmental evidence supports a homeotic frameshift of digit identity in the evolution of the bird wing. Frontiers in Zoology 11: 33.

Smithwick, F. M., Mayr, G., Saitta, E. T., Benton, M. J. & Vinther, J. 2017. On the purported presence of fossilized collagen fibres in an ichthyosaur and a theropod dinosaur. Palaeontology 60, 409-422.

Stewart, T. A., Liang, C., Cotney, J. L., Noonan, J. P., Snager, T. J.& Wagner, G. P. 2019. Evidence against tetrapod-wide digit identities and for a limited frame shift in bird wings. Nature Communications 10: 3244.

Taylor, M. P., Wedel, M. J. & Naish, D. 2009. Head and neck posture in sauropod dinosaurs inferred from extant animals. Acta Palaeontologica Polonica 54, 213-220.

Vargas, A. O. & Fallon, J. F. 2005. The digits of the wing of birds are 1, 2 and 3. A review. Journal of Experimental Zoology (Molecular and Developmental Evolution) 304B, 206-219.

Vidal, D., Mocho, P., Aberasturi, A., Sanz, J. L. & Ortega, F. 2020a. High browsing skeletal adaptations in Spinophorosaurus reveal an evolutionary innovation in sauropod dinosaurs. Scientific Reports 10 (6638).

Vidal, D., Mocho, P., Páramo, A., Sanz, J. L. & Ortega, F. 2020b. Ontogenetic similarities between giraffe and sauropod neck osteological mobility. PLoS ONE 15 (1): e0227537.

Zhang, F., Kearns, S. L., Orr, P. J., Benton M. J., Zhou, Z., Johnson, D., Xu, X. & Wang, X. 2010. Fossilized melanosomes and the colour of Cretaceous dinosaurs and birds. Nature 463, 1075-1078.

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A great many ducks are remarkable. Among them is the Muscovy duck Cairina moschata. Let’s just consider for a moment how remarkable Muscovy ducks are…

**Caption:** my hope with respect to this image is that it takes you a few moments to work out what’s going on. It’s a male Muscovy duck, eyes closed, preening his chest with his head inverted. This was a captive bird photographed on La Palma in the Canary Islands in 2014. Image: Darren Naish.

Before I continue, some caveats: the Muscovy duck occurs as a wild species in the southern USA, Mexico, and then south to Argentina and Uruguay. However, most of us know it as a domestic species, and all my images here are of the domestic form. The name ‘Muscovy duck’ is usually said to reflect a confused connection to the Moscow region, one suggestion being that this is due to shipment of the birds by a European trading company called the Muscovite or Muscovy company. But maybe this is nonsense, and there are other suggestions that the name is a corrupted reference to an indigenous South American culture or location once associated with the bird. Another suggestion is that the bird got its English name merely to mark it as unusual and foreign. The equally wrong name Barbary duck is also used for the species, especially in cookery.

The biggest duck. The Muscovy duck is big and probably the biggest of all ducks, with large males reaching 86 cm in length and exceeding 7 kg. It’s highly dimorphic in size, females sometimes weighing half as much as males. Males are extravagant in terms of secondary sexual characteristics, combining a feather crest on the crown, a pronounced knob at the bill base, and (often red, sometimes black) carunculated, naked skin across the face. An especially curved, large claw is present on the second toe. It’s used in fighting: a Muscovy duck will hold another duck (by the neck) with its bill while beating it with its wings and raking with the claws. The Muscovy is quite pneumatic in the skeleton, with pneumatic invasions of the coracoid that are otherwise not common in ducks (O’Connor 2004).

**Caption:** a domestic male Muscovy duck revealing key facial features of this species. Note the naked facial skin (in this individual, there are few carunculations, maybe because he’s young), prominent knob at the base of the bill, and feather crest. Image: Darren Naish.

What sort of a duck are you, exactly? On the issue of phylogenetics and correct taxonomic placement, the Muscovy duck has proved a bit of a problem. I definitely prefer (because that’s how this works…) the hypothesis that it’s a tadornine: part of the group that includes shelducks and sheldgeese (Livezey 1997, Johnson & Sorenson 1999, Sun et al. 2017). This placement would explain a few unusual features of Muscovy duck biology, since they’re like shelducks and unlike most anatine ducks in being polygynous with a prominent degree of sexual size dimorphism, and in being cavity nesters that produce relatively large clutches (9-11 eggs) (Livezey 1996).

**Caption:** numerous competing views of duck phylogeny exist. In this highly simplified version (based on **Sun *et al*. 2017**), tadornines are closer to dabbling ducks than to seaducks. Muscovy ducks are part of the *Tadorna* lineage in the study concerned. The images used here were created for my in-prep textbook project, which is still underway and **can be supported at my patreon**. Image: Darren Naish.

This suggested placement has been disputed on anatomical grounds, however, where studies have tended to find Muscovy ducks to be anatines close to pygmy geese (Nettapus) and within a clade that includes diving ducks (aythyins), dabbling ducks (anatins) and others (Livezey 1997). Some molecular studies also find Muscovy ducks to be part of the aythyin + anatin lineage rather than the Tadorna one (Donne-Goussé et al. 2002). An older idea, proposed mostly on the basis of behaviour, is that Muscovy ducks are close to perching ducks (the mostly extravagant [in males] Aix ducks, Brazilian teal Amazonetta brasiliensis and so on), since they all share weak or absent pair bonds, reduced or absent precopulatory displays and have similar looking ducklings (Johnsgard 1961). There’s a long tradition of paying lots of attention to mating displays and so on in wildfowl and emphasising their significance in elucidating relatedness, but I think it’s fair to say that we need to combine these traits with other lines of evidence and not rely on them as being all that meaningful… not that anyone does these days, mind you.

Anyway… it might be that Muscovy ducks are related to both perching ducks and tadornines, since some molecular studies find Aix and Cairina to belong in a clade with Tadorna (Sun et al. 2017).

**Caption:** the sexually aggressive proclivities of the Muscovy aren’t in evidence *all* the time. This male (photographed in the Welsh Mountain Zoo in 2014) was allopreening his female companion. Image: Darren Naish.

Domestication. Another reason that the Muscovy duck is special is because it’s been domesticated, this obviously being a wholly separate, South American event from the Eurasian domestication of the Mallard Anas platyrhynchos. Artistic evidence (both images on pottery, and sculpture) demonstrates that the domestic Muscovy was present in Peru round about a thousand years ago (Gamboa 2019) and equally old evidence comes from Ecuador (Stahl et al. 2006); especially old Muscovy bones revealing signs of domestication are known from Bolivia and date to something like the 10th century (Gamboa 2019). So far as I can tell, a specific place of domestication hasn’t been identified, and all indications are that the birds were traded across South America.

Europeans took Muscovy ducks back to Europe (where they were first written about in the 1550s), and here they were bred and crossed with domestic Mallards. They’ve also been taken to African countries, and to India and China, where new local forms have been bred. Genetic studies of these African and Asian variants are underway.

**Caption:** Muscovy duck sexual behaviour is aggressive, with most (maybe all?) events involving rape. Male Muscovies will rape other birds, as evidenced by this photo taken on a Welsh farm in 2014. Image: Darren Naish.

Hybrids combine the traits of both species, being fast-growing like Mallards but reaching the large size typical of the Muscovy. They’re sometimes infertile (hence the name ‘mulard’, originating from ‘mule mallard’) but sometimes not. If Muscovy ducks are tadornines as discussed above, the very existence of these hybrids is remarkable since we’re talking about genetic pairing between the members of groups that have been separate for tens of millions of years (at least since the Early Miocene if certain fossils are tadornines as proposed: Worthy & Lee 2008).

I’ve never eaten Muscovy duck meat but apparently it tastes something like beef and is often much darker than that of other domestic ducks.

As ever, there is much more that could be said but my time is up. For previous Tetrapod Zoology articles on duck and other wildfowl/waterfowl, see…

Attack of the flying steamer ducks, December 2008
Lo, for I have seen the Meller’s duck, and it was good, August 2009
The Madagascar pochard returns (again), August 2009
Pink-headed duck and Red-crested pochard: who would win in a fight?, September 2009
Duck sex: to interfere, or to watch?, March 2010
Can you raise reindeer on goose shit? Amazing waterfowl facts part I, June 2010
Death by toxic goose. Amazing waterfowl facts part II, June 2010
Detachable wing-daggers. Amazing waterfowl facts part III, June 2010
Stinky seal-ducks. Amazing waterfowl facts part IV, June 2010
Laysan Ducks: Not as Degenerate as People Used to Think, August 2023

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

Refs - -

Donne-Goussé, C., Laudet, V. & Hänni, C. 2002. A molecular phylogeny of anseriforms based on mitochondrial DNA analysis. Molecular Phylogenetics and Evolution 23, 339-356.

Gamboa, J. 2019. The modern ontological natures of the Cairina moschata (Linnaeus, 1758) duck. Cases from Perú, the northern hemisphere, and digital communities. Anthropozoologica 54, 123-139.

Johnsgard, P. A. 1961. The taxonomy of the Anatidae – a behavioural analysis. Ibis 103, 71-85.

Johnson, K. P. & Sorenson, M. D. 1999. Phylogeny and biogeography of dabbling ducks (genus: Anas): a comparison of molecular and morphological evidence. The Auk 116, 792-805.

Livezey, B. C. 1996. A phylogenetic reassessment of the tadornine-anatine divergence (Aves: Anseriformes: Anatidae). Annals of Carnegie Museum 65, 27-88.

Livezey, B. C. 1997. A phylogenetic classification of waterfowl (Aves: Anseriformes), including selected fossil species. Annals of Carnegie Museum 66, 457-496.

O’Connor, P. M. 2004. Pulmonary pneumaticity in the postcranial skeleton of extant Aves: a case study examining Anseriformes. Journal of Morphology 261, 141-161.

Stahl, P. W., Muse, M. C. & Delgado-Espinoza, F. 2006. New Evidence for Precolumbian Muscovy Duck Cairina moschata from Ecuador. Ibis 148, 657-663.

Sun, Z., Pan, T., Hu, C., Sun, L., Ding, H., Wang, H., Zhang, C., Jin, H., Chang, Q., Kan, X. & Zhang, B. 2017. Rapid and recent diversification patterns in Anseriformes birds: Inferred from molecular phylogeny and diversification analyses. PLoS ONE 12 (9): e0184529.

Worthy, T. H. & Lee, M. S. Y. 2008. Affinities of Miocene waterfowl (Anatidae: Manuherikia, Dunstanetta and Miotadorna) from the St Bathans Fauna, New Zealand. Palaeontology 51, 677-708.

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Time for the second part in my Minnesota iceman series. The article you’re able to read was originally published at ver 3 (the Sci Am years) in two separate parts. For the first part on the Minnesota iceman, go here…

In the previous article – itself based on a section of text included in my 2016/2017 book *Hunting Monsters* – I mentioned the existence of a whole book devoted to the Minnesota iceman, published in 2016: *Neanderthal: the Strange Saga of the Minnesota Iceman* (Heuvelmans 2016). *Neanderthal* is a most curious and interesting book, and here is what I have to say about it….

**Caption:** cover of **Heuvelmans (2016)**, showing the composite image that Heuvelmans assembled himself. Credit: Darren Naish.

The main story of the iceman is familiar. If you need a primer, please see the previous article. As if the initial circumstances were not suspicious enough (a travelling exhibit very clearly of the ‘Is it real? You won’t believe your eyes!’ tradition of fairground gaffs, initially marketed as the ‘Siberskoya creature’ or ‘Siberskoye creature’, and viewable for a small fee to the public), we even have the name of a person stated to have been the model-maker (Howard Ball), and know of a model (today on display at the Museum of the Weird, Austin, Texas) that most people say exactly matches their recollection of what the original looked like. As justifiable as it might be to regard the whole case as ridiculous and unworthy of scientific consideration from the off, the key factor that transformed the iceman story into an international incident (see Regal 2013) is that several knowledgeable people either became convinced that it was real, or at least became interested in the possibility that it might be.

**Caption:** Bernard Heuvelmans with one of the Crystal Palace pterosaur models. The photo dates to some time in the 1950s. Credit: ***Living Wonders* Thames & Hudson Ltd; New edition edition (25 April 1983)**.

Among those convinced by its reality was the late ‘father of cryptozoology’ Bernard Heuvelmans (1916-2001) who, together with colleague Ivan T. Sanderson – also a formative character as goes writings on alleged mystery animals – examined the iceman in person in 1968. In 1969, Heuvelmans published a brief technical paper on the creature (it’s telling that the paper is sole-authored and not co-written with Sanderson; more on that in a moment) (Heuvelmans 1969). Even better, in 1974 he co-authored an entire book – L’Homme Néanderthal est Toujours Vivant (Neanderthal Man is Still Alive) – on the whole story. While mentioned in every published discussion of the iceman, this book (despite seeing a 2011 reprint) has long been both hard to obtain*, and untranslated from the original French.

Like many of Heuvelmans’s works, it has become a sought-after and expensive collector’s item.

It was thus a pleasant surprise to hear in 2015 that cryptozoologist and author Loren Coleman had learnt of Paul LeBlond’s completion of a translation, and of Coleman and LeBlond’s successful venture to see said translation published in the United States.

**Caption:** different covers of different editions. At left, the **2011 L'Oeil du Sphinx version**; at right, the **2016 English translation by Anomalist Books**. Credit: **l'Oeil du Sphinx** (left); **Anomalist Books** (right).

LeBlond – an oceanographer by profession but well known for his interest in sea and lake monsters (see previous Tet Zoo articles on the Cadborosaurus Wars) – had translated the volume in his own spare time. Retitled Neanderthal: the Strange Saga of the Minnesota Iceman (Heuvelmans 2016), the work does not include a separate section penned by Russian economist, historian and biologist Boris Porchnev, but is nonetheless very welcome. It is an inexpensive, well designed softback, and it’s most interesting – at last – to hear Heuvelmans’s full version of events.

Across 12 chapters, Heuvelmans describes all the steps in the story. It’s immediately apparent just from the titles of the chapters that he believed in a conspiracy of silence that affected his research and how it was received (there are chapters titled ‘Cloak and Dagger’ and ‘The Wall of Incredulity’). The last few chapters promise to provide his specific interpretation of the creature’s zoological and evolutionary significance (‘What it Really Was’ and ‘A History of Man-Beasts’), and it was those I was looking forward to reading the most. The book ends with a series of appendices and an afterword by Loren Coleman.

**Caption:** by far the best known of Bernard Heuvelmans’s books is his 1959 *On the Track of Unknown Animals*, republished several times and translated into many different languages. The images here show (left to right) the 1959 original, the 1965 reprint, and the 1995 edition.

Heuvelmans’s writing style is quite different from that encountered in the works most familiar to English-speaking readers (On the Track and In the Wake). He is less formal, faster-paced, angrier. It is also evident throughout the book that the whole iceman deal led to a falling-out between Heuvelmans and his friend and colleague Ivan Sanderson. Heuvelmans states on several occasions that he regarded certain of Sanderson’s statements as unwise or based on poor judgement; some of the actions concerned are well established in the public record and already noted by others as weird given that Sanderson was apparently trying to drum up official acceptance of the thing as a real carcass (Naish 2016). There are good reasons for thinking that Sanderson was playing this whole episode like a showman because, basically, that’s what he was. I’ve gradually come to the conclusion that he deliberately engaged in mystery-mongering and hype because that’s how he made a living; he was never interested in any of this stuff for honest, scientific reasons. Incidentally, Sanderson disagreed with Heuvelmans’s main premise on the iceman (that it was a Neanderthal); this disagreement is not covered in the book.

**Caption:** Sanderson's views on the iceman were somewhat different from those of Heuvelmans. At left, his ape-like rendition of the iceman (Heuvelmans is critical of this version in his book). At right, one of Sanderson's several books on animals and natural history. As usual, I am amazed how little imagery relevant to Ivan Sanderson (read: none) has been released online via creative commons. Credit:Ivan Sanderson (left); **Ivan Sanderson's *Book of Great Jungles* Julian Messner, A division of Pocket Books** (right).

Anatomy of a corpse, or not. One thing the book did not deliver on – to my disappointment, I regret – is the anatomy of the alleged ‘corpse’ itself. Don’t get me wrong – I don’t expect a popular book to include a heavy, technical discussion of anatomical minutiae. But I was expecting there to be a reasonable discussion of the various convincing anatomical details that Heuvelmans and Sanderson were reportedly impressed by. Instead we just get some brief references to vegetation seen stuck in the teeth, parasites on the skin and some discussion of head size and the anatomy of the hands and feet, not detailed disclosure and documentation of these features.

**Caption:** Frank Hansen – owner of the iceman – with the object itself. Credit: **Costello 1984 *Creatures from Elsewhere*, Imprint unknown**.

That the remarkable proportions of the iceman are not consistent with Heuvelmans’s argument that it represented a relict Neanderthal are explained in two ways. In some cases, the differences concerned (in skull size and overall height, for example) are put down to a continuing evolutionary trajectory not recorded in the fossil record (Heuvelmans even posits the specimen to be the result of “the final extreme result of accelerated Neanderthal evolution”).

**Caption:** Heuvelmans's primary hypothesis was that the iceman represented a population that descended from the Neanderthals of the Pleistocene. Here is the classic La Chapelle aux Saints Neanderthal skull. Credit: **Luna04 Wikimedia (CC BY-SA 3.0)**.

And in others (the proportionally long, slender iceman thumb: a real contrast with the Neanderthal thumb known from fossils*), Heuvelmans deliberately emphasises the minority opinions of certain scholars, positing that consensus opinion might be mistaken (he notes at least twice in the book that the iceman’s thumb is seemingly a neotenous character, an interesting hypothesis were the iceman real but one that contradicts established data on the actual Neanderthal hand). Ultimately, it’s clear – contrary to what I had always expected – that Heuvelmans did not intend this book to include the full, detailed, anatomical discussion he always promised… more on that later.

Based on Boule’s work of the early 1900s it was long thought that Neanderthals had shorter, more massive thumbs than those of modern humans. More recent work indicates that their thumbs were, relative to the rest of the hand, proportioned much like ours (e.g., Aiello & Dean 2002, Lorenzo 2015).

Cloak and dagger. Indeed, a significant part of the story concerns the remarkable ‘cloak and dagger’ tale (to use Heuvelmans’s own words) in which it is alleged that the body was smuggled out of Vietnam via an illicit drug-trafficking route. Seeing as stories about the iceman being discovered frozen in sea ice or encountered and killed in the wild woods of Minnesota are, so Heuvelmans argues, untenable, he instead contends that the body was collected by its ward – Frank Hansen – and smuggled to the USA in a remarkable instance of complex, far-sighted enterprise. Citing the 1972 book The Politics of Heroin in Southeast Asia and various political acts, events and statements relevant to Vietnam, smuggling and trafficking, Heuvelmans goes as far as suggesting as the entire drug-running operation may even have originated from the specific event involving the iceman: that it established the plan that then became standard procedure in drug-smuggling operations. As interesting a tale as this is, I found it incredibly elaborate and based wholly on freewheeling speculation.

**Caption:** like many British people of my approximate age, one of my first introductions to the iceman was its coverage in the (somewhat credulously written) PG tips *Unexplained Mysteries of the World*, published in 1987. The photo of the iceman (taken by Loren Coleman) shows exposed teeth and thus looks different from the images recorded by Heuvelmans and Sanderson. Credit: Darren Naish.

To return to the anatomy of the iceman, it appears that Heuvelmans wrote a full-length monograph on the specimen, a few hundred pages long and unpublished so far as I’m aware. Given his ability to rapidly publish the initial technical paper on the iceman in an apparently sound scientific journal (Heuvelmans 1969) it is slightly surprising that he never got this grand work into print. Presumably it sits, unpublished, in the Heuvelmans archives, and presumably it explains why this book does not include the detailed anatomical data I thought it did.

**Caption:** portrait of Bernard Heuvelmans, here shown reviewing sea monster accounts from the Oudemans archive. Credit: Hill And Wang, 1 Jan. 1969.

Heuvelmans (2016) also describes spending an entire year working on the generation of an enormously detailed illustration of the iceman. He does acknowledge that such effort might transpire to be a waste of time, but this is because he hoped that the body would one day fall into the hands of a zoological institution, not because he considered it probable that it might be a hoax.

**Caption:** just in case you've forgotten, here – again – is a depiction of the iceman as it appeared (at left), and (at right) as reconstructed by Alika Lindbergh (= Monique Watteau). Images in public domain. Credit: Darren Naish.

In some of the literature denouncing Heuvelmans’ endorsement of the iceman it is proposed that he was in an especially fragile and susceptible condition during his time in the United States due to the sad and sudden death of his daughter. I had no knowledge of these circumstances prior to reading the book and was upset by the events he described: he was away from home, unable to rapidly return, and informed that his daughter had literally weeks to live. Heuvelmans (2016) argues – and I’m inclined to agree – that such personal circumstances must not be seen as relevant to his thoughts and conclusions on an alleged frozen Neanderthal. I’m not a psychologist and others who are also not should avoid making drive-by accusations of this sort.

No replacement model. A well known aspect of the iceman tale concerns the clear anatomical difference present between the object as first examined by Heuvelmans and Sanderson, and the object as observed and photographed after the furore of 1968-69. Teeth are clearly visible in the younger photographs; a contrast with the closed mouth in the Heuvelmans and Sanderson originals. The explanation offered for this discrepancy is that the original corpse was removed and hidden (either by Hansen, or by an unhappy top-tier owner, whoever that was) and then replaced by a less realistic model. The story of the ‘replacement model’ is as much a part of the iceman legend as is the discovery of the frozen corpse in the first place. However… most surprising to me was Heuvelmans’ opinion – expressed without equivocation – that there was no replacement model, and that the object said to be such was actually the original carcass, reposed after thawing and re-freezing, and ‘hidden in plain sight’. This view flatly contradicts popular cryptozoological lore in which it is widely stated that the iceman viewed by people after the Heuvelmans-Sanderson pronouncements of 1969 was a replica and not the original.

**Caption:** a small amount of Minnesota iceman merchandise exists. At left, we have the Jean St. Jean 2020 scale figure in one of my display cabinet, a kind gift from Loren Coleman’s International Cryptozoology Museum. At right, a t-shirt from the Museum of the Weird in Austin, a gift from John Conway. Images: Darren Naish.

Here is what Heuvelmans (2016) says: “There was only one point on which my views diverged from Sanderson’s, as well as from all others who had looked into the matter, and that was on the nature of the specimen exhibited by Hansen after April 20 (1969). I was the only one to believe that it was still the actual corpse [emphasis in the original]. True, I had a definite advantage over everyone else – I was the only one to have many excellent photos of the original exhibit … I had been sent a few color slides of Hansen’s new exhibit. After a comparison with my own, I had to agree with the evidence: it was the same and only specimen [emphasis in original].”

Moving on – what was that about “non-standard evolutionary hypotheses”? This book covers two such hypotheses and discusses them in sufficient depth to make it required reading for those interested in such things.

**Caption:** Neanderthals have been depicted in many different ways over the years. This illustration is by Charles Knight and dates to 1920. Credit: Charles Knight Wikimedia.

To begin with, Heuvelmans makes statements about our views on the pattern and detail of hominin evolution that I did not find objectionable. Anyone familiar with the literature on fossil hominins will be aware of arguments whereby Neanderthals can be made to look a certain way according to the bias of whomever produced the reconstruction. Heuvelmans states, and I quite agree, that our views on hominin life appearance have frequently been influenced by our own social and cultural biases, by the way in which certain species have been framed in the evolutionary narrative – hero or villain, brutish peasant or high-born – and by our expectation of what a given animal should look like within the context of the evolutionary model favoured at the time. We increasingly pride ourselves on abandonment of the erroneous ‘march of progress’ view of evolution where members of a given lineage are perceived as half-formed intermediates heading in the direction of a given goal, or where humans are considered ‘more evolved’ than other hominins, hominids and primates. Heuvelmans has quite a modern take on this issue. So far, so good.

All of this is marred, however, by a view of Neanderthals that – while undeniably interesting (a la All Yesterdays) – is surely erroneous, and I’m left wondering whether Heuvelmans developed this view only because of his hypothesis on Neanderthal survival.

**Caption:** integral to the concept of pongoid man is the idea that Neanderthals ('late Neanderthals', anyway) had an elevated snub nose where the nostrils pointed forwards. At left, we see Heuvelmans' reconstruction of the La Chapelle aux Saints Neanderthal with a postulated outline. At right, a reconstruction of the iceman in profile. Credit: **Heuvelmans 2016**.

Neanderthals, so he explains, were likely covered by a hairy pelt (p. 173), possessed a remarkable ‘ultra-human’, upturned nose in which the nostrils pointed directly forwards (p. 179) (something like that of snub-nosed monkeys), “had no lips at all and a widely stretched mouth” (p. 180), had hands in which the thumb was both more elongate and “less readily opposable” than that of H. sapiens (pp. 182-185), had extraordinarily broad feet with curled toes that functioned in rock-climbing (p. 186), were probably capable of accruing fat stores and of indulging in a semi-hibernation (p. 211) and had “bigger eyes” that gave them “the option of vanishing into the night” (p. 211). It is also argued that Neanderthals were seen and depicted by our species as ‘beasts’ fit for hunting, extermination or even domestication as beasts of burden.

**Caption:** the cover of Danny Vendramini's book on the 'Neanderthal predation hypothesis'. Tetrapod Zoology does not endorse this book. Credit: Kardoorair Press.

The view of Neanderthal appearance and biology endorsed in the book is thus vaguely reminiscent of Vendramini’s notorious (and also erroneous) view in which, so it’s proposed, Neanderthals were black-skinned, big-eyed, hunchbacked uber-predators utterly unlike the sophisticated people of current mainstream palaeoanthropology. These ‘bestial’ views of Neanderthals might be jarring if new to you: as I might have said previously, they are, however, a mainstay of the cryptozoological literature (e.g., Loof-Wissowa 1994, Bayanov 1996, de Sarre 1996, Raynal 2001). To clarify, however, Heuvelmans does not promote his particular view of Neanderthals because he regards it as the typical condition for the species (unlike Vendramini): rather, he argues that Neanderthals became this way after evolving from ancestors more like H. sapiens. Let’s look at this idea in more detail…

De-hominisation. Heuvelmans’s view is that Neanderthals underwent a profound change as they abandoned material culture and took to a more ‘bestial’ way of life, the evolutionary process involved being termed de-hominisation. De-hominisation, as a supposed reverting to a more bestial form, is typically imagined as a sort of ‘de-evolution’. It is of course no such thing given the redundancy of that term: evolution means heritable change occurring across generations, it does not mean ‘evolution towards the specific form we have in mind as the best or most recently evolved’. Regardless, the de-hominisation hypothesis is a familiar trope of the cryptozoological literature, integral to the popular idea (within the cryptozoological research community) that Neanderthals have persisted as relict forms of remote, forested or mountainous regions where they avoid the attention of their cousin H. sapiens by being secretive, nocturnal and mostly solitary.

**Caption:** pongoid man as imagined in life by Alika Lindbergh (= Monique Watteau). The original painting features a different look for the penis and shows straggly head hair and 'ear tufts'. Credit: *Creatures from Elsewhere*; Imprint unknown.

The primary reason for the existence of the hypothesis is an attempted rationalisation of those abundant anecdotes and stories relating to hairy wild people across Eurasia. It never had a firm grounding, and it is like so many other evolutionary hypotheses in the cryptozoological literature in that it requires the existence of an entire new phase in evolutionary history – one involving profound ecomorphological novelty – for which we have no material evidence (Conway et al. 2013). Assuming for the moment that it might be worth taking seriously, it is – to repeat points made above – flatly at odds with everything we’ve learnt about Neanderthals in recent years. They have become more sophisticated, more technologically advanced and capable, more socially complex the more we have discovered; they might still have looked quite distinct from us but a view that they were, or were becoming, less like us over time is very much at odds with the evidence we have.

**Caption:** the idea that ape-type hominids evolved from human-type hominids has been put forward quite a few times in both the technical and popular literature. This *New Scientist* cover accompanied an article on that subject (Gribbin & Cherfas 1981). Credit: *New Scientist*.

Moving on, what might also surprise some are Heuvelmans’s (2016) references to evidence that hominins did not evolve from ape-like forms, but that the converse was more likely true; that anthropoid apes and humans did not descend from “some kind of archaic pongid ape like Dryopithecus. It had to be more like man than like a brachiating ape. It was probably some kind of infra-pygmy, a round-headed gnome, walking upright, in other words, the Eoanthropus imagined by leading anthropologists such as Marcellin Boule in France and Henry F. Osborn in the USA” (p. 41). The view that hominin-like proportions and posture evolved deep in hominid history – that the living non-human apes and their fossil relatives are the specialised descendants of such forms – has been revisited many times since and has at least a few modern champions. However….

**Caption:** the erroneous 'march of progress' - with humans at far right - is a familiar image. Less familiar is the idea that quadrupedal hominids evolved from bipedal, human-like ones. What we see in the illustration here is, no doubt, just as erroneous as the 'march of progress', but the general trend depicted here is indeed out there in the literature. *Homo pongoides* – a 'bestial' hominin – is second from the left. This image is inspired by an illustration by François de Sarre. Credit: Darren Naish.

Initial Bipedalism. Long-term readers of this blog and of the arcane cryptozoological literature will recall that pongoid man is one of several icons of cryptozoology mentioned at times within the context of initial bipedalism, a hypothesis which proposes that the human body shape and habit of erect walking are not recent evolutionary innovations, but ancient ones primitive not only to hominoids or primates but perhaps to mammals and even to a far more inclusive clade of vertebrates (the model was covered here on Tet Zoo ver 2, back in 2008). The hypothesis has predominantly been promoted by ichthyologist François de Sarre whose writings have often made reference to Bernard Heuvelmans and his work (e.g., de Sarre 1996, 1997).

Indications that Heuvelmans was a proponent of this hypothesis have always been evident in his better-known works. In On the Track of Unknown Animals, there is a curious passage in the yeti chapter wherein Heuvelmans (1995) states that “man has retained the plantigrade feet of a primitive mammal … that cannot have evolved from the apes’ prehensile feet … It is the other way round: apes’ feet seem to have evolved from feet like man’s” (p. 171 of 1995 edition).

**Caption:** initial bipedalism posits the existence of human ancestors that look somewhat... different relative to the sort of interpretations you might be used to. This reconstruction – showing early proto-human invaders of the land – is one of my favourites. Credit: de Sarre 1997.

Neanderthal provides the full exposé, the denouement. After discussing de-hominisation, Heuvelmans (2016) states “In this work, which challenges such a solid anthropological belief as the extinction of the Neanderthals, I would have preferred not to also bring in a rather heretical theory of human origins. But that can’t be avoided. It should have been expected from the pen of a discipline of Dr. Serge Frechkop. Those who are familiar with his work are aware of my former master’s preference for non-ape theories of human origin, including those of Ranke, Kollman and Osborn, and especially Max Westernhöfer’s theory of initial bipedalism. For over thirty years I have mulled over these ideas … and find that every new discovery in paleontolology has confirmed their soundness. I am well aware that my insolence in defending these theories here will bring as many sarcasms, critiques, and even insults as my candid description of the frozen specimen of a contemporary Neanderthal” (p. 224).

**Caption:** this cartoon – by initial bipedalism proponent François de Sarre – depicts the idea that humans (and/or human-like animals) evolved directly from aquatic ancestors, and that ape-like primates are the specialised 'de-hominised' descendants of human-like forms. Credit: de Sarre (1997).

Final Thoughts. Neanderthal is well illustrated throughout with both black and white photos and diagrams. A colour montage depicting the iceman itself – Heuvelmans made this montage and evidently took care to avoid distortion when photographing the specimen from slightly different angles – appears on the cover. Footnotes are a mix of Heuvelmans’s own notes combined with translator notes on the various weird turns of phrase that do not translate well. A number of typos have slipped through (‘Homo abilis’ is used several times). The lack of an index is most unfortunate and makes the volume very difficult to navigate.

Also at the back of the volume is an afterword by Loren Coleman; it is essentially a personal take on his own encounters with the iceman and extensively relates the thoughts of Coleman’s late friend and colleague Mark Hall. Here there is yet more theorising and speculating about the iceman; there’s lot of talk of models being created, and even photos of the model as displayed today at the Museum of the Weird in Austin. But it’s mixed with the idea that the object was originally a genuine corpse: like Sanderson, Hall did not think that the iceman was a Neanderthal, but instead a surviving member of the erectus lineage.

**Caption:** **Brian Regal’s 2013 book** – probably the best scholarly investigation of sasquatch research out there (it is about the researchers, not the research itself) – includes documentation of the behind the scenes story on the Minnesota iceman. Credit: Palgrave Macmillan.

I regret that I did not find myself agreeing with several of the points made in this section. Firstly, for all the evidence indicating that it was a hoax all along, the text ends in open-ended fashion (“The parasites seen on the body, the vegetable matter viewed in the teeth […] all point to the Minnesota Iceman having been an actual carcass. Maybe it was”; p. 246). Secondly, the entire section endorses the viewpoint that the later (post-April 1969) images were of the supposed replacement model, Heuvelmans’s strongly worded pronouncement to the contrary being ignored. And, thirdly, Coleman implies that Heuvelmans was a victim of “the scientific establishment” in that “he never managed to stir up the interest of professional anthropologists and paleoanthropologists” (p. 246). This is patently untrue. We know that several prominent workers of the time did look into the story – John Napier at the Smithsonian among them – and even took it seriously enough to contact the FBI, eventually concluding for good reason that the object was not a real body at all (Regal 2013).

Neanderthal is a weird book. It provides the full backstory to the case from the horse’s mouth, so to speak, and elucidates the author’s favoured hypotheses on topics touched on in his other works but not previously discussed at length. Bernard Heuvelmans inhabited what many modern researchers would consider an unusual intellectual landscape. As if the promotion of almost 140 unknown animal species was not unusual enough on its own (Heuvelmans 1986), he imagined these creatures within the context of evolutionary scenarios that were decidedly heterodox and at odds with the data accepted by the majority of his peers. Indeed, this book probably provides more insight on Heuvelmans’s opinions and interpretations of evolutionary patterns than any other (the caveat being that there are several of his books that I have never read since they are yet to be translated from the original French). For these reasons the book is of great value to those interested in the history of cryptozoological thought and speculation, on arcane evolutionary hypothesising, and also potentially to those researching the history of 20th century thought on hominin evolution.

For previous articles relevant to issues discussed here, see...

The Cryptozoologicon (Volume I): here, at last, December 2014
Loxton and Prothero's Abominable Science! Origins of the Yeti, Nessie, and Other Famous Cryptids; the Tet Zoo review (review of Loxton & Prothero 2013), September 2014
My New Book Hunting Monsters: Cryptozoology and the Reality Behind the Myths, February 2015
If Bigfoot Were Real, June 2016
Usborne’s All About Monsters, April 2019
Sea Monster Sightings and the ‘Plesiosaur Effect’, April 2019
Books on the Loch Ness Monster 3: The Man Who Filmed Nessie: Tim Dinsdale and the Enigma of Loch Ness, August 2019
The 1972 Loch Ness Monster Flipper Photos, August 2020
Monsters of the Deep, a Ground-Breaking Exhibition at the National Maritime Museum, Cornwall, October 2020
The Lake Dakataua ‘Migo’ Lake Monster Footage of 1994, February 2021
What Was the Montauk Monster? A Look Back to 2008, October 2021
Santa Cruz’s Duck-Billed Elephant Monster, Definitively Identified, November 2021
Legend of the Black Dog, August 2022
The Strange Case of the Minnesota Iceman, Part 1, August 2023

Articles like this are possible because of the support I receive at patreon. Please consider supporting my research and writing if you don’t already, thank you so much.

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Aiello, L. & Dean, C. 2002. An Introduction to Human Evolutionary Anatomy. Elsevier, Amsterdam.

Bayanov, D. 1996. In the Footsteps of the Russian Snowman. Crypto-Logos, Moscow.

Conway, J., Kosemen, C. M. & Naish, D. 2013. Cryptozoologicon Volume I. Irregular Books.

Costello, P. 1984. Mysterious man-beasts 2. In Brookesmith, P. (ed) Creatures From Elsewhere: Weird Animals That No-One Can Explain. Orbis Publishing, London, pp. 72-78.

de Sarre, F. 1996. About the survival of relict hominoids from the point of view of a zoologist. In Downes, J. (ed) CFZ Yearbook 1996. CFZ (Exeter), pp. 98-111.

de Sarre, F. 1997. Were aquatic pre-humans the first vertebrates to enter the land? In Downes, J. (ed) The CFZ Yearbook 1997. CFZ (Exeter), pp. 142-156.

Gribbin, J. & Cherfas, J. 1981. Descent of man – or ascent of ape? New Scientist 91 (1269), 592-595.

Heuvelmans, B 1968. In the Wake of the Sea-Serpents. Hill and Wang, New York.

Heuvelmans, B. 1969. Note preliminaire sur un specimen conserve dans la glace, d’une forme encore inconnue d’hominide vivant Homo pongoides (sp. seu subsp. nov.). Bulletin de I’Institut Royal des Science Naturelles de Belgique 45, 1-24.

Heuvelmans, B. 1986. Annotated checklist of apparently unknown animals with which cryptozoology is concerned. Cryptozoology 5, 1-26.

Heuvelmans, B. 1995. On the Track of Unknown Animals. Kegan Paul International, London.

Heuvelmans, B. 2016. Neanderthal: the Strange Saga of the Minnesota Iceman. Anomalist Books, San Antonio, Tx.

Loofs-Wissowa, H. 1994. The penic rectus as a marker in human palaeontology? Human Evolution 9, 343-356.

Lorenzo, C. 2015. The hand of the Neandertals: dexterous or handicapped? Journal of Anthropological Sciences 93, 181-183.

Michell, J. & Rickard, R. J. M. 1982. Living Wonders: Mysteries and Curiosities of the Animal World. Thames and Hudson, London.

Naish, D. 2016. Hunting Monsters: Cryptozoology and the Reality Behind the Myths. Arcturus, London.

Raynal, M. 2001. Jordi Magraner’s field research on the bar-manu: evidence for the authenticity of Heuvelmans’ Homo pongoides. In Heinselman, C. (ed) Hominology Special Number 1. Craig Heinselman (Francestown, New Hampshire), unpaginated.

Regal, B. 2013. Searching for Sasquatch: Crackpots, Eggheads, and Cryptozoology. Palgrave-Macmillan, London.

Searching for Sasquatch: Crackpots, Eggheads, and Cryptozoology (Palgrave Studies in the History of Science and Technology)

All Yesterdays: Unique and Speculative Views of Dinosaurs and Other Prehistoric Animals

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The modern-day corpse of a human-like hominid, preserved in a block of ice, encountered by researchers in the 1960s, you say? Surely the zoological discovery of the century!

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I really like ducks, and my god there’s a lot to say about them. Here, we look at a small Hawaiian duck that once stood on the very precipice of extinction, some sources saying that it was reduced to a single individual at one point. I speak, of course, of the famous Laysan duck Anas laysanensis…

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Today is July 31st, 2023 which means….

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Over the past day (20th July 2023), social media accounts have been sharing a brief snippet of mobile phone footage purporting to show a lion (specifically, a lioness) walking at the edge of woodland in the Berlin suburb of Kleinmachnow…

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Regular readers of Tetrapod Zoology will be aware that I’m a big fan of cassowaries….

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In spare time, I’m reviewing Alan Feduccia’s 2020 Romancing the Birds and Dinosaurs: Forays in Postmodern Paleontology. It’s an interesting book for sure, and I have quite a lot to say about it….

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Back in October 2022, we looked at the diversity and evolution of plains zebras, a group that includes the Quagga. Here, we carry on with the zebra series, this time looking at Grevy’s zebra….

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Back in December 2022, I had the remarkable privilege of visiting Portland, Oregon, for a conference. I’ve longed to visit the Pacific Northwest for my entire life and this was my first ever visit there. It was a dream trip and I had an amazing time, and while there’s a lot I could talk about… today we’re here to discuss THE BIRDS...

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Over recent years, myself and colleagues at the University of Southampton have published a series of studies on the spinosaurid theropods of the English Wealden Supergroup (Barker et al. 2021, 2022, 2023a). And we’ve succeeded in improving our knowledge of Wealden spinosaurid diversity. But wait — there’s more!

**Caption:** a Hastings spinosaurid montage, featuring images of the specimen discussed further below and some of the analyses we did on it.

Firstly, what’s the Wealden? It’s a Lower Cretaceous sedimentary unit famous for the great number of dinosaurs and other fossils it reveals. Today sees the latest of our publications on Wealden spinosaurids; it’s by Chris Barker, Neil Gostling and myself, and appears in the open access journal PeerJ (Barker et al. 2023b).

And while the results are interesting, arguably newsworthy… don’t get your hopes up. We aren’t (this time) talking about fossils that can be considered spectacular or impressive.

A Hastings spinosaurid specimen. This latest study concerns the spinosaurid tooth HASMG G369a, accessioned at Hastings Museum and Art Gallery in East Sussex, England and part of a collection gifted by Reverend Pierre Teilhard de Chardin during the first half of the 20th century. Teilhard is best known for his metaphysical writings on humanity’s place in the cosmos (incidentally, an area relevant to my research on Dale Russell and the dinosauroid), but his interest in, and discovery of, Wealden fossils is well known among specialists.

**Caption:** the Hastings spinosaurid tooth HASMG G369a, in (A) lingual, (B) basal, (C) mesial, (D) distal and (E) labial views, with close-ups of its surface texture on the labial side (F-H). Scale bar = 10 mm, 1 mm in F-H. Image: **Barker *et al*. (2023b)**.

The precise provenance of the Hastings tooth is a little murky but it almost certainly came from the Wadhurst Clay or Tunbridge Wells Sands formations close to Hastings (Barker et al. 2023b), which in turns means it comes from the Valanginian stage of the Lower Cretaceous*. Theropods from this part of the Wealden (and, indeed, this part of the Cretaceous) are rare, so any additional information on their diversity is welcome.

I should add as an aside that we initially working on the assumption that the specimen was from the Purbeck (way down at the base of the Cretaceous) – an idea arrived at thanks to an associated hand-written note – but ultimately decided that a Purbeck origin was based on a misunderstanding.

Caption: geographical distribution of the Wealden spinosaurid discussed here, from Barker et al. (2023b). 1, 2 = Upper Weald Clay specimens (the Baryonyx walkeri holotype and indeterminate Ewhurst Brickworks specimen); 3 = Suchosaurus cultridens from the Grinstead Clay Formation; 4 = indeterminate spinosaurid specimens from the Wadhurst Clay Formation of Bexhill.

We’re confident that the Hastings tooth is from a spinosaurid, since it possesses the cone-like form and proportionally tiny denticles unique to this group (Barker et al. 2023b). We can eliminate the only other group of animals worthy of consideration – crocodyliforms – since they differ from spinosaurids (and the Hastings tooth specifically) in denticle form, crown ornamentation, size and other features too (Barker et al. 2023b). The tooth’s crown is only 1.3 cm long, which means that it most likely originated from the rear parts of the jaws.

For decades now, people have thought it likely that isolated Wealden spinosaurid teeth should be regarded as belonging to Baryonyx, the iconic English spinosaurid found in Surrey in 1983 and named in 1986 (Charig & Milner 1986, 1997). But many of the teeth in question are (in geological terms) quite a bit older than the remains of Baryonyx, and also different from its teeth in detailed anatomy. For these reasons, it’s been suggested that at least some Wealden spinosaurid teeth might not be from Baryonyx at all (Naish & Martill 2007, Buffetaut 2010, Naish 2011), and in fact they might show that Wealden spinosaurid diversity is higher than otherwise thought. How do we go further with this hypothesis? How do we test it?

**Caption:** the approximate stratigraphic distribution of spinosaurids in the Wealden Supergroup as understood right now. The Hastings specimen discussed here is from the Hastings Group, and thus from the Valanginian. Establishing the presence of spinosaurids here is not novel at all, but working out what kind of spinosaurid we’re dealing with is a worthy endeavour. Image: Darren Naish.

Theropod teeth are complex objects, differing in their ratios of base length to crown height, in their degree of curvature, their surface texture, in the distribution, shape, proportional size and spacing of their serrations, and much else. There are, in short, numerous details that can be codified, and (in cases) quantified and analysed statistically. Today, it’s easier than ever before (though still – by no means – actually ‘easy’) to use statistical tests of different sorts in the analysis of a given fossil specimen or specimens. It’s also generally agreed that the combined use of different tests increases the robustness of a conclusion. Inspired by recent studies that have also been devoted to the identification of theropod teeth (Hendrickx & Mateus 2014, Hendrickx et al. 2015, 2019, 2020), we did exactly this, subjecting the Hastings tooth to separate phylogenetic, discriminant and cluster analyses (Barker et al. 2023b).

Baryonyx in the Valanginian? Probably not. What did we find? When included in phylogenetic tests, the Hastings tooth is nothing to do with Baryonyx, and in fact occupies a position outside both Spinosaurinae and Baryonychinae, the two constituent clades within the group (Barker et al. 2023b). When included in statistical tests that compare tooth proportions and other features, it again never grouped with Baryonyx, but was closer to baryonychines than to spinosaurines (Barker et al. 2023b).

**Caption:** position for the Hastings spinosaurid tooth recovered in our phylogenetic analysis (strict consensus at left, reduced consensus at right). It never grouped with *Baryonyx*, and indeed was not recovered as a baryonychine. Image: **Barker *et al*. (2023b)**.

While we really shouldn’t go any further at this point – we’re talking of a single tooth, after all – the takehome is that the Hastings tooth is not from Baryonyx, and whether it is or isn’t referable to that genus is the primary hypothesis we were aiming to test. Here is another indication, then, that isolated Wealden spinosaurid teeth shouldn’t be referred to Baryonyx by default. And the specimen at least hints at the presence of additional diversity within Wealden spinosaurid that’s yet to be properly documented: in other words, that isolated teeth demonstrate a sort of previously ‘hidden’ diversity within this group (Barker et al. 2023b).

**Caption:** results of one of the several discriminant analyses we applied to the Hastings specimen (visible at far right) and other spinosaurid specimens. HASMG G369a didn’t group with *Baryonyx*, though is closer to baryonychine clusters than to spinosaurine ones. Of incidental interest is that the ‘cf. Baryonychinae’ specimen at upper right (a Santonian tooth from the Majiacun Formation of China, suggested to be an especially late-occurring spinosaurid) never grouped with spinosaurids in our study, meaning that it’s likely not part of this group. Image: **Barker *et al*. (2023b)**.

Pending the discovery of additional Wealden spinosaurid remains that include teeth of the same sort, here’s where things end on this particular specimen. However, this study was sort of a test-run. Numerous other isolated spinosaurid teeth are known from the Wealden, and ‘all’ we have to do now is run similar tests on them too…

As ever, thanks to my co-authors Chris and Neil for collaboration on this study, and to staff at the Hastings Museum and Art Gallery for assistance, access, and continued curation of the specimen.

For previous TetZoo articles on spinosaurids, British theropods and associated issues (some links here are to wayback machine versions due to destruction or paywalling of everything at versions 2 and 3), see…

Of Becklespinax and Valdoraptor, October 2007
The world’s most amazing sauropod, November 2007
Oh no, not another new Wealden theropod!, June 2009
Concavenator: an incredible allosauroid with a weird sail (or hump)... and proto-feathers?, September 2010
A truly tiny Cretaceous theropod... from England?, May 2011
The Wealden Bible: English Wealden Fossils, 2011, November 2011
Ostrich dinosaurs invade Europe! Or do they?, June 2014 (every archived version of this article lacks the original illustrations, sorry)
The Sensitive Face of a Big Predatory Dinosaur, June 2017
Theropod Dinosaurs of the English Wealden, Some Questions (Part 1), March 2020
Introducing ‘Unexpected Isle of Wight Air-Filled Hunter’, a New English Theropod Dinosaur, September 2020
Dr Angela Milner and the Discovery of Baryonyx, August 2021
Two New Spinosaurid Dinosaurs from the English Cretaceous, September 2021
A Giant Spinosaurid Dinosaur from the Cretaceous of the Isle of Wight, June 2022
A brain for Baryonyx: using CT-scanning to examine British spinosaurid brains, February 2023

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Barker, C. T., Naish, D. & Gostling, N. J. 2023b. Isolated tooth reveals hidden spinosaurid dinosaur diversity in the British Wealden Supergroup (Lower Cretaceous). PeerJ 11: e15453.

Buffetaut, E. 2010. Spinosaurs before Stromer: early finds of spinosaurid dinosaurs and their interpretation. In Moody, R. T. J., Buffetaut, E., Naish, D. & Martill, D. M. (eds) Dinosaurs and Other Extinct Saurians: A Historical Perspective. Geological Society, London, Special Publications 343, pp. 175-188.

Charig, A. J. & Milner, A. C. 1986. Baryonyx, a remarkable new theropod dinosaur. Nature 324, 359-361.

Charig, A. J. & Milner, A. C. 1997. Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey. Bulletin of the Natural History Museum 53, 11-70.

Hendrickx, C. & Mateus, O. 2014. Abelisauridae (Dinosauria: Theropoda) from the Late Jurassic of Portugal and dentition-based phylogeny as a contribution for the identification of isolated theropod teeth. Zootaxa 3759, 1-74.

Hendrickx, C., Mateus, O. & Araújo, R. 2015. The dentition of megalosaurid theropods. Acta Palaeontologica Polonica 60, 627-642.

Hendrickx, C., Mateus, O., Araújo, R. & Choiniere, J. 2019. The distribution of dental features in non-avian theropod dinosaurs: Taxonomic potential, degree of homoplasy, and major evolutionary trends. Palaeontologia Electronica 22, 1-110.

Hendrickx, C., Tschopp, E. & Ezcurra, M. d. 2020. Taxonomic identification of isolated theropod teeth: the case of the shed tooth crown associated with Aerosteon (Theropoda: Megaraptora) and the dentition of Abelisauridae. Cretaceous Research 108, 104312.

Naish, D. 2011. Theropod dinosaurs. In Batten, D. J. (ed.) English Wealden Fossils. The Palaeontological Association (London), pp. 526-559.

Naish, D. & Martill, D. M. 2007. Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: basal Dinosauria and Saurischia. Journal of the Geological Society, London 164, 493-510.

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Regular readers of Tetrapod Zoology content will be aware of the long-standing interest round these parts in the dinosauroid, a hypothetical humanoid theropod dinosaur posited to evolve in an alternative timeline where the end-Cretaceous extinction event never happened…

**Caption:** what have we here? Read on…

The dinosauroid was devised in the early 1980s by Ottawa-based palaeontologist Dale Russell and model-maker Ron Séguin, and the most complete rendition of its construction and the story behind it was published in 1982 (Russell & Séguin 1982). Dale – an extraordinary scientist whose work and ideas impacted many of his peers and colleagues – died in 2019 and it’s fitting that a special tribute volume of Canadian Journal of Earth Sciences was devoted to him in 2021. I’m pleased to say that I was invited to contribute a special paper on the dinosauroid for that volume (Naish & Tattersdill 2021), and for more details on the paper concerned see the 2021 Tetrapod Zoology article Humanoid Dinosaurs Revisited Again: Russell and Séguin’s Dinosauroid at (Nearly) 40 Years Old.

**Caption:** my long-standing interest in the dinosauroid can be said to have culminated in this, the Naish & Tattersdill (2021) paper in this special tribute issue of *Canadian Journal of Earth Sciences*. But little was I to know that peak dinosauroid was yet to be achieved.

The dinosauroid has been discussed in a huge number of popular and semi-popular venues, including books and magazine articles and even comics and graphic novels, and a consequence of its popularity is that it’s been realised for TV more than once. Many readers of this blog will remember the 1991 TV series Dinosaur!, presented by Walter Cronkite and co-produced by Granada Television in the UK, Primedia in Canada and Satel across continental Europe. The series was accompanied by a book of the same name, authored by British dinosaur expert Dr David Norman (Norman 1991).

The last episode of Dinosaur! takes an unusual turn in that it ends with the revelation that its narrator is not human, but a dinosauroid from an alternative timeline. The dinosauroid we see is not quite the same as the one depicted by Russell & Séguin (1982) but arguably more interesting in appearance: it’s green with red stripes, and has large, orange, plate-like scales across its chest. You might be disappointed to hear that it isn’t really a dinosauroid from an alternative timeline, but a person wearing a suit, and we learn from the book (Norman 1991) that said person is Emma Norman, Dr David Norman’s daughter.

**Caption:** the Norman/Minister *Dinosaur!* dinosauroid, here ripped mercilessly from **an article at Love in the Time of Chasmosaurs**. Don’t tell them.

  **Caption:** the colourful dinosauroid that appears in the 1991 TV series *Dinosaur!*, narrated by Walter Cronkite. Both images come from David Norman’s 1991 book of the same name, which is not as bad as I remember.

Here's where things take an unusual personal turn. Over the past couple of decades I’ve worked on and off as an author, consultant and editor on various dinosaur-themed books published by Dorling Kindersley, and one thing I’ve done a lot of is provide advice and information on digitally crafted artwork. I’ve worked with several artists while doing this work, and among them is Peter Minister. Peter is a man of many talents. He’s produced no end of digital dinosaurs and other animals (living, extinct and fantastical) – mostly crafted in zbrush – but is also a skilled maker of more traditional models, and to that end has made pieces for TV and film, and for display in museums and exhibitions.

So imagine my surprise on checking the 1991 Dinosaur! book and seeing that Peter is credited as the maker of the dinosauroid from the abovementioned TV series (Norman 1991, p. 187). In conversation, Peter confirmed that he was guilty as charged. It might not be much of a secret that I maintain a large, ahem, ‘working collection’ of animal models, figures and toys. Peter produces zbrush models and once made a dinosauroid. Had he considered, I mused, idly, the creation of a zbrush dinosauroid model?

**Caption:** in which Peter Minister creates what might be the world’s first zbrush model of Russell & Séguin’s dinosauroid. Images: (c) Peter Minister.

Before you could say Stenonychosaurus inequalis, said model was finished and in my inbox. And like SID 6.7 in the 1995 movie Virtuosity, all it needed was access to a convenient 3D printer. Here I owe substantial thanks to my friend and colleague Luke Muscutt (of robotic swimming plesiosaur fame) who kindly did the printing on my behalf. And thus, world, I give you physical models of the dinosauroid that now occupy a position on the cherished top shelf of my finest display case…

**Caption:** two differently sized dinosauroid figures, printed attractively in translucent blue and metallic grey, in a display cabinet. Thanks to Peter and Luke for bringing the dream to life. Images: Darren Naish.

I have no doubt that you want such a model of your own, so let’s see where we go from here.

For previous Tetrapod Zoology articles on the dinosauroid and related issues, see…

Dinosauroids revisited, November 2006
Dinosauroid cave art discovered, March 2007
How intelligent dinosaurs conquered the world, March 2008
Belatedly, Nemoramjetia (= Avisapiens), November 2008
Richard Dawkins and the crappy ‘humanoid dinosaurs’ that just won’t die, November 2009
Dinosauroids revisited, revisited, October 2012
Speculative Zoology at Tet Zoo, The Story So Far, May 2015
Speculative Zoology, a Discussion, July 2018
Alternative Timeline Dinosaurs, the View From 2019 (Part 1), December 2019
Alternative Timeline Dinosaurs, the View From 2019 (Part 2), December 2019
Alternative Timeline Dinosaurs, the View From 2019 (Part 3): the Dinosauroid and its Chums, December 2019
Humanoid Dinosaurs Revisited Again: Russell and Séguin’s Dinosauroid at (Nearly) 40 Years Old, August 2021

Help support Tetrapod Zoology and my research, and see stuff behind the scenes before it gets released to the proletariat.

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Naish, D. & Tattersdill, W. 2021. Art, anatomy and the stars: Russell and Séguin’s dinosauroid. Canadian Journal of Earth Sciences 58, 968-979.

Norman, D. 1991. Dinosaur! Boxtree, London.

Russell, D. A. & Séguin, R. 1982. Reconstruction of the small Cretaceous theropod Stenonychosaurus inequalis and a hypothetical dinosauroid. Syllogeus 37, 1-43.

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Way back in 2009 (the ScienceBlogs years) I published a Tetrapod Zoology article titled ‘Tell me something new about basilisks, puh-lease’. Aware that there’s all too little squamate content here at Tetrapod Zoology ver 4, I here present a much augmented and updated version of that article…

**Caption:** captive Emerald or Plumed basilisk, showing the forked cranial crest and tall, serrated dorsal sail and serrated caudal sail of this species, though more ‘extreme’ individuals of this species exist... The light green ground colour and blue markings are characteristic. Note the massive length of the toes. Image: Tina Whitlock, used with permission.

Thanks to their striking appearance, basilisks are often featured in books and on TV, but people only ever say the same two things about them: (1) that they have striking display structures like those so obvious here, and (2) that they can run (bipedally) across the surface of water. Yeah, I’ve heard all that before. Tell me something new!

Basilisks are a group of large, mostly green iguanians from Central and South America*, the biggest specimens of which reach 90 cm in total length. As a generalisation, they’re animals of mature, densely vegetated forests – preferably with running streams – and prefer high temperatures and humidities. They’re strongly arboreal and adapted to climbing on boughs and branches, sometimes many metres from the ground (to 20 m at least; Mora & Escobar-Anleu 2017). They rely on water as a means of escaping from and retreating from predators; more on that below.

In 1976, the Brown or Striped basilisk Basiliscus vittatus was documented in Florida, its presence here being almost certainly due to escape or intentional release from the pet trade. The species is now well established at several location in the south of the state.

Caption: like many iguanians, basilisks are good at clinging to, and climbing on, vertical and near-vertical trunks and branches, and typically adopt a pose where the palms and soles face inwards. These images show (left to right) two Common basilisks and a Western or Red-headed basilisk B. galeritus. Images: James J. Sharp, CC BY-SA 4.0 (original here); Pavel Kirillov, CC BY-SA 2.0 (original here); Daniel van der Post, open access (original here).

Exactly how much of their foraging and feeding is done in the arboreal environment versus the ground is still unclear as far as I can tell, and some sources imply that their use of the arboreal environment is more to do with predator avoidance and/or basking than specialisation. However, their food preferences and anatomy (read on) mean that they really should be regarded as specialised climbers. It’s also clear that basilisks do a lot of things on the ground. So… arboreal, terrestrial, which is it? And what about the semi-aquatic behaviour (discussed further below)? I just don’t think we have a term for animals that combine habitat use like this. Or do we?

Basilisks sleep on branches and leaves but rare reports of young animals using large riverside boulders as night-time sleeping locations are on record too (Mora & Escobar-Anleu 2017).

**Caption:** montage featuring three different Common basilisks. The juxtaposition of longitudinal white stripes and transverse dark ones is obvious in this species, and note also the variation in sail size and shape. Images (clockwise from upper left): The Rambling Man, CC BY-SA 3.0 (**original here**); Daniel Hincapie, CC BY 4.0 (**original here**); Anthony Batista, CC BY 4.0 (**original here**).

No, basilisks are not aquatic. And having mentioned riverside boulders… basilisks are (mostly) associated with a specific kind of tropical forest: that adjacent to bodies of water, including ponds, streams, rivers, sinkholes and even the sea. Individuals are very often found within 1-2 m of the water’s edge (Lattanzio & LaDuke 2012) but they can’t be regarded as limited to water-edge microhabitats given that adult Brown basilisks are very often observed some distance from water. Furthermore, the Emerald or Plumed basilisk B. plumifrons of Mexico and Central America is seemingly less tied to water than the other species and often occurs well away from it.

**Caption:** Western or Red-headed basilisk photographed in a stream. Basilisks of most (but not all) populations forage close to, or in, streams and other water bodies, meaning that it *might* be right to describe them as semi-aquatic. But it might not. Image: desertnaturalist, CC BY 4.0 (**original here**).

It absolutely isn’t correct to describe basilisks as ‘aquatic’ (seeing as they don’t do anything like live in water or on its surface), but they have been reported to forage in streams and other bodies of water for aquatic insect larvae and crayfish, a consequence being that some authors have described them as ‘semi-aquatic’. They’re also capable divers and the Brown basilisk has been reported to disappear underwater as a predator avoidance tactic. The long hindlimbs, overall proportions and long, heavy tail mean that they can run bipedally if speed and space allows, and here’s where we come to the water-running behaviour.

On being startled or alarmed, a basilisk will drop into the water and quickly skitter across the surface, its body inclined upwards and forelimbs mostly clear of the water. This water-running behaviour works best among small, lightweight juveniles who are able to stay more elevated from the water surface than are large, heavy adults (who can weigh up to 600 g). The key to water-running is the presence of rectangular fringes on the toes that trap air and prevent the foot from being pushed too far into the water, a swift, circular recovery stroke that allows the limb to be extracted and returned to starting position with minimal disruption of the water, and a stable body position that keeps the centre of mass between the supporting limbs (Glasheen & McMahon 1991, 1992, Hsieh & Lauder 2004).

**Caption:** the famous water dashes performed by basilisks happen so quickly, and occur over such short distances, that it’s hard to capture them adequately without high-speed photography. Image: Ted, CC BY-NC 2.0 (**original here**).

How did such a remarkable set of features evolve, given that intermediate stages would disallow water-running behaviour and thus make basilisks vulnerable to predation from some of the animals they most want to avoid (like fishes and large birds)? A good argument has been made that basilisk toe fringes are an excellent example of exaptation, this being the evolutionary phenomenon whereby features that evolve in the context of one behaviour later prove advantageous for another, and hence get co-opted. Rectangular toe fringes have evolved at least five times in arboreal lizards specialised for fast movement on leaves, including in agamids, tropidurines, lacertids and teiids, and they’re present in corytophanids other than basilisks too (Vitt in Pianka & Vitt 2003). Toe fringes are widespread in lizards, though more often involve triangular or conical scales (Luke 1986), not the unusual rectangular scales relevant here.

**Caption:** rectangular toe fringes (actually formed of projecting scales that are in close contact) as depicted in several non-basilisk lizards by Luke (1986). (A) the agamid *Hydrosaurus*, (B) the African lacertid *Holaspis*, (C) the teiid *Kentropyx*, and (D) cross-section of a *Hydrosaurus* fringe, showing how one of the scales forming the fringe projects way out horizontally relative to the rest of the toe. Scale bars = 10 mm. Image: Luke (1986).

The conclusion has to be that the fringes are there due to arboreal specialisation, but later provided an advantage in a specific escape strategy involving water. Basilisks now rely on this strategy and use it routinely; other fringe-toed taxa can do the water-running thing as well but are not as good at it and only use it opportunistically (Vitt in Pianka & Vitt 2003).

**Caption:** close view of the face of a Common basilisk. Not sure why this individual has a white crest. Image: Brian Gratwicke, CC BY 2.0 (**original here**).

Taxonomy, phylogeny, diversity. Four basilisk species are presently recognised: the Common basilisk B. basiliscus, the Western or Red-headed basilisk B. galeritus, the Emerald or Plumed basilisk B. plumifrons and the Brown or Striped basilisk B. vittatus. Subspecies have been recognised for some of them and it’s suspected that there are additional ‘species-worthy’ populations here. I’m thinking in particular of a Colombian population currently included in the Red-headed basilisk. It looks unique due to its especially large, rounded cranial crest.

**Caption:** the Western or Red-headed basilisk, probably the least familiar and most unusual of basilisks (at least, to those of us outside the American tropics). Note the serrated dorsal crest and very long, slender limbs. Images (left to right): Sandy Martinez, CC BY 4.0 (**original here**); desertnaturalist, CC BY 4.0 (**original here**).

A phylogenetic study that incorporated molecular, anatomical and sperm morphology data found the Red-headed basilisk to be outside a clade that contained all the others, and the Emerald and Common basilisks to be sister-taxa (Vieira et al. 2005). The latter two are the most ‘extreme’ of basilisks in morphology (the Red-headed and Brown lack the massive dorsal and tail frills of the Emerald and Common basilisks, and are smaller than them as well), so this makes sense. However, a more recent study (Taylor et al. 2017) found Brown and Common basilisks to be sister-taxa, in which case frills evolved twice, or were lost in the lineage leading to the Brown basilisk.

**Caption:** male Common and Emerald basilisks compared, both photographed in Costa Rica. Note the different attitude of the dorsal sail in these two individuals. Because the sails are supported by neural spines, they aren’t under muscular control. Images: Derkarts, CC BY-SA 3.0 (**original here**); Connor Long, CC BY-SA 4.0 (**original here**).

  **Caption:** highly simplified hypothesis of phylogeny relating to corytophanids and their close relatives. For much of the late 20th century, all of these groups were lumped together within ‘Iguanidae’, though the trend today is to recognise the main constituent clades as ‘family level’ groups. This cladogram and its illustrations are from my in-prep textbook, though that green iguana has broken proportions and needs replacing. Image: Darren Naish.

In the larger picture, basilisks are part of the iguanian group Corytophanidae, a clade that includes three extant genera (casquehead iguanas Laemanctus and helmeted iguanas Corytophanes are the other two) (Lang 1989, Frost et al. 2001, Vieira et al. 2005) (the same group was termed Basiliscinae in the older literature: see Lang 1989). Experts have disagreed on whether corytophanids originated in South America (and ultimately in Gondwana) and moved north, or whether they came from the Mexican Plateau or thereabouts and moved south. By combining a phylogenetic hypothesis with distribution data, Vieira et al. (2005) argued that corytophanid evolution was a Central American event and that basilisks invaded South America at least twice, once in or after the middle Pliocene (giving rise to the endemic South American Red-headed basilisk) and at some point more recently (giving rise to South American populations of the Common basilisk).

A very different hypothesis was put forward by Taylor et al. (2017) who found all species-level splits within Basiliscus to have occurred in the Early Miocene or the preceding Oligocene (between about 35 and 20 million years ago).

**Caption:** time-calibrated corytophanid iguanian cladogram from Taylor *et al*. (2017). The North American fossil taxon *Babibasiliscus* is nested within the crown (though the European *Geiseltaliellus* is not). Within basilisks, Brown and Common basilisks are sister-taxa. Image: **Taylor *et al*. (2017)**, CC.

But then… fossils! Fossils further complicate things. 2015 saw the publication of the fossil corytophanid Babibasiliscus alxi from the Lower Eocene Bridger Formation of Wyoming. Babibasiliscus was argued by Conrad (2015), its describer, to be a crown-corytophanid, closer to Laemanctus than are Corytophanes and Basiliscus. Another Eocene lizard – Geiseltaliellus from Germany, France and Belgium (named back in 1944) – was found to be part of the crown too, and closer to Corytophanes and Laemanctus than Basiliscus. Other studies have, however, found Geiseltaliellus to be outside crown-Corytophanidae (Taylor et al. 2017).

**Caption:** time-calibrated corytophanid phylogeny from **Conrad (2015)**, showing skulls of the relevant taxa. Image: **Conrad (2015)**.

These fossils – found so far from the Central American region otherwise considered integral to corytophanid history – are important, since they could mean that Central America simply represents that region where corytophanids ‘ended up’. It isn’t, perhaps, their ancestral ‘home’ or ‘site of origin’. Two main ways of explaining corytophanid distribution seem to exist. One is to stick with the conventional view that they originated and diversified in Central America, but to posit that one or two lineages moved away from that region and colonised the north and – to the far east – Europe (via the famous/infamous De Greer land bridge). And another is to think that corytophanids were ancestrally present across much of Europe and North America. If the latter is so, the living taxa are relicts or southerly occurring members of a previously more ‘northerly’ clade.

Worth mentioning here as an aside is that a group of extinct iguanians from the Eocene and Oligocene of Europe and Oligocene of North America – the messelosaurines – are apparently part of Corytophanidae too (Rossmann 1999a, b, 2005). Rossmann (1999a) proposed that corytophanids originated in Europe and ended up in South America following dispersal to North America.

**Caption:** female Common basilisk photographed on the banks of the Chagres River, Panama. The prominent white striping and great length and slender nature of the tail are really obvious here. Image: Charles J. Sharp, CC BY-SA 4.0 (**original here**).

Anatomy. Basilisks are highly distinctive anatomically. The cranial crest is unique, as are the tall, serrated dorsal and caudal fins. The cranial crest is supported internally by a sheet-like outgrowth of the parietal and is largest in adult males, though only Emerald and Brown basilisks are truly sexually dimorphic when size-related differences are accounted for (Taylor et al. 2017).

The dorsal and caudal fins are supported by long, slender neural spines that extend all the way to the serrated fin margins, with those of the dorsal region sometimes having triangular bases that are distinct from the spine-like, more dorsal parts (Lang 1989, Sereno et al. 2022). The superficial similarity that these spines have with the tall neural spines of the Cretaceous dinosaur Spinosaurus has not been missed and the fact that basilisk fins are irrelevant to aquatic behaviour has been used by those arguing that Spinosaurus was not aquatic (Sereno et al. 2022).

**Caption:** mounted and posed basilisk skeleton – I presume of Emerald or Plumed basilisk – made by The Skeleton Factory. Image: (c) The Skeletal Factory.

An interesting aside about their flamboyant appearance is that captive basilisks tend not to develop the crests and sails to a prominent degree. This has been blamed on lighting, diet and stress based on captivity, transportation and/or proximity to people. I don’t know if this issue has been solved, but it might have been given that there are at least some captive males with fairly impressive crests and fins.

**Caption:** basilisk neural spines involved in support of the dorsal and caudal spines, from **Sereno *et al*. (2022)**. The life photo shows an Emerald basilisk; **Sereno *et al*. (2022)** CT-scanned a member of this species, and I think the diagram also represents that species. D = dorsal neural spines. CA = caudal neural spines. Image: **Sereno *et al*. (2022)**.

Basilisks are capable of caudal autotomy, which is surprising in view of how important the tail appears to be in locomotion and display.

Behaviour and ecology. As is typical for so many tropical lizards, relatively little is known about basilisk ecology and diet. They are omnivorous and eat certain leaves, flowers and fruit in addition to arthropods. Insects form the bulk of their diet, these variously including grasshoppers, beetles, ants, wasps and caterpillars (Hirth 1963, Fleet & Fitch 1974). Harold Hirth’s 1959-1961 study of basilisks living on the Caribbean coast of Costa Rica is really interesting because he found ants to be the commonest food items recovered from basilisk stomachs, in some cases (6 males and 4 females) filling the stomachs entirely. Basilisks don’t look like ant-eaters and you might not predict that ants are that important to them, yet here we are. Additional interesting arthropod prey include freshwater crayfish, beach-dwelling amphipods, and barnacles that were being opened and dismembered by ghost crabs (Hirth 1963).

**Caption:** close-up profile view of a captive Common basilisk photographed at Buffalo Zoo. As ever for lizards, the complexity of scale arrangement is bewildering. The massive size, bright yellow colour and bulging nature of the eye is also interesting. Image: Becker1999, CC BY 2.0 (**original here**).

Basilisks are large and powerful enough to sometimes consume vertebrates, and frogs, snakes, other lizards, and even birds and rodents are eaten on occasion. A Common basilisk was observed swallowing “a hummingbird nest complete with young hummingbirds” (Glander 1979, p. 235). Because basilisks sometimes forage in streams, pools and on beaches, they also prey on aquatic animals and small fish have been reported as prey in both juvenile Common and Banded basilisks (Echelle et al. 1972, Fleet & Fitch 1974). There are photos online showing basilisks consuming juvenile slider turtles.

**Caption:** Emerald basilisk apparently consuming a juvenile green iguana. I know nothing of the circumstances of this photo (which has been widely shared online) and assume that it’s genuine. Image: (c) Randy Alvarado/Reptilehunter.

Cannibalism is a recorded part of their behaviour too and might even be important given that juveniles and adults sometimes inhabit the same parts of a habitat and sometimes occur in high densities (Lattanzio & LaDuke 2012). Another interesting aside is that sand has repeatedly been discovered within the stomachs of Costa Rican Common basilisks, and in fact occurs so consistently that it’s been regarded as a digestive aid by some authors (Fleet & Fitch 1974). Elsewhere in tetrapods, the ingestion of sand has been linked to insectivory, the idea being that it helps break up the chitinous bodies of insects like ants. Might that be what’s going on here? Or is the ingestion of sand an accidental and inevitable consequence of capturing arthropods on beach sand?

Basilisks of course live alongside diverse other forest-dwelling vertebrates, including other lizards as well as birds, primates and more, and may compete with them for resources. We don’t have much direct data on how the relevant interactions play out but basilisks and birds like thrushes don’t like one another and use chasing and surprise tactics to deter the other from feeding sites (Young 2014). Among non-aggressive observations, Glander (1979) reported Common basilisks feeding on fruit (specifically, on pedicels attached to the fruit) dropped by foraging howler monkeys and Hirth (1963) described basilisks sharing foraging sites (beaches) and sleeping locations with ameivas and green iguanas.

Basilisks themselves are prey for a list of animals, including snakes (like cornsnakes, racers and indigo snakes), herons, black hawks and ghost crabs. Largemouth bass and snooks have been seen to predate on basilisks that were running across water (Flaherty & Friers 2014).

**Caption:** I mentioned at the top of the article that some Emerald or Plumed basilisks are even more ‘extreme’ than some of the individuals we’re already seen. And here’s an incredibly flamboyant individual, with a super-sized cranial crest and dorsal and caudal sails of *Spinosaurus*-like proportions. Image: Bernard Dupont, CC BY-SA 2.0 (**original here**).

Courtship and reproduction. As elaborately ornamented species with (sometimes) obvious sexual dimorphism, we would predict that basilisks are competitive animals where males display during the breeding season and vie for female attention, perhaps in arboreal display areas of the sort known for green iguanas. Males are indeed highly competitive and reported to fight fiercely at times (Fitch 1981) but I’m not aware of observations pointing to lek-type displaying or anything like that. Head bobs are used to signal sexual interest. A size-based dominance hierarchy is established in an area, and smaller males are prevented from breeding by the aggressive actions of larger ones.

**Caption:** two male Common basilisks in an aggressive encounter, photographed in Manuel Antonio National Park, Costa Rica. Both lizards are elaborate and note the variation in striping. Also relevant is that both animals are on the ground, not in trees. Image: Michelle Reback, CC BY-SA 3.0 (**original here**).

Basilisks are oviparous and females produce clutches of between 5 and 18 eggs. At least one species (the Common basilisk, specifically a Colombian population of the subspecies B. basiliscus barbouri) appears to be capable of parthenogenesis (Böhme 1975, Lang 1989). Several females, kept captive for a prolonged period and without access to males, successfully reproduced and all offspring were female. As expected, basilisk eggs are buried in moist soil on the forest floor: the nest isn’t just a pit, but a tunnel more than 30 cm long. There are anecdotes in the pet trade (go here) describing female basilisks observing their nests after backfilling it to ensure that the nest area matches the look of the surrounding forest floor sediment. That’s a sophisticated piece of behaviour that might seem beyond the cognitive abilities of most lizards, but on the other hand I wouldn’t say that it’s unbelievable.

Hatchlings emerge after about two months of incubation and live together for some time after hatching, even sleeping piled up on top of one another. Similar behaviour is known for other iguanians (see the Tetrapod Zoology article Amazing Social Life of the Green Iguana).

**Caption:** all too little study has been done on the behaviour of iguanian hatchlings, but it might be that the sociality and group-living described for green iguanas is widespread within the group. For more information on the images here, see **Amazing Social Life of the Green Iguana**.

And that just about wraps up everything I wanted to say. I’m very fond of these large, attractive lizards and hope to see them in the wild some day. As should be clear, there’s a lot to say about basilisks beyond the water-running behaviour, and even that turns out to be more complex and nuanced than usually explained. As ever, it’s also obvious that a great many questions remain unanswered or (at best) only superficially or anecdotally answered about these lizards, including certain issues of ecology, habitat use and diet.

**Caption:** I adore elaborately ornamented lizards, but good scale figures or toys of them are scarcely available. I own a few basilisk toys and they’re all pretty terrible. This one (which is quite large) looks like it was made by the same people who make those ‘roaring’ dinosaurs with projecting point teeth. Image: Darren Naish.

Iguanian lizards have been covered a few times before at Tetrapod Zoology. For previous articles, see…

Harduns and toad-heads; a tale of arenicoly and over-looked convergence, December 2006
Ermentrude the liolaemine, February 2008
‘Cryptic intermediates’ and the evolution of chameleons, June 2008
Amazing social life of the Green iguana, September 2012
The Squamozoic actually happened (kind of): giant herbivorous lizards in the Paleogene, June 2013
The enormous liolaemine radiation: paradoxical herbivory, viviparity, evolutionary cul-de-sacs and the impending mass extinction, June 2013
Leiosaurus: big heads, bold patterns, October 2013
Grassland earless dragons, January 2014
Australia, land of dragons (by which I mean: agamids) (part I), January 2014
Australia, land of dragons (part II), February 2014
What's With All These New Chameleon Names?, Part 1, February 2016
By the Horns of Trioceros, the Casque of Calumma, the Brood of Bradypodion--Chameleons, Part 2, February 2016
Palleon, Archaius, Kinyongia, Nadzikambia--The Last Chameleons, Part 3, March 2016
The Tropidurine Treerunners, December 2017
The Tet Zoo Guide to Mastigures, August 2018

Help support the production of new articles at Tetrapod Zoology and get to see in-prep material behind the scenes.

Refs - -

Böhme, W. 1975. Indizien für natürliche Parthenogenese beim Helmbasilisken, Basiliscus basiliscus (LINNAEUS 1758) (Sauria: Iguanidae). Salamandra 11, 77-83.

Conrad, J. L. 2015. A new Eocene casquehead lizard (Reptilia, Corytophanidae) from North America. PLoS ONE 10 (7): e0127900.

Echelle, A. A., Echelle, A. & Fitch, H. S. 1972. Observations of fish-eating and maintenance behavior in two species of Basiliscus. Copeia 2, 387-389.

Flaherty, J. P. & Friers, J. 2014. Predation on the Brown Basilisk (Basiliscus vittatus) in South Florida. Southeastern Naturalist 13, 57-58.

Fleet, R. R. & Fitch, H. S. 1974. Food habits of Basiliscus basiliscus in Costa Rica. Journal of Herpetology 8, 260-262.

Frost, D. R., Etheridge, R., Janies, D. & Titus, T. A. 2001. Total evidence, sequence alignment, evolution of polychrotid lizards, and a reclassification of the Iguania (Squamata: Iguania). American Museum Novitates 3343, 1-38.

Glander, K. E. 1979. Feeding associations between howling monkeys and basilisk lizards. Biotropica 3, 235-236.

Glasheen, J. W. & McMahon, T. A. 1992. A hydrodynamic model of locomotion in the basilisk lizard. Nature 380, 340-342.

Glasheen, J. W. & McMahon, T. A. 1992. An analysis of aquatic bipedalism in basilisk lizards. American Zoologist 32, 144.

Hirth, H. F. 1963. The ecology of two lizards on a tropical beach. Ecological Monographs 33, 83-112.

Hsieh, S. T. & Lauder, G. V. 2004. Running on water: three-dimensional force generation by basilisk lizards. Biophysics and Computational Biology 101, 16784-16788.

Lang, M. 1989. Phylogenetic and biogeographic patterns of basiliscine iguanians (Reptilia: Squamata: “Iguanidae”). Bonner Zoologische Monographien 28, 1-172.

Lattanzio, M. S. & LaDuke, T. C. 2012. Habitat use and activity budgets of Emerald basilisks (Basiliscus plumifrons) in northeast Costa Rica. Copeia 2012, 465-471.

Luke, C. 1986. Convergent evolution of lizard toe fringes. Biological Journal of the Linnean Society 27, 1-16.

Mora, J. M. & Escobar-Anleu, B. I. 2017. River rocks as sleeping perches for Norops oxylophus and Basiliscus plumifrons in the Cordillera de Talamanca, Costa Rica. Mesoamerican Herpetology 4, 418-422.

Pianka, E. R. & Vitt, L. J. 2003. Lizards: Windows to the Evolution of Diversity. University of California Press, Berkeley.

Rossmann, T. 1999a. Messelosaurine lacertilians (Squamata: Iguanoides) from the Palaeogene of France and North America. Neues Jahrbuch fur Geologie und Paläontologie, Monatshefte 1999, 577-592.

Rossmann, T. 1999b. "Crotaphytus" oligocenicus (Holman, 1972), (Squamata: Iguanoidea) from the Oligocene of Saskatchewan, reinterpretation and some paleobiogeographical implications. Neues Jahrbuch fur Geologie und Paläontologie, Monatshefte 1999, 186-192.

Rossmann, T. 2005. Nachtrag zur Osteologie und Palaobiologie von Geiseltaliellus longicaudus Kuhn (Lacertilia, Iguanoidea) aus dem Mittleren Eozän (MP 11) der Grube Messel, nahe Darmstadt. Courier Forschungsinstitut Senckenberg 255, 225-236.

Sereno, P. C., Myhrvold, N., Henderson, D. M., Fish, F. E., Vidal, D., Baumgart, S. L., Keillor, T. M., Formoso, K. K. & Conroy, L. L. 2022. Spinosaurus is not an aquatic dinosaur. eLife 11: e80092.

Taylor, G. W., Santos, J. C., Perrault, B. J., Morando, M., Vásquez Almazán, C. R. & Sites, J. W. 2017. Sexual dimorphism, phenotypic integration, and the evolution of head structure in casque‐headed lizards. Ecology and Evolution 7, 8989-8998.

Young, A. M. 2014. Basiliscus plumifrons (double-crested basilisk lizard) antagonistic behavior. Herpetological Review 45, 694-695.

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Regular readers will be aware that my latest book – Ancient Sea Reptiles (Natural History Museum Publishing in the UK, Smithsonian Books in the US) – is now out and on sale…

Massive thanks to those who’ve bought the book and shared images or thoughts on it. A consequence of the book’s newness is that I’ve been giving talks on it, most recently at the Lyme Regis Fossil Festival in late April. I love going to Lyme Regis, and the talk went alright I think (space was very limited, so quite a few people unfortunately had to be turned away). In this brief article, I thought I’d share a few of the slides from the talk, and accompany them with a bit of extra information.

**Caption:** scenes from the Lyme Regis Fossil Festival 2023.

The many diverse groups. Ancient Sea Reptiles devotes appropriate space to the five ‘main’ marine reptile groups of the Mesozoic: the ichthyosaurs or fish-lizards, the mostly long-necked plesiosaurs, the thalattosuchians or ‘sea crocs’ (which are not crocodiles at all, hence my use of the informal ‘crocs’), those great sea-lizards the mosasaurs, and the sea turtles. There’s a lot to say about all of these groups (Naish 2023).

However, there are numerous other relevant groups as well, and I did my best to give them fair coverage too. Many of these were alive during the Triassic, including the long-tailed thalattosaurs, the very peculiar hupehsuchians, the (mostly) shellfish-eating placodonts, the fang-toothed helveticosaurs, the nothosaurs, the pistosaurs, and so on. Shared anatomical traits show that some of these groups are close cousins of plesiosaurs and belong with them in a large group termed Sauropterygia; hupehsuchians appear to be close kin of ichthyosaurs (Motani 1999, Motani *et al*. 2015). Entirely different groups whose affinities lie elsewhere evolved during the Jurassic and Cretaceous, including the marine pachyophiid snakes.

Finally, it’s becoming increasingly inaccurate to refer to these animals collectively as ‘Mesozoic marine reptiles’, given that there are several relevant groups from the Permian (the age before the Mesozoic), and evidence that some of the famous ‘Mesozoic’ groups – I’m thinking here of ichthyosaurs – actually originated during Permian times (Kear et al. 2023).

**Caption:** map of the world during Early Jurassic times, with a focus on far Western Tethys and flooded Europe. The Viking Corridor connected this region to the Boreal Ocean/Sea in the north, and the Hispanic Corridor connected it to Panthalassa in the west. Image: Naish (2023), after **Korte *et al*. 2015**.

A stereotype that’s still perpetuated is that the Mesozoic was monotonously hot, stable, and hence perfect for reptile evolution and perpetuation. It’s true that parts of the Mesozoic were hot and stable. But it’s absolutely not right to think that this was true of the whole of Mesozoic time. Geological and isotopic data shows that temperate (O’Brien et al. 2017, Alberti et al. 2019), cool and even cold conditions were present during parts of the Jurassic and Cretaceous; in fact, sea surface temperatures were close to freezing during parts of the Early Jurassic (Korte et al. 2015). Marine reptile groups like plesiosaurs and ichthyosaurs were swimming in these seas. The fossil record also shows that plesiosaurs, ichthyosaurs and mosasaurs were in polar waters at times when these regions were cool and seasonally dark. Some of these animals were insulated by blubber and it seems that endothermy was present in all of the main groups (Motani 2005, 2010, Martineau et al. 2010, Harrell et al. 2016, Wintrich et al. 2017, Fleischle et al. 2018).

Their world was also geographically and tectonically complex, with active volcanic regions, narrow marine corridors, and semi-enclosed seas. During the Jurassic, the marine faunas of western Europe were connected via the Viking Corridor to the Boreal Ocean (also called the Boreal Sea or Arctic Sea) in the far north, and via the Hispanic Corridor or Caribbean Seaway in the west to the great ocean Panthalassa. Marine reptiles thrived in all of these places and the presence of related species shows that groups moved around via these various seaways. However, the exact timings and directions of their movements remain enigmatic and several possibilities exist.

**On Eurocentricism.** Those of us living in western Europe are in an ideal location when it comes to fossil marine reptiles: there are a great number of significant fossil-bearing sites from which an extraordinary number of spectacular fossils have been extracted. The montage above shows specimens on show in the spectacular palaeontology corridor at the Natural History Museum in London, and all the specimens you see here are British. Germany, France, Luxembourg and other western European countries also have excellent marine reptile fossils.

A consequence of this fact is that our views on these animals – I’m mostly talking here about Jurassic groups – tend to be highly Eurocentric. Today, we know that animals similar to those of Europe occurred around South America, in the Caribbean, on the western coast of North America, and elsewhere too, so the view that these animals originated in Europe or should be considered strongly associated with Europe could well be very wrong.

Much work remains to be done and it’s hoped that new discoveries will improve our understanding of how these animals were distributed and how they moved about across their histories. It does, however, remain the case that a great many groups are essentially known from Europe and nowhere else. We assume that this is an artifact of the geological record and the history of scientific collecting, but bring on those new fossils.

Finally, many people are fascinated by the inferred, imagined or reconstructed behaviour of fossil animals and want to hear more about it. The problem is that we never know anywhere near as much as we’d like. For ancient sea reptiles we know a fair bit about dietary preferences and inferred hunting behaviour, and we’ve also done a lot of work on locomotory behaviour (e.g., Godfrey 1984, Lingham-Soliar 2000, Motani 2002, 2005, Carpenter *et al*. 2010, Liu *et al*. 2015, Muscutt *et al*. 2017); this explains the unusual image at top right of the montage above: see this article on plesiosaur locomotion to have it explained. Grooves and other marks on preserved seafloor sediment suggests that some of these animals ploughed or dug in the mud (Geister 1998), and of course the idea that giant predatory species hunted along shores and grabbed terrestrial animals from the water’s edge is irresistible. The fact that this strategy is present in several groups of living aquatic predators make it likely – yes, *likely* – that this strategy was used by at least some species.

But when it comes to social behaviour, reproductive behaviour and so on, we have but brief snippets. A few fossils provide possible evidence for intraspecific combat in such groups as ichthyosaurs, plesiosaurs and mosasaurs, and limited evidence for social behaviour and maybe even group-living also exists for some (e.g., O’Keefe & Chiappe 2011).

All of the issues touched on here are elaborated and discussed at great length in Ancient Sea Reptiles. Tetrapod Zoology is, even after all these years of operation, actually still quite deficient in marine reptile content… maybe this is something I should rectify over the course of the year. Thanks again to those who’ve bought the book already!

Buy *Ancient Sea Reptiles* here from NHM Publishing and here from Smithsonian Books.

For previous Tetrapod Zoology articles on the groups and subjects mentioned here, see…

The skin of ichthyosaurs, September 2008
Who made the giant Jurassic sea-floor gutters?, December 2009
Prediction confirmed: plesiosaurs were viviparous, August 2011
Can’t get me enough of that sweet, sweet Temnodontosaurus, January 2014
Plesiosaur Peril — the lifestyles and behaviours of ancient marine reptiles, March 2014
Ancient Marine Reptiles Had Absurd, Complex Nostrils, July 2014
‘Proto-Ichthyosaur' Sheds Light on Fish-Lizard Beginnings, November 2014
The Unique and Efficient 4-Flipper Locomotion of Plesiosaurs, August 2017
The Fall and Rise of Protoichthyosaurus, October 2017
The Ichthyosaurs of the Kimmeridge Clay, March 2021
History of the Iraqi Ichthyosaur, April 2021

Support Tetrapod Zoology at patreon (for as little as $1 per month) and get to see behind-the-scenes and in-prep stuff AND help support me in what I do here and elsewhere.

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Alberti, M., Fürsich, F. T. & Andersen, N. 2019. First steps in reconstructing Early Jurassic sea water temperatures in the Andean Basin of northern Chile based on stable isotope analyses of oyster and brachiopod shells. Journal of Palaeogeography 8: 33.

Carpenter, K., Sanders, F., Reed, B., Reed, J. & Larson P. 2010. Plesiosaur swimming as interpreted from skeletal analysis and experimental results. Transactions of the Kansas Academy of Science 113/2, 1-34.

Fleischle, C. V., Wintrich, T. & Sander, P. M. 2018. Quantitative histological models suggest endothermy in plesiosaurs. PeerJ 6: e4955.

Geister, J. 1998. Lebensspuren made by marine reptiles and their prey in the Middle Jurassic (Callovian) of Liesberg, Switzerland. Facies 39, 105-124.

Godfrey, S. J. 1984. Plesiosaur subaqueous locomotion, a reappraisal. Neues Jahrbuch fur Geologie und Paläontologie, Monatshefte 1984, 661-672.

Harrell, T. L., Pérez-Huerta, A. & Suarez, C. A. 2016. Endothermic mosasaurs? Possible thermoregulation of Late Cretaceous mosasaurs (Reptilia, Squamata) indicated by stable oxygen isotopes in fossil bioapatite in comparison with coeval marine fish and pelagic seabirds. Palaeontology 69, 351-363.

Lingham-Soliar, T. 2000. Plesiosaur locomotion: is the four-wing problem real of merely an atheoretical exercise? Neues Jahrbuch fur Geologie und Paläontologie, Abhandlungen 217, 45-87.

Liu, S., Smith, A. S., Gu, Y., Tan, J., Liu, C. K. & Turk, G. 2015. Computer simulations imply forelimb-dominated underwater flight in plesiosaurs. PLoS Computational Biology 11, e1004.

Kear, B. P., Engelschiøn, V. S., Hammer, Ø., Roberts, A. J. & Hurum, J. H. 2023. Earliest Triassic ichthyosaur fossils push back oceanic reptile origins. Current Biology 33, 178-179.

Korte, C., Hesselbo, S. P., Ullmann, C. V., Dietl, G., Ruhl, M., Schweigert, G. & Thibault, N. 2015. Jurassic climate mode governed by ocean gateway. Nature Communications 6: 10015.

Martineau, F., Mazin, J.-M. & Prieur, A. 2010. Regulation of body temperature by some Mesozoic marine reptiles. Science 328,1379-1382.

Motani, R. 1999. Phylogeny of the Ichthyopterygia. Journal of Vertebrate Paleontology 19, 473-496.

Motani, R. 2002. Scaling effects in caudal fin propulsion and the speed of ichthyosaurs. Nature 415, 309-312.

Motani, R. 2005. Evolution of fish-shaped reptiles (Reptilia: Ichthyopterygia) in their physical environments and constraints. Annual Review of Earth and Planetary Sciences 33, 395-420.

Motani, R. 2010. Warm-blooded “sea-dragons”? Science 328, 1361-1362.

Motani, R., Jiang, D.-y., Chen, G.-B., Tintori, A., Rieppel, O., Ji, C. & Huang, J.-D. 2015. A basal ichthyosauriform with a short snout from the Lower Triassic of China. Nature 517, 485-488.

Muscutt, L. E., Dyke, G., Weymouth, G. D., Naish, D., Palmer, C. & Ganapathisubramani, B. 2017. The four-flipper swimming method of plesiosaurs enabled efficient and effective locomotion.

Naish, D. 2023. Ancient Sea Reptiles. Natural History Museum, London.

O’Brien, C. L., Robinson, S. A., Pancost, R. D., Sinninghe Damsté, J. S., Schouten, S., Lunt, D. J., Alsenz, H., Bornemann, A., Bottini, C., Brassell, S. C., Farnsworth, A., Forster, A., Huber, B. T., Inglis, G. N., Jenkyns, H. C., Linnert, C., Littler, K., Markwick, P., McAnena, A., Mutterlose, J., Naafs, B. D. A., Püttmen, W., Sluijs, A., van Helmond, N. A. G. M., Vellekoop, J., Wagner, T. & Wrobel, N. E. 2017. Cretaceous sea-surface temperature evolution: Constraints from TEX86 and planktonic foraminiferal oxygen isotopes. Earth Science Reviews 172, 224-247.

O’Keefe, F. R. & Chiappe, L. M. 2011. Viviparity and K-selected life history in a Mesozoic marine plesiosaur (Reptilia, Sauropterygia). Science 333, 870-873.

Wintrich, T., Hayashi, S., Houssaye, A., Nakajima, Y., Sander P. M. 2017. Triassic plesiosaurian skeleton and bone histology inform on evolution of a unique body plan and survival of end-Triassic extinctions. Science Advances 3: e1701144.

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In the previous article, we looked at the European, scientific discovery of the Okapi Okapia johnstoni…

**Caption:** we shouldn’t take it for granted that one of the greatest zoological discoveries of the 20th century is now a relatively accessible animal to those of us able to visit zoological collections. I always take time to look at captive Okapis (here, at Marwell Wildlife, UK). Images: Darren Naish.

We finished with the official recognition of the Okapi as a new living giraffid species in 1901, culminating with the monographing of the species by E. Ray Lankester and others in the first decade of the 20th century. Many of the events in this tale are well known and often repeated in texts that discuss the Okapi’s discovery.

Less well known and far less frequently discussed is the fact that Europeans might actually have been aware of the Okapi’s existence – albeit in the vaguest sense – prior to Harry Johnston’s recovery of those initial strips of skin in 1900. And also poorly known is the fact that the Okapi’s species-level taxonomy has been slightly more controversial than is generally known. As we’ll see, Okapis are quite variable both anatomically and genetically, and it’s worth considering what implications this might have.

A ’pre-Johnston’ European Okapi specimen. Harry Johnston is credited with being the first European to obtain – and submit to scientific investigation – physical remains of the Okapi (see Part 1). But today we know that he was not, actually, ‘the first’. A different skin strip, about 1 m long, was obtained in June 1899 by the Belgian official Lieutenant Léon Vincart from a local chief, who (once back in Belgium) passed it to his brother Alphonse Vincart at the Maredsous Abbey, near Dinant in Belgium. That might seem like an odd place to deposit a strip of Okapi skin, but it held a natural history museum at the time. The specimen is still retained at the abbey today (Raynal 2023). George A. Boulenger – best known for his abundant pioneering publications on reptiles and amphibians – put the specimen on record in a brief note of 1902 (Boulenger 1902).

**Caption:** the Okapi skin strip collected by Lieutenant Léon Vincart in 1899. Had he succeeding in reporting this specimen in the literature during the year in which he’d found it, he might have won acclaim for its recognition. Image: Jean-Claude Genard, from Raynal (2023).

Incidentally, while Vincart’s Okapi skin specimen is mentioned here and there in literature on the Okapi (e.g., Spinage 1968), the most complete article on it appeared coincidentally within the past few weeks (Raynal 2023). My thanks to Michel Raynal for bringing it to my attention.

While Vincart’s specimen failed to have the same scientific impact that Johnston’s specimens – sent to the Zoological Society of London – most certainly did, it was definitely obtained by a European ‘pre-Johnston’.

**Caption:** there aren’t that many books that provide good, deep information on Okapis and other giraffids. Here are some that I’ve had reason to consult. Image: Darren Naish.

Other Okapi species. Another Belgian official – Lieutenant Leoni, based in the Haut-Ituri District in what’s now north-eastern Democratic Republic of the Congo – obtained the skin of an adult female and near-complete skeleton of an adult male in 1902, and sent both to Brussels. The local people in the region where Leoni was based knew the animal as the N’dumbe, this being another reminder that ‘Okapi’ was likely a specific term used in one small region and not necessarily the best choice of name for the species as a whole (other names mentioned in the literature of the time include M’Boote and Kenghe; Major 1902, p. 344). The Belgians invited Charles Forsyth Major from the British Museum to examine these specimens. He concluded that they represented a different species from the one identified by Johnston and named it Okapia liebrechtsi after a Commandant Liebrechts… though exactly who Commandant Liebrechts is or what contribution they had to the research wasn’t recorded (Major 1902, Spinage 1968).

A third Okapi species was named in 1902 for the skull and skin of a young adult female that was also deemed different from the original skin strips that Johnston had brought to England. E. Ray Lankester – responsible for publishing the generic name Okapia (Lankester 1902a) (see Part 1) – named this Okapia erikssoni after Lieutenant Karl Eriksson (Lankester 1902b), who had otherwise not been credited for his role in procuring the original specimens (again, see Part 1).

**Caption:** the *Okapia erikssoni* holotype as displayed at the British Museum and featured in Lankester (1910). This specimen is otherwise regarded as the first ‘complete’ specimen of *O. johnstoni*, and today is again treated as a member of that species.

Lankester was seemingly still endorsing the existence of Eriksson’s Okapi when he compiled his 1910 atlas of Okapi anatomy, though it’s hard from that work to find any specific statement on how many Okapi species Lankester thought there might be. In the volume’s preface, Sidney Harmer (Keeper of Zoology at the British Museum) drew attention to Lankester’s figuring of distinct broad- and narrow-skulled forms, the implication being that they might belong to two distinct forms (Harmer said “races”). Lankester himself referred to the O. erikssoni holotype without commentary (Lankester 1910), the implication being that he was still regarding it as valid at that time.

Even as early as 1911, however, the idea that there might be more than one species of Okapi was doubted by others (e.g., Lucas 1911) and the concept was essentially abandoned before the 1920s.

**Caption:** Lankester (1910) included numerous diagrams of Okapi skulls and used different width : length ratios of various parts of the skull to demonstrate variation. It is difficult to determine from the images alone what this variation represents, since it could be ontogenetic or sexual rather than anything taxonomically significant. Skull L here (in the middle) is a cast of the holotype skull of *Okapia liebrechti*, named by Charles Forsyth Major in 1902.

The variable Okapi. An additional alleged species – the Kibali okapi Ocuapia kibalensis [sic] – was named in a 1936 book by the Italian explorer and author Attilio Gatti. This animal was said by Gatti (1936) to be especially big, robust and with shorter horns and a different skull shape from other Okapi, though all of this was based on superficial comparisons and there were no formal attempts to establish a type specimen or publish appropriate data. This means that the alleged species has no formal standing.

The generic name ‘Ocuapia’ was not intended as a new genus, since Gatti (1936) used the same spelling was used for O. johnstoni. I think that Gatti just got it wrong.

**Caption:** an Okapi skull montage provided by Gatti (1936), showing females at left and males and right, and with the supposedly new ‘*Ocuapia kibalensis*’ forming the upper row. This montage actually comes from Arment (2018); in the only copy of Gatti (1936) I’ve seen, the male skulls alone are featured.

Needless to say, neither O. liebrechti, O. erikssoni nor ‘Ocuapia kibalensis’ are recognised today. However, could it be that at least one of them might warrant taxonomic distinction after all? One of the points emphasised by Lankester in his 1910 atlas of Okapi anatomy is that the animals are surprisingly variable in skull proportions and pigmentation. Stanton et al. (2014) collected DNA from Okapi faeces, skin clippings, museum specimens and artefacts in Congolese villages and found substantial genetic variation within this animal.

**Caption:** phylogenetic tree from **Stanton *et al*. (2014)**, showing genetic variation in *Giraffa* giraffes (where all eight clades shown here are conventionally recognised as subspecies, and sometimes as species) and Okapi. Note that *Okapia* consists of two main clades, both of which appear to be genetically diverse. Image: **Stanton *et al*. (2014)**.

Two main lineages are present within living Okapis, both of which are estimated to have diverged around 1.7 million years ago, and several major divergences between younger lineages on the tree appear to have occurred more than 1 million years ago (Stanton et al. 2014). Other African artiodactyls with lineage divergence dates of this age are mostly (albeit not universally) thought to consist of several subspecies, or even deserve to be split into several or many species.

Stanton et al. (2014) compared Okapi phylogeny and divergence data with that of bushbuck and giraffe subspecies (or species, take your pick) and with duiker species too, the point being that Okapia johnstoni as currently recognised is more genetically variable than artiodactyls universally considered polytypic. Presumably this reflects a history of being split up into refugia during Pliocene and Pleistocene times, but does it mean that there might be some or many Okapi subspecies or even species? Does this genetic variation map well (or at all) with the anatomical variation?

A lot of difficult work would be required to properly sort this out. What this genetic work indicates is that certain Okapi populations require special emphasis in conservation and management. For an animal that’s already in a dire situation with respect to conservation requirements, this is not good news.

**Caption:** I became interested in seeing how much anatomical variation might exist in the captive Okapis I’ve seen and made a few comparisons. They prove very similar: but that’s expected, since zoo animals (especially within the same geographical area) tend to come from the same source stock, and be closely related. The female Okapi at left is currently on show at London Zoo, the male at right was photographed at Marwell Wildlife in 2022. Images: Darren Naish.

Other claimed early ‘discoveries’. As I mentioned in the previous article, the basics of the Okapi discovery story are well known and often recounted. Less well known is that a few other European people claimed to have encountered Okapis prior to Johnston’s findings of 1900 and 1901. If they’re genuine, they might demonstrate some European awareness of the animal one or two decades earlier. But… are they genuine? I owe my knowledge of these cases to Clive Spinage and his 1968 The Book of the Giraffe for discussion of these accounts.

In 1883, the Russian explorer Wilhelm Junker was in the Nepo Distinct of the Congo when he obtained the complete skin (albeit missing the head and feet) of a hoofed mammal said to be about similar in size to a dwarf antelope, known to some of the locals as the makapi, and suggested by Junker to be a new kind of chevrotain. Junker argued that this was an Okapi skin – surely that of a baby one – which, if true, means that Okapi remains were procured by a European 17 years prior to Johnston’s obtaining the skin bandoliers. Alas, I don’t think we have any way today of testing this claim.

What appears to be a fictitious early European ‘observation’ of the Okapi was provided by Captain James Baptiste Marchand, an officer in the French army, in 1905. Marchand was apparently in the vicinity of Bahr el Ghazel in South Sudan in June 1898 where he was sailing along a tributary of the White Nile in the direction of Kodok, known at the time as Fashoda.

**Caption:** in case you need help visualising the location of the Bahr el Ghazel region in South Sudan, these maps should help. At left: South Sudan (in green), located to the north-east of Democratic Republic of Congo. At right: the Bahr el Ghazel region (in red) within South Sudan. Images: Martin 23230, CC BY-SA 3.0 (original **here**); NordNordWest, CC BY-SA 3.0 (original **here**).

Within the broader context, Marchand’s presence in the region was due to British-French conflict and the colonial Scramble for Africa, and Marchand himself was a veteran of the conquest of both Senegal and what was then known as French Sudan (Mali). Here’s another powerful reminder – as if it were needed – that the European discovery of the Okapi is tightly linked with colonial exploitation and the subjugation of Africans.

Marchard was watching the shore from the deck of his vessel when he observed a remarkable and unusual, beautiful animal seen among a group of antelopes at the edge of a river. It had enormous drooping ears – first likened by Marchand to “horns like the mouflon of Kashmir” – and small horns, and Marchand recognised immediately that it represented a kind of animal as yet unknown to formal zoology. The animal ran away, and Marchand then decided to shoot it, but missed. On seeing a mounted specimen in Paris, he later realised that his mystery Sudanese ‘antelope’ had been one at the same.

But… an Okapi in South Sudan, seen at the edge of the river among a group of antelope? Spinage (1968, p. 144) said that the tale “is far too precise to have a ring of truth about it” and wrote it off as a hoax. I have to concur.

**Caption:** a very beautiful, very dark male Okapi at ZooParc de Beauval in France. Image: Daniel Jolivet, CC BY 2.0 (original **here**).

And that’s where we’ll end things. While some Europeans claim to have seen live Okapis, or even obtain physical remains, prior to Harry Johnston’s discovery of 1900, these are of dubious standing, the exception being Léon Vincart’s recovery of a skin segment in 1899. And while early arguments about Okapi species-level taxonomy have mostly been forgotten in recent decades, phylogenetic work on Okapi diversity implies that there may come a time when this issue needs revisiting.

Needless to say, much more could be said about Okapis: in particular, about their ecology, behaviour and conservation status. I would like to come back to those issues in time, so hold that thought.

For previous mentions of Okapis at Tet Zoo, and other giraffid-themed writings, see…

Giraffes: set for change, January 2006
More on what I saw at the zoo, June 2006
Stuffed megamammal week, day 3: Okapi, April 2009
Inside Nature’s Giants part IV: the incredible anatomy of the giraffe, July 2009
Because giraffes are heartless creatures, and other musings, January 2012
Burning Question for World Giraffe Day: Can They Swim?, June 2016
The Discovery of the Okapi, Part 1, April 2023

Tetrapod Zoology is dependent on funds raised at patreon. Please help support this blog if you consider it worthwhile and want to see me continue doing it, thanks!

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Arment, C. 2018. Profiles in cryptozoology: Commander Attilio Gatti. BioFortean Notes 6, unpaginated.

Boulenger, G. A. 1902. Fresh record of the okapi. The Field 99, 823.

Gatti, A. 1936. Great Mother Forest. Hodder and Stoughton, London.

Lankester, E. R. 1902a. On Okapia, a new genus of Giraffidae, from Central Africa. Transactions of the Zoological Society of London 16, 279-307.

Lankester, E. R. 1902b. The specific name of the Okapi presented by Sir Harry Johnston to the British Museum. Annals and Magazine of Natural History, series 7, 10, 417-418.

Lankester, E. R. 1910. Monograph of the Okapi. Trustees of the British Museum, London.

Lucas, F. A. 1911. [Review of] Monograph of the Okapi. Science 33, 65-66.

Major, C. I. F. 1902. On the okapi. Proceedings of the Zoological Society of London 1902, 73-79.

Raynal, M. 2023. Eine Okapi-Haut im Jahr 1899. Jahrbuch für Kryptozoologie 3 (3), 279-282.

Spinage, C. A. 1968. The Book of the Giraffe. Collins, London.

Stanton, D. W. G., Hart, J., Galbusera, P., Helsen, P., Shephard, J., Kümpel, N. F., Wang. J., Ewen, J. G. & Bruford, M. W. 2014. Distinct and diverse: range-wide phylogeography reveals ancient lineages and high genetic variation in the endangered Okapi (Okapia johnstoni). PLoS ONE 9 (7): e101081.

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If you’ve heard of the Okapi Okapia johnstoni – and if you’re at all interested in the history of zoology – chances are high that you’ve heard the story of this animal’s scientific discovery…

**Caption:** taxiderm mount of the complete Okapi skin obtained by Sir Harry Johnston, photographed in the British Museum (Natural History). This specimen was key in the naming of Ray Lankester’s naming of the genus *Okapia* in 1901. Early depictions of the Okapi – including this one – gave them slimmer limbs than those present on the live animal. Image in public domain.

The story of the Okapi’s discovery has been told several times (Lankester 1902, Wendt 1959, Spinage 1968, Ley 1987, Heuvelmans 1995, Shuker 2012), most frequently in books written by authors especially interested in cryptozoology, the implication being that the relatively late scientific recognition of the Okapi might provide justification for the continuing existence of other large, mysterious terrestrial animals. My retelling here will be familiar to those who know the same books that I do. I have, however, augmented things with tangents and considerations that haven’t been included before, though I have to say that the novel stuff mostly appears in Part 2.

**Caption:** I’m lucky enough to have seen live Okapis on many occasions, though always in zoos of course. These males were photographed at Marwell Wildlife, UK. Images: Darren Naish.

As ever, we should make it clear that we’re talking here about European or scientific discovery, since the Okapi was known for sure to people of the Congo region.

On that note, the overwhelming take-home from the story of the Okapi’s discovery is that it goes hand-in-hand with colonial exploitation and the numerous suppressions of rebellions, administrative reorganisations and negotiations of land ownership that occurred throughout the so-called Scramble for Africa. It doesn’t take much digging to link at least some of the British and Belgian officials concerned to problematic events, all of which means that the story of European science in the Congo is ripe for reframing and a more accurate retelling. I apologise for any shortcomings in the text here.

Today, the area inhabited by the Okapi is within the Democratic Republic of the Congo (or DRC), previously (between 1971 and 2006) known as the Republic of Zaire, before that (between 1908 and 1964) termed Belgian Congo, and before that (between 1885 and 1908) known as Congo Free State. The proximity of the Okapi’s range to the extreme north-east border of DRC means that this area was sometimes within the borders of adjacent Uganda.

**Caption:** current range of the Okapi as depicted by the IUCN. It is entirely within DRC. From **Mallon *et al*. (2015)**.

Europeans learn of the Atti. The first instance of European awareness of the Okapi came in 1890 when author, soldier, journalist and colonial administrator Henry M. Stanley made fleeting reference to a forest ‘donkey’ in volume 2 of his Darkest Africa*. The Wambutti people, said Stanley, knew it as the Atti. Actually, it’s not entirely clear that this was the word they had for the animal, and this might instead have been their word for ‘ass’ or ‘horse-type animal’. At some point, the alleged existence of this ‘Atti’ became known to explorer, author and colonial administrator Harry Johnston (made Sir Harry in 1896), though whether he learnt this from a meeting with Stanley or from reading Stanley’s book isn’t clear, since he claimed both sets of events at different times.

Herodotus mentioned a ‘horned ass’ that inhabited the forests of Libya, and there have been suggestions that this might have been a reference to the Okapi. But it’s really not good enough to say that it was. Other claims of early European awareness of the Okapi have been made; we’ll get to them later.

Caption: the two British people mostly associated with the discovery of the Okapi. At left, Welsh explorer, author, colonial administrator Henry M. Stanley (1841-1904) as he looked in the 1870s. At right, English artist, linguist, colonial administrator Harry Johnston (1858-1927) photographed prior to 1895. Both images are in the public domain.

Descriptions that appear to pertain to the Okapi had been obtained by the Belgians by 1897 when an official learnt from members of the Momvus tribe (in north-east Congo) of a creature termed the Ndumbe. It was taller than a buffalo, maroon-brown across the body, striped with white across the rump and legs, and graceful like a zebra in “form and finish” (Spinage 1968, p. 146).

**Caption:** such is the fame and impact of the Okapi’s discovery that the animal still remains associated with surprising and potential zoological finds. It’s been used as the symbol for at least two cryptozoological organisations, and has a prominent role on the cover of Karl Shuker’s 2012 *The Encyclopaedia of New and Rediscovered Animals*. The other animal on the cover is the Saola *Pseudoryx nghetinhensis*, named in 1993 and often inferred to be ‘the Okapi of the late 20th century’.

A new forest zebra. In 1899/1900, Harry Johnston was tasked with returning a group of Mbuti pygmies – abducted by a German and planned for display in Paris (my god, Europeans do not come out of this looking at all good) – to their home in the Congo. Via an interpreter, Johnston learnt what he could from them and asked them about the horse-like Atti; they termed it o’api (pronounced ‘ocwapi’) (Spinage 1968, p. 147) and conveyed to him the idea that it was like a donkey but equipped with the stripes of a zebra.

Once in the Congo, Johnston travelled to the Belgian outpost known as Fort Mbeni. There, he was told by Lieutenant Meura that the animal was familiar, as carcasses were often bought in by the locals. I’m not sure how remarkable we should find it that Meura apparently regarded the retrieval of deceased Okapi as fairly commonplace, and Spinage (1968) speculated that Meura and his people might have been scolded by their superiors by having apparent ready access to this significant zoological discovery and letting outsiders come in and claim it for themselves.

**Caption:** at left, the two original bandoliers on which the zoological reality of the Okapi was established, as described by Sclater in 1901. At right, a plate from Lankester’s Okapi ‘atlas’ of 1910 in which the two bandoliers were matched up to complete Okapi skins. Numerous of these bandoliers were collected and retained over the years. Images: Sclater (1901); Lankester (1910).

There might not have been a carcass on hand at Fort Mbeni, but there had recently been a skin. Alas, soldiers (part of a local militia, not Belgian soldiers) had cut it into strips, each of which was about 3 ft long and intended for use as a belt or bandolier. Johnston was allowed to take two of these, and he sent them to Philip Sclater of the Zoological Society of London in the November of 1900, who then exhibited them as a meeting of the society in December (Lankester 1902).

**Caption:** Sclater’s initial naming of the Okapi, wherein it’s a new species of horse.

Sclater published a brief description of them in February 1901, interpreting them as the first evidence for a new living horse that he named Equus(?) johnstoni (Sclater 1901), the question mark denoting the provisional nature of this classification. Hairs from the skin sections were examined under the microscope by W. G. Ridewood and Prof. J. C. Ewart. Ridewood found them to be indistinguishable from those of zebras while Ewart concluded that the hairs were more like those of zebras than those of antelopes.

The view that the Okapi was an equid was consistent with Johnston’s view at this time, but references to more than one hoof on each foot led him to think initially that it might be a late-surviving horse like those known from the fossil record, in particular the three-toed Hipparion.

**Caption:** numerous species of the extinct horse *Hipparion* have been named from the fossil record of Europe, Asia, Africa and North America, and today it seems remarkable that Johnston thought that he might have been on the track of living specimens of this animal. As is evident from this reconstruction (showing the North American *H. forcei*), *Hipparion* species were mostly small relative to living horses. If the Okapi was really imagined as a living *Hipparion*, it would have to have been a giant one. Image: Nobu Tamura, CC BY-SA 4.0 (original **here**).

Remarkably, Johnston – supplied with guides provided by Lieutenant Meura – was shown apparent Okapi tracks in the Ituri Rainforest. Because they were made by a cloven-hoofed animal, he concluded that they couldn’t be Okapi tracks but must be those made a big antelope, like an eland (a forest-dwelling form of which would still have been a noteworthy discovery), and he refused to follow the trail. The expedition was a disaster anyway, with many members of the group contracting malaria and requiring rescue by the Belgians based at Mbeni. Johnston complained of the humid, oppressive forest interior.

The Okapi is a new living giraffid. Johnston might not have been successful on that trip, but his fortunes changed in March 1901 when Lieutenant Karl Eriksson – second in command at Mbeni – sent Johnston an Okapi skin with a skull, as well as a second, smaller skull. Eriksson broke official Belgian rules in sending this material to Johnston, since specimens of interest were supposed to go to Tervuren Museum* near Brussels (Spinage 1968). Again, the Belgians lost out on getting credit for this find, but – in saying this today – it’s impossible to once again avoid thinking of the colonial exploitation that was happening here and the absence of specific Congolese people from the narrative.

Today, the Royal Museum for Central Africa or AfricaMuseum.

Caption: Johnston was a skilled artist, and produced a number of black and white and colour illustrations of the Okapi based on descriptions and his viewing of the complete skin he obtained. I feel that these illustrations are too pointy-snouted and slender in the limbs, but they’re otherwise highly accurate. Images in public domain.

The skin that Eriksson acquired had cloven hooves, thereby confirming what Johnston had heard. Unfortunately, they were lost by the time the remains reached him, apparently because they became detached and fell away. A few anatomical features (like a bilobed lower canine and an absence of false hoofs) showed immediately that the Okapi was a giraffid. Johnston sent this material to E. Ray Lankester in England, suggesting that it might best be named as a living species of the European fossil giraffid Helladotherium for which he proposed the name H. tigrinum (Lankester 1902).

**Caption:** a key giraffid character recognised immediately once an Okapi skull was obtained is its bilobed lower canine tooth. This montage – from Lankester (1902) – compares the Okapi lower canine with that of other giraffids living and fossil.

Johnston was a painter, and produced several illustrations, the best known of which is a colour piece depicting two Okapis, both imagined based on his understanding of the skin and skulls. His renditions proved pretty accurate to the look of Okapis in life, one point of interest being that Johnston opted to show the upper margin of the neck continuing in the same approximate line as the dorsal margin of the back, rather than being angled sharply upwards as is typical for artiodactyls like deer and antelopes.

**Caption:** larger version of the fine Okapi illustration produced by Harry Johnston at the same time as he passed the complete skin to Lankester. Johnston wrote a letter on this painting in March 1901, addressed to Philip Sclater, and explained therein how he had taken great care to properly convey the form and colour of the animal in life. He was concerned that the colour of the coat might have faded by the time it reached England and emphasised its reddish nature in some regions. Image in public domain.

Okapia sees print. The remains sent to England by Johnston did not arrive until July 1901. Lankester, however, knew enough to make a taxonomic decision already, and his announcement of the new generic name Okapia Lankester, 1901 was made on June 18th at a meeting of the Zoological Society. His first, initial paper was then published in Proceedings of the Zoological Society of London (Lankester 1901), this being generally regarded ever since as the first official published outing of the name (even though it appeared in other venues at about the same time due to publicity surrounding the Zoological Society meeting). Sclater’s ‘forest horse’ Equus johnstoni clearly pertained to the same species, so the full name had to be Okapia johnstoni* (Sclater, 1901). Actually, Lankester later changed his mind on this, a point we’ll return to in the next article.

The dates provided here come from Lankester’s own publications on the Okapi (Lankester 1902, p. 282) and from the Zoological Society’s published records. They’re inconsistent with sources which state that the Zoological Society meeting happened on May 7th, and even that the skin and skulls were on display at this time too.

Lankester then published a longer, more detailed description of the Okapi in Transactions of the Zoological Society of London (Lankester 1902), and then a comprehensive atlas of Okapi images and illustrations (Lankester 1910). It has been implied that he planned to write a full descriptive monograph but the appearance of two other works – a monograph written by Julien Fraipont, published in 1907 (Fraipont 1907), and a second by Maurice de Rothschild and Henri Neuville which appeared in 1910 (Rothschild & Neuveille 1910) – apparently caused him to abandon these plans.

**Caption:** most major European museums obtained Okapi specimens at some point during the 20th century. This one is on show at the National Museum of Ireland in Dublin. Its fading is typical for dark animals on show in collections. The prominent scars visible on its coat are said to have been caused when it was killed. The animals were trapped in pits and then speared. Image: Darren Naish.

This is far from the end of the Okapi story, but here is where I’ll end for now. Other Okapi species were named after 1901, and claims of earlier ‘discoveries’ of the species were made too. These and other topics will be covered in the next article….

I’ve been saying since 2009 that I’d one day cover the discovery of the Okapi. For previous mentions of Okapis at Tet Zoo, and other giraffid-themed writings, see…

Giraffes: set for change, January 2006
More on what I saw at the zoo, June 2006
Stuffed megamammal week, day 3: Okapi, April 2009
Inside Nature’s Giants part IV: the incredible anatomy of the giraffe, July 2009
Because giraffes are heartless creatures, and other musings, January 2012
Burning Question for World Giraffe Day: Can They Swim?, June 2016

Tetrapod Zoology is dependent on funds raised at patreon. Please help support this blog if you consider it worthwhile and want to see me continue doing it, thanks!

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Rothschild, M. de & Neuville, H. 1910. Recherches sur l’okapi et les girafes de l’Est Africain. Annales des Sciences Naturelles Zoologie 9, 1-93.

Fraipont, J. 1907. L’okapi. Ses affinités avec les giraffides vivants et fossils. Bulletin de l’Académie Royale de Belgique 12, 1097-1130.

Heuvelmans, B. 1995. On the Track of Unknown Animals. Kegan Paul International, London.

Lankester, E. R. 1901. On Okapia johnstoni. Proceedings of the Zoological Society of London 1901 (2), 279-281.

Lankester, E. R. 1902. On Okapia, a new genus of Giraffidae, from Central Africa. Transactions of the Zoological Society of London 16, 279-307.

Ley, W. 1987. Exotic Zoology. Bonanza Books, New York.

Mallon, D., Kümpel, N., Quinn, A., Shurter, S., Lukas, J., Hart, J. A., Mapilanga, J., Beyers, R. & Maisels, F. 2015. Okapia johnstoni. The IUCN Red List of Threatened Species 2015: e.T15188A51140517.

Sclater, P. L. 1901. On an apparently new species of zebra from the Semliki Forest. Proceedings of the Zoological Society 1901 (1), 50-52.

Shuker, K. P. N. 2012. The Encyclopaedia of New and Rediscovered Animals. Coachwhip Publications, Landisville, Pennsylvania.

Spinage, C. A. 1968. The Book of the Giraffe. Collins, London.

Wendt, H. 1959. Out of Noah’s Ark. Weidenfeld & Nicolson, London.

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Time to engage in more lark-themed musings of a phylogenetic nature….

**Caption:** lark images from the previous lark article. Images (clockwise from left): H. E. Dresser, in public domain; Darren Naish; J. G. Keulemans, in public domain.

You’ll no doubt have read the recent Tetrapod Zoology article on larks – A History of Larks: Twists and Turns and Overlooked Convergence – and thus assigned its various arguments and contentions to memory. Here, I want to circle back to what I said in that article about certain lark ‘species’ being species complexes. Buckle in, it’s quite a ride.

If you’re at all familiar with the diversity and taxonomy of living birds, you’ll know that a great many species are polytypic, consisting of some or many ‘subspecies’ that are often about as different from the nominal ‘subspecies’ as are supposedly distinct, related species within the relevant group. The subspecies issue is often treated as a sort of joke, as if subspecies have been invented by bureaucrats for book-keeping purposes or by birdwatchers desperate to increase their list of ticks in a notebook.

**Caption:** you might get the impression from all the complaining that subspecies are some irksome ever-present annoyance that you can’t escape from. They’re really not. Books like these only ever include data on a handful of them, and if you actually want to know what’s going on you have to dig deep into the technical literature. Image: Darren Naish.

‘Subspecies’ 1: mostly a waste of time. On the negative side of things, it is indeed the case that quite a few ‘subspecies’ don’t prove worthy of recognition when we study the relevant populations in detail. Those supposed subspecies that can be differentiated due to slightly larger or smaller size, or lighter or darker plumage, sometimes seem instead to be end members of a cline, and they prove unsupported as distinct entities when their genetics are examined. I’m not old enough to really remember a time when British birds were treated as endemic ‘subspecies’, but I do own books from the 1950s and 60s where at least some British populations of Eurasian species are treated as such. Today, we mostly laugh at that idea (Hebridean wrens excepted).

**Caption:** Charles Tunnicliffe’s 1965 *Wild Birds of Britain* is a wonderful piece of work. But it was written at a time when people were still thinking that even the most minor of anatomical differences were sufficient for a population to be differentiated as a ‘subspecies’. Are British Goldcrests *Regulus regulus* really distinct enough from those of mainland Europe to be worthy of subspecific recognition? Today, we say no.

Examples in larks of defunct subspecies include the many supposed local forms of Greater short-toed lark Calandrella brachydactyla and Red-capped lark C. cinerea, most of which have not been found to be supported by cytochrome b analysis (Stervander et al. 2016).

‘Subspecies’ 2: actually good, worthy, and deserving of recognition. Having said all that, the fact remains that some and perhaps many subspecies are actually ‘distinct’. If you adhere to a ‘diagnostic species concept’ that’s combined with studies of phylogeny, those entities we term ‘species’ are those biological units that can (1) be defined and differentiated on the basis of diagnostic traits and (2) form ‘distinct lineages’ in the phylogeny, which is to say that they’re not nested within another unit already recognised as a species and diverged at an estimated point in geological history consistent with species status. Take this as far as it can go, and many ‘subspecies’ should actually be regarded as ‘species’: a decision that might require us to approximately double the recognised number of living bird species (Zink 1996, 2004, Zink & McKitrick 1995, Barrowclough et al. 2016).

For all those claims that increasing the number of ‘species’ is due merely to ‘taxonomic inflation’ and changing species concepts, the counter-argument is that the relevant decisions have been based on appropriate study and newly acquired data (Sangster 2009). That’s obvious if you go to the trouble of digging into the technical literature. Modern papers that make taxonomic recommendations about species boundaries involve a substantial amount of hard work.

**Caption:** mostly gone are the days where you can just dismiss subspecies as local variants invented by people who never leave museums. Modern analyses of subspecies-level variation involve morphometrics, DNA and data on range and ecology. The new Red-capped lark subspecies *C. cinerea rufipecta* was named by Stervander *et al*. (2020).

Larks: cryptic species and rampant paraphyly. Let’s take this back to larks again. Recent studies have shown that at least some taxa once considered ‘subspecies’ warrant recognition as species, both because they’re diagnostic, and because they’ve been found to be ‘distinct lineages’ as explained above.

Stervander et al. (2016, 2020) looked at populations within Calandrella larks, and found Greater short-toed lark C. brachydactyla, Red-capped lark C. cinerea, Hume’s short-toed lark C. acutirostris and others to have “unexpected deep divergences” (meaning that they originated a surprisingly long time ago within the geological history of the group), and to sometimes be paraphyletic with respect to other Calandrella species. The data revealing “unexpected deep divergences” requires that certain of the lineages conventionally included within the species concerned diverged during the Zanclean Age of the Pliocene (about 4 million years ago), or even before this, meaning that they almost certainly represent cryptic species.

**Caption:** Greater short-toed lark photographed in Maharashtra, India. At right, *Calandrella* lark phylogeny from Stervander *et al*. (2016), based on mitochondrial gene *b*. Of note here is that some subspecies (like *C. brachydactyla dukhunensis*, shown in orange) don’t group with the rest of their species, and that some subspecies-level divergences (like that between *C. b. dukunensis* and the rest of *C. brachydactyla*, and those within *C. blanfordi*) are posited to have occurred during the Pliocene. Images: Shantanu Kuveskar, CC BY-SA 4.0 (**original here**); Stervander *et al*. (2016).

An east Asian lark conventionally included within the Greater short-toed lark has been shown in several studies to be closer to Hume’s short-toed lark and be very distant on the tree from the Greater short-toed lark. It’s now generally regarded as a distinct species: the Mongolian short-toed lark C. dukhunensis (Alström et al. 2013, 2020).

Ghorbani et al. (2020) found populations conventionally grouped together within the Lesser short-toed lark Alaudala rufescens to form four clades that diverged from one another between about 1.6 and 3.2 million years ago, their conclusion being that there are probably four cryptic species here. Furthermore, the Sand lark Al. raytal is nested within this complex, meaning that the traditional version of the Lesser short-toed lark is paraphyletic (Ghorbani et al*. 2020).

Alaudala rufescens was conventionally termed the Lesser short-toed lark. But since the publication of the studies discussed here, it’s become increasingly popular to refer to A. rufescens in the new, restricted sense as the Mediterranean short-toed lark.

Caption: you don’t need to see the whole cladogram (its top is chopped off) to get the point here. Ghorbani et al. (2020) found the animals conventionally included together within Alaudala rufescens to consist of four clades, with the Sand lark Al. raytal nested in among them. Once again, note that the divergence times of these clades occurred at times ordinarily deemed ‘about right’ for species-level divergences. Image: Ghorbani et al. (2020).

Finally, Stervander et al. (2020) found deep evolutionary divergences within species of Eremophila (horned larks), Galerida (crested larks), Alauda (skylarks), Alaemon (hoopoe-larks), Eremopterix (sparrow-larks), and in others. Species in all of these groups include ‘subspecies’ that diverged more than 2, or more than 3, or even more than 5 million years ago. These divergences corresponded to biogeographical barriers, are “mirrored by several concordantly distributed mammalian savanna specialists” (Stervander et al. 2020, p. 17), and sometimes concern isolated populations that appear to be relicts* of occurrence across a broader, ancestral habitat.

Stervander et al. (2020) used the word ‘relic’ but that doesn’t seem right to me. Isn’t ‘relic’ more associated with the leftovers of human civilisations? I’m more used to seeing organisms being referred to as ‘relictual’.

Caption: at left, a Mediterranean short-toed lark Alaudala rufescens photographed in Israel. At right, a Sand lark Al. raytal photographed in India. Genetics indicates that Al. raytal is nested within the traditional version of Al. rufescens where it’s one of five deep, distinct lineages. The solution most consistent with decisions made elsewhere in passerine taxonomy is that all five should be retained as species or elevated to species level. Larks of many sorts can raise and lower the feathers on the crown as desired, and thus end up with different profiles one moment to the next. Images: Lior Kislev, CC BY-SA 4.0 (original here); Savithri Singh, CC BY-SA 4.0 (original here).

Take-home. I know this can be hard to follow if you don’t know all the species/subspecies in question, but the take-home messages here are pretty simple. And while I’ve concentrated here on larks, they’re echoed in numerous other studies of passerine groups. Despite what you’ve heard and read about zoologists – and ornithologists in particular – being trigger-happy splitters, overly keen to recognise ‘species’ when a simpler, lazier, lumpier approach will suffice, the fact is that many (perhaps most) polytypic ‘species’ aren’t monophyletic when studied in detail. The more distinct of their supposed ‘subspecies’ don’t belong with the nominal form and go elsewhere in the tree.

Reticulation and hybridisation introduce further complexity to this issue, for sure, but are not immediately relevant to the cases discussed in this article.

**Caption:** here’s a hypothetical time-calibrated cladogram that illustrates the sorts of issues we’re dealing with. How do we treat (in the taxonomic sense) taxa B, C and E? The primary assumption here is that taxa that have been separate for more than c 2 Ma are ‘different enough’ to be species. B seems here not to belong to the same species as D (as had previously been thought), so is it a new species, or is it part of the same species as A? Is C a distinct species, or is it part of the same species as D and E? And is even E ‘distinct enough’ from D to be a distinct species? After all, the two diverged in the Pliocene.

It’s not just ‘species’ that are non-monophyletic. A vast number of studies now (referring to passerine birds here) show that traditional genera are paraphyletic or monophyletic, and are little more than ‘form taxa’ when analysed rigorously. Do we adapt to this by adopting a very ‘lumpy’ taxonomy, or do we do the opposite and resurrect many old names and coin many new ones? Here, the former approach is mostly winning out.

In which we end by beating up Carl Linnaeus. I’ll end by saying that all of this constant chopping and changing to taxonomy requires that writers and researchers have to check which species- or genus-level name is the ‘correct’ one at the time of writing. And a lot of people don’t like this, the main complaint being that they’d prefer if it taxonomic labels just stayed the same.

I have two points in response to that. Point 1 is that the Linnaean binomial system is a bad one, and that we’ve done ourselves no favours by sticking with it. Linking taxonomy (the naming of things) with phylogeny (the determining of the relatedness of things) was always a bad idea, since our phylogenetic models are constantly in flux. They always have been, and they always will be. We’re thus stuck with a system where taxa will always be moving around, and where names will need to be changed.

**Caption:** oh Carl, what have you done? Image: Nationalmuseum (Stockholm), public domain (**original here**).

Point 2 is that this changing is a consequence of people learning new stuff and doing new studies. So long as we keep learning new stuff about phylogeny and thus about the phylogenetic position of organisms, we’ll need to keep modifying the taxonomy. If you’re interested in nature and science, you presumably want people to keep doing scientific research. Ergo, complaining about the labile nature of taxonomy (see Point 1) can come across as anti-scientific (see Point 2).

Is there an alternative? Yes. Devise a new system where taxonomy is not connected to phylogeny. Could such a system replace the existing one, and is it really worth the hassle at this point? Dear reader, I leave you to decide.

For previous Tet Zoo articles on passerines, see…

Great tits: still murderous, rapacious, flesh-rending predators!, February 2013
For The Love of Crows, October 2015
Thoughts on the Passerine Tree, 2016, October 2016
A Battle Among Blue Tits, February 2018
Cocks-of-the-Rock, Extreme Cotingas, April 2019
Birdwatching in Suburban China, May 2019
Tell Me Something Interesting About Dunnocks, July 2019
Suburban Birdwatching in Queensland, Australia, November 2019
Kirk W. Johnson’s 2018 The Feather Thief, a Review, February 2021
The Locustella Warblers, May 2021
Chiffchaffs and What Are 'Old World Warblers' Anyway?, February 2022
Surprising Diet of the House Sparrow, August 2022
A History of Larks: Twists and Turns and Overlooked Convergence, February 2023

Support this blog AND the newly revived podcast at my patreon, where you also get to see tons of behind-the-scenes material that hasn’t yet been released anywhere else.

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Alström, P., Barnes, K. N., Olsson, U. Barker, F. K., Bloomer, P., Khan, A. A., Qureshi, M. A., Guillaumet, A., Crochet, P. A. & Ryan, P. G. 2013. Multilocus phylogeny of the avian family Alaudidae (larks) reveals complex morphological evolution, non-monophyletic genera and hidden species diversity. Molecular Phylogenetics and Evolution 69, 1043-1056.

Alström, P. & Sundev, G. 2020. Mongolian short-toed lark Calandrella dukhunensis, an overlooked East Asian species. Journal of Ornithology 162, 165-177.

Barrowclough, G. F., Cracraft, J., Klicka, J. & Zink, R. M. 2016. How many kinds of birds are there and why does it matter? PLoS ONE 11, e0166307.

Ghorbani, F., Aliabadian, M., Zhang, R., Irestedt, M., Hao, Y., Sundev, G., Lei, F., Ma, M., Olsson, U. & Alström, P. 2020. Densely sampled phylogenetic analyses of the Lesser Short-toed lark (Alaudala rufescens) — Sand lark (A. raytal) species complex (Aves, Passeriformes) reveal cryptic diversity. Zoologica Scripta 49, 427-439.

Sangster, G. 2009. Increasing numbers of bird species result from taxonomic progress, not taxonomic inflation. Proceedings of the Royal Society of Biological Sciences, Series B 276, 3185-3191.

Stervander, M., Alström, P., Olsson, U., Ottosson, U., Hansson, B. & Bensch, S. 2016. Multiple instances of paraphyletic species and cryptic taxa revealed by mitochondrial and nuclear RAD data for Calandrella larks (Aves: Alaudidae). Molecular Phylogenetics and Evolution 102, 233-245.

Stervander, M., Hansson, B., Olsson, U., Hulme, M. F., Ottosson, U. & Alström, P. 2020. Molecular species delimitation of larks (Aves: Alaudidae), and integrative taxonomy of the genus Calandrella, with the description of a range-restricted African relic taxon. Diversity 12 (11), 428.

Zink, R. M. 1996. Bird species diversity. Nature 381, 566.

Zink, R. M. 2004. The role of subspecies in obscuring avian biological diversity and misleading conservation policy. Proceedings of the Royal Society of London B 271, 561-564.

Zink, R. M. & McKitrick, M. C. 1995. The debate over species concepts and its implications for ornithology. The Auk 112, 701-719.

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A revisiting of a Tetrapod Zoology classic, originally published in 2015…

**Caption:** one of my favourite reconstructions of Piltdown man. It's by Margaret Flinsch and shows supposedly associated proboscideans and horses in a surprisingly tropical Plio-Pleistocene England. Note the (in)famous tool being held by the hominin on the left.

One of the most fascinating episodes in the history of palaeontology is that of Piltdown man, an alleged human ancestor discovered in 1908 at Piltdown in Sussex, England. Formally named Eoanthropus dawsoni in 1912, Piltdown man matched early 20th century expectations of what a human ancestor might be like. It combined a large brain with an ape-like jaw (therefore confirming ideas that the evolution of big brains led the way in hominin evolution), and it lived in Europe (confirming ideas that hominin evolution was a Eurasian event, the hominins of Africa and tropical Asia being divergent irrelevancies or side-branches). The African australopithecines had yet to be discovered, nor had scarcely any of the wealth of fossil African hominins we know of today.

**Caption:** the case against Dawson is really strong. I thought this ever since learning of the Booth Museum 'toad in the hole' when I was about 12. Anyway, the full case was put together by Russell in 2003.

The one thing that every single person who’s heard of Piltdown man knows is that it was eventually determined to be a hoax. This didn’t happen until 1953 when Kenneth P. Oakley, Wilfrid E. Le Gros Clark and Joseph Weiner showed how an orangutan jaw and human skull fragments had all been combined to fool the scientists of the day (Weiner et al. 1953). Exactly who the perpetrator was and what their motives were remain debated, and an enormous amount of work has been published on this subject. This article is not about that particular part of the story, though I will say that amateur archaeologist and palaeontologist Charles Dawson was (in my view) the person most likely to have been behind the hoax.

What’s discussed rather less frequently is that early 20th century views on Piltdown man were far more complex than popularly portrayed. Acceptance of Eoanthropus as a valid proto-human might have been the ‘mainstream’ view that made it into textbooks and encyclopedias, but it certainly wasn’t the only one, nor was this acceptance wholesale or uncontroversial.

Also not especially well known is that Piltdown man was not a singleton. Two specimens were found at locations about 3 km apart. These are known as Piltdown I and Piltdown II, though the Piltdown II specimen (sometimes called the Sheffield Park specimen, and consisting of skull fragments and a tooth) was never properly described, nor was the site identified with precision.

**Caption:** John Cooke's famous 1915 'Piltdown gang' painting. Arthur Keith sits in the middle. Standing at the back (left to right): Barlow, Elliot Smith, Dawson, Woodward. Sitting on the left: Underwood. Sitting at right (left to right): Lankester, Pycraft. This is a big, imposing piece of work - see the photo below. Image in the public domain.

Piltdown man’s describer was Arthur Smith Woodward of the (then) British Museum (Natural History). But Woodward was an ichthyologist and many have wondered if he did himself a favour in devoting so much time and energy to an area that was not his speciality. Regardless, these concerns over Woodward’s expertise did not affect acceptance of Eoanthropus as an ancestor to modern humans among British palaeontologists. Several of Woodward’s colleagues – most notably zoologist William Pycraft (best known for his ornithological work) – were vocal and aggressive supporters of Woodward’s interpretation.

**Caption:** James McGregor's *Eoanthropus* skull reconstruction of 1921. Image in public domain.

Dualism and the dualists. Long prior to 1953 however, certain other anthropologists, primatologists and mammalogists were of the opinion that the cranium and jaw of Piltdown I did not go together, and that while the cranium was human, the jaw was from a chimpanzee or some other non-human ape. In fact, some workers voiced doubts about the authenticity of Woodward’s reconstruction within just two or three years of 1912. We might even go as far as saying that quite a few anthropologists and mammalogists of the early 1900s would not have been surprised on learning that it was a hoax, and some might even have suspected that this is exactly what it was.

This idea that the material was of a ‘dual’ nature became known as dualism, its supporters the dualists. Martin Hinton, a zoologist at the British Museum (Natural History) often linked with the hoax (Gardiner 2003), claimed that he had been aware of this duality since 1912 at least, and he noted that Oldfield Thomas, the museum’s leading mammalogist, was of the same opinion. Of course Hinton’s apparent brilliance takes on a different light if he was the hoaxer as some claim, and he did not come forward with this view until the 1950s.

**Caption:** there's a Piltdown man memorial: I really must go and see it some time. There's a Piltdown man pub too. We should arrange a fieldtrip. Image by Nick Woolley (**original here**), CC BY-SA 2.5.

In 1913, King’s College anatomist David Waterston argued that the Piltdown I jaw was emphatically not that of a human, and that associating it with a human-like cranium was as incongruous as articulating a chimpanzee foot with a human leg (Waterston 1913). This view was also championed by Gerrit Smith Miller – assistant curator of mammals at the United Stated National Museum in Washington – in 1915 (Miller 1915), and by both French palaeontologist Marcellin Boule and German anthropologist Franz Weidenreich in 1923 (Spencer 1990, Walsh 1996, MacRitchie 2011).

**Caption:** portrait of G. S. Miller, taken in 1897 or thereabouts. Miller had an extremely broad knowledge of the world’s mammals, published over 400 scientific contributions, named around 100 new mammal species, and is best known for his work on bats and rodents. Image in public domain.

Miller’s views and comments were especially well argued. He showed how chimpanzee jaws could, if broken in exactly the right places, be made to look almost identical to the jaw of Piltdown I, and he also argued that various details of the cranium indicated that it and the jaw simply couldn’t belong to the same one animal (Miller 1915). He revisited the topic of Piltdown man on a few later occasions (Miller 1918, 1920).

Miller (1915) concluded that the jaw represented a European chimp which he dubbed Pan vetus. The cranium, he thought, should be referred to the genus Homo and the name Eoanthropus, he proposed, should be discarded. Because the jaw and cranium were missing those parts that would ordinarily reveal the way in which they articulated, Miller (1915) noted that “Deliberate malice would hardly have been more successful than the hazards of deposition in so breaking the fossils as to give free scope to individual judgement in fitting the parts together”.

**Caption:** the Piltdown man partial cranium (shown in three views at the top) and mandible (shown in four views at the bottom).

It has been claimed that Miller suspected fraud right from the start (T. D. McCown, in Oakley & Groves 1970) but that he was persuaded to tread lightly given the seriousness of the accusations. We also know that, in 1930, Miller asked Remington Kellogg (best known for his work on fossil whales) to look at the Piltdown I jaw in order to see if the teeth had been artificially modified (Oakley & Groves 1970).

Some of Miller’s colleagues agreed with him, including the American Museum of Natural History’s William King Gregory (1876-1970). But the reaction from others was extremely negative, most notably Pycraft. Furthermore, such noted American workers as Henry Fairfield Osborn and William Diller Matthew accepted Woodward’s interpretation, not Miller’s. Indeed Osborn became a prominent Piltdown proponent after viewing the material in 1921 (Osborn even visited Barkham Manor – the discovery site – with Woodward). Perhaps because of this weight of consensus, Gregory eventually changed his mind as goes agreeing with Miller, and did so at about the same time that Osborn and Matthew were voicing their support for Woodward’s interpretation.

**Caption:** Aleš Hrdlička, photographed in 1930. Image in public domain.

At least one researcher remained on Miller’s side however. Aleš Hrdlička (1869-1943), curator of physical anthropology at the National Museum of Natural History (Washington D.C.), also opposed Woodward’s reconstruction, and also became a dualist. In articles published in 1923 and 1924 he argued that the braincase was essentially that of a modern human while the jaw was from some sort of chimp-like ape, perhaps a dryopithecine (Hrdlička 1923, 1924). Given the poor stratigraphic data recorded from the Piltdown excavation, Hrdlička even intimated that the cranium might be a modern burial that had been incorporated into older strata. We know that Hrdlička and Miller corresponded extensively on Piltdown and it’s possible that Hrdlička instigated Miller’s further research into the subject. As an aside, in his 1990 book on the Piltdown saga, Frank Spencer drew attention to Hrdlička’s efforts of 1912 and before to critically assess Florentino Ameghino’s (1854-1911) startling claims of South American ancestry for humans (Hrdlička and colleagues published a book on this subject in 1912), noting that “Hrdlička was distressed to find his Anglo-Saxon colleagues headed in the same direction” (Spencer 1990).

**Caption:** it's not the original, but it's good enough.

Hrdlička spent time with the Piltdown fossils in 1922. He was troubled by how impossibly similar the molars of Piltdown I and Piltdown II were to one another in every detail, and he wondered if the Piltdown II tooth might have been mislabelled: could it actually be part of the Piltdown I remains? With the benefit of hindsight we can say now that this was another obvious clue indicating the fraudulent nature of the remains. Of course Hrdlička was not about to make such a bold claim.

Why were the dualists ignored? The dualist arguments were there right throughout the sorry history of one of palaeontology’s greatest and most notorious hoaxes, yet were ignored or waved away by several prominent workers who held influential positions. Piltdown’s proponents (like Pycraft) used the weight of authority and special access to the material to dismiss or quash the dualist’s arguments. This is despite the fact that those who made the dualist arguments were actually more qualified, more experienced, and more ‘relevant’ (as goes their qualifications and areas of expertise) when compared to the proponents.

Dualism is discussed at length in all the good books on Piltdown and also formed the focus of one of Stephen Gould’s essays. I would say the same as Gould and other commentators: if leading workers in England, the USA and elsewhere had not been so invested, so committed, in the supposed authenticity of Piltdown man, if only they had listened to their detractors – to Miller, Hrdlička and others – they might well have accepted and understood the very reasonable, very well supported arguments of the dualists. Alarm bells were ringing right from the start.

**Caption:** here's that painting by John Cooke, in its home at Burlington House, the Geological Society of London. Humans for scale. Photo by Darren Naish.

Long-time readers might know that this article has been mentioned on and off since 2006, and the version you’ve just read represents a version first published at ver 3 (the Sci Am years) back in 2015. The original version is here. For previous Tet Zoo articles on hominins and other primates, see...

Zihlman’s ‘pygmy chimpanzee hypothesis’, October 2012
Marmosets and tamarins: dwarfed monkeys of the South American tropics, November 2012
The amazing swimming Proboscis monkey (part I), November 2012
Nasalis among the odd-nosed colobines or The “Nasalis Paradox” (proboscis monkeys part II), December 2012
Old World monkeys of choice, February 2014
De Loys’ Ape and what to do with it, July 2014
Why Humans Are Important to Studies of Primate Diversity, July 2014

Huge thanks to those who help support this blog at patreon. You can see behind-the-scenes and in-prep stuff relevant to projects for as little as $1 per month.

Refs - -

Gardiner, B. G. 2003. The Piltdown forgery: a re-statement of the case against Hinton. Zoological Journal of the Linnean Society 139, 315-335.

Hrdlička, A. 1923. Variation in the dimension of lower molars in man and anthropoid apes. American Journal of Physical Anthropology 6, 195-216.

Hrdlička, A. 1924. New data on teeth of early man and certain other fossil European apes. American Journal of Physical Anthropology 7, 109-132.

MacRitchie, F. 2011. Scientific Research as a Career. CRC Press, Boca Raton, USA.

Miller, G. S. 1915. The jaw of Piltdown man. Smithsonian Miscellaneous Collections 65, 1-31.

Miller, G. S. 1918. The Piltdown jaw. American Journal of Physical Anthropology 1, 25-52.

Miller, G. S. 1920. The Piltdown problem. American Journal of Physical Anthropology 3, 585-586.

Oakley, K. P. & Groves, C. P. 1970. Piltdown man: the realization of fraudulence. Science 169, 789.

Spencer, F. 1990. Piltdown: A Scientific Forgery. British Museum (Natural History) & Oxford University Press.

Walsh, J. E. 1996. Unravelling Piltdown. Random House, New York.

Waterston, D. 1913. The Piltdown mandible. Nature 92, 319.

Weiner, J. S., Oakley, K. P. & Le Gros Clark, W. E. 1953. The solution of the Piltdown problem. Bulletin of The British Museum (Natural History) Geology 2, 141-146.

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Plesiosaurs, ichthyosaurs, mosasaurs and the other sea-going reptiles of the Mesozoic seas are among the most fascinating and awesome animals of all time. But if you want a comprehensive, well-illustrated book that reviews them all… well, you’re out of luck, since no such volume exists. UNTIL NOW.

Yes, today see the publication of my new book *Ancient Sea Reptiles* (Natural History Museum in the UK; Smithsonian Books in the USA), and getting it published marks a major personal achievement. I’ve been trying for years to get such a book off the ground, but it’s the success of *Dinosaurs: How They Lived and Evolved* (Naish & Barrett 2016) that’s allowed things to go forward. I can’t express how pleased I am that things have finally worked out.

Ancient Sea Reptiles is approximately similar in style and format to Dinosaurs: How They Lived and Evolved but is substantially more focused on a ‘group-by-group’ arrangement of the animals it covers. The main chapters cover ‘lesser known’ marine reptile groups (including the Triassic hupehsuchians, thalattosaurs, placodonts, and nothosaurs, the Jurassic pleurosaurs, and the marine snakes of the Cretaceous), then the geologically longer-lived, physically more imposing ichthyosaurs, the diverse and often long-necked plesiosaurs, those phenomenal giant lizards the mosasaurs and, finally, the sea turtles.

**Caption:** if you want to know about a relatively obscure group like the thalattosaurs, where do you go? That’s a problem, since such animals are generally not covered in the available non-specialist literature. *Ancient Sea Reptiles* discusses this group and so many others. These illustrations show (at left, top to bottom) the skulls of an unnamed Chinese taxon, *Nectosaurus* and *Hescheleria* and (at right) *Hescheleria* as it might have looked in life. Images: Tosha Hollman, from Naish (2023).

Introductory chapters cover Mesozoic marine reptiles more generally. Chapter 1 reviews our understanding of Mesozoic marine palaeogeography and climate as well as the scientific discovery of Mesozoic marine reptiles. Chapter 2 covers extinction events and broad evolutionary relationships. And chapter 3 covers anatomy, functional morphology and biomechanics, the focus being on swimming, feeding biology, the plesiosaur neck, and skin anatomy and life appearance.

**Caption:** our view of what Mesozoic marine reptiles were like in life have changed substantially the more we’ve learnt. The Crystal Palace models from the 1850s represent interpretations that - in cases - persisted until surprisingly recently. Clockwise from upper left: the thalattosuchian *Steneosaurus*, the mosasaur *Mosasaurus*, the ichthyosaur *Ichthyosaurus*, the plesiosaur *Plesiosaurus*. Images: Darren Naish.

An inevitable personal perspective. I began my academic career as someone hoping to go into Mesozoic marine reptile studies. Ultimately, it wasn’t to be (I got side-tracked into dinosaurs), but I very much remember the 1990s-era realisation that things were changing fast. The stereotypical view mostly conveyed beforehand – at least, so far as I understood it from books – was that these animals were static in evolutionary terms, that they weren’t remarkable with respect to biomechanics, and that they’d spent their history in stable, constantly warm conditions.

Clear signs of change came from a 1993 article by Robert Bakker in which he argued for a complex, dynamic view of Mesozoic marine reptile evolution that turned the then ‘traditional’ view of plesiosaur phylogeny on its head (Bakker 1993). Meanwhile, a series of studies published by Arthur Cruickshank, Mike A. Taylor and their colleagues in the UK showed that plesiosaurian skulls were sophisticated and specialised with respect to biting, stress dissipation and perhaps sensory abilities too (e.g., Cruickshank et al. 1991, Cruickshank 1994, Storrs & Taylor 1996, Taylor 1992).

**Caption:** a montage depicting various of the publications and proposals that made it clear (speaking from a personal perspective) that things in the world of Mesozoic marine reptiles were becoming more interesting during the late 1980s and early-and-mid 1990s. Clockwise from upper left: one of Mike Taylor’s diagrams of plesiosaurian jaw musculature, from his 1992 paper on *Rhomaleosaurus*; one of Riess & Frey‘s 1991 depictions of the alternating downstroke model of plesiosaur locomotion; my redrawing of the (still controversial) hydrodynamically driven underwater olfaction model of Cruickshank *et al*. (1991); and one of Taylor’s diagrams of ichthyosaur buoyancy and propulsion, from a 1987 paper in *Palaeontology*.

Renewed interest in the swimming biology of plesiosaurs and ichthyosaurs – something that kicked off in the mid-1970s (Robinson 1975) – was present by the early 1990s and had even snuck into popular venues like New Scientist. And new specimens and species from Antarctica, Niger, Nigeria, Angola, Mexico, Cuba, Chile and Argentina were making it clear that Mesozoic marine reptile research was very much a global event: the traditional Eurocentric view of marine reptile diversity and evolution (on which more in a moment) was very likely highly misleading.

A Mesozoic Marine Reptile Renaissance. While not fronted by a bombastic individual or a specific pop-sci piece – there’s no ‘Dinosaur Renaissance’ article in Scientific American for Mesozoic marine reptiles – it was obvious by the mid-1990s that a quiet revolution was happening, and that more and more people were being attracted to the study of plesiosaurs, ichthyosaurs and the others. This wasn’t due to a specific claim or contention made about the animals, but instead to the realisation that a substantial number of interesting questions about them had essentially never been asked. I see the publication of the 1997 book Ancient Marine Reptiles, edited by Jack Callaway and Elizabeth Nicholls, as proof that a Mesozoic Marine Reptile Renaissance was underway and obvious by the mid-1990s, even to non-specialists.

**Caption:** Callaway & Nicholls’s 1997 *Ancient Marine Reptiles*, the first substantial 20th century book devoted to Mesozoic marine reptiles since Samuel Williston’s 1902 *Water Reptiles of the Past and Present*. It’s multi-authored and includes 17 separate contributions. Like so many academic books, it’s prohibitively expensive should you wish to obtain it for yourself today.

In my writings on the history of dinosaur research (Naish & Barrett 2016, Naish 2021), I’ve been accused of talking about the ‘Dinosaur Renaissance’ too much (this being the sociocultural event, mostly led by Bakker and other vociferous scientists in the USA, in which it was argued that the biology, diversity and evolution of dinosaurs was substantially more interesting than the ‘conventional’ narrative). But, alas, I’m unable to ignore the fact that I’ve seen a major scientific and cultural paradigm change occur in real time. It’s such a big deal.

I feel the same way about Mesozoic marine reptiles. It might not be obvious to young researchers and enthusiasts today – they’re living in the Shiny Future of the 2020s where everything is different – but things really have changed a lot over the past several decades, and this is a story that mostly hasn’t been conveyed.

**Caption:** among the many interesting phylogenetic hypotheses pertinent to *Ancient Sea Reptiles* are (as shown at left) those positing how thalattosaurs, hupehsuchians, ichthyosaurs, saurosphargids and sauropterygians might all be close kin, and (at right) the alliance of cryptoclidids and xenopsarians within Plesiosauria. The book includes numerous cladograms that I designed and illustrated myself. Images: Darren Naish.

Ancient Sea Reptiles aims to make this point obvious. The evolution of Mesozoic marine reptiles wasn’t static and about long stretches of conservatism, but dynamic and complex, with major overturns and innovations happening right to the end. Plesiosaur phylogeny, it turns out, is hilariously complicated, the several fully pelagic mosasaur groups might have evolved their pelagic specialisations independently from less specialised ancestors, and pelagic sea turtles – similarly – might have evolved twice, from different ancestors.

Mesozoic marine reptiles didn’t live in a stable hothouse world, but a changeable one affected by major tectonic and volcanic events, and they had to deal with significant environmental changes and even ice ages.

**Caption:** the idea that Mesozoic climates were eternally equable and stable hasn’t been valid or defensible for decades now. This graph shows isotopic data recovered from Early and Middle Jurassic shelly fossils from Europe. The data reveal very high sea surface temperatures in parts of the Toarcian (that big red ‘low point’) but cool and even cold temperatures in the Aalenian in particular. Image: **Korte *et al*. (2015)**.

And far from being uninteresting from the point of view of anatomy and functional morphology, there are reasons for thinking that Mesozoic marine reptiles include some of the most extreme animals that have ever evolved, with incredible innovations in jaw and tooth morphology, neck anatomy, propulsion and more. For elaboration on that fairly nebulous point, I’ll have to direct you to the book!

On Eurocentricism and going beyond it. For all this talk of newness and paradigm shifts, one aspect of Mesozoic marine reptile research that makes the subject both eternally frustrating and fascinating is the historical 17th to 19th century angle that ties the topic to the geological locations of western Europe. Will we ever stop talking about the Dorset coast and Mary Anning, the Solnhofen Limestone, Monte San Giorgio in Switzerland, the German Posidonia Shale, Holzmaden and the Oxford Clay of the English midlands?

It seems not. The fossils of Mesozoic marine reptiles might, by now, have been found worldwide but it remains the case that western Europe remains important. A great paradox of the Mesozoic marine reptile fossil record is that a good number of groups remain almost unique to this region, this creating the impression that they scarcely occurred elsewhere. Were these animals really provincial (something that seems unlikely given their adaptation for pelagic life)? Or is it that our knowledge is still very much in its infancy?

**Caption:** if an animal like *Temnodontosaurus* is known only from western Europe and western Chile, what should we infer about its actual distribution when it was alive? Images: (c) Trustees of the Natural History Museum, London; (c) **Deep Time Maps**.

In the book, I present reasons for thinking that the latter is the case. My favourite example is provided by the long-bodied temnodontosaurian ichthyosaurs of the Early Jurassic. These large to stupendously large, wholly pelagic predatory ichthyosaurs are exclusive to western Europe… bar a single chunk of jaw from Chile. If we look at an Early Jurassic palaeomap, an occurrence in western Europe and western South America implies a near-global geographical range. Oh, for a better understanding.

Having mentioned Mary Anning: I do, of course, provide a dedicated section on the most famous fossilist of all time, a woman considered by many a patron saint of palaeontology. Anning’s impact on our understanding of Jurassic marine life was revolutionary. If only she had some way of knowing, when alive, how important she would be in future decades. A few aspects of the Anning story, however, have very often been mis-framed, among them the claim that she’s been “overlooked” or “forgotten”.

**Caption:** just how many palaeontologists get immortalised as action figures? Yes, I own a small plastic Mary Anning, and you should get your own too. It was made by **splendidhand toys** but it looks like they’re not selling that specific figure right now.

Art and imagery. As should be obvious from this blog, and from the look of my various books and other publications, I really care about pictures. I’m pleased to say that Ancient Sea Reptiles is extremely well illustrated, featuring numerous specimen photos, life reconstructions, skeletal diagrams, cladograms and so on throughout. It goes without saying that we used what photos we could of specimens that are on show at the Natural History Museum, or in its collections.

**Caption:** huge thanks to Brian Choo for allowing use of these excellent Triassic marine scenes, depicting life in the Guizhou (left) and Guanling faunas of the Middle and Late Triassic, respectively. *Tanystropheus*, nothosaurs, pachypleurosaurs and thalattosaurs are visible in the image at left; archaic ichthyosaurs, thalattosaurs, placodonts and turtles are visible in the one at right. Oh, also fishes. Images: Brian Choo, used with permission.

But we also went to some trouble to feature reconstructions produced by some very skilled artists, among them Davide Bonnadonna, Brian Choo, Julius Csotonyi, Tosha Hollmann, Joschua Knüppe, Julia Lacerda, Robert Nicholls, Júlia d’Oliveria, Scott Reid, Gabriel Ugueto and Esther van Hulsen. Huge thanks to everyone who submitted work to the book, and I hope you’re happy with the way in which it’s presented. The UK and US issues have different covers, but both are spectacular. The UK cover is by Haider Jaffri and the US one by Robert Nicholls.

While I could say a lot more, that’s where I’ll end. Already several publications have appeared that outdate a few of the contentions made in the book, but such is the nature of writing about a fast-moving, dynamic field of research. Hopefully they can be incorporated into the second edition.

*Ancient Sea Reptiles* is available from the Natural History Museum (at £20) if you’re in the UK, and from Smithsonian Books (at $29.95) if you’re in North America. And I’ll say again that finally seeing this book in print is a dream come true. I hope those of you that buy it enjoy it.

For previous articles on Mesozoic marine reptiles, see…

The skin of ichthyosaurs, September 2008
Sea Dragons of Avalon, an Arthurian adventure (part I), August 2009
Who made the giant Jurassic sea-floor gutters?, December 2009
Prediction confirmed: plesiosaurs were viviparous, August 2011
Rigid Swimmer and the Cretaceous Ichthyosaur Revolution (part I), January 2012
Plesiosaurs and the repeated invasion of freshwater habitats: late-surviving relicts or evolutionary novelties?, January 2013
Malawania from Iraq and the Cretaceous Ichthyosaur Revolution (part II), May 2013
Can’t get me enough of that sweet, sweet Temnodontosaurus, January 2014
Plesiosaur Peril — the lifestyles and behaviours of ancient marine reptiles, March 2014
Ancient Marine Reptiles Had Absurd, Complex Nostrils, July 2014
The Unique and Efficient 4-Flipper Locomotion of Plesiosaurs, August 2017
The Fall and Rise of Protoichthyosaurus, October 2017
Up Close and Personal With the Crystal Palace Dinosaurs, December 2018
The Ichthyosaurs of the Kimmeridge Clay, March 2021
History of the Iraqi Ichthyosaur, April 2021

Here's your regular reminder that this blog relies on support via patreon, thank you to those providing support already.

Refs - -

Bakker, R. T. 1993. Plesiosaur extinction cycles - events that mark the beginning, middle and end of the Cretaceous. In Caldwell, W. G. E. & Kauffman, E. G. (eds) Evolution of the Western Interior Basin. Geological Association of Canada, Special Paper 39, 641-664.

Cruickshank, A. R. I. 1994. Cranial anatomy of the Lower Jurassic pliosaur Rhomaleosaurus megacephalus (Stutchbury) (Reptilia: Plesiosauria). Philosophical Transactions of the Royal Society of London B 343, 247-260.

Cruickshank, A. R. I., Small, P. G. & Taylor, M. A. 1991. Dorsal nostrils and hydrodynamically driven underwater olfaction in plesiosaurs. Nature 352, 62-64.

Korte, C., Hesselbo, S. P., Ullmann, C. V., Dietl, G., Ruhl, M., Schweigert, G. & Thibault, N. 2015. Jurassic climate mode governed by ocean gateway. Nature Communications 6 (10015).

Naish, D. 2021. Dinopedia: A Brief Compendium of Dinosaur Lore. Princeton University Press, Princeton NJ.

Naish, D. 2023. Ancient Sea Reptiles. Natural History Museum, London.

Naish, D. & Barrett, P. M. 2016. Dinosaurs: How They Lived and Evolved. The Natural History Museum, London.

Robinson, J. A. 1975. The locomotion of plesiosaurs. Neues Jahrbuch fur Geologie und Paläontologie, Abhandlungen 149, 286-332.

Storrs, G. W. & Taylor, M. A. 1996. Cranial anatomy of a new plesiosaur genus from the lowermost Lias (Rhaetian/Hettangian) of Street, Somerset, England. Journal of Vertebrate Paleontology 16, 403-420.

Taylor, M. A. 1992. Functional anatomy of the head of the large aquatic predator Rhomaleosaurus zetlandicus (Plesiosauria, Reptilia) from the Toarcian (Lower Jurassic) of Yorkshire, England. Philosophical Transactions of the Royal Society of London B 335, 247-280.

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A group of passerine birds often framed as a bit boring and samey are no such thing…

**Caption:** the Eurasian skylark *Alauda arvensis*, the ‘classic lark’ for those of us in Europe. Image: Frebeck, CC BY-SA 3.0 (**original here**).

Among the many passerine groups of the world are the larks, known technically as alaudids. Larks are hugely widespread, occurring on all continents excepting South America and the polar regions, and all are strongly associated with deserts, semi-deserts, steppes and savannahs. A few species – most famously the Eurasian Skylark Alauda arvensis – also exploit farmland, since monotone fields obviously have a strong similarity with the steppe-like environments these birds prefer.

The caveat on lark distribution is that it’s a little misleading to describe them as denizens of (nearly) all continents given that only a single species (the Horned or Shore lark Eremophila alpestris) occurs in North America, while another singleton (Horsfield’s bush lark or Australasian lark Mirafra javanica) is the sole representative of the group in Australia. The caveat to that caveat is that the Horned or Shore lark is likely a species complex, and the same might be true of the variable Horsfield’s bush lark as well. Another caveat on distribution is that the group is predominantly African, 80% or more of the c 100 extant species occurring on the continent.

**Caption:** a lark montage, showing members of groups that might be considered ‘typical’ within the family. Left to right: Horsfield’s bush lark or Australasian lark *Mirafra javanica*, Temminck’s lark *Eremophila bilopha*, Rufous-tailed Lark *Ammomanes phoenicurus*. Images: JJ Harison, CC BY-SA 3.0 (**original here**); Francesco Veronesi, CC BY-SA 2.0 (**original here**); J. M. Garg, CC BY-SA 4.0 (**original here**).

Having just written about ‘bush larks’ I have to note here the inconsistency that exists in the names of these birds. Certain species have common names that are often or usually concatenated. Examples include woodlark, skylark and bushlark. The same names, however, are sometimes written as two words: wood lark, bush lark and so on. Bird vernacular names are more standardised than the names of virtually all other animal groups (largely because bird experts are a very…. thorough lot). Even so, there’s frustrating inconsistency here.

Lark basics. Larks are generally brown, pale ventrally, highly cryptic and good at concealing themselves when on the ground. Head crests have evolved several times and take the form of elongate, erectile feathers on the top of the head (as in the Skylark and Galerida crested larks) or paired, horn-like structures on either side of the crown (as in the Eremophila horned larks, where the crests form part of a boldly patterned series of facial markings). Larks are generally long-legged and good at running and walking. A long hallux claw – an often prominent feature of ground-adapted passerines – is present in many, and the other claws are often long and fairly straight as well.

**Caption:** among my favourite larks is the Greater hoopoe-lark *Alaemon alaudipes*, and I’m lucky enough to have seen it in life. That explains the photo shown here on the left: it was taken by my friend Richard Hing while we were on a palaeontological expedition in Morocco (a region great for larks). At right, a sharper image of the same species (albeit a different subspecies), this time photographed in Gujarat, India. Images: Richard Hing; Sumeet Moghe, CC BY-SA 3.0 (**original here**).

  **Caption:** I distinctly remember the few occasions in which I saw hoopoe-larks in the Sahara, and this photo is included because it gives some idea of the habitat in which our sightings mostly occurred. The birds were in flat, mostly open areas with scant nearby scrubby vegetation. This one was actually photographed about 10 m away from the tents of a group of nomadic people. Image: Richard Hing.

You might assume that larks are dull, samey, brown birds. But that’s not true or fair since the group include a list of really interesting, exotic species that rank among the most notable and unusual of passerines. My favourites include the large, leggy, tall, long-billed Greater hoopoe-lark Alaemon alaudipes of northern Africa, the Middle East and western Asia, and the fabulously thick-billed, cursorial, err, Thick-billed lark Ramphocoris clotbey of north-west Africa and the Middle East. A novice to passerine classification and diversity would be surprised that these finch-like and vaguely hoopoe-like birds are included in the same family as ‘classic’ larks like skylarks.

**Caption:** an artistic rendition of a Thick-billed lark *Ramphocoris clotbey* that doesn’t really do a good job of showing it in a natural posture. This image is from H. E. Dresser’s 1871 *A History of the Birds of Europe, Volume 4*. I really like Dresser’s books and just looked into the possibility of buying them. But I just learned that you can’t really buy them for less than £850. Guess I’ll stick with the pdfs then.

Here in the UK, larks like skylarks are famously associated with ‘the countryside’ and their melodious and energetic song-flights are characteristic of healthy farmland, meadows and heaths. It’s easy to hear them in such places but finding them in the sky can be hard. Unsurprisingly, skylarks are referenced in various old poems and rhymes, and they were formerly kept frequently as cage birds for their song (Greenoak 1997).

**Caption:** Alan Harris’s skylark painting from the cover of Paul Donald’s 2004 book *The Skylark*. Image: (c) Alan Harris.

Placing larks in the oscine tree. It’s long been obvious that larks are oscines: part of the huge passerine clade that includes crows and their allies as well as Passerida, the warbler-sparrow-flycatcher group. The traditional view most familiar to me during my student years is that larks are anatomically nondescript, even ‘primitive’, relative to oscines as a whole, and potentially something like ‘living fossils’ within the group. This view is owed to the fact that larks look anatomically archaic relative to most other oscines: there’s only a single pneumatic opening in the humerus (as opposed to two or more), the tarsus is latiplantar (meaning rounded on its posterior surface, rather than acutiplantar) and with scutes on both its posterior and anterior surfaces, and the syrinx lacks – or possesses in only rudimentary form – the midline structure termed the pessulus otherwise common in oscines (Sibley & Ahlquist 1990).

It's for these reasons that larks have ‘conventionally’ been placed or listed outside the clade that contains all warbler-like, sparrow-like and flycatcher-like oscines. Alexander Wetmore promoted this view in his highly influential classification of birds published in 1960, and it explains why larks – sometimes together with hirundines (swallows and martins) and also motacillids (pipits and wagtails) – are virtually always placed first among oscines in books that arrange passerines by family. In those books that include the Australasian lyrebirds and scrubbirds, larks are listed after those two ‘more archaic’ families.

**Caption:** a representation of oscine phylogeny as it was generally imagined throughout most of the 20th century. Lyrebirds and scrubbirds were posited as the earliest of the oscine lineages to diverge (a position they still occupy today), while larks, swallows and wagtails were all considered close, close to the ancestry of all ‘more advanced’ passerines, and also as ‘early-diverging’ in the tree. Several bird books from the 1960s, 70s and 80s depict this sort of thing and it’s also reflected in the way families are listed in fieldguides. The images you see here were produced for my in-prep textbook, the completion of which can be supported **via my patreon**.

A number of contradictions always implied that this ‘archaic position’ was wrong. The finch-like bills of certain larks mean that a few authors once considered them allied to weavers or similar birds, and a reduced 10th primary led some to regard larks as close to the ‘nine-primaried oscine’ group, some members of which have finch-like bills. It has to be said that these proposals were themselves contradicted by a list of other anatomical features, however.

Today, molecular results show that larks are deeply embedded within oscines – not located at or close to its root – and part of the ‘flycatcher-like’ clade Sylvioidea or Sylviida (Cracraft 2014, Fregin et al. 2012, Selvatii et al. 2015). Within that clade, they’re close to, and surrounded by, groups like babblers, acrocephalid and locustellid warblers and – surprisingly – are close to reedlings (Panuridae) (Cracraft 2014, Fregin et al. 2012, Selvatii et al. 2015).

**Caption:** increasingly complex versions of my passerine phylogeny image have appeared at Tetrapod Zoology over the years, and here’s the latest iteration. The topology here is based on that of several recent studies (e.g., Selvatii *et al*. 2015) and the taxonomy partly follows that of Cracraft (2014). The images you see here were produced for my in-prep textbook, the completion of which can be supported **via my patreon**.

Abundant surprises in lark phylogeny. Moving now to phylogenetics within the lark family, it turns out that things are more complex, and more interesting, than conventionally thought. Few studies of lark phylogeny exist. Sibley & Ahlquist (1990) included data from a few taxa and found Ammomanes (desert larks), Certhilauda (long-billed larks) and Mirafra (bush larks) to be part of early-diverging lineages outside a clade that includes Eremophila (horned larks), Galerida (crested larks) and Alauda (skylarks). A far more comprehensive job of analysing larks was published by Alström et al. (2013); they recovered a phylogeny similar in approximate structure to that of Sibley & Ahlquist (1990) but reported the following novelties.

**Caption:** simplified lark phylogeny, based on that presented by Sibley & Ahlquist (1990). Images (in branching order): J. M. Garg, CC BY-SA 4.0 (**original here**); JJ Harison, CC BY-SA 3.0 (**original here**); Francesco Veronesi, CC BY-SA 2.0 (**original here**); Bob Loveridge, used with permission.

Firstly, several genera proved non-monophyletic, Ammomanes and Mirafra among them. Clades considered good honest genera are located in ‘between’ the newly separated segments of the respective former genera, meaning that new (or resurrected) genus-level names are needed. For Ammomanes, this has meant resurrection of Ammomanopsis (originally coined in 1905) for Gray’s lark A. grayi from south-west Africa; for Mirafra, it’s meant that some species are best considered to belong to other genera, like Eremopterix (Alström et al. 2013). Here, it’s worth making the point that many (dare I say… most?) polytypic passerine genera have proved non-monophyletic when sufficiently well sampled, which isn’t a surprise given that most passerine genera have spent the bulk of their lives in the literature as ill-defined dumping grounds for species that look only approximately alike.

**Caption:** Gray’s lark might look like it should be included in *Ammomanes*, but genetics shows that it’s not. It’s not close to the *Ammomanes* species at all, but is instead part of the clade that otherwise includes the slender-billed *Chersomanes* and *Certhilauda* larks. Image: J. G. Keulemans, in public domain.

Secondly, certain taxa previously considered close allies based on morphology proved to be distant relatives. A good example is provided by the African Spizocorys larks and the mostly Asian Calandrella short-toed larks, two genera that are incredibly similar and have mostly been considered congeneric over the past several decades. But Spizocorys isn’t close to Calandrella at all: the former is part of the same clade as crested larks and skylarks, while short-toed larks are close kin of horned larks and the Bimaculated lark Melanocorypha bimaculata and kin (Alström et al. 2013). This discovery is pretty disturbing if you think that the external look of an animal is a good guide to its affinities.

What this means is that highly similar species classified within these disparate genera occur as geographically distinct ‘species pairs’. Alström et al. (2013) pointed to Dunn’s lark Eremalauda dunni (of northern Africa and western Asia) and Stark’s lark S. starki (of south-west Africa) as one such convergently similar, disjunct yet distantly related pair, and a similar situation exists for Dupont’s lark Chersophilus duponti of the Iberian Peninsula and northern Africa and the Certhilauda larks of southern Africa. We’re familiar with the idea of surprising convergence explaining the similar appearance of well-separated distant cousins – marsupial possums and squirrely rodents, say – yet here’s surprising convergence happening in the same family-level group.

**Caption:** *Eremalauda* and *Spizocorys* species look sufficiently alike that they’ve mostly been considered congeneric. But molecular data indicates that they’re not especially closely related. Images: Opisska, OA (**original here**); Alastair Rae, CC BY-SA 2.0 (**original here**).

Thirdly (and finally), taxa that do seem to be close relatives are very different in anatomy. Spizocorys might not be close to Calandrella, but it is close to the Woodlark Lullula arborea, which is a surprise given the slim, straight form of the Woodlark bill relative to the far stouter, deeper, Calandrella-like bill of Spizocorys. Meanwhile, Ramphocoris – the uberthickbill we looked at earlier – is close to the skylark-like Pinarocorys and Ammomanes desert larks, which is again not what you’d expect. I’d previously imagined an affinity with the Melanocorypha larks given a few similarities, and indeed Ramphocoris was originally included in Melanocorypha when first scientifically named in 1850.

**Caption:** one of the several lark clades identified by Alström *et al*. (2013). Note that *Ramphocoris* is close to desert larks and to superficially skylark-like taxa. Meanwhile, hoopoe-larks and other slender-billed taxa form a sub-clade within the same clade... as is *Ammomanopsis*. Image: Alström *et al*. (2013).

Rather than being predictable and conforming to guesses based on gross morphology, it turns out that lark phylogeny is complex, with unexpected groupings and convergences aplenty. In fact, the study that’s most responsible for establishing this – Alström et al. (2013) – has made larks into potential poster-kids for ‘cryptic complexity’ (if I may), and one of the best examples of this sort of thing in bird history. Which is a fine turn-up for the books in a group so often considered samey and conservative.

And that’s not all. More on larks coming soon.

For previous Tet Zoo articles on passerines, see…

Great tits: still murderous, rapacious, flesh-rending predators!, February 2013
For The Love of Crows, October 2015
Thoughts on the Passerine Tree, 2016, October 2016
A Battle Among Blue Tits, February 2018
Cocks-of-the-Rock, Extreme Cotingas, April 2019
Birdwatching in Suburban China, May 2019
Tell Me Something Interesting About Dunnocks, July 2019
Suburban Birdwatching in Queensland, Australia, November 2019
Kirk W. Johnson’s 2018 The Feather Thief, a Review, February 2021
The Locustella Warblers, May 2021
Chiffchaffs and What Are 'Old World Warblers' Anyway?, February 2022
Surprising Diet of the House Sparrow, August 2022

Support this blog AND the newly revived podcast at my patreon, where you also get to see tons of behind-the-scenes material that hasn’t yet been released anywhere else.

Refs - -

Cracraft, J. 2014. Avian higher-level relationships and classification: Passeriformes. In Dickinson, E.C. & Christidis, L. (eds) The Howard and Moore Complete Checklist of the Birds of the World (fourth edition), Volume 2: Passerines. Aves Press, Eastbourne, pp. xvii-xlv.

Fregin, S., Haase, M., Olsson, U. & Alström, P. 2012. New insights into family relationships within the avian superfamily Sylvioidea (Passeriformes) based on seven molecular markers. BMC Evolutionary Biology 12, 157.

Greenoak, F. 1997. British Birds: their Folklore, Names and Literature. Christopher Helm/A & C Black, London.

Selvatti, A. P., Gonzaga, L. P. & Russo, C. A. M. 2015. A Paleogene origin for crown passerines and the diversification of the Oscines in the New World. Molecular Phylogenetics and Evolution 88, 1-15.

Sibley & Ahlquist 1990. Phylogeny and Classification of Birds: a Study in Molecular Evolution. Yale University Press, New Haven and London.

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The rumours are true! At last, we have data on baryonychine brains…

**Caption:** the Isle of Wight baryonychine spinosaurid *Ceratosuchops* interacts with another animal, with the position and anatomy of its brain (and associated structures) shown in the inset. Image: Anthony Hutchings.

Long, long ago – in 2021 – myself and a team of colleagues (mostly associated with the University of Southampton) published our initial descriptive volley on the new British spinosaurid dinosaurs Ceratosuchops inferodios and Riparovenator milnerae (Barker et al. 2021). The study was led by Chris Barker, and analysis of the two animals forms the bulk of his PhD work.

Ceratosuchops and Riparovenator are both from the Lower Cretaceous Wessex Formation (part of the famous, and famously complex, Wealden Supergroup) of the Isle of Wight, and both lived during the Barremian age of the Early Cretaceous, round about 127 million years ago. Neither is complete nor even close to it, but both include well-preserved sections of skeletal anatomy. The braincases of both are three-dimensional and well preserved, as is also the case for the related Baryonyx walkeri from the Upper Weald Clay Formation of the English mainland (Charig & Milner 1986, 1997). All three are united within the spinosaurid group Baryonychinae.

**Caption:** it should be obvious how well-preserved the remains of *Ceratosuchops* and *Riparovenator* are. These photos both show posterior surfaces of the braincases: at left is the original of *Ceratosuchops* (scale bar = 50 mm); at right, a 3D print of *Riparovenator*. Images: **Barker *et al*. (2021)**; Darren Naish.

I’ve mentioned a few times that additional work on the Isle of Wight baryonychines is due to appear this year and beyond, and today sees another of our publications on these dinosaurs: ‘Modified skulls but conservative brains? The palaeoneurology and endocranial anatomy of baryonychine dinosaurs (Theropoda: Spinosauridae)’; it’s published in Journal of Anatomy (Barker et al. 2023). Chris and his PhD supervisor Neil Gostling led this research, and the contributors in addition to myself are Jacob Trend, Lysanne Veerle Michels, Larry Witmer, Ryan Ridgley, Katy Rankin, Claire Clarkin and Philipp Schneider.

What work did we do, and what are the conclusions?

Let’s CT-scan some baryonychines. Anyone paying attention to the scientific study of extinct dinosaurs will be aware of the revolution that computed tomography (or CT) has brought to the field. Thanks to advances made in CT-scanning technology and in the availability of CT-scanners themselves, scientists interested in the biology, behaviour, function and evolution of extinct animals now regularly place fossils in CT-scanners and analyse the results (Witmer et al. 2008). We’re especially blessed at the University of Southampton since we have the giant walk-in machine at the μ-VIS X-Ray Imaging Centre.

**Caption:** working with giant CT-scanners is not for for the feint of heart. This photos isn’t from the baryonychine project, but from another dating to 2012. Image: Darren Naish.

A caveat worth pointing out is that fossils from certain stratigraphic units tend not to scan well: the composition and density of the sediment that often surrounds and infills the fossils ruins the scanner’s ability to pick out fossil bone. This concern is especially relevant to Wealden fossils, since some of the respective sediments are full of minerals that disrupt easy scanning efforts. Exactly this affects one of the two Isle of Wight spinosaurids: the Riparovenator specimen is chock full of radio-opaque material that essentially make it impenetrable for this sort of work.

But I’m pleased to report that this is not the case for all Wealden spinosaurid specimens. We scanned the Ceratosuchops holotype in Southampton and the Baryonyx holotype at the Ohio Health O'Bleness Hospital in Athens, Ohio, and the key novelty of our study is that we provide the first CT-scan data yet reported for baryonychines (Barker et al. 2023). Both yield good results, and we reasoned that the good data on brain anatomy that they revealed would prove informative with respect to baryonychine biology and lifestyle. As discussed below, these aren’t the first spinosaurids to be CT-scanned, but they are the first baryonychines.

**Caption:** image showing the brain of *Baryonyx* in-situ inside the braincase. The braincases of baryonychines are narrow and deep, with basipterygoid processes that descend way below the rest of the structure. The section of the brain we’re seeing here (it doesn’t include the olfactory bulbs and tract) is about 12.8 cm long. Image: WitmerLab and Chris Barker.

Baryonychines vs other theropods. What do the scans mean with respect to how baryonychine brains compare to those of other theropods? We do at least have brain data for other, related theropods, including one other spinosaurid, as well as members of the more distantly related Allosauroidea. Here I should remind you that megalosaurids and spinosaurids are close kin within the theropod clade Tetanurae, both forming the clade Megalosauroidea. Views differ as to whether allosauroids are more closely related to birds than megalosauroids are, or whether there’s a megalosauroid + allosauroid clade.

**Caption:** the first of two competing theropod cladograms, highly simplified. This tree shows the more ‘traditional’ topology (at least, in these post-Gauthier 1986 times) in which allosauroids are closer to coelurosaurs than to megalosauroids (e.g., Holtz *et al*. 2004, Carrano *et al*. 2012). Image: Darren Naish.

  **Caption:** a tree found in some studies (e.g., Rauhut 2003, Naish & Cau 2022), where megalosauroids and allosauroids form a clade. Image: Darren Naish.

In overall structure and proportion, baryonychine brains conform to what we’d expect for theropods of this sort. The forebrain and hindbrain are in line with one another in the horizontal plane, there’s an elongate, narrow olfactory tract that projects forwards to the olfactory bulbs, the optic lobes are difficult to differentiate from the rest of the forebrain, the floccular lobes are present as projecting tabs surrounded by the semicircular canals of the inner ear, and the cranial nerves are in the expected position. The braincases of both animals are pneumatic, but not to the degree present in coelurosaurs (Barker et al. 2023).

The brains of Ceratosuchops and Baryonyx differ in several respects. Ceratosuchops has a midline groove along its upper surface, the two differ in terms of the position and extent of the peaked section at the top of the cerebrum (it’s less prominent and located a bit further back in Ceratosuchops), the Ceratosuchops forebrain is inclined downwards (whereas that of Baryonyx is more horizontal), the Baryonyx inner ear is proportionally larger than that of Ceratosuchops, and both differ in the shape of the lateral semicircular canal (Barker et al. 2023).

**Caption:** a baryonychine brain CT-scan composite, featuring a brain scan of *Baryonyx walkeri* at left, and *Ceratosuchops inferodios* at right. Image: **Barker *et al*. (2023)**.

Why so conservative? Overall, these brains aren’t tremendously different from those of other large tetanuran theropods. If anything, baryonychine brains appear anatomically conservative, certainly so relative to the bony anatomy of their skulls. That’s a surprising result, since the modification and specialisation of the spinosaurid skull – the rostrum in particular – might lead you to predict a modified brain too.

There are several ways to interpret these results. One possibility is that brain evolution was ‘lagging behind’ the evolution of the bony parts of the skull, either because brains and the nervous system are slow to evolve or conservative overall, or because the ancestral morphology was ‘good enough’ and that further modification wasn’t needed. Another possibility is that the megalosaurid-like theropods ancestral to spinosaurids already possessed the sensory and somatosensory features required for a successful baryonychine lifestyle. In other words, megalosauroids were pre-adapted for aquatic predation and an amphibious lifestyle, and ‘all’ they needed to do to become adept at it was modify the snout, jaws and teeth (Barker et al. 2023).

Other possible explanations exist though. A third possibility is that changes had happened, but concern details of brain structure that aren’t knowable via CT-scan data. And a fourth possibility is that baryonychines just didn’t need specialised brains relative to other theropods because all this speculation about them being heron-like waders is overblown. Maybe they were ecological generalists, predating on all manner of aquatic and terrestrial prey, and not much different in foraging behaviour from megalosaurids and the like. That proposal does a poor job of explaining why their skulls are otherwise so modified, however, so I have a hard time buying it.

**Caption:** a reconstructed, mounted skeleton of the spinosaurine *Irritator*, as previously displayed at the National Museum of Rio de Janeiro, and posed with an anhangeurid pterosaur as a prey item. This photo was taken in 2013. Image: Darren Naish.

I mentioned earlier than we have brain data from one other spinosaurid. In 2020, Marco Schade and colleagues reported CT-scan data for the brain of Irritator, a South American spinosaurine spinosaurid (Schade et al. 2020). One aspect in which it differs from the baryonychines (and other related theropods too) is that the floccular lobe is larger. Because the floccular lobe is relevant to gaze stabilisation via coordination of head, eye and neck movement, the large lobes of Irritator could be a specialisation for aquatic predation (Schade et al. 2020). Conversely, the more ‘typical’ floccular lobe of baryonychines could mean that they were less specialised, or non-specialised, for aquatic predation (though, another caveat: there’s some controversy as to how reliable a guide floccular lobe is for ecology and behaviour).

**Caption:** the hindbrain region of (a) *Baryonyx* and (b) *Ceratosuchops*, showing the arrangement of cranial nerves (in yellow), the semicircular canals (in pink), and the associated segment of cerebrum (in blue). The right side’s floccular lobe – labelled fl – is tab-shaped and projects into the semicircular canals. Image: **Barker *et al*. (2023)**.

Some specifics, and some implications. As should be obvious from the paper, we did just about everything we could from the data we gleaned. We estimated encephalisation quotient (EQ)*, hearing range, olfactory acuity and possible head pose.

Because it’s been shown that vertebrates as a whole don’t fit on the same brain : body size coefficient, experts today use different ones for the different vertebrate group. That for non-bird reptiles is termed REQ (Hurlburt et al. 2013).

By comparing estimated brain size to body size, we can work out the REQ of our sampled baryonychines, and it’s between 1.2 and 1.6 (Barker et al. 2023). That’s in the same approximate range as that of theropods like Carnotaurus and Allosaurus and could mean that these animals were similar in cognitive abilities and behavioural complexity (and here I’ll avoid any further discussion of how reliable a guide REQ is to ‘intelligence’, for reasons).

**Caption:** these days, there’s often a fairly rapid turnover time between the publication of a new dinosaur taxon and the appearance of good, commercially available figures of said taxon. New for 2023 is this *Ceratosuchops* figure from CollectA which I’m very pleased to own in my collection. Images: Darren Naish.

Moving on… based on cochlear duct size relative to basicranial length, it looks like Ceratosuchops and Baryonyx had mean hearing frequencies in the 1400-1600 Hz range (Barker et al. 2023). That’s very approximately in the crocodylian range but also – for those who might think that this is significant – in about the same range as a great many birds. On olfaction, there’s been some suggestion that spinosaurids had reduced olfactory bulbs and thus a reduced sense of smell, but this doesn’t seem to be the case in baryonychines at all. Their olfactory bulbs are ‘normal’ for theropods of their size, and their sense of smell was likely pretty good, and again about similar to that of abelisaurids and Allosaurus (Barker et al. 2023). Finally, both the nature of the condyle at the back of the skull and orientation of the semicircular canals imply that baryonychines were ‘standard’ as theropods go (Barker et al. 2023). There’s no indication that they maintained the skull at a dipped-down angle or anything like that, as has been inferred for Irritator (Schade et al. 2020).

All in all, our data “indicates that baryonychines did not deviate substantially from the cognitive, auditory or olfactory capabilities of ‘typical’ non-coelurosaurian theropods” (Barker et al. 2023, p. 17). Interesting, and weird.

**Caption:** life reconstruction of *Ceratosuchops* (at left) and *Riparovenator*, shown sharing a flooded depression on the Wessex Formation floodplain. Image: Anthony Hutchings.

And right now that sums up just about everything that’s worth saying at this point. Just successfully obtaining CT data on brain anatomy – palaeoneurological data – from baryonychine spinosaurids is a big deal, and using it to infer sensory abilities and so on is also a worthy endeavour. Finding that baryonychines appear surprisingly samey relative to non-spinosaurid theropods is a curious result, but as knowledge improves we’ll understand better just what this means. I want to thank Chris, Neil, Jacob and my other coauthors for their co-operation on such an interesting study, and I’m pleased that we’ve succeeded in contributing once again to our growing knowledge of Wealden spinosaurids.

On that note -- more Wealden spinosaurid news is coming!

Of Becklespinax and Valdoraptor, October 2007
The world’s most amazing sauropod, November 2007
Oh no, not another new Wealden theropod!, June 2009
Concavenator: an incredible allosauroid with a weird sail (or hump)... and proto-feathers?, September 2010
A truly tiny Cretaceous theropod... from England?, May 2011
The Wealden Bible: English Wealden Fossils, 2011, November 2011
Ostrich dinosaurs invade Europe! Or do they?, June 2014 (every archived version of this article lacks the original illustrations, sorry)
The Sensitive Face of a Big Predatory Dinosaur, June 2017
Theropod Dinosaurs of the English Wealden, Some Questions (Part 1), March 2020
Introducing ‘Unexpected Isle of Wight Air-Filled Hunter’, a New English Theropod Dinosaur, September 2020
Dr Angela Milner and the Discovery of Baryonyx, August 2021
Two New Spinosaurid Dinosaurs from the English Cretaceous, September 2021
A Giant Spinosaurid Dinosaur from the Cretaceous of the Isle of Wight, June 2022

Refs - -

Barker, C. T., Naish, D., Trend, J., Michels, L. V., Witmer, L., Ridgley, R., Rankin, K., Clarkin, C. E., Schneider, L. & Gostling, N. J. 2023. Modified skulls but conservative brains? The palaeoneurology and endocranial anatomy of baryonychine dinosaurs (Theropoda: Spinosauridae). Journal of Anatomy doi: 10.1111/joa.13837

Carrano, M. T., Benson, R. B. J. & Sampson, S. D. 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda), Journal of Systematic Palaeontology 10, 211-300.

Charig, A. J. & Milner, A. C. 1986. Baryonyx, a remarkable new theropod dinosaur. Nature 324, 359-361.

Charig, A. J. & Milner, A. C. 1997. Baryonyx walkeri, a fish-eating dinosaur from the Wealden of Surrey. Bulletin of the Natural History Museum 53, 11-70.

Holtz, T. R., Molnar, R. E. & Currie, P. J. 2004. Basal Tetanurae. In Weishampel, D. B., Dodson, P. & Osmólska, H. (eds) The Dinosauria, Second Edition. University of California Press (Berkeley), pp. 71-110.

Hurlburt, G. R., Ridgely, R. C. & Witmer, L. M. 2013. Relative size of brain and cerebrum in tyrannosaurid dinosaurs: an analysis using brain endocast quantitative relationships in extant alligators. In Parrish, J. M., Molnar, R. E., Currie, P. J. & Koppelhus, E. B. (eds.) Tyrannosaurid Paleobiology. Bloomington, Indiana University Press, pp. 134-154.

Naish, D. & Cau, A. 2022. The osteology and affinities of Eotyrannus lengi, a tyrannosauroid theropod from the Wealden Supergroup of southern England. PeerJ 10:e12727.

Rauhut, O. W. M. 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology 69, 1-213.

Schade, M., Rauhut, O. W. & Evers, S. W. 2020. Neuroanatomy of the spinosaurid Irritator challengeri (Dinosauria: Theropoda) indicates potential adaptations for piscivory. Scientific Reports 10, 1-9.

Witmer, L. M., Ridgely, R. C., Dufeau, D. L. & Semones, M. C. 2008. Using CT to peer into the past: 3D visualization of the brain and ear regions of birds, crocodiles, and nonavian dinosaurs. In Endo, H. & Frey, R. (eds) Anatomical Imaging. Berlin, Springer, pp. 67-87.

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I knew that 2022 would be a busy year, but… my god.

**Caption:** images from Portsmouth Comic Con 2022, and the coast of Oregon, respectively. Images: Darren Naish.

Tetrapod Zoology the blog is now 17 years old (the blog’s birthday, or blogoversary if you must, is on January 21st, and the blog started operation in 2006). Meaning that – once more – it’s time to look back at the year that’s passed from my highly biased, personal, TetZooniverse-focused perspective. 2022 was a year that involved a reasonable amount of travel, the publication of some significant studies with long backstories, attendance at various exciting events, and a respectable amount of animal-watching.

As ever, stop reading now if you’re not keen on the prospect of hearing me talking about my own adventures ad nauseum, since that’s all I’ll be doing in this article.

**Captions:** a fair number of new animal figures entered my collection in 2022. Those shown here include some of the *Prehistoric Planet* figures made by Marco Makes, some Beasts of the Mesozoic ceratopsians, and an assortment of CollectA prehistoric animals (released 2021). Images: Darren Naish.

Our story begins in late January with the celebration of Tetrapod Zoology’s 16th birthday, though I didn’t actually get to publish the article on said celebration until late February. Most of my plans for the first half of the year revolved around the release of the Apple TV+ series Prehistoric Planet. I haven’t written much about Prehistoric Planet here at Tet Zoo – I can’t – but if you go back to my review of 2022 knowing that Prehistoric Planet was in production at the time, various cryptic statements therein make sense.

**Caption:** I had quite a few chances during 2022 to wander around the Natural History Museum, London, before before and after hours. At left we see the marine reptile display; at right, Hope the Blue whale in Hintze Hall. Images: Darren Naish.

On that note, I stayed in London across late January and early February as we did the final bits of filming. This meant being in the Natural History Museum both before and after closing – what a privilege – as well as hanging out with David Attenborough; it’s not a coincidence that he now owns a copy (actually, several) of All Yesterdays.

TetZoo articles of January and February included those on my reminiscences of the putting together of the 2001 Dorling Kindersley *Encyclopedia of Dinosaurs and Prehistoric Life*, and on leaf warblers. I went to Marwell – my local zoo – to look at penguins, lemurs and okapis.

**Caption:** at left, a somewhat unusual photo of a Black-and-white ruffed lemur *Varecia variegata* at Marwell. At right, one of Marwell’s several Okapi *Okapia johnstoni*. Images: Darren Naish.

The Hum Hole pond renovation. I’ve mentioned here, on and off over the years, how I’ve assisted local conservation and habitat improvement efforts. One of these concerns the plan to renovate and restore the tiered pond system – part of so-called Hum Hole – close to where I live in Southampton. The ponds are no longer ponds but have become entirely silt-filled and are now entering their willow-dominated marsh phase. Succession of this sort is normal, but we want them to be maintained as ponds seeing as their loss means that various pond-using animals (most notably Palmate newts Lissotriton helveticus) no longer have useable breeding sites in the area.

**Caption:** habitat creation has led to a year-on-year increase in the number of Common frogs *Rana temporaria* that spawn in the ponds here during February. For 2021, I observed 13 adult frogs engaged in spawning. For 2022, there were 25 frogs. Image: Darren Naish.

Efforts to get habitat restoration projects off the ground are always slow-moving since they involve coordination between local authorities and volunteer groups, and events didn’t get underway late February. So far, we’ve cleared a lot of the willow and scrub that surrounds the ponds, the next phase being to excavate the substantial amount of silt and dump it where the scrub was. It’s a big project and will take years.

**Caption:** it might be hard to believe, but the image at left shows a pond. A team of volunteers and council workers began renovating the site in early 2022. Images: Darren Naish.

Loch Ness. I travelled to Loch Ness (again) in late February, again for a Loch Ness monster documentary, one of two I helped with during the year. At the time of writing (November 2022), neither have been screened but I look forward to the time when they are. I can say from extensive experience that TV documentaries often seem impressive and potentially impactful when you’re working on them, only to be unimpressive and wholly forgettable when broadcast. Right now, the projects seem noteworthy. One of them is a two or three-part series that looks more focused on the history of the search for the Loch Ness Monster than any previously broadcast series, and I look forward to seeing it out.

**Caption:** one of two trips to Loch Ness I made during 2022. We weren’t close to the loch during the filming of this one, but high up on a hillside. It’s important to remember that the Loch Ness area is bleak and mostly stripped of its original flora; it’s the very opposite of a pristine, attractive landscape. Images: Darren Naish.

For those unfamiliar with my writings on monsters: no, I don’t ‘believe’ in the Loch Ness Monster (nor indeed in any of the famous cryptids), but I am interested in the history of research on it, whether that research involves ‘believers’ or ‘critics’, on those who approach the monster as a sociocultural phenomenon, and on those who approach the monster as a real and undiscovered animal (Naish 2017, 2022).

**Caption:** argh, Loch Ness Monster souvenirs — my one weakness! And Highland cow figures, I guess. Image: Darren Naish.

My ‘spare’ time during this part of the year involved my (ultimately successful) writing of the text for Mesozoic Art (on which more later) and Ancient Sea Reptiles (more on that later too). It isn’t that I want there to be multiple projects running concurrently, it’s that there are only ever certain windows in which these things can be done.

**Caption:** at far left, Chessel Bay Nature Reserve during 2022. The discarded fridge is an especially nice touch. At right, litter-picking in Sholing Valley, Southampton. Images: Darren Naish.

I assisted in a local beach clean at the end of March and I was meant to be speaking (about dinosaurs, with a connection to my 2021 book Dinopedia) at the Oxford Literary Festival at about the same time. Illness meant that I had to cancel. I took multiple covid tests and all were negative, but my doctor said that it was likely covid anyway and I believe her. I also had to turn down involvement in a new dinosaur-themed TV series – ultimately broadcast on the UK’s Channel 5 late in 2022 as Dinosaur – since the filming schedule clashed with other events and I can’t do everything.

**Caption:** blurry but still acceptable photo of Crocodile monitor at Marwell Wildlife. On some visits, this animal can be seen resting or hiding on a ledge (itself part of the replica cliff-face in its enclosure). But on others, it swims, it walks, and basically moves around a lot. I like all monitors, but *V. salvadorii* is one of my favourites. Image: Darren Naish.

An April trip to Marwell resulted in observation of Banteng Bos javanicus, Crocodile monitor Varanus salvadorii and more, and also the latest of their Lego prehistoric animals. These were surprisingly novel in terms of the animals chosen, their number including such stem-mammals as Estemmenosuchus and gorgonopsians of some sort, the hadrosaur Olorotitan, and a Styracosaurus with a Beasts of the Mesozoic colour scheme.

**Caption:** well, there’s something you don’t see everyday. A Lego *Estemmenosuchus* at Marwell Wildlife in April 2022. I took photos of all the other constructions too. Images: Darren Naish.

I published a revised version of an old cassowary-themed article at TetZoo.

Tring. Also during April, I visited the Natural History Museum at Tring (formerly the Walter Rothschild Museum) to meet up with Todd Green – the cassowary kid himself, you know – to discuss cassowary-themed plans. Tring is an absolute mecca if you’re a zoology or museum nerd: it houses an important collection (of birds in particular), has a huge number of interesting and significant specimens on display, and is one of those museums that’s effectively a ‘museum of a museum’. They’ve deliberately kept things largely as they were during the mid-1900s.

**Caption:** Tring is well known for its bird collection, but it has a ton of other things too, including a few marine mammals on show. The model here depicts Sowerby’s beaked whale *Mesoplodon bidens* (its skeleton is in the background); the skeleton below is that of a Southern elephant seal *Mirounga leonina*. Images: Darren Naish.

I’ve had reason to mention Tring several times at Tetrapod Zoology but haven’t ever written about the museum itself; the closest I’ve come to this is when I reviewed Kirk Wallace Johnson’s 2018 book The Feather Thief, since that’s very much a Tring-based story. While in Tring, Todd and I walked the full distance from the train station to the museum – the same trip made on foot by Edwin Rist during his despicable heist of 2009 – and visited the (now partly fortified) side alley and stone wall next to the museum, both of which were used by Rist as well.

**Caption:** Naish and Green at Tring, with various of the cassowary specimens (and cassowary art) they have on show. The big bird at far right is a Northern or Single-wattled cassowary *Casuarius unappendiculatus*.

Todd and I met up later in the year at the Society of Vertebrate Paleontology meeting in Toronto and we always end up talking about cassowaries a lot. A lot is happening that’ll be of interest to those who want to know more about these amazing birds, stay tuned.

Prehistoric Planet Premiere Month. May was one of my busiest months. After several years of production – during which it was keep entirely secret – the new Apple TV+/BBC Studios series Prehistoric Planet was released to the world, and what a triumph. The audience reaction was overwhelmingly positive, and those journalists, media pundits, authors and TV personalities made aware of it were generally enthused and keen to do their bit in promoting and reacting. Together with series producer/showrunner Tim Walker and executive producer Mike Gunton, I did numerous interviews, both in the UK and in Los Angeles.

**Caption:** a *Prehistoric Planet* promotion event in Los Angeles, May 2022. From left to right: directors Andy Jones and Adam Valdez, executive producer Mike Gunton, lead scientific consultant Darren Naish, executive producer Jon Favreau, series producer Tim Walker, and musicians Anže Rozman, Kara Talve and Russell Emanuel. The original image is **here**.

In LA, the three of us – together with Jon Favreau and other people involved in the series – attended screenings at the Santa Monica IMAX, and then various press junkets, press conferences and red carpet events.

**Caption:** scenes from various of the Prehistoric Park premiere events. At upper left, part of the landscape built for the Los Angeles press event. At lower left, myself and Nigel Marven at the Bristol IMAX event. At right, an audience scene at the BFI IMAX event. Those on stage are (left to right) our host Stephen Armstrong, executor producer Mike Gunton, series producer Tim Walker, and myself.

We then came back to the UK and did the same thing in London (the screening this time being the BFI IMAX), then again at the Bristol IMAX, and then in London again. Episode 1 of Prehistoric Planet was released on Apple TV+ on May 23rd, and I went to one more promotional London event – this time a special meal at the Natural History Museum, held partly to celebrate the return to the NHM of Dippy the Diplodocus – at the end of the month. It was a total whirlwind and phenomenal fun.

**Caption:** a special evening event at the Natural History Museum. I don’t really hold *Mantellisaurus* – or any iguanodontian – in disdain, unlike many of my colleagues. Dippy as displayed in its new, *Prehistoric Planet* sponsored gallery is now illuminated by unusual greenish lighting. Images: Darren Naish.

Also at the end of May, Dave Hone and I met up for a trip to the Battersea Park Children’s Zoo, a small zoo I’ve never visited before. I’ll write up thoughts properly sometime (for my zoo review series), but personal highlights included Chinese crocodile lizard Shinisaurus crocodilurus and Chacoan mara Pediolagus salinicola.

**Caption:** Chinese crocodile lizard photographed at Battersea Park Children’s Zoo. Some of these animals have a lot of light green and white on them, while others are more reddish. Check out the big keeled scales on the neck and the substantial amount of sclera this individual is showing. Crocodile lizards (sometimes called crocodile-tailed lizards) are viviparous anguimorphs and part of a group that extends back to the Cretaceous. Image: Darren Naish.

We then went to St James’s Park for their impressive bird collection. The park is famous for its pelicans (noted for their occasional grabbing and eating of feral pigeons) but is home to numerous wildfowl, as you can see from the photo here.

**Caption:** a St James’s Park wildfowl montage, albeit arguably a boring one since all the species shown are highly familiar. The park is also home to numerous exotics, including several Eurasian geese, mergansers and diving and dabbling ducks. Image: Darren Naish.

The White Rock spinosaurid. June saw the release of the first of my technical papers for 2022, this being the PeerJ article on the giant Vectis Formation spinosaurid from the Isle of Wight (Barker et al. 2022). This project was led by Chris Baker and Neil Gostling of the University of Southampton and is the second of our group’s publications on Wealden spinosaurid dinosaurs.

The animal concerned – it’s from a part of the Vectis Formation called the White Rock and is thus informally known as the White Rock spinosaurid – isn’t known from sufficient diagnostic remains to warrant being formally named as a new species, but it was very obviously enormous, and potentially one of the largest theropods known. Predictably enough, journalists went wild for it and stories on this fossil appeared throughout the global press. A TetZoo article on the study is here. Other Wealden spinosaurid studies are in preparation and will appear during 2023.

**Caption:** the White Rock spinosaurid material isn’t as good as we might like it to be, but it’s certainly good enough to show that (1) it’s a spinosaurid, and (2) it was a huge animal, certainly over 10 m when complete. Our paper on this new (but as yet unnamed) species is open access **and here**.

Panthera Britannia. I appeared in the Dragonfly Films documentary Panthera Britannia, released in June and devoted to the British big cat phenomenon. Yes, I do believe that wild-living, non-native British felids are a thing, but that’s not why I get used in these TV shows. Instead, it's because I can provide both background information on the subject (pertaining to prehistory and archaeology) and some theoretical content on how non-native cats might survive in the UK. Incidentally, Panthera Britannia has just been released for free online.

I still haven’t published as much on the big cat phenomenon as I’d like to: there’s the Blake et al. (2014) paper on the lynx specimen accessioned at Bristol Museum and Art Gallery, but one day I hope to do a book on the subject.

Eotyrannus: theropod of 2022. Several relevant things happened in July. For starters, the Eotyrannus monograph (Naish & Cau 2022) – long in review at PeerJ – saw publication. As should be well known to readers of this blog, this work has been in preparation for years, and finally getting it into print was and is a significant event. My inability to finish it sooner is a consequence of my non-standard relationship with academic publishing. I don’t work in academia and only do research and writing ‘in spare time’, so it takes years to get anything done. Taking months or even years to respond to reviewer comments is generally disastrous, since editors mostly want reviews to be current and timely, and hence will send a submission out for review again if you don’t turn it around quickly. You thus become locked into a cycle: manuscript comes back from review, you take months to get it back to the editorial team, it goes out for review again, you take months to get it back to the editorial team, it goes out for review again. Some editors, it should be said, are very much sympathetic to this. Others are not.

**Caption:** the *Eotyrannus* monograph (**Naish & Cau 2022**) arrives at last. The skeletal reconstruction by Dan Folkes, shown at lower right, appears in the paper. The life reconstruction by Loana Riboli (copyright, used with permission), shown at upper right, was produced to assist in promotion.

Enough complaining. The point is that I got it done, and a TetZoo article on that event is here. I fully intended to put out a press release and all that but, in the end, I couldn’t pull it together in time. I also planned to throw a party – a physical event, in a hired venue and everything – since getting the study published means that much to me, but I couldn’t get this sorted in time, either. The monograph is not the end of work on Eotyrannus. There’s more to do and more papers to come. Eventually.

Also during July, Chris Barker and I visited various of the spinosaur-bearing Wessex Formation locations on the Isle of Wight. Did we find any new material? That would be telling…

**Caption:** *thar be baryonychine spinosaurs in them there rocks*… A scene from an Isle of Wight fossil-finding trip of July 2022. Image: Darren Naish.

Goodbye podcats. I realised on July 31st that Tetrapod Zoology ver 4 (the version you’re looking at now) had been going for four years. So I knocked up a quick article summarising the highlights of four years of blogging. Hey, it’s good to provide regular reviews and reminders and whatnot when you’re a blogger, or that’s what they tell you anyway. John Conway and I released episode 83 of the podcast in late July, which had sat in John’s files since late 2021. Obviously, we both struggle to make time to record the podcast, let alone edit and publish it (these days, Sharon Hill does the editing), and episode 83 is the most recent one we’ve recorded. I’ve been telling people that the podcast is dead. That’s not strictly true but I feel it may as well be. It’s just another thing I can’t fit into the schedule anymore.

**Caption:** part of the interior of Brighton Aquarium, opened to the public in March 1872. In case this image creates the wrong impression, the aquarium isn’t poorly designed or cramped and has a good selection of aquatic animals. Among the tetrapods there are Roti Island snake-necked turtles *Chelodina mccordi*, a species now kept in quite a few collections. Images: Darren Naish.

TetZoo articles of August included those on spider-frog relationships, Asian elephants, and legendary black dogs. I visited Brighton (and went to Brighton Aquarium, the world’s oldest extant aquarium), went glamping in Buckinghamshire, and paid attention to the newly metamorphosed frogs leaving the pond. My advance copy of Mesozoic Art arrived, and – wow – is it a very fine piece of work if I say so myself.

**Caption:** glamping and a CollectA *Lythronax* figure, what could be funner? A photo from August 2022. Image: Darren Naish.

My BBC/Prehistoric Planet credentials meant that I was able to attend an exclusive event in Soho hosted by Hans Zimmer and members of his Bleeding Fingers group. It was about the musical work of these people in general, but Prehistoric Planet got mentioned a couple of times. It was great to again meet musicians Anže Rozman and Kara Talve (we attended the same meetings in Los Angeles during May, but never had the chance to talk there).

**Caption:** in which I leverage the votes at the 2022 Fortean Film Festival.

September began with my attendance at the second Fortean Film Festival in Gloucester (England), held again at the Sherborne Cinema. This is becoming a new annual meeting for people involved both in paranormal research (whatever we mean by that term) and its portrayal in the media. While I have no real interest or involvement in ‘the paranormal’, I’m certainly interested in knowing what people think about monsters, cryptids and the study of them. We went on a guided ghost tour of Gloucester and I visited the Beatrix Potter Museum. I was interviewed for Fortean News Podcast (the relevant episode is here). This happened in a bar late at night, so I said a bunch of stuff that would have benefitted from greater planning and preparation. But isn’t that always the case.

**Caption:** Gloucester is good for secondhand books, I’ll say that. Very pleased to get hold of these while in the city for the Fortean Film Festival during September 2022. Image: Darren Naish.

Cryptozoology: in print, on a podcast, in a virtual discussion. I published an article in The Biologist in September. It’s called A Cultural Phenomenon (Naish 2022) and discusses my (by now very familiar) argument (espoused in my 2017 book Hunting Monsters and elsewhere) that many so-called cryptids are products of culture and lore, not new animal species awaiting discovery (Naish 2022: digital version here). Nevertheless, discussions of these alleged creatures are so entwined with writings on zoology and natural history that they are very much part of zoological history, real or not. A slightly controversial claim made in the article is that cryptozoology as currently conceived is creeping away from the zoology-based version of the subject favoured by Bernard Heuvelmans and his associates, and is instead becoming a vague, pseudoscientific topic where people think that bigfoot and Nessie should be discussed alongside demons, floating lights, UFOs and portals to other dimensions. There’s evidence that this is indeed the case within the popular sphere, but the pushback view is that ‘the popular sphere’ doesn’t represent the views and actions of authors and researchers.

**Caption:** I’m pretty pleased that my article in *The Biologist* made the cover. The article itself includes images of cryptids provided by Tyler Stone.

Another cryptozoology-themed thing happened soon afterwards in the month when I spoke with Blake Smith and Karen Stollznow for an episode of the Monstertalk podcast. It’s called Bernard’s Beasts and was inspired by my long-running (and still nowhere close to being finished) project on Bernard Heuvelmans’s definitive – and fairly hilarious, sorry Bernie – 1986 list of cryptids. As will be obvious if you’ve listened to it, I did no preparation whatsoever and there are things I should have known but didn’t. Anyway, the relevant episode can be listened to here.

**Caption:** cover image for the cryptozoology-themed Monster Talk podcast episode I recorded in September 2022.

And another cryptozoology event happened in September too: on the 16th, I took part in an online roundtable event on the subject, itself part of the Animals in the American Popular Imagination conference hosted by anthrozoologist Margo Demello of Carroll College in Montana. The other panel members were Richard Fallon, Samantha Hurn and Greg Morrow. Topics covered included the connection between colonialism and cryptozoology, British big cats, and the notion of cryptids as cultural icons. The discussion was recorded and there are plans to post it to YouTube (maybe this has already happened, I don’t know). There’s also mention of an edited volume resulting from the meeting; we will see.

**Caption:** a scene from SVPCA 2022; specifically, from Romain Pintore’s talk on theropod femoral evolution. I’m taking this as further evidence that *Prehistoric Planet* has well and truly embedded itself in popular consciousness. Image: Darren Naish.

SVPCA in London. In late September I was at the Natural History Museum in London again, this time for the 70th Symposium on Vertebrate Paleontology and Evolution (SVPCA). I did think about giving a talk (it would have been on the results of the Eotyrannus monograph), but my abstract failed to make it through review. The meeting was good fun and I succeeded in winning at least a few of the things I wanted from the auction, including the poster version of the fabled Brooke Bond Prehistoric Animals card set (art by Maurice Wilson, text by Alan Charig).

**Caption:** at left, in which I get to hang out with the legend that is Dr Tori Herridge. At right: finally, my own Brooke Bond Picture Cards *Prehistoric Animals* poster.

Tetrapod Zoology at the United Nations Science Summit. A significant event happened later in September, when I was among a group of scientists and natural historians who presented at the United Nations Science Summit. This event was organised by Seabird McKeon and Michele Weber, and involved talks by Ursula Valdez, Tom Fleischner, Dita Cahyani, Nalini Nadkarni and myself. The physical version of the summit happened in New York, so our original plan was to meet there. But, alas, we couldn’t get the funds together for this, and zoom it was.

**Caption:** my cover slide from the United Nations Science Summit of September 2022.

My talk was Prehistoric Animals are Ambassadors for Science, and Learning About Them Arms us With Respect to Sustainability, which is pretty self-explanatory. The other talks were variously about natural-history-based initiatives that have a positive impact on human lives (and foster an improved connection with the natural world) or were about conservation efforts that involve cooperation with local people. My main takehome was that human health and wellbeing are proving tightly connected to our access to nature and natural places, something I find myself thinking about more and more the older I get. We finished with a Q&A session that focused on our own experiences, our feelings for the future and more. I think that everything was recorded but I don’t know what the plans are on sharing it. Stay tuned.

**Caption:** a moment from our natural history event at the United Nations Science Summit, specifically from Nalini Nadkarni’s talk.

It was a huge privilege to be part of this event and I’m extremely grateful to Sea and the other team members for making it happen. There are plans for this to be the start of something permanent and embedded in the function of the United Nations, but it’s early days and discussion is ongoing. A brief TetZoo report on the event – I think published on the day that it happened – is here.

**Caption:** a September scene from Pickett Post in the New Forest. The temperatures were of the sort we used to expect for summer. Image: Darren Naish.

Lyme Regis and Mesozoic Art. In October, I visited Lyme Regis Museum to see the Mary Anning portrait by B. J. Donne (painted in 1850, and thus after Mary’s death in 1847) on its last weekend of display. This is the first time it’s been outside of London ever.

**Caption:** at left, the B. J. Donne portrait of Mary Anning, outside of London for the first time ever. Donne lived in Lyme Regis while Mary was alive and it’s considered likely that he knew her before her death in 1850. At right, an ichthyosaur display at Lyme Regis Museum. The big skull belongs to *Temnodontosaurus*. Images: Darren Naish.

This was also my chance to see the Mary Anning statue, unveiled earlier in 2022. We can’t know for sure that the statue represents much of a likeness to the real Mary Anning, but it’s a fascinating statue whatever you think, with belemnites, ichthyosaur and plesiosaur bones, and the actinopterygian fish Dapedium built into its base. Numerous ammonites decorate it here and there.

**Caption:** the Mary Anning status at Lyme Regis, unveiled in early 2022. The base is decorated with several fossils relevant to Mary Anning’s career, and to the palaeontology of the Lyme Regis area. Images: Darren Naish.

The museum itself has had something of a revamp in recent years, for the better. Many new marine reptile fossils are on display, making the museum more of a Mesozoic marine reptile mecca than it was before. I’m due to be speaking there in 2023 as part of the promotional tour for my Ancient Sea Reptiles book, on which more below.

Mesozoic Art and Ancient Sea Reptiles. October saw the publication of Mesozoic Art, the big, lavishly illustrated art-themed book compiled by Steve White and myself, and published by Bloomsbury (White & Naish 2022). I’ve already written about the book at TetZoo. It showcases the work of 20 palaeoartists, many of them relatively new to the field and doing more than their fair share of innovation. Palaeoart today – as seen from my unusual position in the north-east Atlantic maritime provinces – is becoming more of an international endeavour, with more diversity in style and texture. We hope that this is reflected in the book.

**Caption:** our new book *Mesozoic Art* isn’t technically part of the same series as its predecessors, but it’s sufficiently similar in design to look like it might be. Image: Darren Naish.

While we planned various signing events (at museums, shops and so on), none panned out for various reasons, so the closest thing we had to a launch was the signing event at TetZooCon… on which, read on. Already, Mesozoic Art has done well in terms of sales and positive response, all of which makes it likely that follow-up volumes might appear.

Another new book for 2022 – the aforementioned Ancient Sea Reptiles, published by the Natural History Museum – also appeared in my hands in October, though it’s not yet officially on sale and won’t be for a few weeks yet. I’ll be talking about it a lot when the time is right.

**Caption:** I have two copies of Ancient Sea Reptiles in my hands; I’ll have more soon. This is the cover of the UK edition (published by the Natural History Museum); the US edition (published by the Smithsonian Books) is different and features a Bob Nicholls painting. A few images from the interior are shown at right.

Articles on caecilians and zebras appeared at the blog at this time.

To Toronto. November was a packed month. For work reasons, I attended the Society of Vertebrate Paleontology meeting in Toronto, this being the 82nd annual meeting of the society. This was my first ever trip to Canada. As is usually the case when I visit an area for conference-related reasons, I didn’t much get to see the city, let alone the surrounding area: airport to hotel, hotel to conference venue, and repeat. Nevertheless, it was good to see friends old and new and catch up on the science. Many people (the society’s president Dr Jessica Theodor among them) wanted to talk about Prehistoric Planet and how much they enjoyed it, which was good.

**Caption:** an SVP Toronto montage. The excellent dromaeosaur head (part of an under-construction suit) is by Sam Stanton; at lower left we have a collection of Late Cretaceous dinosaur skulls at the ROM; in the middle, I’m signing a copy of *Mesozoic Art* for the montage; at far right we see Anthony Maltese with his favourite fossil, *Xiphactinus*. Images: Darren Naish.

Thanks to Jordan Mallon, I got hold of a physical copy of the issue of Canadian Journal of Earth Sciences that includes the paper on Dale Russell’s dinosauroid that Will Tattersdill and I published in 2021 (Naish & Tattersdill 2021). Incidentally, an original reprint of the Russell & Séguin (1982) paper on troodontids and dinosauroids appeared for sale in the SVP auction but I didn’t know this and someone else won it. While on the subject of the dinosauroid, I have to mention that Will – the Tattersdill of Naish & Tattersdill (2021) – was interviewed for an excellent dinosauroid-themed episode of the Palaeocast Podcast during September.

**Caption:** here’s me and Jordan Mallon at the Toronto SVP, with the Dale Russell special issue of *Canadian Journal of Earth Sciences*. At right: the cover (the published version looks different from the preview online).

Back to SVP: an added bonus was our conference visit to the Royal Ontario Museum (ROM), famous for its palaeontological displays. My phone ran out of charge, so I didn’t get to take as many photos as I hoped to. But you get at least some idea of what they have from the photos here. Highlights included the Paleogene and Pleistocene mammals, the mounted Pachycephalosaurus and the Jurassic dinosaur display. I donated a signed copy of Mesozoic Art to the SVP auction and am pleased to say that it fetched several hundred dollars.

**Caption:** mostly Pleistocene fossils at the Royal Ontario Museum, albeit with Mesosoic dinosaurs in the distance at right. Image: Darren Naish.

  **Caption:** more mostly Pleistocene fossils at the Royal Ontario Museum, including the giant sloth *Eremotherium*, the pampathere *Holmesina* and glyptodont *Glyptodon* (both are giant armadillos), and the cat *Smilodon*. Image: Darren Naish.

SVP done, I had time to do a bit of sight-seeing and visited Ripley’s Aquarium of Canada with Albert Chen. It’s not a bad aquarium but wasn’t as impressive as I was hoping. Highlights included the giant groupers, a gar and paddlefish exhibit, and the very nice ray exhibit.

**Caption:** familiar birds of the Toronto area, both new to me. Ring-billed gull *Larus delawarensis* and Black-capped chickadee *Poecile atricapillus*. Toronto is home to what’s said to be the world’s largest population of Ring-billed gulls. Images: Darren Naish.

Albert and I also went birdwatching. If you’re in a country for the first time ever, even seeing mundane and common ‘street birds’ can be exciting. New species for me included Double-crested cormorant Nannopterum auritum, Long-tailed duck Clangula hyemalis, Ring-billed gull and Black-capped chickadee. I was also interested to learn that Toronto is home to an entirely black form of the Eastern grey squirrel Sciurus carolinensis. It was possible to get quite close to these, and I photographed them whenever I could.

**Caption:** black squirrels of Toronto. These images show two different individuals. Images: Darren Naish.

Oregon and the aesthetics of dinosaurs. I was back home for a little while before it was time to disappear overseas once again, this time for Oregon and the 80th annual meeting of the American Society for Aesthetics in Portland. I’ve always wanted to visit the Pacific Northwest, and the trip did not disappoint. Even the flight in was a phenomenal experience that I’ve found myself recounting on many occasions. The photos show what I mean.

**Caption:** views from the air over what I think are parts of southern Alberta, and either Washington or Oregon. Incredible scenery, viewed in perfect conditions. Image: Darren Naish.

  **Caption:** I was thinking to myself “I wonder if I’ll get to see Mount St. Helens” when, suddenly… I think that’s Mount Rainier in the background. Image: Darren Naish.

The ‘aesthetics’ of the American Society for Aesthetics involves the philosophy of art criticism, and certain of the presentations involved jargon and argumentation way beyond my cognitive limits. I was there for a panel event devoted to the aesthetics of dinosaurs – yes, really – organised by Michel-Antoine Xhignesse. Besides Michel and myself, Zoë Lescaze and Derek Turner also spoke. We all approached dinosaur aesthetics from different angles. My talk was on the idea that those artists depicting Mesozoic animals in art today are mostly functioning as time-travelling wildlife artists: I reject the idea – which I’ve now seen implied or stated by philosophers several times – that palaeoart has some sort of symbolic or allegorical function. It was great and we got a lot of positive feedback from our audience.

**Caption:** my cover slide for the dinosaur aesthetics events of the 80th annual meeting of the American Society for Aesthetics, December 2022. I haven’t discussed the contents of the talk outside of the relevant event itself. Maybe sometime I will.

  **Caption:** the four speakers of the dinosaur aesthetics events of the 80th annual meeting of the American Society for Aesthetics, December 2022. From left to right: Derek Turner, Zoë Lescaze, Michel-Antoine Xhignesse and Darren Naish.

Portland itself was fun. I walked part of the length of the Willamette River and saw a good number of birds that were (again) new to me, including assorted geese, gulls, eagles and a ton of passerines. I don’t want to talk about them here as I’m saving that for another article. I also discovered Powell’s Books – where I had to limit myself in terms of the number of books I obtained – and a few vintage toy shops.

**Caption:** argh, books on animals — my one weakness! Powell’s Books, Portland, is home to hundreds of natural history books, a great many of which I would obtain if I had unlimited money and space. As it happens, I purchased a hardback original of Paul’s *Predatory Dinosaurs of the World* (previously, I only owned the softback) and only a couple of other things. Images: Darren Naish.

Going squatchin’. The ASA meeting done, I headed east in the direction of Mount Hood, specifically to the town of Rhododendron where I stayed with my friends Nico and Haley Spadafore. A whole week dedicated to exploring the forests, mountains and coasts of the state of Oregon. Thanks to Nico, I also visited Cliff Barackman’s North American Bigfoot Center in Boring.

**Caption:** myself and Cliff Barackman (at left) at the North American Bigfoot Center, with Murphy the sasquatch in the background. Wow, I sure do like that jumper.

At the risk of repeating what I said about the Loch Ness Monster earlier in this article, regular readers of this blog will know that I’m seriously interested in bigfoot whether it’s a sociocultural phenomenon (as I currently suspect) or a biological one (oh, how I wish it were so), and with that in mind I found it a welcoming and fascinating museum. It’s dedicated, of course, to the idea that bigfoot is a real primate species awaiting scientific discovery, with tons of content on eyewitness experiences, field evidence, and bigfoot history and lore. It also has the best shop dedicated to bigfoot-themed material I’ve ever seen. I bought way too much stuff.

Cliff himself was gracious enough to spend time with me, and kindly showed me his collection of bigfoot print casts. I can now say that I’ve seen copies of quite a few of the famous bigfoot, yeti, almas and yowie tracks.

**Caption:** an ambition fulfilled — I finally get to visit the spectacular forests of the North American Pacific Northwest. It wasn’t especially cold, but it was wet. Image: Darren Naish.

I always knew that going squatchin’ would be a rewarding experience whatever the outcome. We travelled to two confidential, essentially private stretches of woodland where people have reported bigfoot encounters. I heard coyotes in the wild for the first time. Nico is a good tracker, and showed me bobcat, coyote and elk tracks. At one of the locations we visited, Nico, Cliff and I heard several unusual sounds – they seemed to be animal vocalisations, and what sounded like a wood knock – that defied easy explanation. I’m not saying that we heard bigfoot. I’m not saying that bigfoot is real. I’m saying that we experienced some unusual stuff.

**Caption:** I really, really enjoyed visiting sites of possible interest in the Oregonian wilderness. Had lots of wildlife experiences, and loved the scenery and environments. Image: Darren Naish.

Other places I visited included the spectacular Multnomah Falls in the Columbia River Gorge (where I saw my first chipmunk), the Bonneville fish hatchery (home to a great number of spectacular sturgeon), and the Oregon Coast Aquarium at Newport. It has some stupendous exhibits that house Pacific marine wildlife, including rockfishes, sharks, harbour seals and sea otters. The aviary was closed due to the current avian flu outbreak but I at least got to see swimming puffins.

**Caption:** some of the many sturgeon on show at the Sturgeon Viewing and Interpretive Center at Bonneville Hatchery, adjacent to the Columbia River in Oregon. Several of the sturgeon there are big, the largest being Herman. He’s 3.35 m long and weighs over 220 kg. Herman was stolen in 1983 – yes, I said stolen – and there has been more than one occasion in which people have jumped into the water and stabbed him. Image: Darren Naish.

  **Caption:** a few more Oregon scenes. From left to right: Multnomah Falls, a stretch of scrubby ground at the edge of the Sandy River in the town of Rhododendron, and a late evening scene taken from Cliff’s vehicle while deep in the woods. Images: Darren Naish.

  **Caption:** the Oregon Coast Aquarium at Newport. It houses numerous really nice marine exhibits and has various pinnipeds on show. I was surprised to see that harbour seals and fur seals are kept together, but apparently they’re friends and it’s not an issue. Images: Darren Naish.

  **Caption:** views of the Oregon coast. At left, a Pacific Harbour seal surfing waves at Siletz Bay. At right, gulls and and other birds at Yaquina Bay. Images: Darren Naish.

We also visited Oregon Zoo. That’s something I also intend to write about it in a separate article so won’t say much about here, but it’s especially strong on North American wildlife, highlights for me including California condor Gymnogyps californianus and Rocky mountain goat Oreamnos americanus. Incidentally, neither the aquarium nor zoo had good shops – at least, not in terms of what I hope for in a shop attached to a place devoted to the displaying of animals. Huh.

**Caption:** a montage of things from Oregon Zoo, a great zoo that has an excellent collection of North American animals as well as much else too. I’ll be talking in detail about what the zoo is like in a later article. Images: Darren Naish.

TetZooCon 2023. After returning from Oregon it was time for TetZooCon 2022, the 9th Tetrapod Zoology Convention. It was the biggest so far (thereby continuing the trend in which the meetings have gotten bigger year on year) and with a packed schedule: if you want to know more go see the article here. This year’s TetZooCon was held at Bush House, King’s College, a venue we owe to Chris Manias, whose Popularising Palaeontology event was combined with TetZooCon this year.

It looks like the same deal will be in operation for 2023, in which case we’ll see you at Bush House again for the TENTH TetZooCon. Stay tuned – it should be something special.

**Caption:** a good number of TetZooCon 2022 photos were included **in the relevant article**, but here’s one that wasn’t. This image was taken during John Conway’s talk *A History of Painting (With Dinosaurs)*. Image: Darren Naish.

Also in December, I finally outed the long-running ‘Plot bone’ project I worked on with Martin Simpson, Paul Stewart and Hilary Ketchum. We did discuss at one point writing the whole thing up as a paper – despite the negative results – but in the end I opted to write the story up as a TetZoo article; it’s here. Inspired by the release of the movie Avatar: the Way of Water, I republished the TetZoo guide to the creatures of the first Avatar movie. It’s here.

And so to January 2023…

Monsters of the Deep says goodbye to Cornwall. Long-term readers here will know of my Monsters of the Deep (MOTD) exhibition at the National Maritime Museum, Cornwall (if you don’t, an article announcing its opening is here). January is a slightly sad month because this is when MOTD’s tenure at NMMC comes to a close. I had planned to lead a tour of the exhibition prior to the final day, but it just wasn’t doable.

**Caption:** exterior of the National Maritime Museum, Cornwall, in Falmouth. Formerly the home of the Monsters of the Deep exhibition…. which is now moving elsewhere! Image: Darren Naish.

Anyway, I’m now allowed to say that MOTD isn’t finished for good: it’s travelling and is currently being installed at Chatham Historic Dockyard in Kent. This is great news and means that a huge number of people in the south-east of England (including the London area) will be able to see it. More news on this soon.

Dinosaurs in Dorchester. I visited the Dinosaur Museum at Dorchester. I haven’t been there for years and was interested in seeing what had changed. They’ve replaced or modified various of the displays on the walls, a life-sized Allosaurus model has gone, and a Tyrannnosaurus statue – based, I think, on the Invicta Tyrannosaurus model from 1977 – has been given a new head. But many of the older things are still there.

**Caption:** the Dinosaur Museum at Dorchester is certainly interesting if you’re interested in historical depictions of dinosaurs, since it houses quite a few unusual models and other objects. The images here show a *Corythosaurus* costume/model said to be connected to Pertwee-era Dr Who (though I’ve seen this contested), a model of Richard Owen (for once, not depicted as ‘old man Owen’) and a very familiar replica theropod skeleton (with a model *Megalosaurus* behind). Images: Darren Naish.

This museum holds a special place in my memory because it’s the first place where I saw – saw, wasn’t able to buy – several dinosaur-themed books that proved crucial to my nascent interest, namely David Lambert’s Collins Guide to Dinosaurs and Peter Zallinger’s Dinosaurs and Other Archosaurs.

But the big event of January was my research trip to the Fort Pierce area of Florida. I assure you that I was in Florida for very interesting reasons, but I can’t yet say what they were. So don’t ask. Whatever, I saw tons of stuff. I’ve been to the American tropics before (I visited Brazil for a pterosaur conference in 2013), but this was my first time in Florida and about everything I saw wildlife-wise was new. Yet again I’m not going to say much here since I’ll discuss it all later (some of what I saw has been shared via Twitter already).

**Caption:** just a little of the wildlife I saw while in Florida. Yes, that’s a Limpkin *Aramus guarana*. Images: Darren Naish.

What happened at Tetrapod Zoology during January? I published a modified version of my article on Tripoli Zoo in Libya and another on the laryngeal sacs of baleen whales. And that brings us up to the present.

So…..

Reviewing 2022. As you’ll know if you’ve read these birthday articles before, one thing I like to do is list the year’s blog articles (the year extending from Jan 21st 2022 to Jan 21st 2023, of course) in order to see what taxonomic coverage was like. And if you’ve read these articles before, you’ll know how things pan out: I find that charismatic megafauna and weird stuff have been covered disproportionately, and I then get cross about it. So let’s start with a list of all the year’s articles, grouped by category…

Miscellaneous

A Look Back at Dorling Kindersley’s 2001 Encyclopedia of Dinosaurs & Prehistoric Life
Happy 16th Birthday, Tetrapod Zoology
Announcing the All Yesterdays Range of Collectible Figures
Four Years of Tetrapod Zoology ver 4
TetZooCon 2022 Is Go
Tetrapod Zoology at the United Nations Science Summit 2022
Announcing Mesozoic Art, a Lavish New Volume on Modern Palaeoart
The Ninth and Largest of the Tetrapod Zoology Conventions
The Tet Zoo Guide to the Creatures of Avatar, Updated for 2022
TetZoo Reviews Zoos: Tripoli Zoo in Libya

Amphibians

Tiny Frogs and Giant Spiders: Best of Friends
The Remarkably Weird Skeletons of Frogs
The Amazing Caecilians

Mammals

Kogia, Shark-Mouthed Horror
The Incredible South American Maned Wolf
Meeting the Hayling Island Jungle cat
Suddenly -- Duikers!
Hartebeest, Long-Faced Antelopes of Many Forms
Domestic Horses of Africa
The Domes of Wisdom: an Asian Elephant Tale
The Slightly Surprising Diversity of Zebras, Part 1
Pouches, pockets and sacs in the heads, necks and chests of baleen whales

Non-bird dinosaurs

A Giant Spinosaurid Dinosaur from the Cretaceous of the Isle of Wight
The Eotyrannus Monograph Is Finally Published
Robert Plot’s Lost Dinosaur Bone

Birds

Chiffchaffs and What Are 'Old World Warblers' Anyway?
Inside the Cassowary's Casque, Updated for 2022
Otidiphaps the Pheasant-Pigeons
Surprising Diet of the House Sparrow

Cryptozoology

Legend of the Black Dog

What’s obvious for 2022’s coverage is that most tetrapod groups got no coverage at all. I put that down to me not really blogging much over the year. The fact that mammals are so well represented is (as ever) a surprise; the fact that miscellaneous material is out in front probably reflects increasing coverage of parish notices, and reminiscences about books and such. As ever, I’m not pleased with the lack of balance, but at least amphibians got a look in. Seems bizarre that non-dinosaurian reptiles got no coverage, but there you go.

Caption: it should be obvious from these birthday articles that I photograph animals as and when I can, generally quite badly. Birds are among the animals we see most often here in the UK, and here are two passerines photographed at Milford-on-Sea in September 2000. The blob at left is a European rock pipit Anthus petrosus; the corvid at right is a Rook Corvus frugilegus carrying a potato… or a rock, I can’t remember. Images: Darren Naish.

Wrapping it up. So there we have it: the TetZooniverse year in review. A lot happened, but I survived. The Eotyrannus monograph published. The book Mesozoic Art published, and a second – Ancient Sea Reptiles – finished, though not yet released for sale. The biggest and likely best TetZooCon so far. The TV series Prehistoric Planet released to the world. Two technical papers published (Barker et al. 2022, Naish & Cau 2022). I think that those are substantial and satisfying wins. But, as ever, I’m frustrated by the fact that there are those grand projects where little to no progress was made, most notable of these being the textbook. I can’t say more about it or I’ll get angry.

So, happy 17th birthday Tetrapod Zoology. 17 years of this blog, at four different hosting sites. As ever, I want to thank those of you who read and visit Tet Zoo; especially to those of you who comment and interact here, and to those who provide financial support and help make it a worthwhile venture. Thank you, thank you all.

Welcome to the 18th year of Tetrapod Zoology.

**Caption:** Mount Hood, Oregon, December 2022. Image: Darren Naish.

For previous Tet Zoo birthday articles, see...

Happy first birthday Tetrapod Zoology (part I), January 2007
Happy first birthday Tetrapod Zoology (part II), January 2007
Happy second birthday Tetrapod Zoology (part I), January 2008
Tetrapods of 2007 (happy birthday Tet Zoo part II), January 2008
Happy THIRD birthday Tet Zoo, January 2009
Tet Zoo = 4 years old today, January 2010
2009, a year of Tet Zooery, January 2010
Four years of Tet Zoo: to infinity... and beyond!, April 2010
It is with some dismay that I announce Tet Zoo's first hemi-decade, January 2011
Tet Zoo 5th birthday extravaganza, part II, January 2011
Happy Birthday Tetrapod Zoology: SIX YEARS of blogging, January 2012
Happy 6th Birthday, Tetrapod Zoology (part II), January 2012
Tetrapod Zoology enters its 8th year of operation, January 2013 (I cannot find any intact versions of this article, thanks SciAm)
Today marks NINE YEARS of Tetrapod Zoology, January 2015
Tetrapod Zoology 10th-Birthday Extravaganza, Part 1: 2015 in Review, January 2016
Tetrapod Zoology 10th Birthday Extravaganza, Part II: the Rest of 2015 Reviewed, January 2016
Tetrapod Zoology 10th-Birthday Extravaganza, Part 3: Tet Zoo's Tetrapod Treatment in 2015, January 2016
Today Is Tet Zoo's 11th Birthday, January 2017
The 12th Year of Tet Zoo, January 2018
The Tet Zoo 12th-Birthday Event, Part 2, January 2018
The Much Belated Final Part of the Tetrapod Zoology 12th Birthday Event, December 2018
Tetrapod Zoology Is A Teenager Now, January 2019
Tetrapod Zoology's 14th Year of Operation, 2019 in Review, January 2020
On Tetrapod Zoology’s 15th Birthday, the Year in Review, January 2021
Happy 16th Birthday, Tetrapod Zoology, February 2022

Huge thanks to those who help support this blog at patreon. You can see behind-the-scenes and in-prep stuff relevant to projects for as little as $1 per month.

Refs - -

Blake, M., Naish, D., Larson, G., King, C. L., Nowell, G., Sakamoto, M. & Barnett, R. 2014. Multidisciplinary investigation of a ‘British big cat’: a lynx killed in southern England c. 1903. Historical Biology 26, 441-448.

Naish, D. 2017. Hunting Monsters. Arcturus, London.

Naish, D. 2022. A cultural phenomenon. The Biologist 69 (3), 16-21.

Naish, D. & Cau, A. 2022. The osteology and affinities of Eotyrannus lengi, a tyrannosauroid theropod from the Wealden Supergroup of southern England. PeerJ 10:e12727.

Naish, D. & Tattersdill, W. 2021. Art, anatomy and the stars: Russell and Séguin’s dinosauroid. Canadian Journal of Earth Sciences 58, 968-979.

Russell, D. A. & Séguin, R. 1982. Reconstruction of the small Cretaceous theropod Stenonychosaurus inequalis and a hypothetical dinosauroid. Syllogeus 37, 1-43.

White, S. & Naish, D. 2022. Mesozoic Art: Dinosaurs and Other Ancient Animals in Art. Bloomsbury Wildlife, London.

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Way back in 2010, I published a series of articles on the various pouches, pockets and sacs (virtually all of which are laryngeal diverticula of one sort or another) that exist in the heads, necks and chests of mammals. I never finished that series.

**Caption:** screengrabs of parts I and II of the pouches, pockets and sacs series.

Here’s one of the articles from that series: the one on baleen whales. The original is here. I haven’t really updated it relative to the 2010 original, but if corrections are required I’ll add them when time allows.

**Caption:** cartoon from Desray Reeb's 1997 thesis, the bagpipe here being a clever reference to the potential role of the laryngeal sac as a vocal organ… on which read on.

Like the primates we looked at previously, mysticetes have enlarged laryngeal ventricles that (mostly) meet along the ventral midline of the throat and form a single large laryngeal pouch or sac. The presence of a raphe along the sac's ventral midline seems to mark the line of fusion between the two ancestral, bilateral sacs. It's probably understandable that few of us are aware of the presence of inflatable laryngeal sacs in mysticetes, but people have known about the existence of these structures for a long time: pioneering scientist and surgeon John Hunter, for example, wrote about their presence in Northern/Common minke whales Balaenoptera acutorostrata* in 1787, and they've frequently been remarked on in the cetacean anatomical literature.

The laryngeal ventricles are normally bilateral recesses, located on either sides of the glottis (if you need help with the terminology, see the relevant section in the first article).

Caption: everything about whales is amazing. Here’s a minke whale skeleton, probably belonging to a Northern or Common minke. I should mention in passing that minke whales appear to be a species complex. Image: User BS Thurner Hof, CC BY-Sa 3.0 (original here).

The sac is surrounded by muscle and is very flexible: when it's contracted, a series of folds in the side walls provide additional surface area that facilitates expansion (Reidenberg & Laitman 2007a). Some authors have suggested that the sac can be filled with air when the whale surfaces, and that they sac can then be used as an oxygen store during diving (Negus 1962). The idea here is that the whales are able to push this stored gas from the laryngeal sac into the lungs and thereby 'take a breath' while at depth (and 'at depth' should - based on what we know about dive depths of whales and on the distribution of possible prey - be well below 100 m).

**Caption:** it should be obvious from this article where the laryngeal sac is located, but this dissection image of Fin whale anatomy (from Brodie & Påsche 2001, p. 357) helps. Here. you’re standing in the location of the lungs, looking towards the position of the tip of the snout. The trachea (t) is huge and wide (c 40 cm); the oesophagus (e) is above it. The laryngeal sac (s) is massive, heavily muscular, and beneath the trachea.

The problem with this idea is that it can't really work at depth at all, given that the lungs are thought to collapse at depths greater than 100 m, thereby making any gas exchange impossible (Reeb & Best 1999). Also possibly counting against the idea of an extra 'gas store' in mysticetes is that it seems inconsistent with the surprisingly low theoretical aerobic dive limits (TADLs) present in some species (namely the rorquals): I covered the topic of mysticete TADLs, and other neat aspects of their diving and feeding behaviour, here.

However, while respiratory exchange might not be possible at depth, it's still theoretically conceivable that the laryngeal sac might work as a gas store in shallow water... even though this would put the whales close to the surface and hence in relatively ready reach of the sea surface. Also possible is that the sac does indeed serve as a 'gas store', but that the role of this gas is not a respiratory one: maybe the whales move gases from the sac into other parts of the respiratory tract to prevent lung collapse, or ear damage, during diving (Reidenberg & Laitman 2010)

**Caption:** Blue whale exhaling at the surface, "the greatest respiratory action in history". Image: NOAA, public domain (original **here**).

A role in dynamic exhalation? Another really interesting possibility is that the sac doesn't function as a 'store' at all, but that it plays a crucial role in the dynamic exhalation practised by these animals: an event described by Brodie & Påsche (2001) as "the greatest respiratory action in history" (p. 353). What is it with cetaceans and superlatives? Brodie & Påsche (2001) proposed that – in its inflated condition – the laryngeal sac becomes forced up against the glottis when the whale is at depth, and thereby forces the arms of the paired arytenoid cartilages together to form a tight seal. The arms of the arytenoid cartilages seem to be similar in all mysticetes, and have often been referred to in the descriptive literature as 'lips' located at the opening of the laryngeal sac.

Increasing the pressure within the sac forces the arytenoid cartilage arms to push even closer together: they seem to act as a self-sealing valve. In another memorable quote, Brodie & Påsche (2001) said that "the remarkable size and robust nature of this entire tracheal/laryngeal mechanism appears more reminiscent of some man-made industrial device, than a component of the respiratory tract of a mammal" (p. 359).

It might be that, as pressure is relieved in the trachea (through the opening of the blowhole), the seal formed by the arytenoid cartilages relaxes, causing the laryngeal sac to deflate and fall away from its close contact with the trachea. This, in turn, allows the gases in the bronchi and lungs to be suddenly released; they then rush outwards at high speed, forming the characteristically explosive blow. Brodie & Påsche (2001) performed a number of experiments with the laryngeal sacs of fin and sei whales at an Icelandic whaling station, and found that the structures operated as hypothesised. Their proposal that the sac facilitates high velocity exhalation is a neat one... the problem is that it seems to contradict other possible functions, as we'll see.

A vocal role. Moving away from a role in exhalation, it's also been argued that the sac is used in vocalising. It's well known that mysticetes are able to generate low-frequency noises, and there's no doubt that these noises come from the lower and lateral parts of the throat (a contrast to odontocetes [toothed whales], where noises are mostly – but perhaps not entirely – generated by apparatus around and within the nasal region). Accordingly, it has been suggested that the comparatively enormous mysticete larynx is the source of these sounds (e.g,, Quayle 1991, Reidenberg & Laitman 2007a). Recent spectral analyses of Humpback Megaptera novaeangliae songs seem to show that the noises are indeed generated here, with resonating air chambers probably contributing to sound production (Mercado et al. 2010).

**Caption:** diagram (from Reidenberg & Laitman 2007b) showing how the laryngeal sac (marked 's' in upper diagram) might partly function as an air store, receive air from the lungs, and have a role in the production of a bubble cloud.

How might the sac contribute to vocalisation? Perhaps it enables air to be 'recycled' between the larynx and lungs during vocalising. And perhaps it (also?) acts as a resonating chamber when inflated (some pinnipeds may use laryngeal diverticula in the same way: we'll be looking at pinnipeds, and other carnivorans, later on). The adjacent diagram (from Reidenberg & Laitman 2007b) shows a Humpback using the laryngeal sac in concert with its lungs and mouth during the release of a bubble cloud (the ability of humpback whales to release bubbles is itself a fascinating and complex operation).

The massive size of the mysticete larynx, the large size of the expandable laryngeal sac, and the large size of these animals in general seem, combined, to allow mysticetes to generate some of the loudest, longest and lowest sounds produced by any mammal. As Reidenberg & Laitman (2010) said, they are vocal athletes.

**Caption:** most of us know that humpbacks sing, and make a variety of other sounds too. But the idea that laryngeal structures have a possible resonating role isn’t well known outside of specialists on cetacean anatomy. Image: Christopher Michel, CC BY 2.0 (original **here**).

That contradiction I mentioned earlier. You might have noticed that the possible use of the laryngeal sac in seal-forming and dynamic exhalation (as proposed by Brodie & Påsche (2001)) is, apparently, at odds with the possible use of the sac in relieving pressure in the respiratory tract or ear region, as a gas store, or in vocalising. If the sac works in maintaining a tight seal at the arytenoid cartilages, it has to (I presume) remain inflated: if it becomes deflated, high pressure in the bronchi and lungs is lost, and the relaxation of the proposed laryngeal seal cannot be the 'trigger' responsible for dynamic exhalation. Yet, if the sac is used in vocalising or as a store allowing deeper and/or longer dives, gas simply must be passed back and forth between the sac and the trachea at least. It seems to me that the possible role of the laryngeal sac in vocalising is likely and backed by experimental support. Ergo, the presence of a permanently inflated sac that help form a seal against the glottis cannot be permitted. The proposals seem to contradict one another.

However... is it possible that the sac might still work in sealing the glottis at particular times (such as immediately prior to exhalation)? Maybe the sac can be used as a gas store and/or as a resonating chamber when the whale is at depth (albeit not at depths greater than 100 m, as discussed above), but can be pressurised and used to seal the throat when the whale approaches the surface for exhalation? If mysticetes can blow and vocalise at the same time, the 'glottal seal' hypothesis is dead. Probably.

Caperea: weird weird weird, like you didn't already know. While the laryngeal sacs of most mysticetes are located on the midline, that of the weird little Pygmy right whale Caperea marginata is positioned on the animal's right side (Reeb & Best 1999). The sac undergoes extensive expansion and contraction: possibly more than that of any other mysticete.

**Caption:** *Caperea* lungs and associated laryngeal sac (in the human hand).

It's possible that the unusual laryngeal sac anatomy of Caperea explains the unusually long thorax and weird, partially overlapping, strangely flattened ribs of this whale: this long thorax, and the weird rib shape, might provide a lot more space for expansion of the laryngeal sac, and help to protect and maintain the sac's shape when fully inflated (Reeb & Best 1999). The ambiguity in this explanation – for which I apologise – results from the fact that the proposed correlation here is a vague, speculative and un-tested. It's also been said that the ribs of Caperea* are rather loosely connected to the transverse processes of the vertebrae (Beddard 1901), perhaps implying that the ribcage is more flexible than is usual for a mysticete. Those weird ribs have always been the source of interesting speculation: there's the idea that they function as some sort of armour, and the peculiar notion that they perhaps help support the whale's weight when it lies on the sea floor... yeah, because whales are always lying down on the seafloor, right?

The thoracic vertebrae make up about 39-45% the length of the vertebral column, whereas they make up less than 33% the length of the column in all other mysticetes (Reeb & Best 1999).

Caption: image from a Caperea dissection, showing the distinctive, partly overlapping ribs of this most unusual whale.

As every Caperea fan knows, the dissection of a juvenile that stranded in New Zealand in 2007 was live-blogged at Te Papa's Blog. A team of international experts, including Catherine Kemper, Ewan Fordyce, Joy Reidenberg and Sentiel Rommel, assisted in the dissection: the image above of the lungs and (juvenile!) laryngeal sac is borrowed from the event, as is the adjacent picture of the peculiar, overlapping ribs (the size of the laryngeal sac increases substantially during ontogeny: the adult sac is about five times bigger than that of the juvenile (Reeb & Best 1999)).

**Caption:** *Caperea* skeleton, photographed in New Zealand. Other images of the same skeleton appear in **this Tet Zoo article from 2010**. Image: Joy Riedenberg, used with permission.

We'll finish on cetaceans by noting that laryngeal sacs are not unique to mysticetes: they're also present in such odontocetes as the Sperm whale Physeter macrocephalus, Beluga Delphinapterus leucas and Risso's dolphin Grampus griseus.

For the previous article on pouches, pockets and sacs in mammal heads, necks and chests, see...

Pouches, pockets and sacs in the heads, necks and chests of mammals, part I: primates, October 2010
Pouches, pockets and sacs in the heads, necks and chests of mammals, part II: elephants have a pouch in the throat... or do they?, October 2010

If you're interested in tracheae and their role in respiration, vocalising and such, or on any of the associated structures in the neck, check out...

Deer oh deer, this joke gets worse every time I use it, November 2007
Dissecting an emu, April 2009
Ridiculous super-elongate, coiled windpipes allow some birds to function like trombones - - or is it violins?, April 2009
Inside Nature's Giants part IV: the incredible anatomy of the giraffe, July 2009
Dissecting lions and tigers: Inside Nature's Giants series 2, part III, September 2010

And for more on mysticete anatomy, evolution and diversity, see...

A 6 ton model, and a baby that puts on 90 kg a day: rorquals part I, October 2006
From cigar to elongated, bloated tadpole: rorquals part II, October 2006
Lunging is expensive, jaws can be noisy, and what’s with the asymmetry? Rorquals part III, October 2006
Inside Nature's Giants part II: whale guts and hindlimbs ahoy, July 2009
When GREY WHALES - you know, from the PACIFIC OCEAN - crossed the Atlantic, May 2010
Did I mention that Caperea is really, really weird?, October 2010
Once more on little Caperea, October 2010
Caperea alive!, November 2010
Cetacean Heresies: How the Chromatic Truthometer Busts the Monochromatic Paradigm, April 2015 (but now lacking all images)
Remembering Lyall Watson’s Whales of the World, June 2019
Whale Watching in the Bay of Biscay, August 2019

If you enjoyed this article and want to assist me in what I do — and see behind-the-scenes stuff in development — please support me at patreon. Click anywhere on this text in bold.
Refs - -

Beddard, F. E. 1901. Contribution towards a knowledge of the osteology of the pygmy whale (Neobalaena marginata). Transactions of the Zoological Society of London 16, 87-114.

Brodie, P. F. & Påsche, A. J. 2001. The mechanics of cetacean respiration: the significance of rapid gas exchanges in a selectively tuned system, with emphasis on the rorquals (Balaenoptera sp.). In Mazin, J.-M. & de Buffrénil, V. (eds) Secondary Adaptation of Tetrapods to Life in Water. Dr Friedrich Pfeil (München), pp. 353-362.

Ellis, R. 1982. The Book of Whales. Alfred Knopf, New York.

Mercado, E., Schneider, J. N., Pack, A. A. & Herman, L. M. 2010. Sound production by singing humpback whales. The Journal of the Acoustical Society of America 127 2678-2691.

Negus, V. E. 1962. The Comparative Anatomy and Physiology of the Larynx. New York, Hafner.

Quayle, C. J. 1991. A dissection of the larynx of a humpback whale calf with a review of its functional morphology. Memoirs of the Queensland Museum 30, 351-354.

Reeb, D. 1997. Comparative anatomy of the larynx of the minke whale, Balaenoptera acutorostrata and the pygmy right whale, Caperea marginata. MSc disertation, University of Pretoria, Pretoria.

Reeb, D. & Best, P. B. 1999. Anatomy of the laryngeal apparatus of the pygmy right whale, Caperea marginata (Gray 1846). Journal of Morphology 242, 67-81.

Reidenberg , J. S. & Laitman , J. T. 2007a. Discovery of a low frequency sound source in Mysticeti (baleen whales): anatomical establishment of a vocal fold homolog. Anatomical Record 290, 745-760.

Reidenberg , J. S. & Laitman , J. T. 2007b. Blowing bubbles: an aquatic adaptation that risks protection of the respiratory tract in Humpback whales (Megaptera novaeangliae). The Anatomical Record 290, 569-580.

Reidenberg , J. S. & Laitman , J. T. 2010. Generation of sound in marine mammals. In Brudzynski, S. M. (ed) Handbook of Mammalian Vocalization. Elsevier, pp. 451-466.

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In 2009, I visited a zoo in Libya, and here are thoughts on it…

**Caption:** Tripoli Zoo as of 2009, home to bears, gazelles and more. Read on…

Regular readers of Tetrapod Zoology will know that I occasionally write reviews of zoos I’ve visited. I’ve long intended for this ‘zoo review’ series to be useful and even comprehensive, but the fact remains that I only rarely get round to writing them. Anyway, find a list of what I’ve done so far at the bottom of this article.

Here, let’s look at a zoo you may not have heard of, or indeed be aware of: Tripoli Zoo in Libya. I was fortunate enough to visit Libya in October 2009 for fieldwork reasons, and on one of my final days in the country I made a special trip to the zoo. Here are my thoughts. They partly repeat things I published back at Tet Zoo ver 2 in 2009, but are augmented and modified quite a bit, and include new and more images.

**Caption:** I enjoyed my time in Libya. It’s a huge country with some incredible landscapes. I was lucky enough to visit several rocky desert locations. Image: Darren Naish.

  **Caption:** the walls surrounding Tripoli Zoo feature a series of bass reliefs, mostly depicting African wildlife. This one has an Indian rhino at upper left but otherwise shows African rhinos, giraffes, hippos and elephants. Image: Darren Naish.

By way of introduction I have to say that I know little about Tripoli Zoo and, despite checking, haven’t succeeded in finding out much about it. The Muslim world has a long and noble tradition of menageries and other animal collections dating back to the 10th century at least, and Islamic writings make it clear that animals kept in captivity must be cared for properly and kept in good surroundings. I didn’t get the impression from the structure of Tripoli Zoo that it had a history extending beyond, say, the 1970s but I’d love to know more.

Information about the zoo online centres around two stories: one being that it and its animals were mostly abandoned during the Arab Spring of 2011 (which resulted in the Libyan Civil War), another being that it was part-used as a detention centre in 2013. The 2011 Libyan Civil War – mostly under-reported in Europe and North America – was a huge event in the country’s history, the death toll probably exceeding 10,000.

**Caption:** just two of the many news stories that have covered Tripoli Zoo since 2011.

So far as I understand, the zoo and its animals ultimately weathered these events and the zoo persists – with a mostly intact collection of animals – today. This is in part thanks to the efforts of Amir Khalil, an extraordinary man associated with animal rescue events in Bulgaria, Iraq, Palestine, Pakistan and elsewhere. Obviously, my understanding of issues here is rudimentary and skeletal and I apologise for being scant on details. If you know more, feel free to add comments below.

Another point I have to add is that a large section of the zoo was under renovation at the time of my 2009 visit, and as a result there was less to see than usual. Ironically, neither Addax Addax nasomaculatus nor Cheetah Acinonyx jubatus were there, despite featuring on the zoo’s entrance sign. In fact, having read what I can on the zoo’s history, it’s clear that it previously included many more animals than it did in October 2009, including elephants, rhinos, giraffes, ostriches, hippos, zebras, hyenas, lions, tigers, lynxes, tapirs, kangaroos, monkeys, water buffalo and more. I know from photos online that at least some of these animals were back there once the renovation was finished. Despite this, I was fairly impressed. The animals were in good condition, the enclosures were reasonably large, and the selection of animals was pretty good. It certainly was not a bad zoo. The complete lack of signage did, however, reduce any educational value zoo-going might have for the public, to put it mildly.

Crowned cranes, raptors and more. We’ll start with birds. The zoo had a fairly good collection, including several species I’ve never seen before. Many zoos today have crowned cranes, but these are virtually always Grey crowned cranes Balearica regulorum, the larger of the two species, and the one that’s light grey and with white cheeks. Tripoli Zoo had this species but also the Black crowned crane B. pavonina, which is darker, smaller and has pink cheeks.

**Caption:** Black crowned cranes of Tripoli Zoo (still the only examples of this species I’ve seen), with White stork in the background. This species occurs east to west across central Africa. Images: Darren Naish.

Greater flamingos Phoenicopterus roseus, White stork Ciconia ciconia, Muscovy Cairina moschata, Black swan Cygnus atratus and Emu Dromaius novaehollandiae were also on display, as were various parrots, including Green-winged macaw (aka Red-and-green macaw or Red-blue-and-green macaw) Ara chloroptera. There were also aviaries with Red kite Milvus milvus, Indian peacock Pavo cristatus and what I think were female Golden pheasant Chrysolophus pictus.

**Caption:** a Tripoli Zoo bird montage, showing (clockwise from far left) Green-winged macaw, Golden eagle, Golden pheasant, and Red kite. Images: Darren Naish.

An additional large aviary was devoted to big raptors, namely Egyptian vulture Neophron percnopterus, Eurasian griffon Gyps fulvus and Golden eagle Aquila chrysaetos. I don’t get to see Egyptian vultures much, so spent a lot of time taking pictures of the one they had.

**Caption:** I’m really interested in what this Egyptian vulture is doing with its feet. Note here how it seems to be partly supporting its weight with flexed toes — but, look at the next photo…

  **Caption:** it turns out that the vulture is holding something in its strongly flexed toes. It’s now switched from doing whatever it was doing with its right foot to its left one. It’s not surprising that a raptor can do this, but it’s still interesting to see it. Image: Darren Naish.

Cats and bears. Moving now to mammals, let’s start with the carnivorans. As mentioned above, Tripoli Zoo has been (and is now?) home to tigers and lions – and even a liger – but none were present when I visited. However, several Puma Puma concolor were there, as were several black Leopard Panthera pardus. The camera I owned at the time wasn’t of the sort that allows you to take photos of animals that are at distance, and kept behind dense-mesh fencing, so I don’t have any photos of the leopards.

**Caption:** a pretty poor photo, but the best I could do in view of limitations. There are three pumas in this photo; two are reclining on the platform. Presumably all three are related. Image: Darren Naish.

On the subject of cats, I have to mention that a number of feral cats Felis catus were hanging out in the zoo, sometimes literally in the enclosures. That’s not great in that they can act as disease vectors to other carnivoran species and even predate on zoo animals (various of the birds there could potentially be on a feral cat’s menu). It’s difficult to think of a way of excluding them from the zoo without potentially endangering the zoo’s formal inhabitants.

**Caption:** Brown bears of Tripoli Zoo. I had the impression that the larger, broader-headed individual at right (with the paler fur and unusual left ear) was male and the other female, but I could be very wrong. Image: Darren Naish.

Among other carnivorans, I certainly remember the Brown bears Ursus arctos. Both were adults of dark brown, typical appearance (I mean: they weren’t reddish or unusually patterned like certain Middle Eastern and east Asian brown bear forms) but I don’t know anything about their origins. Sadly, both bears were displaying abnormal behaviour. It’s obvious that bears, like primates, need stuff to keep them constantly occupied. Fail in that and they go insane.

**Caption:** I think it’s obvious just from my few photos here that these were (and still are?) unfulfilled, bored bears. In case you’re wondering, the bears aren’t separated from the public merely by a short metal fence. As you can see from the image at right, there’s actually a deep moat around the whole enclosure. Image: Darren Naish.

Cattle and antelopes. The zoo’s collection of hoofstock – by which I mean artiodactyls, or even-toed hoofed mammals – was impressive, again including a number of species I’ve never seen in captivity or at all. I’ll say to start with that they had several familiar domestic species, including Domestic cattle Bos taurus, Dromedary Camelus dromedarius and Llama Lama glama.

**Caption:** camelids of Tripoli. The dromedaries are interesting in including white and piebald individuals (you might also be able to see the chest callous on the reclining camel in the middle). Images: Darren Naish.

Two cattle breeds were kept at the time of my visit (2009). One was Ankole, but I’ve been unable to identify the second. The adult females were short-horned, black and brown across the body, and white-muzzled, while two young males in the same enclosure were entirely black. I’ve wondered on and off over the years whether these were reconstituted Aurochs but that doesn’t seem to have been the case. Whatever they were, I’d like to know.

**Caption:** when you go to the zoo, you sure want to see some cows! Image: Darren Naish.

A group of Scimitar-horned oryx Oryx dammah included a youngster with short horns, so perhaps this was an indication that captive breeding had occurred. A large Blackbuck Antilope cervicapra group consisted of at least six males and numerous females (at least 12), and only one of the males was adult. This is the typical herd structure in this species: a single mature male typically leading the group. The substantial extent of the white on the male shows that these blackbuck were of the Rajputan subspecies A. c. rajputanae.

**Caption:** Blackbuck group, with the sole adult male adopting a straddle while peeing. I guess antelopes don’t like urine splashing on to their hooves, but then who does. Image: Darren Naish.

  **Caption:** Blackbuck females and non-adult males. A surprisingly large group. I haven’t yet seen this species in any other zoological collection. Image: Darren Naish.

A small group of Nilgai Boselaphus tragocamelus were present, as was a lone Brindled or Blue wildebeest Connochaetes taurinus. As you might know, the ‘Brindled gnu’ of tradition has recently been split into several species (I wrote about this at Tet Zoo ver 3 here). The overly dark grey colour, black beard and mostly dark forehead and muzzle of the Tripoli Zoo animal indicate that it’s probably a member of the southern African C. taurinus after all, but light areas around the eyes and a few other things leave me unsure. You might know that a good number of wildebeest in captivity are hybrids between members of the brindled wildebeest complex and the Black wildebeest C. gnou, though I’m not saying that that’s necessarily relevant here.

**Caption:** reclining wildebeest or gnu. I assume that live lawns are difficult to maintain on the zoo’s budget, hence the use across the enclosures of sandy substrates. This can be linked to insufficient hoof wear in certain hoofstock species. Images: Darren Naish.

The zoo also had Slender-horned, Sand, Rhim or Rheem gazelle Gazella leptoceros, a poorly known, particularly pale, mid-sized gazelle of the Sahara, one of two species included by some authors in the (probably not monophyletic) ‘subgenus’ Trachelocele. The other Trachelocele species is the Asian Goitered gazelle G. subgutturosa. The ‘rhim’/'rheem’ name comes from Hebrew for ‘wild ox’ (used in that part of the Bible that also discusses the leviathan) and has thus been argued to be inappropriate (Spinage 1986). A true desert specialist, the Slender-horned gazelle exhibits several features convergently present in some other desert antelopes (like Arabian oryx O. leucoryx): an unusually pale coat, enlarged hooves that help spread its weight on sand, and an ability to get most (or even all) of its water from plants and dew. As is the case with most other bovids from northern Africa, it has been strongly reduced in numbers by trophy hunters and is possibly endangered.

Aoudad aplenty. A big and pleasant surprise for me were the many Aoudad, Arrui or Barbary sheep Ammotragus lervia at the zoo. The Aoudad is a north African wild sheep (though read on), best known for its splendid mane, which is formed of long, straight hairs that grow from the underside of the neck and can reach the front hooves or even the ground. Aoudads are not sheep in the strict sense, since phylogenetic studies variously find them to be close to the base of the goat-sheep clade, or even close to Arabian tahr or part of the ovibovine goat-antelope clade (e.g., Lalueza-Fox et al. 2002, Calamari 2021).

**Caption:** an aoudad montage, deliberately included to show rams, ewes and kids… though these terms could be completely inappropriate given that aoudad are not really sheep. At left, the animal staring at the viewer is an adult male, and the animal looking in our direction in the image at right is an adult female. Images: Darren Naish.

Aoudads are light brown with a pale muzzle and possess ridged horns that curve outward and slightly backwards. The Aoudad of tradition is a polytypic species with several different forms, conventionally regarded as subspecies. Thanks to Ridgely S. Wren (who checked at the IUCN page), I know that the Tripoli Zoo animals are part of the locally occurring Libyan subspecies A. l. fassini, though both the Saharan aoudad A. l. sahariensis and Kordofan aoudad A. l. blainei apparently also occur in the south-east of the country. The nominate subspecies – the Atlas aoudad A. l. lervia – is an animal of Morocco, Algeria and Tunisia while the Egyptian aoudad A. l. ornata was thought extinct until evidence for its persistence was reported in 2002.

The zoo’s group included adult ewes, at least one ram, and some number of youngsters. At the time of my visit (October 2009), there were at least 17 individuals there. They were very personable and came right up to the walls and fences – sometimes standing on their back legs, frontlimbs on the wall – to interact.

**Caption:** suggest to an aoudad (or a group of them) that they might talk to you, and you can get images like this. Note the anteroposteriorly short hooves with steep anterior faces: these are hooves of a sort suited for agile movement on rocky surfaces with small areas of support. Image: Darren Naish.

Deer of Tripoli. Tripoli Zoo also had deer of at least two species during my visit. Chital, Axis or Spotted deer Axis axis have been (and likely still are) kept at various zoos in the UK but, despite that, I don’t recall seeing them before: Tripoli had a group of males and females together.

**Caption:** Chital group at Tripoli Zoo, featuring what I think are two males and four females. Chital are mid-sized cervine deer of the Indian subcontinent (occurring as far south as the Andaman and Nicobar Islands; they occur on Sri Lanka too), but they’ve been introduced to Australia, the USA and Croatia. Images: Darren Naish.

They also had a group of Fallow deer Dama dama, some of which were extremely dark. It’s often said that the coat variation present in Fallow deer is connected to a history of semi-domestication and the Tripoli Zoo animals certainly looked more variable than the wild Fallow I’m familiar with in the UK. Again, you’d need to know specific details of the history of these animals to know what’s going on here.

**Caption:** dark Fallow deer at Tripoli Zoo. The individual at left – which wasn’t the darkest one they had – might be described as semi-melanistic. Note the strong antler asymmetry that’s common in this species. Images: Darren Naish.

The grounds, and some final thoughts. And that about bring things to a close as goes discussion of Tripoli Zoo’s animals. As mentioned, the zoo was moderately spacious. I can’t pretend that the fencing or enclosure walls were attractive (the walls were drab concrete overall) but the grounds were as well planted as space allowed and the animals had sufficient cover. The toilets – housed in a temporary building of the sort you hire for festivals and the like – didn’t seem to have ever been cleaned at the time of my visit (note caveat), and a layer of mush and slurry on the floor meant that visiting was not an experience I’d want to repeat. I don’t recall there being a shop or a restaurant or anything like that, but that could be because my time was limited and I was only there to see the animals. The carpark was really big, presumably indicating high visitor count during at least part of the year (remember that I was visiting in October).

**Caption:** another of the bass relief artworks on the walls of the zoo, this one depicting monkeys (macaques of some sort, perhaps), lions, what might be a cheetah, and zebras. It looks like the artist made a mistake at some point and ended up turning some of the monkeys into lions, or vice versa. Image: Darren Naish.

Another thing I certainly remember about the zoo grounds is the presence of several bass relief pieces of art on the walls surrounding the zoo. I enjoyed these and was amused to see that they combined a number of prehistoric animal reconstructions – heavily inspired by the paintings of Zdeněk Burian – in addition to images of living species.

And that is where we end. I hope you enjoyed this review, and if you know more about Tripoli Zoo and the animals it has (or had), please don’t hesitate to add information in the comments. More reviews are set to appear in time, oh yes.

**Caption:** a mix of Permian and Mesozoic animals on another bass relief at Tripoli Zoo, with a 2009 Darren Naish for scale. The animals include representations of *Edaphosaurus*, *Stegosaurus* and *Moschops*. Not sure what the animal at upper left is though.

As ever, here are my entirely subjective scores…

Selection of species: 7 out of 10
Zoo nerd highlights: Black crowned crane, Slender-horned gazelle, Libyan aoudad
Quality of signage: 0 out of 10
Value for money: 7 out of 10 (but note caveats about zoo being renovated during 2009)
Overall worthiness: 7 out of 10

For previous articles in my zoo reviews series, and articles relevant to some of the topics touched on here, see…

Britain: wildlife theme-park, December 2007
Tet Zoo Reviews Zoos: Edinburgh Zoo, August 2016
Tet Zoo Reviews Zoos: Colchester Zoo, October 2017
TetZoo Reviews Zoos: the Isle of Wight Zoo, February 2020
TetZoo Reviews Zoos: ZSL Whipsnade Zoo, June 2021
TetZoo Reviews Zoos: Bristol Zoo, July 2021

If you like what I do and want to support it, please consider checking in at patreon - thanks!

Refs - -

Calamari, Z. T. 2021. Total evidence phylogenetic analysis supports new morphological synapomorphies for Bovidae (Mammalia, Artiodactyla). American Museum Novitates 3970, 1-38.

Lalueza-Fox, C., Shapiro, B., Bover, P., Alcover, J. A. & Bertranpetit, J. 2002. Molecular phylogeny and evolution of the extinct bovid Myotragus balearicus. Molecular Phylogenetics and Evolution 25, 501-510.

Spinage, C. A. 1986. The Natural History of Antelopes. Christopher Helm, London.

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After a 13-year hiatus, the long-awaited sequel to James Cameron’s 2009 Avatar is out…

**Caption:** among the many interesting animals of *Avatar: the Way of Water* is the gar-like skimwing. It goes without saying that all the images used in this article are (c) 20th Century Studios, many taken from the **Avatar Wiki**.

I and my family saw it last night. It’s Avatar: the Way of Water, and… my god, it’s visually amazing. James Cameron has made no secret of the fact that these movies are love-letters to the natural world, to environments and spectacular creatures, and to indigenous cultures and peoples. We – as in, all of us, whatever sort of life we lead – need to maintain our connection to the rest of the natural world, to preserve and maintain what remains, and to me it’s obvious that this is the primary theme of these movies. It feels childishly naive to even say it, but colonisation, industrialisation, destruction and exploitation are very much pitted as bad in the two Avatar films; those fighting to save and preserve the natural world and its indigenous peoples are the opposite.

**Caption:** it should be obvious to anyone who follows me on social media that I talk often about those remaining green spaces close to me. We all need to do what we can to see that these places persist. I feel as if almost everything is at threat of destruction or deterioration.

That as it may, I am of course here for the creature design (not for the plot or storyline). Visually, Avatar: TWOW is simply incredible: huge congrats to everyone involved for the awe-inspiring and beautiful animals and environments they designed and built. But, alas, I’m not about to publish an article on these new creatures, at least not today. Instead, I’m using the release of Avatar: TWOW as an excuse to dust off and republish the 2010 Tet Zoo ver 2 article I published on the animals of the first movie, since now is as good a time as any. The article originally appeared here (looks like the reader comments have been saved, which is good). I’ve made a few minor editorial tweaks and corrections, and have also added new images. But here we go…

Tone and I recently went to see Avatar. I’ve been reading up on the movie for months and was really looking forward to seeing it. As implied above, I’m not here to talk about storylines and plot devices… you want to know about the creatures. And a lot of thought and time obviously went into the design of Pandora’s ecosystem and creatures. In part, I’d say that this was a success: a lot of people (even many not that interested in the natural world) have been very much taken in by the movie’s xenobiology, and let’s hope that this inspires them to become interested in, and passionate about, the biology and ecology of the real world. Without further ado, here are assorted musings on Pandora’s creatures. And I’d be interested in your take on them too, so feel free to discuss things in the comments below.

I should add here that the original article included a spoiler warning at this point. Seeing as I’m writing here about a movie that came out 13 years ago, I’m assuming that I no longer need to include one though.

**Caption:** in case there’s any doubt, I had no involvement at all with *Avatar* or its creature design, nor do I have any reason to think that those behind the making of *Avatar: TWOW* are aware of my writings on the first film. I have, however, worked closely with several people connected to the film, in particular animation supervisor Andy Jones.

Banshees and other fliers. In my opinion, the coolest creatures in the movie are the flying beasts: the dragonesque Mountain banshee and the awesome, gigantic Great leonopteryx. Banshees are bluish-green, long-necked creatures with membranous wings and a pair of hindlimbs that also sport flaring membranes (there are two kinds of banshee, but we only get to know Mountain banshee in the movie). Unlike most Pandoran creatures they aren’t hexapodal, and are assumed to have lost one of the rear limb pairs.

A prominent alula-like clawed digit (flanged with another membrane) allows the animal to climb and cling; it looks similar to the large thumb present in megabats. The distal parts of the wings are translucent. The hindlimbs also have a prominent, alula-like, clawed digit. There’s a propatagium, but the main part of the wings are not composed of a single, continuous sheet (as they are in pterosaurs); instead, there are several rods embedded within the main membrane that help it to fold and fan out. The wings thus combine elements of the pterosaur wing with bird primary feathers and bat fingers.

**Caption:** banshees and Na’vi have a long history of association. Na’vi riders of the Omaticaya clan form lifelong bonds with Mountain banshees (known to them as ikran); the image here shows Naytiri’s ikran Seze.

When the animals fly, they stick the hindlimbs out sideways from the body; while the hindlimbs have membranes, these aren’t connected to the body or forelimbs. As a consequence, the flight configuration reminded me of certain pterosaur reconstructions, since some authors have also shown pterosaur hindlimbs projecting outwards and backwards. The banshee tail is long and slim, with a horizontal vane at the tip. The superficial similarity between Ikran and certain pterosaurs have not gone unnoticed by palaeontologists, as is demonstrated by the naming of the Chinese Cretaceous pterosaur Ikrandraco avatar Wang et al., 2020.

**Caption:** several very nice banshee figures are available. These images show the McFarlane Toys banshees, released in 2022 to coincide with *Avatar: the Way of Water*.

The team behind Avatar‘s creatures put some thought into stuff like respiration, aerodynamics and gravity. Gravity is lower on Pandora than on Earth (it’s a moon orbiting the gas giant Polyphemus), and this has facilitated the evolution of giant fliers. It’s also noted that at least some Pandoran creatures have stronger bones (or bone analogues) than what we’re used to, as a sort of carbon fibre reinforces the animal’s tissues, and also makes them lighter. Banshees also have ‘intake valves’ – spiracle-like openings on their thorax – that allow them to take in more air than allowed through the head alone, and they’re said to have a unidirectional respiratory system, with used gas passing outwards via gill-like slits on the posterior thorax. This sort of stuff is fairly plausible and we might honestly expect alien creatures to have respiratory systems like this, rather than the nostrils and whatnot that we’re more familiar with.

**Caption:** banshees recall the Cretaceous dromaeosaurid *Microraptor* in having flight surfaces on two limb sets, rather than one, and in having what might be a lift-creating structure at the end of the tail as well. Spectacular *Microraptor* specimens (like IVPP V13352, shown here at left) have been known since around 2002. At right, a scale model (showing one of several possible hindlimb poses) of *Microraptor* produced for a study on flight performance in this animal, from **Dyke *et al*. (2013)**. Images: Colin Palmer; **Dyke *et al*. (2013)**.

While the banshees look pterosaur-ish, I’m curious to know if their designers were also inspired by the crow-sized, four-winged dromaeosaurid dinosaur Microraptor. Microraptor – famous for having really long feathers on its hindlimbs as well as its arms – has been imagined by some to ‘flap’ with its legs as well as with its arms (this may not be at all possible, but let’s avoid that debate for now), and the double-winged flight motion of the banshees reminded me of this hypothetical flight style. There’s also a scene where the two banshees ridden by Jake and Neytiri glide rapidly downwards in parallel with a vertical cliff face. This reminded me of some of the stunts practised by people who use proximity gliding suits.

**Caption:** great raptorial hooks and recurved, slightly heterodont teeth (or tooth analogues) are among the obvious predaceous specialisation of the Toruk. As is the case in the Thanator and Viperwolf, the teeth are black, which suggests that they are not composed of enamel and other materials familiar on Earth.

Toruk. The Great leonopteryx – known to the Na’vi as Toruk – is neat, and the movie is worth seeing for this creature alone in my opinion. A giant predator of banshees and other animals, it’s a vividly coloured, spectacularly crested hexapodal eagle-like monster, with terminal hooks on its rostrum and a pair of grasping posterior limbs. Some sequences in the film remind me of real-world sequences depicting the behaviour of giant raptors, most especially the incredible South American Harpy eagle Harpia harpyja.

**Caption:** Toruks can open and close the slots between their leading wing elements, and rotate the individual elements as required for vortex formation and so on.

Unlike banshees, the Toruk retains all three limb pairs. Its wings are also more complex than those of banshees: they’re mostly membranous, but there are three feather-like elements along the wing’s leading edge that can separate and create leading-edge slots. Again, there’s an alula-like digit with a huge, curved claw. We see clearly that the digit is used to help support the animal’s weight when it rests, with wings folded, on the ground. Rather than being directed laterally, the digit projects directly forwards when the animal is grounded.

**Caption:** the Toruk is absurdly big, with a wingspan that can exceed 25 m. It preys on other predators (mostly banshees) and is a solitary, large-brained animal with a central role in Na’vi lore.

Dark lateral ridges project from the sides of its head, shading its eyes (like many of the Pandoran creatures, it has more than two eyes: there are two smaller eyes behind the main ones). These ridges make its head superficially raptor-like. Enormous, sheet-like blue sagittal crests project both from the dorsal surface of the head and from the lower jaw. To my eyes, these make the creature superficially similar to a tapejarid pterosaur, in particular to the sail-crested Tupandactylus (for a now very dated ver 2 article on tapejarids, go here).

So – even before I’d seen the movie – I’d decided that the banshees and Great leonopteryx were inspired by (1) microraptors, (2) tapejarids and (3) proximity gliding suits, plus with a bit of raptor and megabat thrown in too. While there might be some truth in this, the creatures are actually more complex, with inspiration for their design apparently drawn from even more diverse sources. Neville Page – mostly responsible for the creature’s design – has noted how he worked to emulate the smooth lines and streamlined shapes of creatures such as Great white sharks. It’s evident from the way their jaws open that banshees have some similarities with teleost fishes: when the animals gape, a maxilla analogue is pulled downwards from the upper jaw. A neat detail is that the entire tooth row in the upper jaw is mobile: when the jaws are closed, the teeth fold posteriorly into slots on the ventral surface of the maxilla analogue, but they swing anteroventrally when the jaws are open.

**Caption:** at left, a proximity gliding suit or wingsuit. Certain of the scenes in *Avatar* are reminiscent of wingsuit flight sequences. At right: the amazingly elaborate skull of the tapejarid pterosaur *Tupandactylus imperator*, showing the extent of the soft tissue and the associated branched, filamentous structures on the rear spar. Images: Wingsuiting, CC BY-SA 4.0 (from **here**); Cincotta *et al*. (2022), CC BY 4.0 (from **here**).

Also on the teleost-like anatomy, we have good views in the movie of a Great leonopteryx skull (the Omaticaya clan of the Na’vi keep one in their giant hometree for symbolic or ceremonial reasons: the creature is important in their mythology). It doesn’t look tetrapod-like, but has the corrugated, gnarly bone texture you normally associate with fish. Having said that, there are some bird-like aspects to the skull as well. The rostral hooks recall the hooked tips of raptor bills, for example. Some articles state that manta rays, skates and plesiosaurs were also inspirational in the design of these creatures, and Wayne Barlowe (who was involved in creature design early on) is on record as saying that he based the banshee’s sleek design on sports cars. The patterns and colours of these animals were apparently inspired by those of birds, poison-dart frogs and monarch butterflies.

Hammerhead titanothere. Pandora is home to at least one mega-herbivore, the spectacular, enormous Hammerhead titanothere. The fact that it’s called a titanothere might explain the origin of its design. Titanotheres, generally called brontotheres (or brontotheriids) these days, are rhino-like Eocene perissodactyls: they’ve been discussed on Tet Zoo a few times. Go here, here and here.

**Caption:** the giant brontothere *Megacerops* (previously known as *Brontotherium*, *Menodus*, *Brontops* or *Titanotherium*) is diverse in horn form, this varying according to age and sex. I like this image (taken at Badlands National Park) because it shows an especially big-horned individual. Image: Dave Fuentes, **from here**.

The hammerhead-like, err, head is a nice idea, but – for my tastes – too ‘familiar’ given that we all recognise this shape. They could have gone for the same idea (after all, a battering-ram head could well be useful to a giant terrestrial herbivore: note that the Hammerhead titanothere does not have eyes on the lateral ends of its ‘hammer’), but made it far freakier and more alien. The animal uses a fan-like cluster of flag-like structures on the top of its head as a signalling device. I liked the fact that (ordinary) guns wouldn’t be much use when confronting an animal this large and formidable: shoot it, and it will still be coming right for you. The stampede scene was awesome and very satisfying.

**Caption:** Hammerhead titanothere from the encounter scene with Jake. Note that the eyes are present posteroventral to the hammer-like cranial structure. Titanotheres are social animals, adults varying in the colour of their fan-like cranial structures. They reach 11 m in length.

Thanator: super-predator. The Thanator – a large hexapodal terrestrial predator – is another of my favourites from the movie. I initially thought from the trailers that its designers had been looking at gorgonopsians, but, nope, no good reason for thinking this. Cameron has said that it was meant to be a sort of super-panther.

**Caption:** Thanator in lateral view, with retracted lips and elevated soft cranial flaps. The opposable nature of the digits on the anterior limbs is obvious here, as is their heavily muscled form.

It’s dark, with smooth, mostly leathery skin, and dextrous muscular limbs with long, sharp claws borne on semi-opposable digits. Its head is long with a sort of naked, protruding rostrum and large anterior fangs. Its eyes are large and it has peculiar flap-like and soft, spike-like structures arranged around the back of its head. Not sure what these are for, though they might be used in visual display. In one scene, it grabs Jake’s back-pack. Jake slips out of the back-pack and escapes, and when the Thanator notices, it opens it jaws, raises its paddle-shaped tail, and flares the flaps and spikes outwards. That’s some nice body language there, implying intelligence and mood.

**Caption:** a Thanator encounters Naytiri, adopts a conciliatory pose, and invites her to ride it into battle. Alas, this doesn’t end well for the Thanator.

While I think that the Thanator is a very neat looking creature, it does exhibit ‘Hollywood super-predator syndrome’ to a degree, and acts like an unstoppable, psychotic whirlwind, smashing through vegetation, tearing tree roots up, and altogether doing everything possible in order to kill and eat the object of its attention. It’s not even deterred by a barrage of automatic gunfire, and almost pursues Jake right off the end of a cliff. It also appears to be incredibly intelligent, grabbing guns and discarding them during combat, for example. We all like predators in movies to be super-predators with super-powers, and on the one hand you might argue that creatures like the Thanator are so over the top that they really create the wrong impression as to what real predators are (mostly) like. One day I want to see a movie where the predator is a conservative coward that faints when confronted with a gun.

On the other hand, we know for sure that certain predators can be unbelievably gung-ho and bold on occasion, Exhibit A being the various tales that involve man-killing big cats that have moved into dwellings and vehicles to kill people. And I should add that I only recently read Stephen Herrero’s 1985 Bear Attacks: Their Causes and Avoidance. Oh. My. God…

**Caption:** I really like this poseable Thanator figure, made by Mattel in 2011. I’d love to own one but have never seen it for sale. I’m not sure it was ever available here in the UK.

I do dislike one other thing about the Thanator: its roars sound exactly the same as the tyrannosaurs in the Jurassic Park movies. Having said all this, it’s an awesome creature and its two major appearances in the film are among the highlights.

Direhorses and viperwolves: not my favourites. As for some of the other creatures, I confess that I found them to be less interesting: they were just too similar in form or behaviour to real animals, or, in other words, too derivative. I refer in particular to direhorses and viperwolves. I didn’t like the Direhorse because it just seemed all too much like, well, a horse, though a very big, alien-ish horse… though still a horse.

**Caption:** direhorses can bond with Na’vi through integration of their neural whips. This image is from a battle scene where a Na’vi cavalry battles a heavily armed RDA squadron involving infantry augmented with an AMP (amplified mobility platform) unit.

To their credit, Cameron et al. did try and make the animal fit in with the rest of Pandora: direhorses have the same thorax spiracles as the banshees, and – in keeping with the planet’s riotous and super-elaborate flora – we see in the movie that direhorses aren’t grazers; instead, they’re nectarivorous, and feed with a long proboscis from flowers. Needless to say, the evolution of gigantic nectar-feeders like this could only work in an ecosystem where flowers are enormous, permanent, and produce huge quantities of nectar.

**Caption:** it should be obvious that the direhorse face superficially recalls that of a giant anteater. Horizontal pupils on the largest eye set are obvious, as is an anterodorsally projecting cranial crest (continuous with a tall ridge present along the midline of the neck) and the two parallel neural whips.

The viperwolves were the least compelling creatures for me. They’re long-bodied, slinky, black-skinned hexapods with hand-like feet and facial tissue that can be retracted right off their scary looking, pointed teeth. They have floppy neural whips at the back of the head. We see a viperwolf group behave in an extremely tenacious way when Jake is alone in the forest for the first time; despite his use of fire and other aggressive tactics, he’s not succeeding in preventing their attack until Neytiri comes to his rescue. I did initially think that this was unrealistic, but I now think that this appraisal is unfair given how social canids will behave when harassing animals the size of people and large deer. Whatever, I didn’t especially like the design. They remind me of small, black versions of Falkor the luck dragon in The Neverending Story.

**Caption:** viperwolves are sharp-toothed, lithely built predators that share highly mobile lips, large and pointed teeth (or tooth homologues) and dextrous digits on their anterior limbs with thanators. Viperwolves are co-operating, pack-hunting predators.

The Na’vi and others. Finally, the creatures that feature most strongly in the movie are of course the blue humanoid Na’vi. In many ways these are the easiest of the creatures to criticise, if – that is – you’re like me and think that the odds of human-like creatures evolving independently of us are vanishingly small and downright improbable. On the one hand, you can argue that successful alien films can work fine when the creatures don’t look at all human. And – given that humans are meant to be remotely piloting genetically modified Na’vi bodies – it wouldn’t have mattered to the story what the Na’vi looked like.

**Caption:** a scene showing several of the main Na’vi characters. It’s hard to make people care about characters in a film where those characters aren’t relatable.

On the other hand, this movie is about warfare between cultures, about allegiances, and – I suppose – about cool-looking shit, so it figures that good looking ‘people’ need to feature large in the story. Originally, the Na’vi were going to look weirder, with gills, fins and other structures, but over time they were made to look more human simply to appeal more to the audience. Female Na’vi have breasts, specifically for this reason, apparently. So, yeah, the Na’vi are nothing more than attractive, semi-naked blue people with cat-like features… let’s let it go. When we’re introduced to their spiritual beliefs and practises, the Na’vi incorporate details inspired by or based on those of various indigenous groups of people, and it’s not really possible to think about the plot without comparing it to the conflicts that have occurred between invading European cultures and indigenous ones.

The idea in the movie is that Pandora includes a lineage of primate-like animals whose evolution has closely paralleled primate evolution on Earth. Early in the movie, we see the hexapodal, arboreal Prolemuris (they also make a guest appearance in Avatar: TWOW). This animal is to the Na’vi what lemurs are to us (distant cousins that share a common ancestor), and this explains why its two more anterior limb pairs are partially fused. Presumably, the arms of the four-limbed Na’vi represent two, fully fused original limb pairs (though, if this is true, it might be odd that the Na’vi have only four fingers).

**Caption:** Prolemuris is a social, arboreal, primate-like Pandoran creature. Unlike the majority of Pandoran animals, they only have a single pair of eyes and only one neural whip. Both features are shared with the Na’vi.

Several other creatures feature in the movie. The hexapede is a deer-like animal with a fan-like cranial structure, while we also see a small, arboreal hexapod that looks like a cross between a leaf-tailed gecko and a frog. When disturbed, it unfolds a giant glowing spiral-shaped structure on its back and takes flight while spinning and emitting bioluminescence. Bioluminescence is a major theme on Pandora, with virtually all of the plants glowing in the dark.

A linkage of some sort unites most (or all?) living things on Pandora, and this proves key to the victory of the Na’vi over the invading humans. The Na’vi ‘store’ their cultural heritage in the memory of special trees, and it seems that memories and even personalities of individuals can be uploaded and downloaded via biological links with these plants. The Na’vi also have neural links with the creatures they ride (direhorses and banshees) and have to plug into them via a tentacle-like structure (termed the queue) on their heads. I don’t want to over-analyse any of this, but the link between the Pandoran creatures and the trees reminded me of the mycorrhizal networks known to involve certain trees and nearby fungi. I wonder if this is what inspired the whole idea?

**Caption:** mycorhizzal fungi have a mutualistic relationship with plants via a subterranean network. Inter-plant chemical messages relating to predation, disease and perhaps environmental stress are sent across the network. Image: Charlotte Roy, Salsero35, Nefronus, CC BY-SA (**original here**).

Finally, I loved the technology featured in the movie. Avatar is set in 2154 and I think the tech looks plausible based on the way things are going (yeah, if our culture lasts that long). The aircraft looked and behaved plausibly. The fat, heavily armed C-21 Dragon Assault Ship has an aesthetic charm, and the gigantic Valkyrie reminded me a lot of the Hercules and other bombers it was probably inspired by. The AMP suits look like evolved, combat versions of the cargo-loaders we know and love from Aliens. Though, I think we can doubt that an exosuit would be kitted out with a giant KNIFE.

**Caption:** Ellen Ripley using a P5000 load-lifter, a scene from *Aliens*, of course. *Aliens* and *Avatar* almost certainly occur in the same universe… though I don’t know if this has been specifically confirmed.

The Avatar movies really are the visual feasts that we’ve been promised. Both the 2009 original and the 2022 sequel look great, and the cool animals – the Mountain banshee, Great leonopteryx, Thanator and so on – make them well worth a viewing, or two. I ended up seeing the first film several times in the cinema. Avatar: the Way of Water is brand-new at the time of writing (December 2022) and my first and only viewing is still fresh in my mind. Additional movies in the series are due to appear over coming years, meaning that the series will be a mainstay in discussions about creature design and speculative biology for some time yet.

That’s where we’ll end things for now. Say what you like about the plots of these films… I cannot help but like them in view of the stupendous creature design they include, and their emphasis of our empathy with the natural world.

**Caption:** we must do all we can to prevent the destruction and deterioration of the natural world. We need it as much as all the other animals do.

The 2022 movie has a website here, and the website for the entire franchise – merch, extra materials, behind the scenes info and more – is here.

For previous Tet Zoo articles on speculative zoology and aliens and so on, see…

The science of Godzilla, February 2007
Speculative Zoology: Wedel throws down the gauntlet, February 2007
Oh no, not another giant predatory flightless bat from the future, March 2007
How intelligent dinosaurs conquered the world, March 2008
Alien para-tetrapods of Snaiad, April 2009
Richard Dawkins and the crappy ‘humanoid dinosaurs’ that just won’t die, November 2009
Dinosauroids revisited, revisited, October 2012
Of After Man, The New Dinosaurs and Greenworld: an interview with Dougal Dixon, April 2014
Speculative Zoology at Tet Zoo, The Story So Far, May 2015
Speculative Zoology, a Discussion, June 2018
Could We Domesticate (Non-Bird) Dinosaurs?, August 2018
The Dougal Dixon After Man Event of September 2018, September 2018
TetZoo Bookshelf, February 2019, Part 1, February 2019
Alternative Timeline Dinosaurs, the View From 2019 (Part 1), December 2019
Alternative Timeline Dinosaurs, the View From 2019 (Part 2), December 2019

Refs - -

Dyke, G., de Kat, R., Palmer, C., van der Kindere, J., Naish, D. & Ganapathisubramani, B. 2013. Aerodynamic performance of the feathered dinosaur Microraptor and the evolution of feathered flight. Nature Communications 4, Article number: 2489 doi:10.1038/ncomms3489

Herrero, S. 1985. Bear Attacks: Their Causes and Avoidance. Nick Lyons Books, New York.

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Have we discovered one of vertebrate palaeontology’s most famous ‘missing’ specimens? Well, read on…

If you’ve ever looked at a book or article that discusses the early history of dinosaur discoveries, chances are high that you’ve read about a large partial bone documented in 1677 by Dr Robert Plot (1640-1696) in his *The Natural History of Oxford-shire*. Plot – a naturalist, antiquarian and scientist – was both Professor of ‘Chymistry’ at Oxford University *and* first Keeper of the Ashmolean Museum in Oxford. In fact, he was the only person to ever hold both positions at the same time. He apparently did this as neither one paid well enough to provide a living wage… *insert here appropriate quip about scientific and museum salaries*.

**Caption:** at left, the best known portrait of Robert Plot of Oxford, naturalist, antiquarian and scientist. At right, his famed book of 1677.

It was in his aforementioned book that Plot chronicled various geological oddities discovered in the Oxfordshire countryside. Plot’s views on the origins and identities of these objects have to be seen within the context of the time: fossils and fossilisation were not understood, and it was suspected that objects dug from the earth were formed naturally by some sort of generative process, a “plastic virtue, latent in the earth” (Plot 1677, p. 112).

Our focus here is on a large, incomplete bone that Plot discussed and illustrated, given to him by “the ingenious Sir Thomas Pennyston”. It has been discovered in a quarry in the parish of Cornwell, Oxfordshire (not to be confused with Cornwall in England’s south-west). The specimen’s exact geological provenance isn’t known but it’s thought that it came from the Middle Jurassic (Bajocian or Bathonian) oolitic limestone of the region (Delair & Sarjeant 2002). Plot recognised – correctly – that the specimen looked like the articular end of a femur, or thigh bone, and wrote that it “has exactly the figure of the lowermost part of the thigh-bone of a man, or at least of some other animal” (Plot 1677, p. 132) before describing its detailed anatomy.

**Caption:** at left, the relevant plate from Plot’s 1677 book *The Natural History of Oxford-shire*. The famed specimen is shown alongside objects discovered in the ground, and thought to represent a petrified heart and human foot, as well as assorted rhomboidal and trapezoidal objects.

Because the bone was larger than that of a horse or ox, Plot wondered if it might have come from an elephant “brought hither during the Government of the Romans in Britain” (Plot 1677, p. 133), but noted that “this opinion too lies under so great difficulties, that it can hardly be admitted”. He was in fact sufficiently sceptical that he wanted to see this idea tested. Remarkably, he managed to examine actual elephant bones in 1676 and found them very different in shape and size from the Cornwell bone.

Plot was left with no option but to conclude that the bone was from a giant person, and he devoted a few pages of his text to chronicling those giant men and women discussed in antiquarian sources (Plot 1677, pp. 136-138). It’s impossible today to appreciate how Plot and other people of his time might have regarded the discovery of physical evidence for the existence of giant humans, some of whom were considered by Plot to be 17 ft tall (Plot 1677, p. 137). It requires a worldview we just can’t appreciate.

**Caption:** it’s been generally agreed for decades now that the Plot bone likely belonged to *Megalosaurus*, the large tetanuran theropod famously associated with the Middle Jurassic sediments of Oxfordshire. Whether this is justifiable is a complex issue, but I’ve essentially followed that assumption here. *Megalosaurus* skeletal reconstruction by Dan Folkes, used with permission.

Brookes and ‘Scrotum Humanum’. The Cornwell bone was written about again by Richard Brookes in his 1763 book The Natural History of Waters, Earths, Stones, Fossils, and Minerals, with their Virtues, Properties, and Medicinal Uses, to which is added the Methods by which Linnaeus has treated these subjects, Vol. 5 of A New and Accurate System of Natural History (Brookes 1763). Brookes’s writings and illustrations of fossils and supposed fossils were very similar to those of Plot and evidently based on them. He depicted the Cornwell bone again, albeit in more schematic fashion and inverted relative to the figure provided by Plot.

Brookes’s (1763) text makes it clear that, like Plot, he regarded the specimen as “the lowermost part of the thigh-bone of a man” (Brookes 1763, p. 317) but his plate (published opposite p. 318 of his book) labels it not as the lower-most part of a thigh bone, but as ‘Scrotum Humanum’. Why the book’s text and plate provide different, competing identifications isn’t obvious, but they do.

**Caption:** plate from Brookes (1763) showing the bone and other objects. It’s obvious that several of the objects illustrated here were redrawn directly from Plot’s plate of 1677. Note that the label ‘Scrotum Humanum’ is written in the same style as such obvious labels as ‘Kidney Stone’ and ‘Petrifaction’.

And here’s the origin of the idea that the object was misidentified as the scrotum of a giant human, and that the bone was awarded the scientific name Scrotum humanum (Halstead 1970, Delair & Sarjeant 1975, Norman 1992, Torrens 1995, Spalding & Sarjeant 2012).

At least a few writers after Brookes took this idea seriously, the French philosopher Jean-Baptiste Robinet going so far as to claim – in 1768 – that the specimen preserved visible traces of musculature and even a urethra (Buffetaut 1979). The late palaeontologist Beverley Halstead argued – apparently seriously – that Scrotum humanum should stand as the first technical name ever attached to a (non-bird) dinosaur, and that it should be considered technically admissible seeing as its publication post-dated the 1758 appearance of the 10th edition of Carl Linnaeus’s Systema Naturae (Halstead 1970), the work responsible for establishment of the binomial system. Halstead died in 1993 but his colleague William Sarjeant (who himself died in 2002) apparently tried petitioning the ICZN (International Commission on Zoological Nomenclature) to have the ‘name’ technically supressed so that it wouldn’t usurp those names it might be considered synonymous with, namely Megalosaurus bucklandii (Halstead & Sarjeant 1993). Spalding & Sarjeant (2012) noted that this “proposition has now been firmly rejected by the cognoscenti of zoological taxonomy” (p. 10).

**Caption:** the *Megalosaurus* display at Oxford University Museum of Natural History (photographed in 2016), showing most of the original bones as described by Buckland in 1824. Note that an illustration of the Plot bone occupies the centre. Image: Darren Naish.

This whole saga always was a silly waste of time and I don’t think Halstead was being serious in the first place. ‘Scrotum Humanum’ is transparently a label on the relevant plate, not an attempt to attach a scientific name to a species. It’s written alongside such other labels as ‘Kidney Stone’ and ‘Petrifaction’.

Where is the Plot bone today? That, then, is the required preamble on the specimen, which I’ll call ‘the Plot bone’ from hereon (the term ‘Cornwell bone’ has been used here and there too). Whatever became of the Plot bone? Where is it today? A certain cartoon notwithstanding (see below), popular history has it that the specimen is lost: that we simply have no idea, today, what happened to it or where it might be.

**Caption:** an illustration reproduced by Torrens (1995) wherein the caption reads “Little girl and *Scrotum humanum*, from the title page of *Dinosaur Bones* by ‘Aliki’, London : A. & C. Black, 1989”.

Over recent years I’ve worked closely (due to my role as scientific consultant for the BBC Studios/Apple TV+ series Prehistoric Planet) with Paul Stewart. Paul is a qualified zoologist (his PhD was on the behaviour of European badgers Meles meles, completed under Professor David Macdonald at the University of Oxford) and has a serious interest in palaeontological history, especially in early ideas about dinosaurs, marine reptiles and Pleistocene mammals.

**Caption:** Dr Paul Stewart and a taxiderm specimen of a European badger at the Oxford University Museum of Natural History. Image: Darren Naish.

The Plot bone found, perhaps. While working with Paul one day, he dropped a bit of a bombshell. He’d noticed that what appeared to be the Plot bone was very much still in existence, and – furthermore – was on show in a museum; specifically, the Ashmolean Museum in Oxford. The Ashmolean is devoted to art and archaeology so it’s conceivable, I suppose, that a specimen as famous as the Plot bone might be on show there but not noticed by the staff (few of which are experts on fossils) nor by visitors. The specimen concerned did not belong to the Ashmolean but to the Oxford University Museum of Natural History, where it’s accessioned as OUMNH PAL J29757. For ease of reading, I’ll call it the ‘Ashmolean bone’ from hereon.

**Caption:** OUMNH PAL J29757, as on display at the Ashmolean Museum in February 2020. At left, the specimen in posterior view. At right, anterior view. Images: Darren Naish.

The Ashmolean bone is obviously the distal end of a dinosaur femur, its prominent condyles and other structures showing that it belongs to a tetanuran theropod dinosaur, and perhaps to Megalosaurus or a close relative. Cracks and areas of damage much resemble features visible in Plot’s original illustration but it was obviously different from the Plot bone in that the hollow interior wasn’t visible. We reasoned that a section of shaft had been glued on to the distal, condyle-bearing end: a crack separating the two was in exactly the right position to make the distal end look like the Plot bone if imagined on its own, and the label on the shaft section suggested a history distinct from that of the distal end. Maybe, we surmised, they’d been discovered at different times and only connected sometime after Plot and Brookes had written about the specimen.

Also relevant is that Plot’s illustration of the specimen was a mirror-image of the real thing, meaning that direct comparison between it and the Ashmolean bone made them look quite different. If we flip Plot’s illustration back to how it ‘should’ look, we see some points of similarity between it and the Ashmolean bone.

**Caption:** a reasonable list of detailed similarities exist between the Ashmolean bone (at left) and the Plot bone (here, flipped back to its original orientation), enough that we became quite confident that the two might be the same thing. If my interpretation of anatomical features is correct, both illustrations here depict the distal end of a right femur.

Antique labelling reveal that the Ashmolean bone originated at Enstone, which is encouraging because this is the former location of a quarry within the parish of Cornwell (which, you’ll recall, is where the Plot bone came from).

Paul seems moderately confident, but I wanted another opinion and so put all of this to Martin Simpson. Martin and I have worked together on Cretaceous pterosaurs and dinosaurs and he’s especially interested in historical palaeontology, owning a substantial amount of relevant material (original publications and so on) himself. By comparing the various cracks, areas of damage and detailed anatomical nuances of the Ashmolean bone with illustrations of the Plot bone, Martin also became confident that the two were indeed a good match. Between the three of us, we began working on a plan to write this up as a technical paper. If we were right, this was a pretty big deal: one of vertebrate palaeontology’s most famous, or infamous, early specimens wasn’t lost, but had been successful relocated.

**Caption:** OUMNH PAL J29757 within the Robert Plot display at the Ashmolean Museum in February 2020. Plot also obtained and wrote about crinoids, bivalves, brachiopods, ammonites and corals. The fossil coral immediately to the left of the theropod femur belongs to a sort suggested by Plot to be identifiable as petrified human hearts. Images: Darren Naish.

Paul, Martin and I met up at the Ashmolean early in 2020 and examined the bone as best as we could. It was (and currently still is) part of a display devoted to the museum’s early history and Plot’s role specifically. It ‘stands in’ for the Plot bone and there’s no clear indication from the labelling that that’s not what it is.

**Caption:** part of the Plot display at the Ashmolean Museum (in February 2020), with three of the primary investigators staring at it in loving fashion. Darren Naish at far left; Martin Simpson standing at right; Paul Stewart at lower right.

Hands-on with the Ashmolean bone. What tests might be possible in order that our suggested identification could be checked? Our knowledge of the Plot bone doesn’t just concern its external appearance: we also know something about its size. Plot (1677) wrote that it was about 15 inches in circumference around the shaft, about 24 inches in circumference around the bulbous distal end, and about 20 lbs in weight. Plot (1677) also provided details on its interior, since its hollow shaft was filled at its bottom with a sparry mineral, supposed by Plot to perhaps be petrified marrow. In order to check these things, we’d need to examine the specimen outside of a display case, and measure and weigh it appropriately.

We approached Hilary Ketchum, Collections Manager at the Oxford University Museum of Natural History. Hilary might not work at the Ashmolean, but her contacts in the Oxford museum world would be essential and she’d know how best to get access to the specimen in person. And she proved so important that we wanted her as part of the team.

Getting access to museum specimens that are on display can be complicated and slow, since it takes a while for the right people to be available at the right time. The Ashmolean is also an exceptionally busy museum, and we were requesting special access to a specimen right at the height of the covid pandemic. We got the ball rolling in February 2020, but any and all plans were thwarted as the museum effectively became closed to research visits for the rest of the year. Thanks to organisation work from Hilary and co-operation from collections staff at the Ashmolean, we finally got the required access to the specimen in December 2021.

**Caption:** OUMNH PAL J29757 on the day of reckoning, the images again emphasising how compressed it is at its preserved proximal end. Images: Paul Stewart.

Denouement. On the day of reckoning, Paul and Hilary met at the Ashmolean to examine, weigh and measure the bone in person. It proved to be a pretty good match for Plot’s bone in rough dimensions. But here’s where things essentially fell apart. The Ashmolean bone is a veritable lightweight relative to the Plot bone, weighing about half of what it ‘should’.

Could there still be an interior hollow filled with a sparry mineral ‘marrow’, as Plot had described in 1677? Well, maybe, but there would be no point in breaking the Ashmolean bone open to find out in view of the very different measurements already evident. The conclusion had to be that the specimen was not the Plot bone after all. And at this point our hope that we might have discovered the original Plot bone were dashed. We hadn’t. It remains of unknown whereabouts.

**Caption:** OUMNH PAL J29757 again on the day of reckoning, this time with Hilary’s hands providing useful indications of scale. From left to right, we see the specimen in posterior view, medial view, and proximal view. The size means that this can’t be the Plot bone after all. Images: Paul Stewart.

Most scientific stories that involve the suspected discovery of a given thing – whether it’s a new animal species, a fossil or a biological phenomenon or event – end with the thing being found: with positive results. But a major part of the scientific process involves negative results; we often fail to find or confirm what we hoped we would, an outcome that feels like failure.

However, scientific results are results however they pan out, and failing to confirm something, or confirming that something is not the thing you hoped or suspected it was, is still important. For that reason, we can feel good about the fact that we investigated a specimen and found that it was not the one we hoped it would be, even though that doesn’t result in the high-profile publication and undoubted fame and fortune we were aiming for. I kid, of course. But here ends our story.

**Caption:** the Plot bone - a mainstay of popular and academic books on dinosaurs - remains lost after all. This montage shows just a few of the many appearances the bone has enjoyed in the literature.

For previous Tet Zoo articles on historical palaeontology, see…

The Crystal Palace monsters, armoured tyrannosaurs and lurking sauropods: a look back at ‘Dinosaurs – A Historical Perspective’ (part I), May 2008
The Dinosaurs of Crystal Palace: Among the Most Accurate Renditions of Prehistoric Life Ever Made, August 2016
Up Close and Personal With the Crystal Palace Dinosaurs, December 2018

Refs - -

Brookes, R. 1763. The Natural History of Waters, Earths, Stones, Fossils, and Minerals, With Their Virtues, Properties and Medicinal Uses, To Which Is Added, the Method in Which Linnaeus Has Treated These Subjects, Vol. 5 of A New and Accurate System of Natural History. Newbery, London.

Buffetaut, E. 1979. A propos du reste de dinosaurien le plus anciennement décrit: l’interprétation de J. B. Robinet (1768). Histoire et Nature 14, 79-84.

Delair, J. B. & Sarjeant, W. A. S. 1975. The earliest discoveries of dinosaurs. Isis 66, 5-25.

Delair, J. B. & Sarjeant, W. A. S. 2002. The earliest discoveries of dinosaurs: the records re-examined. Proceedings of the Geologists’ Association 113, 185-197.

Halstead, L. B. 1970. Scrotum humanum Brookes 1763 – the first named dinosaur. Journal of Insignificant Research 5, 14-15.

Halstead, L. B. & Sarjeant, W. A. S. 1993. Scrotum humanum Brookes – the earliest name for a dinosaur. Modern Geology 18, 221-224.

Norman, D. B. 1992. Dinosaurs past and present. Journal of Zoology 228, 173-181.

Plot, R. 1677. The Natural History of Oxfordshire, being an Essay toward the Natural History of England. Robert Plot, Oxford.

Spalding, D. A. E. & Sarjeant, W. A. S. 2012. Dinosaurs: the earliest discoveries. In Brett-Surman, M. K., Holtz, T. R. & Farlow, J. O. (eds) The Complete Dinosaur (Second Edition). Indiana University Press (Bloomington & Indianapolis), pp. 3-23.

Torrens, H. S. 1995. The dinosaurs and dinomania over 150 years. In Sarjeant, W. A. S. (ed) Vertebrate Fossils and the Evolution of Scientific Concepts. Gordon and Breach Publishers, pp. 255-284.

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TetZooCon 2022 just happened. How did it go? Well, it went pretty well is how it went…

**Caption:** at upper left, a *Prehistoric Planet*-style *Mononykus* plush by Leo; at lower left, an origami *Triceratops* by Joel Colbourne; TetZooCon badges in the middle; and our amazing venue – Bush House – at far right. Images by Darren Naish.

I’ll start by staying that our venue – Bush House, King’s College, The Strand, London – was amazing and just about perfect for our needs. The auditorium was spacious and modern (it even had balconies and a green room) with good acoustics, the reception area was good and appropriately nearby, and there were convenient amenities within very short walking distance. Enormous thanks to Chris Manias of King’s College for working with us in setting up Bush House as a venue: excellent work, and it looks like we’ll be there again in 2023. Before proceeding, I also need to mention that we have an official photographer at TetZooCon: Georgia Witton-Maclean. Huge thanks to her for the photos and for passing them to me in good time.

**Caption:** our banner this year was squamate-themed… for no particular reason at all.

Platypus Matters and Locked in Time. Our opening talk was one of my favourites: Jack Ashby’s Platypus Matters: the Extraordinary Story of Australian Mammals. Jack reviewed the history of platypus science and knowledge before surveying marsupial diversity and some of their remarkable specialisations. Emphasised throughout was the fact that these animals – monotremes and marsupials – shouldn’t be assumed to be, or be framed as, weirdos or primitives, as is tradition within the Eurocentric world. Jack’s talk was accompanied by the selling and signing of his excellent new book: Platypus Matters (Ashby 2022).

**Caption:** Jack Ashby presents *Platypus Matters*, also the name of his new book. Image: Georgia Witton-Maclean.

Jack was followed by Dean Lomax on Locked in Time: Unearthing the Behaviours of Prehistoric Animals, another talk connected to a new book (Lomax 2021). Dean explained his own background and his involvement in research on extraordinary fossils, some of which depict ‘frozen behaviour’ of the sort described in his talk and book. Amazing stuff: think mammoths locked together in combat (I wrote about them back in 2008), fighting dinosaurs, ichthyosaurs giving birth. An amusing connection with Jack’s talk – and Dean drew attention to this – is that both of them used the mass annual death of male antechinuses as an example of an extreme behaviour. Dean’s talk was also followed up by a selling and signing event.

**Caption:** the tangled mammoths. Amazingly, this association has not been properly described in the academic literature (come on, mammoth experts: *what are you doing with your time?*). At right, Dean Lomax’s excellent new book *Locked in Time*, featuring art by Bob Nicholls.

A third speaker of that planned opening session (Hana Ayoob) was unfortunately unable to join us; hopefully we’ll be able to host her in the near future.

All Yesterdays, ten years on. 2022 marks ten years since the publication of the palaeoart-themed book All Yesterdays. Accordingly, it seemed appropriate to have an All Yesterdays event at TetZooCon 2022 whereby the book’s three architects – myself, John Conway and C. M. Kösemen – discussed aspects of its creation and formation on stage. Maybe it was a little less structured that it should have been, but it seemed to be enjoyed, and provided a bit of context and background to the project. One takehome is that its completion was rushed and involved a lot of provisional decisions and short-cuts.

**Caption:** at left, Memo Kösemen at his TetZooCon 2022 stall, with art prints (and originals) for sale. Note the *All Yesterdays* and Squamozoic art (and more) stuck on the column. At right, Naish, Kösemen and Conway on stage during the *All Yesterdays* event. Images: Georgia Witton-Maclean.

I kicked things off by running through – at speed – the talk I delivered at the original launch event of 2012. We had copies of All Yesterdays on sale at TetZooCon 2022 and at least a few people got all three of us to sign them; Memo also sold – yes, sold – a fair amount of original All Yesterdays artwork. I would have bought some but never had time to sort it out.

**Caption:** Memo Kösemen signing and selling art at his TetZooCon 2022 stall. Image: Georgia Witton-Maclean.

Palaeoart Workshop. Next up, our parallel session began. One group of people attended the Palaeoart Workshop, and another a talk session. I didn’t attend the Palaeoart Workshop so I don’t know exactly what happened, but I do know from photos I’ve seen that a whole lot of art was produced. The workshop this year incorporated a meeting of Chris Manias’s Popularising Palaeontology event, the result being a discussion of exhibition design. Led by John Conway, the group also had fun in depicting the radically unusual Triassic reptile Tanystropheus in innovative ways.

**Caption:** a table-top scene from the TetZooCon 2022 palaeoart workshop. Image: Georgia Witton-Maclean.

Cockatoos and other theropods, and the British herpetofauna. The talks happening at the same time kicked off with Could Ancient Theropods Use Tools? from Jennifer Colbourne. Jennifer’s PhD is mostly on cockatoos and their tool-using and problem-solving abilities, and wow it’s interesting. Does Jennifer think that Mesozoic dinosaurs could have used tools? She focused on troodontids in particular – for obvious reasons – and kindly plugged my 2021 Dinopedia as her primary source of information on Mesozoic dinosaurs; ha ha, how kind. Anyway… the tool-use thing is not especially likely based on what we know right now.

**Caption:** Jennifer Colbourne talks about cockatoos and her research on their intelligence and use of tools, and on tool use in non-human animals in general. Image: Georgia Witton-Maclean.

Next up, we had a Mesozoic theropod-focused talk from Cassius Morrison. The idea that spinosaurids were fish-eaters is well known, but what about their relatives the megalosaurids? New data relevant to niche partitioning and diet in Jurassic and Cretaceous theropods of various sorts was discussed; this work forms the focus of Cassius’s PhD and I look forward to seeing it out there in print.

The final talk in the session was by myself and concerned Neglected Natives and the Reptiles and Amphibians of Britain’s Past. I’ve given this talk a few times before and have written about a few of the specific issues here at Tet Zoo over the years: it concerns such things as the loss of the UK’s native Pool frogs Pelophylax lessonae, arguments that the Common treefrog Hyla arborea might be a British native, and the saga of the Bournemouth green lizards.

**Caption:** one of the TetZooCon organisers discusses aspects of the British herpetofauna. I’ve always planned to write up the contents of this talk. But so far I haven’t. Image: Georgia Witton-Maclean.

Palaeoart, cosplay and stalls. Speaking and presenting events over, we moved out of the respective rooms and back to the reception area where our art exhibition and drinks reception happened. I didn’t drink too much wine, but I don’t remember all the art on show: it combined palaeoart (including the work of John Conway, Rebecca Gelernter, Natalia Jagielska, Joschua Knüppe, Joanna Kobierska, Luis Rey, Jed Taylor, Gert van Dijk and Steve White) with some academic-leaning anatomical reconstruction work (Matt Dempsey), book showing and plushy representation.

**Caption:** pictures taken at the TetZooCon 2022 art exhibition. At left, James McKay with a selection of small colour images produced for Palaeontological Association outreach events (children create prehistoric animals from jumbled bits and pieces, and James then illustrates these creations as real-looking animals). At right: Gert van Dijk with a selection of images relevant to his Furaha project. Images: Georgia Witton-Maclean.

  **Caption:** at left, palaeontologist and PhD student Matt Dempsey with his poster on dinosaur reconstructions. At right: a famous internet image incorporating Philip Hood art from *Man After Man*, now existing as a physical Christmas card that was gifted during TetZooCon 2022 to Dougal Dixon. Images: Darren Naish.

  **Caption:** Rebecca Gelernter art at TetZooCon 2022. Very pleased to see her there.

If you’ve been paying attention, I’ve been saying for years that I’d welcome cosplay at TetZooCon (I owe this idea to Rebecca Groom). We’ve never really got it going though. And then along came Meghan Jenkinson, who just happened to have an Ellie Sattler outfit with her. She wore it during the Saturday evening – and almost instantly we had a full Jurassic Park cosplay event on our hands. Which is great. I need a make a few enquiries, but I essentially reckon that we should lean hard into this and have a cosplay event – with accompanying competition – at the TetZooCons of the future. Watch this space.

**Caption:** all of a sudden - - - *Jurassic Park* cosplay! Image: Darren Naish.

I haven’t much mentioned the stalls and merchandise we had at TetZooCon 2022. I think we had a pretty good spread of art (both prints and originals), merch like stickers and pin badges, and books on sale. Sales were good, and I basically hope that we can expand the number and diversity of stalls we have in the future. I took along a good list of books I needed to sell (including several sought-after semi-academic texts) and did so via a silent auction. Thanks to Joanna, Natalia, Friends of the Crystal Palace Dinosaurs, Luis Rey, Jed Taylor, Steve White, Mark Witton and the others who hosted stalls.

**Caption:** thanks to the **Friends of Crystal Palace Dinosaurs**, I discovered and obtained these excellent enamel pin badges. They’re made by Joseph Fells and are available at **BleachedBonesStore.etsy.com**.

  **Caption:** various of the items that featured in the silent auction and quiz. Massive thanks to everyone who played along and took part. Image: Darren Naish.

What happened at the end of day 1, after the drinks reception? I have no idea but I remember waking up in a taxi at some point during the night.

And thus day 2 began. Hangover, what’s a hangover? The day consisted of five separate things: the pterosaur event, the Mesozoic Art signing event and its associated talk, the ‘designing aliens’ roundtable, the Crystal Palace talk, and the quiz.

The pterosaur session. I would have loved it if we’d done, say, a whole day, or a whole half a day, dedicated to pterosaurs – the membranous-winged flying cousins of the dinosaurs – but, in the end, a couple talks were still great. At last, I succeeded in getting my long-serving friend and colleague Liz Martin-Silverstone to attend and speak at TetZooCon (kidding, Liz has been busy or unavailable on the years I’ve asked). Liz’s talk was Why You Should (or Shouldn’t) CT Scan a Pterosaur. She discussed the ups and downs of CT-scanning and her own relevant studies, including the paper we co-authored on locomotor abilities and pterosaur sacral anatomy: Martin-Silverstone et al. (2018).

**Caption:** Liz Martin-Silverstone opens our special pterosaur session and asks the important questions. Image: Georgia Witton-Maclean.

Natalia Jagielska was up next with From Stone to Bone: a Step by Step Guide to Describing Your Own Pterosaur. It was much more than a retelling of the Dearc story (Dearc is a Middle Jurassic pterosaur from Scotland, described by Natalia and colleagues earlier this year: Jagielska et al. 2022) but actually a good guide to the entire process whereby fossil species are discovered, studied, published and incorporated into human knowledge and culture thereafter. Memes, gifs, joke captions… Natalia is a highly entertaining speaker!

**Caption:** the Natalia Jagielska stall. I understand that Natalia sold nearly all of the merch and art she bought along. Image: Agata Stachowiak, used with permission.

Liz and Natalia then joined me on stage – together with Mark Witton and John Conway – for the pterosaur roundtable event. I think it went well. We discussed current thinking on pterosaur integument and wing membrane anatomy, public misconceptions about pterosaurs, and our favourite pterosaurs (interestingly, we essentially all chose different groups) before taking questions from the audience. Natalia was accompanied by her trusty Dearc plushie (made by Rebecca Groom of Palaeoplushies fame) and had the opportunity to use it as a model when discussing pterosaur anatomy.

**Caption:** our pterosaur roundtable, as seen from the audience. Left to right: Darren Naish, Liz Martin-Silverstone, John Conway, Natalia Jagielska, Mark Witton. Image: Agata Stachowiak, used with permission.

  **Caption:** another pterosaur roundtable, showing (l to r) Darren, Liz and John. Image: Georgia Witton-Maclean.

A last-minute schedule modification meant the addition of an extra talk at this point in the schedule: John Conway’s A History of A History of Painting (With Dinosaurs). This accompanied John’s book A History of Painting (With Dinosaurs), officially launched a couple of weeks ago (I missed that launch due to being in the USA or something), and good fun it was too. The book was on sale at the event – I know John sold them all because there were none left when I went to steal one. I mean buy one.

**Caption:** John Conway presents A History of A History of Painting (With Dinosaurs), on his new book *A History of Painting (With Dinosaurs)*. Image: Georgia Witton-Maclean.

Onto the Mesozoic Art section. Steve White – the man, the legend – discussed Herding Cats: How to Publish a Palaeoart Book, a review of Steve’s involvement in palaeoart publishing. Steve might be seen as a ‘community hub’ today, but his initial entry into the area was tenuous and included a lot of luck. He also provided some cautionary tales on what works and what doesn’t when it comes to publishing. Niche books with unusual, complex titles (a hypothetical ‘Palaeoenvironments of the Mesozoic’ was chosen as an example) essentially won’t succeed if you take them mainstream.

**Caption:** Steve White discusses *Mesozoic Art* and its backstory. Image: Agata Stachowiak, used with permission.

The Dinosaur Art books from Titan have to be considered among the most successful palaeoart-themed works ever, and 2022’s Mesozoic Art is a sort of descendant of those books. Steve explained how thanks were owed to Jim Martin at Bloomsbury for setting things up, and I’m pleased to say that Jim was in attendance.

We moved on to the signing event. The fact that seven contributors to the book – Steve, myself, John Conway, Joschua Knüppe, Joanna Kobierska, Jed Taylor and Mark Witton, plus the publishers – were present at TetZooCon 2022 means that it was the closest thing we’ve had to an official launch-type event. Copies were sold throughout the whole of TetZooCon, such that I think all copies were bought (and hopefully all signed) by the end of the signing event.

**Caption:** one of the tables in the *Mesozoic Art* signing event (there were… three such tables, I think). Those doing the signing here are (left to right) John Conway, Mark Witton and Steve White. Image: Georgia Witton-Maclean.

Designing Aliens. Next, a (mostly different) set of seven people gathered on stage for a roundtable discussion on the designing of hypothetical aliens. What do we like, what do we not like, what possibilities exist, and what sort of projects are people working on? The event was co-led by myself and Gert van Dijk (of Furaha) and featured Jennifer Colbourne, Dougal Dixon, C. M. Kösemen, Joschua Knüppe and Adrian Tchaikovsky.

**Caption:** a scene - taken from my position on stage - of the designing aliens roundtable, showing (l to r) Gert van Dijk, Dougal Dixon, Adrian Tchaikovsky and Joschua Knüppe. Dougal is holding a *Greenworld* model; a woman (equipped with animal skins and other products manufactured from endemic wildlife) is riding a large, vaguely conch-like terrestrial animal. Image: Darren Naish.

  **Caption:** the designing aliens panel event as seen from the audience. Left to right: Darren Naish, Jennifer Colbourne, Memo Kösemen, Gert van Dijk, Dougal Dixon, Adrian Tchaikovsky and Joschua Knüppe. Image: Agata Stachowiak, used with permission.

Dougal bought along models constructed for his Greenworld project (still not published in English), Memo had original Snaiad artwork with him, and Adrian told us about his intelligent giant spiders. A question from Frank Drake’s grandson – yes, Drake Equation Drake – inspired us to consider how likely ‘human mimicry’ would really be when it comes to biological diversity in the universe. Interestingly, panel members mostly agreed that we shouldn’t assume a human-like form (even at the superficial level) in an intelligent alien, a view I strongly agree with. I would be interested in knowing whether people would have said similar things in a discussion panel 20 years ago, or 40 years ago.

**Caption:** more from the designing aliens roundtable, this time with images from Dougal Dixon’s *Greenworld* project on the screen behind us. Image: Georgia Witton-Maclean.

Anyway, the panel was over all too quickly – it was obvious that we could have talked for another hour… though it doesn’t necessarily follow that the audience would have been as enthusiastic as we were.

The Crystal Palace talk. Sunday wrapped up with the excellent talk The Art and Science of the Crystal Palace Dinosaurs: Changing the Narrative by Mark Witton and Ellinor Michell, which was again followed by a book signing. The new arguments about the prehistoric animal models of Crystal Palace put forward by Mark and Ellinor are now fairly well known (among those interested in palaeoart, anyway). The models represent a major triumph of design, construction and anatomical depiction, the role of Richard Owen is mysterious and likely minimal at best, and the relationship that the models have with mainstream science and science advocacy is complex and always have been. One of the most exciting discoveries that stems from Mark and Ellinor’s work is that some number of the original models have been entirely lost, a fact wholly unrecognised until now (Witton & Michel 2022).

**Caption:** Mark Witton and Ellinor Michel on stage at TetZooCon 2022. Buy the book, it’s amazing. Image: Georgia Witton-Maclean.

  **Caption:** slide from Mark and Ellinor’s talk, in this case focusing on the anatomy of the anoplotheres on show at Crystal Palace. What’s with the comparison between the anoplothere model and the shaking pet dog? You should check Witton & Michell (2022) for an explanation… Image: Agata Stachowiak, used with permission.

Be sure to get Mark and Ellinor’s book – The Art and Science of the Crystal Palace Dinosaurs (Witton & Michel 2022) – if you haven’t already. I feel compelled to add that I’ve written about the Crystal Palace models a few times here at Tet Zoo myself, most relevantly at Up Close and Personal With the Crystal Palace Dinosaurs.

Next up: time for the quiz. 30 questions of varying complexity, all on matters relating to the TetZooniverse, were put to the audience. They included the storylines of Primeval, zebra subspecies, the dispersal abilities of mink and Dr Alan Grant’s biography. Albert Chen got the highest score and thus first pick of the prizes, Kelvin Britton came next, then Richard Hing (all three have scored extremely highly or won in previous years). A remarkable number of excellent prizes – books, animal figures, plush toys, art prints, replicas and more – were available; massive thanks to those who donated them.

**Caption:** we had a selection of excellent quiz prizes. This large CollectA *Triceratops* and the accompanying book (authored by Erich Fitzgerald) were kindly provided by Jack Perkins and Melbourne Museum. At right: the three highest-scoring quiz players and myself. Images: Darren Naish, Hel Naish.

I should add here – mostly because I don’t know where else to put it – that we had several podcasting teams present at TetZooCon 2022, and they were essentially given free rein to record interviews with people throughout the event. I look forward to see—hearing the results; we couldn’t record the events of TetZooCon 2022 with cameras but at least some material from the meeting will exist in audio form. Having said that things weren’t recorded, there are at least a few brief videos: check out this one. I like it a lot.

**Caption:** Natee Himmapaan kindly provided this print of a *Prehistoric Planet*-inspired *Mononykus* (I also now own one myself). At upper right we see the diminutive, exquisite original in Natee’s drawing book.

We went to a large pub on Sunday night. Gone are the days where we can find a restaurant large enough to cater for us in cost-effective manner. Anyway, I think we had a good time. I got corralled in by a gang of Prehistoric Planet fans who refused to let me escape until I’d answered 240 questions about Prehistoric Planet. Ha ha, I kid, I kid, it was all good and I feel the love.

**Caption:** a beautiful day to visit the Crystal Palace dinosaurs. We see the two *Iguanodon* (which are no longer actually *Iguanodon* at all) at left, the *Hylaeosaurus* at right, and one of the two large pterosaurs (noawadays: *Cimoliopterus*) in the background. Image: Darren Naish.

To Crystal Palace. And so to Monday, the final day: the fieldtrip to Crystal Palace day. Bad weather threatened to ruin things, but in the end we had a bit of light rain and nothing more. More problematic was the fact that tall metal fences, locked gates and other barriers barred all entry to the park when we arrived. Various men in hi-viz coats assured me that entry wouldn’t be possible for a day or two, so it’s fair to say that I was a bit frustrated at this point. In the end, none of it made that much difference: entry was still possible via other routes.

**Caption:** group 2 look at the Crystal Palace *Megalosaurus*. Image: Darren Naish.

I did my best to swot up on the new ideas and discoveries covered in Mark and Ellinor’s book, and thus told my group a bunch of new stuff about anoplotheres and palaeotheres, about the missing Xiphodon and the Megaloceros fawn that isn’t, about the enigma of the megathere’s hands and so on. And if you want to know what I’m talking about, you’ll need to obtain the book (Witton & Michell 2021). The conditions were beautiful for photography, and we were essentially done by around 3pm.

**Caption:** more from the Crystal Palace fieldtrip. At left, the ‘*Megaloceros* fawn’, now known to be the last surviving *Xiphodon*. At right: your blog’s author with the original *Hylaeosaurus* head, today relocated on a mound above the main part of the geological court (the actual *Hylaeosaurus* model has a fibreglass replica for a head). Images: Darren Naish, Hel Naish.

And that’s that. TetZooCon 2022 was the biggest of our meetings so far; I’m so pleased with how it went. It was the NINTH TetZooCon, meaning that the 2023 one has the potential to be something bigger and more unusual than… usual. There’s already talk about things that might be arranged for 2023 but all I can say for now is watch this space, variously here at the blog, at the TetZooCon facebook page, or via # TetZooCon on twitter… or whatever social media platform is appropriate (I’m on mastodon but don’t use it much).

Finally, thanks as ever to my co-organisers and assistants – John, Jenny, Hel, Georgia, the staff at King’s College – and in particular to Chris Manias for access to the venue. Huge thanks to all the speakers, presenters, those with stalls, and of course to all the attendees. Another substantial success, and here’s to the TENTH event next year!

For previous articles on TetZooMCon and TetZooCon, see…

The events of TetZooCon 2014, July 2014
TetZooCon 2015 Is On, July 2015
The Events of TetZooCon 2015, November 2015
Coming Soon: TetZooCon 2016, September 2016
The Day After TetZooCon, October 2016
The Fourth TetZooCon, September 2017
The TetZooCon of 2017, October 2017
Reasons to Attend TetZooCon 2018, September 2018
TetZooCon 2018: Best TetZooCon So Far, October 2018
Announcing TetZooCon 2019 – the Biggest Yet, August 2019
Final Call For TetZooCon 2019, October 2019
The Sixth TetZooCon, October 2019
TetZooCon 2020 + Zoom = TETZOOMCON 2020, December 2020
TetZooMCon 2020 an Unbridled Success, December 2020
Cronch Cats, Beasts of Gévaudan, Dinosauroids, Mesozoic Art and Much More: TetZooMCon 2021 in Review, September 2021

Refs - -

Ashby, J. 2022. Platypus Matters: the Extraordinary Story of Australian Mammals. The University of Chicago Press, Chicago.

Jagielska, N., O’Sullivan, M., Funston, G. F., Butler, I. B., Challands, T. J., Clark, N. D. L., Fraser, N. C., Penny, A., Ross, D. A., Wilkinson, M. & Brusatte, S. L. 2022. A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs. Current Biology 32, 1-8.

Lomax, D. R. 2021. Locked in Time: Animal Behavior Unearthered in 50 Extraordinary Fossils. Columbia University Press, New York.

Martin-Silverstone, E., Sykes, D. & Naish, D. 2018. Does postcranial palaeoneurology provide insight into pterosaur behaviour and lifestyle? New data from the azhdarchoid Vectidraco and the ornithocheirids Coloborhynchus and Anhanguera. Palaeontology 2018, 1-14. doi: 10.1111/pala/12390

Witton, M. P. & Michel, E. 2022. The Art and Science of the Crystal Palace Dinosaurs. The Crowood Press, Marlborough.

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The giant, fantastically illustrated new volume Mesozoic Art: Dinosaurs and Other Ancient Animals in Art, edited and written by Steve White and myself and published by Bloomsbury Wildlife, is now on sale…

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I have elected to republish my articles on caecilians. (1) Because TetZoo always needs more amphibian coverage. (2) Because caecilians are awesome. The text you’re about to read - assuming you choose to continue - first appeared in 2008 (original here); I haven’t much updated it. Here we go…

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Finally, it’s time to talk about the zebras…

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Today (September 27th 2022), I’m speaking at the United Nations Science Summit panel Knowing and Protecting Life on Earth Starts with Natural History and Science Innovation…

I don’t have time today to talk about the presentation itself (that will come later), but I make several points: on education and outreach, on accessibility of knowledge, on the importance of a historical approach to natural history, and on the universal appeal of amazing animals to all people of all backgrounds. I’m one of a team of people on this panel; the others are highly respected experts on biodiversity, conservation and nature writing: namely Dr Ursula Valdez (tropical avian ecologist and trans-cultural educator), Dr Tom Fleischner (author and director emeritus of The Natural History Institute), Dr Dita Cahyani (marine biodiversity scientist and innovator), and Dr Nalini Nadkarni (forest canopy pioneer and conservationist). The panel was arranged by Dr Seabird McKeon and Dr Michele Weber, and will be led and convened by Dr McKeon.

You can register to attend yourself (it’s free) here.

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I’m very happy to announce that tickets for TetZooCon 2022 are now on sale …

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Frogs and toads – anurans – have profoundly modified skeletons and are among the most atypical of tetrapods...

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Time for another article from the extensive TetZoo archives, this time a piece from ver 2, 2008 (original here). We begin with this interesting photo provided by my good friend Markus Bühler (of Bestiarium): it shows a bull Asian elephant Elephas maximus at Hagenbeck Zoo, Hamburg…

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Of the several phantasmal creatures associated with the British and European countryside, among the most widespread and ancient is the Black Dog...

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It seems a good time to republish this article from the archives, first published at ver 2 in 2009, and then again at ver 3 in 2015…

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I have a new camera (thanks Chris), so I’ve been trying to photograph birds (trying to). And look: a success, of sorts…

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Yup, the blog has been here (as in: at TetZoo.com) for FOUR YEARS…

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I’ve written a fair amount about HORSES at TetZoo, predominantly at versions 2 and 3. For reasons, I now aim to reproduce it here at ver 3. We start with a article I first published in 2015 (original version here)…

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After a gestation that’s involved most of the current century, the long-awaited monographic description of the Early Cretaceous theropod dinosaur Eotyrannus lengi has finally appeared in print…

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In another effort to rescue old TetZoo material from the vandalized archives of ver 2 and 3, here’s a slightly revamped version of a 2009 article on hartebeest (the original version is here)…

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It’s time to look at duikers. Again.

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Yup, another new dinosaur from the Cretaceous of the UK…

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No time for anything new here lately – I think some of you will know why – but I have some breaking news...

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I like pigeons….

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Regular readers of Tetrapod Zoology should know that I’m a big fan of cassowaries, and indeed have a serious academic interest in them…

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Once again I’m giving you something from the archives, since there’s just no chance at all to produce anything new right now. Here, then, is a TetZoo ver 3 article that was originally published there in 2013 (that version is here). Ironically, that 2013 article was itself a republishing of a version from 2009…

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Inspired by my recent article on South American wild dogs, I went to the trouble of digging out a very brief TetZoo ver 2 article I published in 2007 on the remarkable and beautiful 'fox on stilts', the Maned wolf Chrysocyon brachyurus (said article is here). My aim was to augment and update that text such that it might be a useful one-stop review on this animal.

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Once more I must resort to plundering stuff from the archives, this time an article from TetZoo ver 2, originally published in July 2008 (and available here at wayback machine). Today: the kogiid sperm whales!

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What happened at Tet Zoo Towers during 2021? Well….

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The wooded places of Europe, Asia and northern Africa are home to a warbler with a distinctive song, a song that explains its English common name…

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Do you know this book? If the answer is yes, you might enjoy the following…

Tetrapod Zoology: Recent Episodes

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