Aldrovandi’s Monstrous Rooster, a 16th Century Dino-Chicken

Chickens are remarkable animals, and I’ve written about them a few times at TetZoo before, albeit always too briefly (see links below). I really need to write about them at length at some point; I actually worked for a few years as a specialist chicken researcher and gathered a lot of interesting information on these birds. Anyway… here, I want to talk about one chicken in particular: a famous individual that has been mentioned several times in the recent literature (e.g., Kaiser 2007). Namely: Aldrovandi’s monstrous rooster.

 Original image of Aldrovandi’s monstrous rooster, from volume 2 of his  Ornithologia . Credit: scan archived by University of Oregon ( original here ).

Original image of Aldrovandi’s monstrous rooster, from volume 2 of his Ornithologia. Credit: scan archived by University of Oregon (original here).

Ulisse Aldrovandi (1522-1605) was an Italian philosopher, physician and naturalist who’s most frequently mentioned for his three-volume Ornithologia of 1600. Therein, he wrote a lot about chickens, and part of his text covers mutants and monsters, predominantly conjoined chickens and chickens with extra limbs. My Latin is not that hot (err… Caecilius est pater), so it’s fortunate that Fernando Civardi transcribed, and Elio Corti translated, the chicken section of this work such that it’s today available to English readers. The resulting volume, published in 2009, is titled The Chicken of Ulisse Aldrovandi (Corti & Civardi 2009). It’s a great read if you like chickens.

For completeness I should note that Aldrovandi’s chicken text was actually translated beforehand (in 1963) by another worker (L. R. Lind) (Aldrovandi 1963). This version is said to be plagued with translation errors (hey, not my opinion) and hence is not preferred by specialists.

 The cover of Lind’s 1963 translation of Aldrovandi’s chicken text. What’s that thing on the cover? Well… that’s a whole ‘nother story. Credit:  amazon .

The cover of Lind’s 1963 translation of Aldrovandi’s chicken text. What’s that thing on the cover? Well… that’s a whole ‘nother story. Credit: amazon.

To return to the monstrous rooster, the book includes a fine depiction of it as well as a good paragraph of description and interpretation. Aldrovandi himself observed this remarkable bird, alive, in the collection of Francesco Medici – the Grand Duke of Tuscany – and described it as so shocking in appearance that “it struck fear into brave men with its terrifying aspect”. It was blackish overall with white bases to its feathers, and its feathers are said to have looked scale-like. Given that clean-edged, metallic feathers in many gallinaceous birds can look superficially scale-like (one of the best examples being the feathering of male Green peacock Pavo muticus, sometimes called the dragonbird), this is perhaps not as surprising as it might sound.

 There are a great many amazing gallinaceous birds, and here’s one of my favourites: the Green peacock. This is a captive individual at Tierpark Berlin. There’s an entire TetZoo article on this species: see links below. Credit: Markus Bühler, used with permission .

There are a great many amazing gallinaceous birds, and here’s one of my favourites: the Green peacock. This is a captive individual at Tierpark Berlin. There’s an entire TetZoo article on this species: see links below. Credit: Markus Bühler, used with permission .

The monstrous rooster did not possess a conventional fleshy comb and paired wattles but had feathers in their place: a large feathery crest projected from the top of the head and two spines – interpreted by Aldrovandi as feather quills lacking barbs – pointed upwards and outwards from the forehead, “as if they were two horns”. Tufts of long, bristle-like structures emerged from either side of the bill (close to the nostrils) and from the back of the neck as well. The legs were feathered down to the ankles and the feet appear not to have been remarkable. The bird appears to have been tall and large, based on the illustration (though no scale was provided). A colourised interpretation of the animal was produced by Corti & Civardi (2009).

 Cover of Corti & Civardi (2009), showing a colourised version of Aldrovandi’s monstrous rooster. Oh to see such a bird in life. Credit: Corti & Civardi (2009).

Cover of Corti & Civardi (2009), showing a colourised version of Aldrovandi’s monstrous rooster. Oh to see such a bird in life. Credit: Corti & Civardi (2009).

Finally, the most remarkable feature was the tail. It emerged from a whitish, rounded mass of feathers, and was long, slender, fleshy and pale blue. A mass of feathers formed a tuft (Aldrovandi called it a “flock”) at the tip. This tail reminded Aldrovandi of that of “quadrupeds”, predumably meaning lizards or mammals like rats or cats. Here I would remind you that a maniraptoran with a long tail ‘should not’ – so we think based on fossils – have a tail superficially recalling that of a lizard, but instead have a resplendently feathered one. The weirdness here perhaps indicates that the embryological development of this animal’s tail was likely different from that of extinct long-tailed birds and other Mesozoic maniraptorans.

Indeed, this bird sounds so weird overall that I sometimes even wonder whether it really was a domestic chicken, and not a member of some other (presumably now extinct) gallinaceous bird. But I don’t think that this is really up there as a possibility: it really was a member of Gallus gallus.

So… wow. What are we to make of all this?

 Agostino Carracci’s portrait of Ulisse Aldrovandi (1522-1605), physician, philosopher and naturalist. Credit: image in public domain, from wikipedia ( original here ).

Agostino Carracci’s portrait of Ulisse Aldrovandi (1522-1605), physician, philosopher and naturalist. Credit: image in public domain, from wikipedia (original here).

For starters, what do we know about Aldrovandi? Well, quite a lot. He was a credible, well trained individual who had studied law, philosophy, mathematics and logic at university; he published on insects and other invertebrates and was even credited by Linnaeus as the ‘father of natural history’. He also wrote extensively about anomalous cases in zoology and medicine and collected enough of them that they were (posthumously) published in the 1640 volume Historiae Serpentum et Draconum and the 1642 Monstrorum Historia. His serpents and dragons book also includes a brief discussion of the monstrous rooster, but doesn’t add information relative to that included in his Ornithologia.

 Damn, chickens are awesome. These birds belonged to a group living semi-wild on Madeira. Credit: Darren Naish.

Damn, chickens are awesome. These birds belonged to a group living semi-wild on Madeira. Credit: Darren Naish.

What we can glean from Aldrovandi’s writings is that he lived within the ethnographic landscape of Renaissance Italy, by which I mean that he seemingly believed in things (like human-like monsters and mythical beasts of far-off lands) that we today know not to exist. But his writings on direct, specific cases show that he was not credulous or prone to endorsing half-truths. He was critical of stories about basilisks, for example (thought at the time to result from the production or brooding of eggs by roosters)*. Furthermore, his illustrations of known animals are often highly accurate, as you can see from the examples shared here. Indeed, his writings on other anomalous specimens – like the ‘Homuncio’ (a short-statured Indian man whose body was draped with massive fleshy growths) – have been interpreted as biologically accurate (Ruggieri & Polizzi 2003).

* For those curious, I’m not exploring the basilisk angle here; I have to avoid it for now. Constraints of time.

 Aldrovandi was a skilled and accurate artist, and his illustrations of known species - familiar and foreign - are usually highly accurate, as is demonstrated by these chickens. His chicken text also includes several illustrations of curassows, and they’re all essentially accurate. Credit: Corti & Civardi (2009).

Aldrovandi was a skilled and accurate artist, and his illustrations of known species - familiar and foreign - are usually highly accurate, as is demonstrated by these chickens. His chicken text also includes several illustrations of curassows, and they’re all essentially accurate. Credit: Corti & Civardi (2009).

The major caveat here is that Aldrovandi’s text and illustration of the monstrous rooster were (so far as we can tell) produced some considerable time after his observation of the bird, in which case all sorts of discrepancies might have crept in. The possibility that it had been modified or fitted with an artificial tail is not out of the question, but can’t be tested and is just an idea I need to mention in passing. We do know of other cases whereby animals have been made to look remarkable to impress or dupe observers, after all.

But, all in all, I’m inclined to think that the case was genuine, and that a long-tailed mutant rooster really was observed at some point in the 1500s by an erudite young man.

 If a ‘dino-chicken’ ever does come to pass, it should be awesome and beautiful — like a real chicken. Err, in which case I don’t think anyone will look at it and think of a connection with the Mesozoic maniraptorans it’s meant to evoke. Whatever. Credit: Rebecca Groom.

If a ‘dino-chicken’ ever does come to pass, it should be awesome and beautiful — like a real chicken. Err, in which case I don’t think anyone will look at it and think of a connection with the Mesozoic maniraptorans it’s meant to evoke. Whatever. Credit: Rebecca Groom.

I will leave you with one final thought. Jack Horner’s ‘dino-chicken’ project seeks to create a mutant fowl with a long, bony tail and other ‘ancestral’ features, all brought into existence via genetic and embryological modification. And research underpinning such efforts has already been published (Rashid et al. 2018). Was Aldrovandi’s rooster a demonstration that some of these developmental changes can occur without modern, deliberate modification? In other words, could it have been a real, ‘natural’ dino-chicken; one that existed four-hundred years before our time? If only the body, or skeleton, of this amazing bird had been preserved.

For other TetZoo articles linked to things mentioned here, see… (note: TetZoo ver 2 articles - the ScienceBlogs ones - are now appearing without their images, yay!)…

Refs - -

Aldrovandi, U. 1963. Aldrovandi on Chickens. Translated by L. R. Lind. University of Oklahoma Press, Norman, OK.

Corti, E. & Civardi, F. 2009. The Chicken of Ulisse Aldrovandi. www.summagallicana.it

Kaiser, G. 2007. The Inner Bird: Anatomy and Evolution. University of British Columbia, Vancouver.

Rashid, D. J., Surya, K., Chiappe, L. M., Carroll, N., Garrett, K. L., Varghese, B., Bailleul, A., O’Connor, J., Chapman, S. C. & Horner, J. R. 2018. Avian tail ontogeny, pygostyle formation, and interpretation of juvenile Mesozoic specimens. Scientific Reports 8: 9014.

Ruggieri, M. & Polizzi, A. 2003. From Aldrovandi’s “Homuncio” (1592) to Buffon’s girl (1749) and the “Wart Man” of Tilesius (1793): antique illustrations of mosaicism in neurofibromatosis? Journal of Medical Genetics 40, 227-232.

Comical Tales From the Animal Kingdom, a Zoological Society of London Meeting

2018 has been a pretty busy year here at TetZooTowers, and still there are a list of things set to happen between now and the start of 2019. What’s next on the list? Why, it’s the Zoological Society of London meeting From Stoned Sloths to Farting Fish: Comical Tales From the Animal Kingdom, happening at the ZSL’s Huxley Lecture Theatre on the evening of Thursday November 15th. Oh my god that’s next week.

 There will be books.

There will be books.

The event sees me, Dani Rabaiotti, Lucy Cooke and Jules Holland discuss weird and wild stories from the animal kingdom, and in particular how said stories relate to the animal-themed books we’ve published. Dani, as you’ll know, is the highly acclaimed co-author of the insanely successful Does It Fart? as well as its hugely crappy follow-up True or Poo? Lucy recently published the brilliant The Unexpected Truth About Animals (she spoke about this book at TetZooCon 2018). Jules’s recent books are Sex on Earth and Death on Earth. I’ll be talking about dinosaurs and publishing books on them. How on earth can I make that at all humorous? Well, we’ll just see.

 Books. Image: Darren Naish.

Books. Image: Darren Naish.

We’re also signing and selling our books (I’m selling copies of the brand-new second edition of Dinosaurs: How They Lived and Evolved, co-authored with the Natural History Museum’s Paul Barrett… PLEASE BRING CASH), and I believe that we’re having a Q&A session after our talks as well. It should be great fun and I’m looking forward to it.

You have to book for this event (tickets are £5); please go here for further information and booking.

 A slide I generated for my talk, showing some (yeah… some) of the dinosaur-themed books I’ve been involved in during my career as a freelancer. What’s the story here? All will be revealed on the 15th. Oh, and I’m talking about dinosaur sex as well. Again.

A slide I generated for my talk, showing some (yeah… some) of the dinosaur-themed books I’ve been involved in during my career as a freelancer. What’s the story here? All will be revealed on the 15th. Oh, and I’m talking about dinosaur sex as well. Again.

Brian Switek’s My Beloved Brontosaurus: A Belated Review

I’ve posted a few reviews of recently-ish published dinosaur books here lately, aaaand I have a few more to publish yet – the backlog is long. So.. it seemed as good a time as any to recycle this review from five years ago, pertaining to a book published in 2013. The review was written for the website produced to accompany the 2013 Walking With Dinosaurs movie but… for reasons that I’m sure make sense to someone, this review and in fact the entire site was later removed from the internet (yeah yeah yeah... nothing is ever really removed, blah blah blah IT’S GONE FOR NORMAL PEOPLE, OK?). I here republish it in full, unmodified form since those far-off, halcyon days of 2013. Which is kind of ironic, given what’s happened to Brontosaurus since 2013 (see links below for more on that)…

 The brilliant cover-art to   Switek (2013)  , by Mark Stutzman. Image:   Switek (2013)  .

The brilliant cover-art to Switek (2013), by Mark Stutzman. Image: Switek (2013).

Dinosaurs are popular. We seem, in fact, to be at an all-time high in terms of our hunger for new dinosaur news, the frequency with which new discoveries are announced, and in the sheer volume of dinosaur-themed books, TV shows and movies.

Brian Switek’s My Beloved Brontosaurus describes the author’s personal journey as he visits various dinosaur-bearing fossil sites and dinosaur-themed museums and other institutions across the United States, all the while discussing the fast-changing pace of our knowledge, the new insights we have into dinosaurs and their biology, and the fondness we retain for the big, fat, swamp-dwelling, pea-brained version of the dinosaur that almost refuses to die. Herein lies the explanation for the book’s title: the concept of ‘Brontosaurus’ embodies a defunct stereotype, a name that ‘shouldn’t’ be in current use [UPDATE: OH THE IRONY] and which remains associated with a version of the dinosaur that pre-dates the Dinosaur Renaissance.

 The iconic diplodocid specimen (AMNH 460) on display in the American Museum of Natural History, New York, with its semi-imaginary skull. Long labelled as  Brontosaurus excelsus , and then  Apatosaurus excelsus , it’s currently of indeterminate status. Image: AMNH, in public domain ( original here ).

The iconic diplodocid specimen (AMNH 460) on display in the American Museum of Natural History, New York, with its semi-imaginary skull. Long labelled as Brontosaurus excelsus, and then Apatosaurus excelsus, it’s currently of indeterminate status. Image: AMNH, in public domain (original here).

The book reads like something of a road trip, arranged such that Switek tells the story of the dinosaurs’ Triassic origins and gradual rise to dominance, explores ideas about their sex lives and reproductive and growth strategies, looks at their evolution of giant size, at dinosaur social lives, at feathers and featheryness, at pathologies and illnesses, and, eventually, at the end-Cretaceous extinction event. It’s a well-written, flowing narrative; the chapters are connected, both to one another and to Switek’s journey across the country, though the connections do sometimes seem a bit contrived. Notes at the back point the reader to technical sources for the information included, illustrations appear throughout, and the book is fully indexed.

 Front cover of   Switek (2013)  . Some apatosaurines truly were gargantuan - the individual shown here might be overly gargantuan, but the image is brilliant anyway. Image:   Switek (2013)  .

Front cover of Switek (2013). Some apatosaurines truly were gargantuan - the individual shown here might be overly gargantuan, but the image is brilliant anyway. Image: Switek (2013).

I really like Mark Stutzman’s cover art so was especially thrilled to see that the dust jacket folds out into a double-sided poster. Extra dino-geek points if you know the inspirations behind either of Stutzman’s illustrations.

By and large the volume is up-to-date and factually accurate, but there are a few things I take issue with. The name Brontosaurus is of course a major fixture throughout the book, the obvious reason for this being that the name loomed large in virtually every single popular dinosaur book prior to... well, prior to some surprisingly recent date. As Switek explains, it’s not true to say – as people sometimes do – that “Brontosaurus never existed”; rather, the species that the name Brontosaurus is tied to was shown in 1903 to be so similar to species included in another genus (Apatosaurus) that separate status for Brontosaurus was no longer defensible. If Brontosaurus ‘died’ in 1903, why has it persisted until so recently? [UPDATE: AGAIN… OH THE IRONY. For the one or two of you that don’t know, the name Brontosaurus was resurrected from synonymy in a comprehensive analysis published in 2015 (Tschopp et al. 2015)].

 Another fantastic apatosaurine: this is  Apatosaurus louisae , photographed at the Carnegie Museum. You might be wondering what the deal is as goes those massive club-like processes on the cervical vertebrae. Yeah, we’re working on that. Really… we are. Image:  Tadek Kurpaski  CC BY 2.0 ( original here ).

Another fantastic apatosaurine: this is Apatosaurus louisae, photographed at the Carnegie Museum. You might be wondering what the deal is as goes those massive club-like processes on the cervical vertebrae. Yeah, we’re working on that. Really… we are. Image: Tadek Kurpaski CC BY 2.0 (original here).

Switek’s answer is that this choice was essentially arbitrary: he suggests that New York’s American Museum of Natural History stuck with the name in their massively popular museum display because they maybe “thought the old name sounded better, or were unsure about rebranding one of the most famous dinosaurs in their halls”.

However, as explained in Paul Brinkman’s 2010 book The Second Jurassic Dinosaur Rush, there’s a good reason why Brontosaurus persisted for so long. That reason: Henry Fairfield Osborn, president of the AMNH between 1908 and 1933. Osborn was hugely influential and highly opinionated, and his insistence on sticking with the name Brontosaurus ensured the use of the name well beyond its time as a popular (rather than technical) moniker. The 1903 sinking of Brontosaurus was suggested by Elmer Riggs. Riggs did good work, but he simply lacked the academic clout to make Osborn change his mind.

 Another excellent volume on the history of Mesozoic dinosaur palaeontology:   Paul Brinkman’s 2010  The Second Jurassic Dinosaur Rush   .   My review of this book can be found here   (at the paywalled and utterly useless no-man’s land that is the SciAm blogs site). Image:   Brinkman (2010)  .

Another excellent volume on the history of Mesozoic dinosaur palaeontology: Paul Brinkman’s 2010 The Second Jurassic Dinosaur Rush. My review of this book can be found here (at the paywalled and utterly useless no-man’s land that is the SciAm blogs site). Image: Brinkman (2010).

Something I think we need to do more when writing about scientific discoveries and hypotheses is discuss the social dimension to science: that is, the reception said discoveries and hypotheses received within the academic community. New and radical notions and proposals are – despite the impression you might get from TV shows and magazine articles – typically not embraced with open arms. Instead, they initially receive cold and even harsh treatment. Since scientists are human, the way they react to such ideas may be influenced by their personal feelings, the research environment they were trained in, who their friends and enemies are, and so on. In the world of palaeontology, it seems that we’re entering a phase where we’re increasingly analysing the work and ideas of the last few decades: in other words, charting the socio-political background to the science.

 It is important, I feel, that we document the history of our changing ideas, something I tried to do in    The Great Dinosaur Discoveries  (Naish 2009)  . Image:   Naish (2009)  .

It is important, I feel, that we document the history of our changing ideas, something I tried to do in The Great Dinosaur Discoveries (Naish 2009). Image: Naish (2009).

I tried to do this myself in my 2009 book The Great Dinosaur Discoveries (Naish 2009), and Switek often does it in My Beloved Brontosaurus since he quotes experts and reports the conversations he has with them. People interested in the history of palaeontology, and in the history of science in general, should read the book for this reason. It helps make the book a snap-shot of where we are now in the world of dinosaur science – it covers the issues we’re currently interested in; the questions and discoveries that we’re talking about.

All in all, My Beloved Brontosaurus is both a fun and absorbing read as well a good, popular guide to our current understanding of dinosaurs and their evolution.

Brian Switek, 2013. My Beloved Brontosaurus: On the Road with Old Bones, New Science, and our Favourite Dinosaurs. Scientific American/Farrar, Straus and Giroux, New York, 256 pp. ISBN 978-0-374-13506-5. Hardback, index, illustrations.

For previous TetZoo articles relevant to this one, see…

Refs - -

Brinkman, P. 2010. The Second Jurassic Dinosaur Rush. Chicago: University of Chicago Press.

Naish, D. 2009. The Great Dinosaur Discoveries. A & C Black, London.

Switek, B. 2013. My Beloved Brontosaurus: On the Road with Old Bones, New Science, and our Favourite Dinosaurs. Scientific American/Farrar, Straus and Giroux, New York.

Tschopp, E., Mateus, O. & Benson, R. B. J. 2015. A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda). PeerJ 3:e857; DOI 10.7717/peerj.857

New Living Animals We Want to Find

As a regular denizen of the TetZooniverse, you may well remember the July 2017 article ‘Fossils We Want to Find’ in which I discussed a list of hypothetical fossil things that we might one day discover but haven’t yet. Wouldn’t it be fun to do the same sort of thing with extant species; that is, with discoveries pertaining to living, breathing animals? Over at the Zoology for Enthusiasts facebook group (a spinoff of the Tetrapod Zoology facebook group), Jordan Fryer suggested doing exactly this, and consequently people have been coming up with their own suggested living animals that might await discovery. Because this seemed like a lot of fun (and a chance to discuss some really neat and unusual stuff), I thought I’d give it a go.

 The 2017 precursor to the article you’re reading here was all about fossil animals. It included this photo, which shows me in the act of discovering a dinosaur bone in the Moroccan Sahara. Image: Richard Hing.

The 2017 precursor to the article you’re reading here was all about fossil animals. It included this photo, which shows me in the act of discovering a dinosaur bone in the Moroccan Sahara. Image: Richard Hing.

Naturally, any list of this sort is horribly subjective, reflecting the interests and biases of the person compiling the list, but so be it. It also seems all too easy to turn any compilation into a ‘list of most discoverable cryptids’: for those of you who don’t know, I have a long-standing interest in cryptozoology and have published on it quite frequently (see Naish (2017) for starters). For the most part, I’ve not done this, though read on.

 Many of my thoughts on mystery animals can be found in my   2017 book  Hunting Monsters   . I am not - sorry - much impressed by the case for such supposed animals as the mokele-mbembe, an artistic reconstruction of which is shown at right. Image: David Miller, in   Mackal (1987)  .

Many of my thoughts on mystery animals can be found in my 2017 book Hunting Monsters. I am not - sorry - much impressed by the case for such supposed animals as the mokele-mbembe, an artistic reconstruction of which is shown at right. Image: David Miller, in Mackal (1987).

I’ve also mostly excluded hypothetical discoveries that are inspired by the creatures of cryptozoology but could arguably be considered independent of the cryptozoological literature. In part this is because I don’t think they’re plausible or worth considering, but it’s also because they’re cliched and the opposite of original. So, no ‘living sauropods from the Congo’ or ‘living plesiosaurs in Loch Ness’, for example.

As for what I have selected: well, some of my suggestions are sillier than others, and some are perhaps not that interesting to non-specialists. But, whatever. Feel free to dissect my suggestions in the comments, and perhaps come up with your own.

 Among my suggested ‘fossils we want to find’ are protobats (like the hypothetical examples shown at left, from Graham (2002)) and a good skeleton of the giant hominid  Gigantopithecus blacki . This ilustration of a lower jaw is from Simons & Ettel's (1970) magazine article. Images: Graham (2002), Simons & Ettel (1970).

Among my suggested ‘fossils we want to find’ are protobats (like the hypothetical examples shown at left, from Graham (2002)) and a good skeleton of the giant hominid Gigantopithecus blacki. This ilustration of a lower jaw is from Simons & Ettel's (1970) magazine article. Images: Graham (2002), Simons & Ettel (1970).

A habitually bipedal, large, non-human hominid. Whatever you think of all those stories, anecdotes and sightings about bigfoot, yeti, almas, orang-pendek, yowie and so on and on, the fact remains that the discovery of a large, bipedal non-human hominid – whether it be a pongine, hominine, or member of another hominid lineage – would be a huge deal. It would not just be one of the most newsworthy creatures to ever be discovered; it would also have enormous ramifications for our understanding of hominid evolution and potentially the human condition itself.

 Are crypto-hominids a cultural phenomenon more than a zoological one? I’ve argued for both possibilities at different times. Whatever… for the purposes of the article you’re reading now, I hope we can agree that the discovery of such an animal would be high on any hypothetical ‘wants’ list. Image: Darren Naish.

Are crypto-hominids a cultural phenomenon more than a zoological one? I’ve argued for both possibilities at different times. Whatever… for the purposes of the article you’re reading now, I hope we can agree that the discovery of such an animal would be high on any hypothetical ‘wants’ list. Image: Darren Naish.

It would also – if relating to North America or northern Eurasia in particular – very likely have a significant impact on economy, land management and land use in those regions… or, you’d hope it would, anyway (who knows, given the current state of environmental protection in the USA). The hypothetical discovery of such an animal would also be regarded by many as one of the biggest ‘wins’ ever scored against ‘establishment science’, and thus could well be a bad thing (viz, “if scientists were wrong about this, what else could they be wrong about?”). And I’ll stop there before we dive into a rabbit-hole of conspiracy theories and coverups.

A big, flightless passerine. The majority of living bird species – over 60% of them – are passerines, or perching birds. This is the great group that includes crows, thrushes, warblers, finches, sparrows and so many others. For all their success, wide distribution and diversity, passerines are generally quite samey. There are no big, long-legged wading passerines, or heavy-bodied diving passerines or flightless running passerines, for example. Why this is so remains mysterious: passerines didn’t take to those niches because… well, they just didn’t. Does this mean that they couldn’t? As usual, we can come up with a few reasons as to why they were ‘constrained’ in evolutionary potential, but any one of those reasons could be overturned by some evolutionary deviant that refuses to pay attention to the rules.

 Passerine birds are diverse, to a degree… here’s just a sample of their diversity. This is part of a giant montage that’s being built for   my in-prep textbook The Vertebrate Fossil Record  . Image: Darren Naish.

Passerine birds are diverse, to a degree… here’s just a sample of their diversity. This is part of a giant montage that’s being built for my in-prep textbook The Vertebrate Fossil Record. Image: Darren Naish.

And thus I submit that a particularly large, wholly flightless, cursorial passerine should make itself known to the world. It should be a record-holder as goes size, but not necessarily be that much bigger than the largest known passerines (like lyrebirds and ravens): I’m talking about a bird that weighs 3-5 kg and is thus similar in size to a large chicken. It should be a big, long-legged rail-babbler, quail-thrush or similar, and hence be a denizen of Wallacea or nearby.

  Eupetes , the Malaysian rail-babbler. A hypothetical big, flightless passerine should be a close relative of this bird. Image: Francesco Verronesi, CC BY-SA 2.0 ( original here ).

Eupetes, the Malaysian rail-babbler. A hypothetical big, flightless passerine should be a close relative of this bird. Image: Francesco Verronesi, CC BY-SA 2.0 (original here).

A few recently extinct, island-dwelling passerines were flightless, so we do know that passerines have the evolutionary potential to follow this pathway. Such species (a bunting and a few New Zealand wrens… and possibly a few others) were all small (less than 40 g).

A western Asian giant salamander. Giant salamanders (cryptobranchids) are restricted today to eastern Asia (where Andrias occurs) and North America (where Cryptobranchus occurs). Hunting, human disturbance, habitat loss and deterioration, climate change and other issues are putting them into perilous decline, right at the same time as we’re discovering that some of them are species complexes. They were more widespread in the past than they are today, since fossils show that Andrias salamanders were widespread across Europe and Asia between about 28 and 2 million years ago.

 An Asian giant salamander ( Andrias ) photographed in captivity. Record-holding specimens of  Andrias  can be 1.8 m long and exceed 60 kg, and some extinct species reached even larger sizes. Image: Markus Bühler.

An Asian giant salamander (Andrias) photographed in captivity. Record-holding specimens of Andrias can be 1.8 m long and exceed 60 kg, and some extinct species reached even larger sizes. Image: Markus Bühler.

While there are very good reasons for the decline and extinction of the animals in the areas concerned, some of the regions where they formerly occurred still have what look like suitable habitat today and are sparsely populated by people. Furthermore, extinct giant salamanders weren’t all denizens of fast-flowing, highly oxygenated streams like those inhabited by the modern populations. Some inhabited ponds and lakes. Ergo: I would really, really like there to be a west Asian cryptobranchid that comes from a habitat considered weird for the other living members of the group. And it doesn’t have to be a giant of 2 metres or more. A hellbender-sized species of 70 cm or so will do fine thank you very much.

 Some extinct cryptobranchids - this is Zdeněk Burian’s reconstruction of  Andrias scheuchzeri  - inhabited European ponds and lakes. I’ve previously criticised this image for showing the animal as terrestrial. Since then, the proposal has been made that some extinct cryptobranchids (albeit not  A. scheuchzeri ) were significantly more terrestrial than living species. Image: (c) Zdeněk Burian.

Some extinct cryptobranchids - this is Zdeněk Burian’s reconstruction of Andrias scheuchzeri - inhabited European ponds and lakes. I’ve previously criticised this image for showing the animal as terrestrial. Since then, the proposal has been made that some extinct cryptobranchids (albeit not A. scheuchzeri) were significantly more terrestrial than living species. Image: (c) Zdeněk Burian.

Again, this is an area of special interest to cryptozoologists, since there have been occasional suggestions that stories, engravings and such from western Asia might reflect folk knowledge of unusually big salamanders in the region. In reality, the images and stories concerned are super-ambiguous and more likely refer to otters and god knows what else.

 At left:  Andrias  skull. Image: Darren Naish. At right: Japanese giant salamander ( A. japonicus ) illustration by Y. de Hoev from 1887. Image: Y. de Hoev, public domain ( original here ).

At left: Andrias skull. Image: Darren Naish. At right: Japanese giant salamander (A. japonicus) illustration by Y. de Hoev from 1887. Image: Y. de Hoev, public domain (original here).

A living albanerpetontid. Everyone knows that there are three main groups of living amphibian: caecilians, salamanders and anurans (frogs and toads). But until (geologically) recently, there was a fourth group: the albanerpetontids, sometimes termed albies by those who work on them. Albanerpetontids were geographically widespread, their range including Eurasia, northern Africa and North America, and they were geologically long-lived too. The oldest are from the Middle Jurassic while the youngest are… well, we’ve known of Miocene fossil albanerpetontids for decades, have known of Pliocene specimens since 2005 (Venczel & Gardner 2005), and now know that at least one species persisted into the Pleistocene (Villa et al. 2018). The fact that their fossil record has been creeping towards the Recent means that the possibility of fossil and even extant Holocene specimens being discovered isn’t ridiculous, especially given the small size of these animals and hence tiny size of their bones.

 New salamander species are occasionally discovered in Europe and Asia even now. It would be amazing if an animal suspected to be a ‘new salamander’ one day turned out to be a living albanerpetontid. These reconstructions were published by McGowan & Evans (1995). They might have erred in implying that the scales would be externally visible as shown here; more likely is that they were concealed by epidermis, as in other scaly fossil amphibians. Image: McGowan & Evans (1995).

New salamander species are occasionally discovered in Europe and Asia even now. It would be amazing if an animal suspected to be a ‘new salamander’ one day turned out to be a living albanerpetontid. These reconstructions were published by McGowan & Evans (1995). They might have erred in implying that the scales would be externally visible as shown here; more likely is that they were concealed by epidermis, as in other scaly fossil amphibians. Image: McGowan & Evans (1995).

To be frank, a live albanerpetontid wouldn’t be a particularly spectacular animal: it would be a tiny, slim, salamander-like amphibian less than 10 cm long, and it wouldn’t be much fun to watch since it would spend most of its time hiding and burrowing in leaf litter. But among herp-nerds it would be a huge deal. Live albanerpetontids were scaly-skinned (though the scales were not necessarily visible externally), with eyelids, and with adaptations in the snout, skull-roof, neck and body shape linked to head-first burrowing (McGowan & Evans 1995).

 An artistic reconstruction of a live albanerpetontid…   produced for my in-prep The Vertebrate Fossil Record  . Image: Darren Naish.

An artistic reconstruction of a live albanerpetontid… produced for my in-prep The Vertebrate Fossil Record. Image: Darren Naish.

A Eurasian palaeognath. Palaeognaths are the big, flightless ratites (ostriches, emus and so on), the superficially gamebird-like, flight-capable tinamous, and their extinct relatives. A huge amount has been written about the evolutionary history and biogeography of these birds, since their distribution is curious and has resulted in all kinds of different models about how they might have spread around the world. I’ve written about this issue at length on previous occasion (the articles concerned being famous for generating the longest-ever comment threads in the history of TetZoo… though all of this is mostly wasted now, what with SciAm’s paywalling of the site, sigh). Living palaeognaths are absent from Eurasia, despite the former present in the region of ancient, flight-capable Paleogene taxa, extinct ostriches and others.

 As this map shows, modern palaeognaths occurred everywhere until recently (except Antarctica) with the exception of northern North America and the cooler parts of Eurasia. Extinctions across Eurasia, Madagascar and New Zealand of course saw the disappearance of various members of the group. Image: Darren Naish.

As this map shows, modern palaeognaths occurred everywhere until recently (except Antarctica) with the exception of northern North America and the cooler parts of Eurasia. Extinctions across Eurasia, Madagascar and New Zealand of course saw the disappearance of various members of the group. Image: Darren Naish.

The fact that Paleogene Europe was home to many bird groups that no longer occur there but are now denizens of tropical regions elsewhere leads me to hope for a living palaeognath – a tinamou- or bustard-sized species – that descends directly from archaic Paleogene taxa and now lives in the Asian tropics. It should be a cryptic generalist with barred plumage and a mid-length bill and a reduced flight ability.

A gigantic, predatory, limbed amphisbaenian. Regular readers of TetZoo might know that I really like amphisbaenians: the mostly limbless, bullet-headed ‘worm lizards’ of the American tropics, Africa, and parts of southern Europe and western Asia. Amphisbaenian evolutionary history and biogeography has become increasingly complex in recent years as we’ve learnt a bunch of new stuff about their fossil history, genetics and anatomy. Among the weirdest of amphisbaenians are the ajolotes (or bipedids), the only extant group to possess limbs. These limbs are not small stumps or flaps (as they are in some other near-limbless, serpentine squamates) but well-developed, clawed forelimbs. According to some phylogenetic models, ajolotes are not the sister-group to limbless amphisbaenians but deeply nested within the limbless clade (Conrad 2008, Videl et al. 2008), in which case their limbedness – if you will – perhaps evolved from limbless ancestors. Add to this the fact that some amphisbaenians are robust-jawed, short-faced predators of vertebrates that ambush prey from beneath the surface and bite chunks from the bodies of surface-dwelling mammals and reptiles.

  Bipes , an ajolote of Mexico (they might occur in parts of the USA as well). Three extant species are recognised. Image: Darren Naish.

Bipes, an ajolote of Mexico (they might occur in parts of the USA as well). Three extant species are recognised. Image: Darren Naish.

So then… where oh where are the giant, limbed, robust-skulled, vertebrate-eating amphisbaenians? By ‘giant’, I am not talking about a graboid-sized monster of several metres (though that would be nice), but a more reasonable animal of a mere 1.5 metres or so. Easily the stuff of nightmares. They could inhabit warm regions of any continent.

 Carl Gans’s illustration of a burrowing ajolote, showing how the large, well-clawed forelimbs function in propulsion. This is clearly a Five-toed worm lizard  Bipes biporus ; the other extant species have four and three digits, respectively. Image:   Gans (1974)  .

Carl Gans’s illustration of a burrowing ajolote, showing how the large, well-clawed forelimbs function in propulsion. This is clearly a Five-toed worm lizard Bipes biporus; the other extant species have four and three digits, respectively. Image: Gans (1974).

Is there any reason to think that gigantic, predatory, limbed amphisbaenians might actually exist and await discovery? Nope. But I wish it were so. Regular readers might recognise that such creatures are denizens of the alternative-timeline Earth of the Squamozoic, but I’m sure that that’s coincidental.

 What would a gigantic, predatory, limbed amphisbaenian look like? Like this, of course. Image: Darren Naish.

What would a gigantic, predatory, limbed amphisbaenian look like? Like this, of course. Image: Darren Naish.

An African or west Eurasian, long-beaked river dolphin. On several occasions within the history of odontocete cetaceans (‘toothed whales’), lineages have moved into brackish and estuarine environments, and eventually made the transition to committed freshwater life. There are the Asian Platanista species, the recently extinct Lipotes of China, and the tropical American Inia species. Once united within Platanistoidea and thought to be close kin, we know today that these animals represent at least three separate transitions to the freshwater environment (the term Platanistoidea is now restricted to the Platanista lineage alone). In addition, members of other groups – I’m thinking of the delphinid Orcaella – occur in rivers within parts of their range. There’s also a fossil beaked whale that might be indicative of freshwater specialisation in yet another odontocete group (Mead 1975).

 River dolphins are pretty special looking. This is a Ganges river dolphin ( Platanista gangetica ). Image: Zahangir Alom / Marine Mammal Commission / National Oceanic and Atmospheric Administration, public domain ( original here ).

River dolphins are pretty special looking. This is a Ganges river dolphin (Platanista gangetica). Image: Zahangir Alom / Marine Mammal Commission / National Oceanic and Atmospheric Administration, public domain (original here).

In view of all this, why aren’t there river-dwelling dolphins in Africa, Europe or western Asia? Again, the answer seems to be… there just aren’t. A few fossil taxa suggest that such animals might have evolved if things had gone another way (there are fossil platanistoids from the Caucasus, for example). But I humbly submit that the great river systems of tropical Africa, the Tigris-Euphrates system of western Asia and the Danube, Po, Ebro, Dniester and others of southern Europe would be much improved if only we knew of their endemic riverine dolphins. I’m talking about a true riverine specialist, convergent with Inia and Platanista, with a long beak, spike-like teeth, reduced eyesight, the works. And if you want to play fast and loose with antiquarian literature and anecdote, there are references in the literature to ‘river dolphins’ in the Nile and there are even one or two eyewitness accounts from central Europe that describe long-beaked ‘dolphins’ seen in rivers and lakes.

 If there are extant west Eurasian or African river dolphins, they should look like this. This is a hypothetical species, modelled on the American  Inia  and Asian  Platanista . Image: Darren Naish.

If there are extant west Eurasian or African river dolphins, they should look like this. This is a hypothetical species, modelled on the American Inia and Asian Platanista. Image: Darren Naish.

An endoparasitic tetrapod. Tetrapods have become parasites on several occasions. Vampire bats are parasites of birds and mammals, and it’s even been argued that some blood- and milk-eating human populations can be considered parasites of the mammals they rely on (though the mammals concerned are domesticated, so it’s complicated). Elsewhere among vertebrates, everyone knows about the parasitic catfishes that invade the gills of other actinopterygian fishes and even the urethras of mammals; less familiar is the fact that other actinopterygians can, on rare occasion, become trapped inside the bodies of other vertebrates and then make a successful living. Yes, you read that right. I have in mind the case where two Snubnosed or Pugnose eels Simenchelys parasitica were discovered living inside the heart of a mako shark (Caira et al. 1997; see also Eagderi et al. 2016). This eel is not – despite its name* – ordinarily an internal parasite: this was a case of facultative endoparasitism!

 At left: a snubnosed eel found living inside the heart of a shark. Eels are not tetrapods, it’s true. But here’s evidence that aquatic vertebrates can become endoparasites. Image: Caira  et al . (1997). At right: an aquatic typhlonectid caecilian. Surely it’s only a matter of time before we discover an endoparasitic one of those as well. Image: Neil Phillips.

At left: a snubnosed eel found living inside the heart of a shark. Eels are not tetrapods, it’s true. But here’s evidence that aquatic vertebrates can become endoparasites. Image: Caira et al. (1997). At right: an aquatic typhlonectid caecilian. Surely it’s only a matter of time before we discover an endoparasitic one of those as well. Image: Neil Phillips.

There are all kinds of reason why a tetrapod couldn’t become an endoparasite, respiration being high on the list. A hypothetical endoparasitic tetrapod would have to be small, with remarkable tolerance of unusual chemical and thermal conditions, with low oxygen requirements, and most likely with the ability to respire cutaneously or via gills. In other words, it should be the world’s weirdest caecilian. As if caecilians aren’t weird enough, I’d love there to be small, endoparasitic caecilians. Given that some caecilians are already aquatic gill-breathers that will consume the tissues of fish (exhibit A: the sequence from River Monsters where Jeremy Wade discovers swarming typhlonectid caecilians in the carcass of a large fish), I predict these animals to be aquatic, South American species that parasitise actinopterygians and aquatic mammals, like Inia the river dolphin.

* Snubnosed eels were given the name ‘parasitica’ because they opportunistically latch on to the bodies of larger fish and eat away at the flesh. They were not thought to ever be proper internal parasites prior to 1992.

And that’s where I’ll stop for now. I actually came up with a list containing numerous additional ‘wish list’ animals but time is against me. Maybe I’ll cover them in another article. Whatever, this was all a bit of fun and I hope you enjoyed it.

For TetZoo articles relevant to the issues covered here, see…

PS I’m going to stop linking to the SciAm run of TetZoo articles soon, because I cannot access them at all and they’re now all but useless. They all need to be relocated to an open-access site.

Thanks to those supporting this work – and the very blog itself – via pledges at patreon. You can support what I do and see works-in-prep behind the scenes, via pledges as small as $1 per month.

Refs - -

Caira, J. N., Benz, G. W., Borucinska, J. & Kohler, N. E. 1997. Pugnose eels, Simenchelys parasiticus (Synaphobranchidae) from the heart of a shortfin mako, Isurus oxyrinchus (Lamnidae). Environmental Biology of Fishes 49, 139-144.

Conrad, J. 2008. Phylogeny and systematics of Squamata (Reptilia) based on morphology. Bulletin of the American Museum of Natural History 310, 1-182.

Eagderi, S., Christiaens, J., Boone, M., Jacobs, P. & Adriaens, D. 2016 Functional morphology of the feeding apparatus in Simenchelys parasitica (Simenchelyinae: Synaphobranchidae), an alleged parasitic eel. Copeia 104, 421-439.

Gans, C. 1974. Biomechanics: An Approach to Vertebrate Biology. J. B. Lippincott Company, Philadelphia, Toronto.

Graham, G. L. 2002. Bats of the World. St. Martin’s Press, New York.

Mackal, R. P. 1987. A Living Dinosaur? In Search of Mokele-Mbembe. E. J. Brill, Leiden.

McGowan, G. J. & Evans, S. E. 1995. Albanerpetontid amphibians from the Cretaceous of Spain. Nature 373, 143-145.

Mead, J. G. 1975. A fossil beaked whale (Cetacea: Ziphiidae) from the Miocene of Kenya. Journal of Paleontology 49, 745-751.

Naish, D. 2017. Hunting Monsters: Cryptozoology and the Reality Behind the Myths. Arcturus, London.

Simons, E. L. & Ettel, P. C. 1970. GigantopithecusScientific American 222 (1), 77-84.

Venczel, M. & Gardner, J. D. 2005. The geologically youngest albanerpetontid amphibian, from the Lower Pliocene of Hungary. Palaeontology 48, 1273-1300.

Vidal, N., Azvolinsky, A., Cruaud, C. & Hedges, S. B. 2008. Origin of tropical American burrowing reptiles by transatlantic rafting. Biology Letters 4, 115-118.

Villa, A., Blain, H.-A. & Delfino, M. 2018. The Early Pleistocene herpetofauna of Rivoli Veronese (Northern Italy) as evidence for humid and forested glacial phases in the Gelasian of Southern Alps. Palaeogeography, Palaeoclimatology, Palaeoecology 490, 393-403.

Whatever Happened to the Kabomani Tapir?

As a regular TetZoo reader, you’ll no doubt be aware of Tapirus kabomani, the (alleged) new South American tapir species named by Mario Cozzuol and colleagues in 2013 (Cozzuol et al. 2013).

  Tapirus : poster-child for TetZoo Park. Image: Patrick Murphy.

Tapirus: poster-child for TetZoo Park. Image: Patrick Murphy.

T. kobamani – popularly termed simply the Kabomani tapir (the names Little black tapir or Black dwarf tapir are also available; read on) – was described as smaller and darker than other South American tapirs, and as possessing a few distinctive osteological features, like a broad forehead and frontal bones that are more inflated than those of other tapirs. Its validity as a distinct species is, however, controversial, as we’ll see.

 There’s TetZoo HAVE YOU HEARD ABOUT THE NEW TAPIR merchandise.   Go here.

There’s TetZoo HAVE YOU HEARD ABOUT THE NEW TAPIR merchandise. Go here.

I wrote about the initial naming and description of T. kabomani back here at TetZoo ver 3 (warning: now paywalled). Immediately there was a reasonable amount of scepticism from other zoologists familiar with tapirs and their biology, distribution and systematics – many of whom argued that T. kabomani was similar enough to the Brazilian or Lowland tapir T. terrestris to be considered conspecific with it – and also a claim that exactly the same sort of tapir had already been described by another zoologist (namely, Marc van Roosmalen).

 One of several  T. kabomani  images captured by remote cameras: from  Cozzuol  et al . (2013) . Image:  Cozzuol  et al . (2013) .

One of several T. kabomani images captured by remote cameras: from Cozzuol et al. (2013). Image: Cozzuol et al. (2013).

But what has happened since 2013? Well, quite a bit: several studies evaluating the status of T. kabomani have been published since Cozzuol et al.’s initial paper of 2013. What do these various studies state, and what do they conclude? Let’s look at each of the studies in turn. I’ve done my best to summarise the relevant papers, and to keep my summaries brief.

  • As noted above, Marc van Roosmalen has been stating right from the time that kabomani was first described that it is identical with another alleged small tapir – T. pygmaeus, termed the Black dwarf tapir by van Roosmalen – that he named (online) in 2002, briefly diagnosed and described in popular books of 2008 and 2013, and formally published within a section of another book of 2013 (Van Roosmalen & Van Hooft 2013). Van Roosmalen (2014) petitioned the ICZN (case 3650) to have the 2013 book in question – Barefoot Through the Amazon – On the Path of Evolution – registered as part of the ‘Official List of Works Approved as Available for Zoological Nomenclature’, and thus to have T. pygmaeus endorsed as a valid, official name with priority over T. kabomani. A ruling on this proposal has not been made, to my knowledge. Van Roosmalen again stated his view that these two forms are synonymous in a 2015 publication (Van Roosmalen 2015).

 The Black dwarf tapir, as illustrated in Marc van Roosmalen’s 2013 book. Image: van Roosmalen (2013).

The Black dwarf tapir, as illustrated in Marc van Roosmalen’s 2013 book. Image: van Roosmalen (2013).

  • Robert Voss et al. (2014) were highly critical of Cozzuol et al.’s (2013) case for the validity of T. kabomani, literally starting their paper with reference to Carl Sagan’s aphorism that “extraordinary claims require extraordinary evidence” (p. 893). To be honest, I’m not sure that the reporting of a new species of tapir is as extraordinary as they think it is but… whatever, they argued that none of the molecular, morphological or ethnological evidence compiled by Cozzuol et al. (2013) withstood scrutiny. Their most interesting contention was that kabomani is not ‘distinct enough’ from T. terrestris to warrant species-level separation (the sequence divergence in Cytb amounting to 1.3%), and that both T. kabomani and the Mountain or Woolly tapir T. pinchaque are not phylogenetically separate from T. terrestris, both failing to be recovered as reciprocally monophyletic (Voss et al. 2014). Remember that this affects T. pinchaque as well as T. kabomani: we’ll be coming back to that point.

 This section of Voss  et al .’s (2014) molecular phylogeny shows  T. pinchaque  and  T. kabomani  as poorly differentiated from  T. terrestris . Image: Voss  et al . (2014).

This section of Voss et al.’s (2014) molecular phylogeny shows T. pinchaque and T. kabomani as poorly differentiated from T. terrestris. Image: Voss et al. (2014).

  • Mario Cozzuol et al. (2014) published a response to Voss et al. (2014). With regard to Voss et al.’s (2014) contention (“Have several generations of Neotropical mammalogists really failed to recognise a species of Recent megafauna that is said to be widely distributed in Amazonia?”), their specific response was: “The answer is, simply, yes. Specifically, they failed, as many others have done for many years, to listen to the local people more carefully; those people have been aware of the existence of this species for a long time” (p. 899). Cozzuol et al. (2014) re-iterated their view that kabomani is morphologically distinct (they noted in particular its sagittal crest), and that it is recognised as distinct by various Amazonian peoples. Regarding Voss et al.’s (2014) finding that kabomani is nested within T. terrestris, Cozzuol et al. (2014) reported that different molecular trees were recovered depending on which tapir taxon was used as the outgroup, and that the nesting of kabomani within T. terrestris did not disprove its status as a distinct species given that T. pinchaque (the Mountain tapir, universally regarded as a distinct species) was found to be intractable from the terrestris-kabomani clade based on genetic data (Cozzuol et al. 2014).

 Portrait of  T. kabomani , produced by G. Braga to accompany  Cozzuol  et al .’s (2013)  original paper. Image:  Cozzuol  et al . (2013) .

Portrait of T. kabomani, produced by G. Braga to accompany Cozzuol et al.’s (2013) original paper. Image: Cozzuol et al. (2013).

  • Manuel Ruiz-García et al. (2015) examined mitochondrial gene diversity across all South American tapirs, their aim being to better understand the genetic history of the group and thus its evolution, systematics and conservation biology. They included data from five kabomani specimens: two were the Brazilian specimens analysed by Cozzuol et al. (2013), and the others were individuals from Colombia, Peru and Ecuador (Ruiz-García et al. 2015). They found kabomani tapirs to be “more closely related to T. terrestris than to the other tapir species”, reported “significant differences with [DN – I think they meant ‘from’] T. terrestris as well as with [DN – surely ‘from’] T. pinchaque” (p. 11), but overall found kabomani tapirs to “yield lower genetic distances with regard to T. terrestris than did T. pinchaque” (p. 14). In other words, kabomani tapirs were not – in their view – genetically ‘distinct enough’ to warrant species status and should more reasonably be considered a distinct population of T. terrestris. This view was modified later in the text where they specifically stated (and depicted) kabomani tapirs as a clade within T. terrestris, the divergence of the kabomani lineage being suggested to have occurred between 1.3 million and 360,000 years ago (p. 18). Such a divergence date is young relative to other divergences between extant tapir species (some of you might recall date of divergence being deemed relevant in the debate surrounding white rhino phylogeny and taxonomy). These authors also provided a lengthy critique of anatomical criteria used by Cozzuol et al. (2013) to differentiate T. kabomani, arguing that some ‘kabomani-type’ tapirs did not have the morphological features supposedly diagnostic for this taxon, and furthermore than some small, ‘kabomani-type’ animals are not of the genetic kabomani group. Bottomline: kabomani “is a particular lineage within T. terrestris” (p. 34), and it is not morphologically well differentiated from the rest of T. terrestris, some other populations of which look kabomani-like.

 Ruiz-García  et al .’s (2015) maximum likelihood tree, incoporated mitochondrial gene data for 93 tapir specimens.  T. pinchaque  is blue,  T. terrestris  is red,  T. kabomani  is green, and  T. bairdii  is purple. Note that  kabomani  is nested within  T. terrestris . Image: Ruiz-García  et al .’s (2015).

Ruiz-García et al.’s (2015) maximum likelihood tree, incoporated mitochondrial gene data for 93 tapir specimens. T. pinchaque is blue, T. terrestris is red, T. kabomani is green, and T. bairdii is purple. Note that kabomani is nested within T. terrestris. Image: Ruiz-García et al.’s (2015).

  • Dumbá et al. (2018) analysed skull shape variation in living tapir species (though they also included a few fossil ones too), and specifically analysed kabomani. They found kabomani skulls to have some overlap in morphospace with T. terrestris though noted that both could still be distinguished, in part because of kabomani’s broad forehead. In some landmark-based analyses, the kabomani sample overlapped almost entirely with the region of morphospace occupied by T. terrestris. The assumption of this study appears to be that kabomani is distinct and only similar to T. terrestris when certain sets of cranial landmarks are used (note that Mario Cozzual is the study’s last author). Another interpretation could be that kabomani overlaps so extensively with T. terrestris that it should be regarded as a poorly differentiated ‘extension’ of the morphospace occupied by T. terrestris.

 The landmark-based morphometric work published recently by Dumbá  et al . (2018) shows  T. kabomani  to overlap quite extensively with the morphospace occupied by  T. terrestris . Image: Dumbá  et al . (2018).

The landmark-based morphometric work published recently by Dumbá et al. (2018) shows T. kabomani to overlap quite extensively with the morphospace occupied by T. terrestris. Image: Dumbá et al. (2018).

What, then, to conclude? By now you’ve surely heard, on a great many occasions, the contention that the entities we call ‘species’ do not have a consistent, well defined definition across all tetrapods, let alone across all animals or all organisms. Whether a given population warrants recognition as a ‘species’ is still, to some considerable degree, a subjective issue. Having gotten that caveat out of the way, most (note: most) mammalogists would agree that the minor molecular and morphological differences separating the Kabomani tapir from T. terrestris are not compelling, and it does appear most likely that it is a variant, morph or lineage of T. terrestris. However…

 The Mountain or Woolly tapir was posited by  Cozzuol  et al . (2013)  as closer to  T. terrestris  than is  T. kabomani , but the inverse was recovered by Ruiz-García  et al . (2015). Image: Just Chaos, CC BY-SA 2.0 (original  here ).

The Mountain or Woolly tapir was posited by Cozzuol et al. (2013) as closer to T. terrestris than is T. kabomani, but the inverse was recovered by Ruiz-García et al. (2015). Image: Just Chaos, CC BY-SA 2.0 (original here).

Problem 1: Voss et al.’s (2014) initial claim that the Mountain tapir is as much as part of T. terrestris as is kabomani was always problematic. After all, everyone agrees that the Mountain tapir should be retained as a valid species; if the Mountain tapir were to be regarded as part of T. terrestris, people would likely use special pleading to keep it distinct… which would, in turn, then negate the claim that kabomani was not worthy of species status. However, the more comprehensive analysis compiled by Ruiz-García et al. (2015) seems to have resolved this issue. Problem 2: the implication that the recovery of kabomani as a lineage within T. terrestris automatically negates its status as a species is not entirely fair or technically correct, since there are a great many animal populations we term ‘species’ that are not monophyletic and/or not outside other populations also termed ‘species’. A classic example is the Polar bear Ursus maritimus (generally found in studies to be nested within the Brown bear U. arctos.) but there are many, many others. In other words, finding kabomani to be a lineage within T. terrestris does not automatically negate a species-level status. But is it ‘distinct enough’ to be regarded as a ‘species’ all its own? My conclusion from all the work discussed above: no, no it is not, alas.

  Tapirus terrestris  is a pretty variable animal, seemingly with a complex evolutionary history and a degree of morphological variation that’s only now beginning to come to light. This captive individual is from Chester Zoo, UK. Image: Darren Naish.

Tapirus terrestris is a pretty variable animal, seemingly with a complex evolutionary history and a degree of morphological variation that’s only now beginning to come to light. This captive individual is from Chester Zoo, UK. Image: Darren Naish.

And that is where we end for now. Tapirs have been covered quite a few times on TetZoo now, though once again I will note that many of the articles concerned are now paywalled due to a recent decision made at Scientific American blogs. If you can access them, the articles are here…

Thanks to those supporting this work – and the very blog itself – via pledges at patreon. You can support what I do, and see works-in-prep behind the scenes, via pledges as small as $1 per month.

Refs - -

Cozzuol , M. A., Clozato, C. L. , Holanda, E. C., Rodrigues, F. H. G., Nienow, S., de Thoisy, B., Redondo, R. A. F. & Santos, F. R. 2013. A new species of tapir from the Amazon. Journal of Mammalogy 94, 1331-1345.

Cozzuol, M. A., de Thoisy, B., Fernandes-Ferreira, H., Rodrigues, F. H. G. & Santos, F. R. 2014. How much evidence is enough evidence for a new species? Journal of Mammalogy 95, 899-905.

Dumbá, L. C. C. S., Parisi Dutra, R. & Cozzuol, M. A. 2018. Cranial geometric morphometric analysis of the genus Tapirus (Mammalia, Perissodactyla). Journal of Mammalian Evolution https://doi.org/10.1007/s10914-018-9432-2

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.

Van Roosmalen, M. G. M. 2014. Case 3650: Tapirus pygmaeus Van Roosmalen & Van Hooft in Van Roosmalen, 2013 (Mammalia, Perissodactyla, TAPIRIDAE): proposed confirmation of availability of the specific name and of the book in which this nominal species was proposed. Bulletin of Zoological Nomenclature 71, 84-87.

Van Roosmalen, M. G. M. 2015. Hotspot of new megafauna found in the Central Amazon (Brazil): the lower Rio Aripuanã Basin. Biodiversity Journal 6, 219-244.

Van Roosmalen, M. G. M. & Van Hooft, P. 2013. New species of living tapir, the dwarf tapir (Mammalia: Tapiridae) from the Brazilian Amazon, in Van Roosmalen, M. G. M. (ed), Barefoot Through the Amazon – On the Path of Evolution. CreateSpace, North Charleston SC, pp. 400-404.

Voss, R. S., Helgen, K. M. & Jansa, S. A. 2014. Extraordinary claims require extraordinary evidence: a comment on Cozzuol et al. (2013). Journal of Mammalogy 95, 893-898.

Avocets in Flight and Phylogeny

Avocets are a small group of long-legged, long-billed wading birds, all four species of which are boldly patterned in black and white. The American avocet Recurvirostra americana and Red-necked avocet R. novaehollandiae also have a reddish or brown head and neck. All avocets are included in the single genus Recurvirostra. Avocets are present on all continents except Antarctica, are absent from cold northern parts of Eurasia and North America, and are associated with habitats ranging from estuaries, lagoons, sewage ponds and salt-pans to ponds and rivers.

 Pied avocets (and Black-tailed godwits) in flight. Image: Bernie Dempsey.

Pied avocets (and Black-tailed godwits) in flight. Image: Bernie Dempsey.

I don’t think I’ve ever written about avocets before. The article here came about because I received permission to use the photos you see here from my long-standing friend Bernie Dempsey. Avocets mostly nest in colonies, though solitary nesting is practised as well; they also tend to be gregarious outside the breeding season. The photos here show large numbers of Pied avocet R. avosetta taking flight at Brownsea, southern England, due to the presence of a hunting peregrine. Black-tailed godwits Limosa limosa (and a single gull) are visible in flight as well. Having mentioned breeding, avocets are famous for performing elaborate social breeding ceremonies where groups form circles, raise and lower their bills in ritualised fashion, and often engage in fights and scuffles.

 Another shot of the same group of Pied avocets (and Black-tailed godwits) in flight. Image: Bernie Dempsey.

Another shot of the same group of Pied avocets (and Black-tailed godwits) in flight. Image: Bernie Dempsey.

The fine, upcurved avocet bill is used in ‘skimming’ or ‘scything’ across the surface of water. The birds do this while wading, typically in water only as deep as their ankles. Small food items (mostly molluscs, crustaceans and aquatic insects) are retained on internal lamellae, and the large, fleshy tongue is (presumably) used to remove said items once enough have been collected. The bill is apparently more strongly curved in females than in males: another example of bill dimorphism of the sort I’ve written about before (see this ver 2 article) and perhaps present as a consequence of resource partitioning. As always with animals, avocets are not limited to this form of foraging alone, however: they will also swim and up-end to reach food from a submerged muddy substrate and will also grab insects from terrestrial plants.

 A Pied avocet, photographed at Minsmere, UK. Note the highly reduced hallux. Image: Tim Felce, CC BY-SA 2.0 ( original here ).

A Pied avocet, photographed at Minsmere, UK. Note the highly reduced hallux. Image: Tim Felce, CC BY-SA 2.0 (original here).

Avocets might best be regarded as weird plovers. Avocets are closely related to a similar group of long-legged waders: the stilts Cladorhynchus and Himantopus. Stilts are very avocet-like but differ in having a straight bill. Anyone familiar with bird taxonomy will know that bird groups that look as different from one another as avocets and stilts do normally get given their own ‘families’, but the tradition here is that both are combined within Recurvirostridae, avocets being Recurvirostrinae and stilts being Himantopodinae. However, not only does a ‘subfamily-level’ classification seem somewhat pointless given that we’re only talking about three genera, it is by no means clear that stilts are monophyletic, since some studies find Cladorhynchus (the banded stilt) to be outside a Himantopus + Recurvirostra clade (Christidis & Schodde 1992, Dove 2000).

Having mentioned Cladorhynchus within the context of phylogeny, I have to note that this bird was used by Olson & Feduccia (1980) to support their idea that flamingos “evolved directly from the Recurvirostridae”. This view (which was never that popular among palaeornithologists anyway) now has to be considered compellingly falsified. I might write about this whole story some other time.

 A selection of birds included within the charadriiform group Charadrii, chosen to represent most (though not all) main lineages. This is a montage produced for   in-prep textbook  The Vertebrate Fossil Record   . Image: Darren Naish.

A selection of birds included within the charadriiform group Charadrii, chosen to represent most (though not all) main lineages. This is a montage produced for in-prep textbook The Vertebrate Fossil Record. Image: Darren Naish.

As goes the larger view, recurvirostrids are charadriiforms (that is, part of the same group as plovers, gulls and auks), and specifically part of Charadrii, the charadriiform group that contains sheathbills, stone-curlews or thick-knees, ibisbills, oystercatchers and plovers. While conventionally considered a distinct ‘family’, note that Recurvirostridae is one of several long-billed groups within Charadrii that’s surrounded in phylogenies by taxa traditionally grouped together within the plover family Charadriidae (Chu 1995, Ericson et al. 2003, Baker et al. 2007, Mayr 2011). No one has acted on this yet, even though the most ‘sensible’ solution is to sink Recurvirostridae and the other affected groups (Ibidorhynchidae and Haematopodidae) into Charadriidae. The alternative solution is to raise several lineages to ‘family' level’ such that oystercatchers, recurvirostrids and so on get to retain ‘family status’. Whatever, the interesting thing here are goes charadriiform evolution is that the very long-billed recurvirostrids, oystercatchers and ibisbills all evolved from short-billed ancestors (or a single ancestor?) that would have been similar to plovers, stone-curlews or sheathbills. Maybe we’d guess this already, but it receives firm support from the topology of cladograms. I’ve written a fair bit about this issue (about the topology and taxonomy of these groups) in my in-prep textbook, and the cladogram you see here is from that work (you can see more from the book and my other projects here at patreon).

 There are several competing cladograms for Charadrii: this one is a simplified version of the topology recovered by Mayr (2011). Like all of these sorts of cladograms, this one was produced for my in-prep textbook…   on which go here  .  Pluvianus  is missing simply because I haven’t drawn it yet. Image: Darren Naish.

There are several competing cladograms for Charadrii: this one is a simplified version of the topology recovered by Mayr (2011). Like all of these sorts of cladograms, this one was produced for my in-prep textbook… on which go here. Pluvianus is missing simply because I haven’t drawn it yet. Image: Darren Naish.

The restoration of Pied avocets in Britain. Here in the UK, Pied avocets (which we typically term just ‘avocets’) are famous for virtually disappearing from the country due to wetland reclamation followed by egg collecting and shooting. They were gone as a breeding species for about 100 years. During the 1940s, the birds began breeding again due to the modification of Minsmere in Suffolk to its original flooded condition (this flooding being a WWII defensive measure). Competition and predation from gulls initially kept breeding success low but removal of gull nests and eggs eventually allowed avocet breeding success to increase; another drop during the 1970s – this time caused by a decline in invertebrate prey – was countered by waterway management (Thomas 1985).

 The modern RSPB logo. Image: fair use.

The modern RSPB logo. Image: fair use.

This determined conservation effort was a great success and a good example of how systems (which were already heavily modified by human action) have to be managed and manipulated extensively for an endangered species to be bought back from the brink. It explains why the Pied avocet was adopted in 1955 as the symbol for the RSPB (Royal Society for the Protection of Birds). This is apparently the first use of a bird image as a symbol for conservation.

 Pied avocets and Black-tailed godwits (and a gull), alarmed by a nearby peregrine. Image: Bernie Dempsey.

Pied avocets and Black-tailed godwits (and a gull), alarmed by a nearby peregrine. Image: Bernie Dempsey.

Charadriiform birds have been covered a few times at TetZoo before (wow… I really thought I’d written about waders - that is, scolopacids - a great deal more than this). My issue with providing lists of links like this is that many of these articles are now paywalled. Support me at patreon and assist my efforts in getting them transferred to ver 4 (where paywalling will never occur)…

Refs - -

Baker, A. J., Pereira, S. L. & Paton, T. A. 2007. Phylogenetic relationships and divergence times of Charadriiformes genera: multigene evidence for the Cretaceous origin of at least 14 clades of shorebirds. Biology Letters 3, 205-209.

Christia, P. D., Christidis, L. & Schodde, R. 1992. Biochemical systematics of the Charadriiformes (shorebirds): relationships between the Charadrii, Scolopaci and Lari. Australian Journal of Zoology 40, 291-302.

Chu, P. C. 1995. Phylogenetic reanalysis of Strauch’s osteological data set for the Charadriiformes. The Condor 97, 174-196.

Dove, C. J. 2000. A descriptive and phylogenetic analysis of plumalaceous feather characters in Charadriiformes. Ornithological Monographs 51, 1-163.

Ericson, P. G. P., Envall, I., Irestadt, M. & Norman, J. A. 2003. Inter-familial relationships of the shorebirds (Aves: Charadriiformes) based on nuclear DNA sequence data. BMC Evolutionary Biology 3: 16.

Mayr, G. 2011. The phylogeny of charadriiform birds (shorebirds and allies) – reassessing the conflict between morphology and molecules. Zoological Journal of the Linnean Society 161, 916-934.

Olson, S. L. & Feduccia, A. 1980. Relationships and evolution of flamingos (Aves: Phoenicopteridae). Smithsonian Contributions to Zoology 316, 1-73.

Thomas, G. 1985. Plovers and sandpipers. In Perrins, C. M. & Middleton, A. L. A. (eds) The Encyclopedia of Birds. Equinox, Oxford, pp. 162-175.

TetZoo ver 3 and a Dark Day for the Dissemination of Knowledge

As most of you will know, the blog you’re reading now is the fourth iteration of Tetrapod Zoology, hence the name ‘Tet Zoo ver 4’.

 Once upon a time, you could look at blogs for free. Imagine that.

Once upon a time, you could look at blogs for free. Imagine that.

And as most of you will also know, ver 3 was hosted at Scientific American blogs where things were good for a while but ended up being not so good eventually. The reason for this short article is that an unusual and very bad thing has just happened: Scientific American blogs have just (within the last several days) switched to a ‘subscription only’ plan whereby readers will only see their blog material if they’re paid-up subscribers to SciAm’s online content. The first I knew of this was when I visited one of my ver 3 articles, only to see this…

 In the background - behind the pop-up with the hilariously appropriate feature about  money  - you should be able to see that I was trying to access my 2015 ver 3 article on cassowaries.

In the background - behind the pop-up with the hilariously appropriate feature about money - you should be able to see that I was trying to access my 2015 ver 3 article on cassowaries.

Other readers report that this does not yet affect them (and that they’re able to see ver 3 material just fine), and others say that they’re seeing messages that they “have x complimentary viewings left”. I consider this a total disaster as goes the dissemination of information and find it shameful that SciAm has gone down this route. I’m not about to argue with anyone there, since it was clear to me during my time at SciAm that decisions like this were made high up and typically by people who have no direct interaction with the team involved with blogging. But at a time when humanity needs all the science-themed communication it can get, this is really bad news.

 A montage depicting things covered in just a few of the TetZoo ver 3 articles, all of which are now safely locked away and only available to you if you own a SciAm subscription.

A montage depicting things covered in just a few of the TetZoo ver 3 articles, all of which are now safely locked away and only available to you if you own a SciAm subscription.

I have yet to dig out my SciAm contract and see exactly what the deal is as goes use of my own material (I’m going through an exceptionally difficult time as goes workload at the moment), but my recollection is that TetZoo material is mine, and that I can do what I want with it once a short (one or two month) grace period has passed. In view of that, my aim is to migrate ver 3 material to here: many thanks to those who’ve helped salvage the material concerned (I now have all the text and images, but not the comments). This takes time that I don’t have… whatever, I’ll prioritise those articles that are ‘most valuable’ as goes online presence, and herein we find the reason for this article: dear reader, which articles are ‘most valuable’? Let me know your thoughts in the comments below. Possibilities high on my agenda include…

  • The multi-part guide to crocodile diversity, evolution and systematics

  • The multi-part guide to Australian agamid lizards

  • Brian J Ford “I’m smarter than everyone else, all palaeontologists are idiots and all dinosaurs were aquatic” smackdown

  • Domestic horses of Africa

  • Raymond Hoser smackdown

  • The “why the world needs to ignore pseudoscientist and self-promoter extraordinaire David Peters and his Pterosaur Heresies and ReptileEvolution sites” article

  • The multi-part guide to the world’s petrels

  • On why humans and other apes are actually monkeys

  • No trunks for sauropods

  • Ornithoscelida debut

  • Piltdown man and the dualist connection

Ok, thoughts appreciated. If all of this turns out to be a mistake, and access to the articles is reverted to its previously open status, I can relax a bit. But for now, it really looks like everything has to be moved. Not good when your stated aim (my stated aim; evidently not that of SciAm) is the dissemination and sharing of knowledge, not its locking away.

 This image appeared in the very first TetZoo ver 3 article of July 2011. Oh god… you mean I have to back up  eight years worth  of old articles? Great. Good job I have nothing else to do. Image: Darren Naish.

This image appeared in the very first TetZoo ver 3 article of July 2011. Oh god… you mean I have to back up eight years worth of old articles? Great. Good job I have nothing else to do. Image: Darren Naish.

Oh – and… to those of you who immediately say “don’t worry, the articles can still be found on wayback machine” or “don’t worry, the articles can still be seen if you open them in incognito mode” or whatever… please remember that these things are not much use to those who want information quickly and will give up immediately if a page doesn’t give them what they want (viz, 99% of internet users).

TetZooCon 2018: Best TetZooCon So Far

We did it… and survived. In fact, it was an all-round success (pretty much; see below for caveats). Yes, the TetZoo-themed event of the year – TetZooCon 2018, organised by myself and World’s Joint Most Influential Palaeoartist John Conway – has just happened and I’m now back at home and buried in all the other work I managed to avoid by organising a grand, two-day conference and associated fieldtrip. TetZooCon has now been going for five years, and as the fifth of these events, this one felt a little bit special.

 TetZooCon 2018: the first one to be more like an actual convention, or conference. Left to right: Caitlin Kight, Hanneke Meijer, Darren Naish. Image: Xane/Michael Lesniowski.

TetZooCon 2018: the first one to be more like an actual convention, or conference. Left to right: Caitlin Kight, Hanneke Meijer, Darren Naish. Image: Xane/Michael Lesniowski.

Indeed, TetZooCon 2018 was ambitious – possibly over-ambitious. In addition to a long list of talks, the event included a Palaeoart Workshop, a special session on talks devoted entirely to bird evolution, a bird evolution roundtable event, a Speculative Biology on-stage discussion, a quiz, a conference meal and drinks reception, and a post-conference fieldtrip. Abundant stalls with merchandise were there too; we also had numerous book signings and a few ‘show and tell’ events (relating to Dougal Dixon’s SpecBio projects and my in-prep textbook, among other things). For the second year running, Beth Windle brought along a TetZoo-themed cake; this one was devoted to the theme of plastic pollution, and a fine thing it was (even though I only got to eat a small bit of its neck and none of the actual cake). It was also a great social event with a lot of networking and pubbing going on.

 It’s a TetZooCon tradition that we make special little icons for our speakers and other presenters, and here’s the haul for 2018. Image: John Conway/Darren Naish.

It’s a TetZooCon tradition that we make special little icons for our speakers and other presenters, and here’s the haul for 2018. Image: John Conway/Darren Naish.

Virtually everything was filmed, but pressures of time and workload mean that I haven’t looked at a single bit of the footage yet. I should also add that my plan to record short interviews with people at the event (not just speakers) never panned out (even though I went round with a dictaphone and spare batteries in my pocket the whole while) as I just never had time. I should have given the job to someone else but never even thought about it. On that point, I enrolled some additional session moderators this year: thanks to Dani Rabaiotti, Beth and Georgia Harper. Georgia Witton-Maclean worked as official photographer.

 What with it being the fifth TetZooCon, the 2018 banner was a special one. Image: Xane/Michael Lesniowski.

What with it being the fifth TetZooCon, the 2018 banner was a special one. Image: Xane/Michael Lesniowski.

I will try and keep my recollections brief, otherwise it will take me several articles to get through everything that happened, and that’s not ideal. Already I’m aware that TetZoo is becoming a report of recent adventures more than a blog about tetrapods.

Our venue was The Venue (part of the University College London complex at Malet Street) once more; attendee count was somewhere around the 150 mark.

 A huge quantity of palaeoart was both on show at TetZooCon 2018, and available for sale. Part of my personal haul from the meeting is shown at right. More on palaeoart below. Images: Darren Naish.

A huge quantity of palaeoart was both on show at TetZooCon 2018, and available for sale. Part of my personal haul from the meeting is shown at right. More on palaeoart below. Images: Darren Naish.

Baleen whales, music in documentaries, Palaeoloxodon. After a brief intro in which John and I bigged up the fact that we were at THE FIFTH TETZOOCON, talks kicked off with polar biologist, geneticist and whale expert Jennifer Jackson. This was a remarkable and very well illustrated review of everything about baleen whale history you might imagine: their origins and phylogeny, competing views on the taxonomy and systematics of extant forms, population biology and phylogeography, biogeography and historical distribution, the impact of humans on their distribution and abundance, and more! Jenn and I have a long-standing disagreement over the interpretation of a particular sea monster sighting. It’s not impossible that this issue will be thrashed-out in detail at TetZooCon one year.

 Jenn Jackson gave the most enthralling presentation on baleen whales. Image: Darren Naish.

Jenn Jackson gave the most enthralling presentation on baleen whales. Image: Darren Naish.

In one of the most innovative and unusual talks of the event, professional composer Fiona Taylor discussed music for wildlife documentaries. This included a background to ideas, disagreements and arguments that have occurred around wildlife documentary soundtracks (“there are no bassoons in the Serengeti”, the Guardian reminds us) as well as demonstrations and explanations of how music can work (and not work), how it can be used (and mis-used), and how it can convey specific emotions or themes. It’s a huge relief to me that everything worked fine with the audio system during Fiona’s talk – it failed to work at all on the Sunday, on which more later. Anyway: my suspicion was high that Fiona’s talk would be really fun and interesting, and I’m pleased that I was right. And I’m not too ashamed to admit that I teared up at the piece of music accompanying the Sad Wolf. Oh… extra points, Fiona, for the Bad Wolf references.

 Fiona Taylor discusses the use of music in wildlife documentaries, and also shows us how it’s done. Image: Darren Naish.

Fiona Taylor discusses the use of music in wildlife documentaries, and also shows us how it’s done. Image: Darren Naish.

Steven Zhang gave us a specialist’s view of where we’re at with thoughts on the taxonomy, phylogeny, anatomy and palaeobiology of the straight-tusked elephant Palaeoloxodon. Coincidentally (…. or was it?), one of the prizes we had for our quiz was the new Eofauna Palaeoloxodon model.

 Steven Zhang talks  Palaeoloxodon  - specifically, at this point, about the alleged survival of this animal into the Holocene (the evidence isn’t great). Image: Georgia Witton-Maclean.

Steven Zhang talks Palaeoloxodon - specifically, at this point, about the alleged survival of this animal into the Holocene (the evidence isn’t great). Image: Georgia Witton-Maclean.

Bird evolution roundtable event. Our next event was the roundtable session on bird evolution. I was joined on stage by Albert Chen, Caitlin Kight, Hanneke Meijer, Robyn Womack and Glyn Young (all of whom were scheduled to give bird-themed talks the following day) as we discussed breaking news, current events, on-going research and future prospects relevant to bird evolution. We will definitely be doing roundtables again. When discussing the evolution of Strisores (the bird clade that includes swifts, hummingbirds and nightjars), Albert mentioned in passing that a hummingbird is “a dinosaur trying to be a butterfly”, an evocative line that struck a chord with the audience and even became a meme during the conference.

 A scene from the bird evolution roundtable event. From left to right: Albert Chen, Caitlin Kight, Hanneke Meijer. Image: Xane/Michael Lesniowski.

A scene from the bird evolution roundtable event. From left to right: Albert Chen, Caitlin Kight, Hanneke Meijer. Image: Xane/Michael Lesniowski.

Lucy Cooke and Katrina van Grouw. The excellent and hilarious Lucy Cooke was up next, speaking about her new book The Truth About Animals … aka The Unexpected Truth About Animals (Cooke 2018), depending on whether you obtain the UK or US edition. Lucy has a background in TV and film-making but switched track to bring attention to amphibian conservation. Today she writes about sloths, giant pandas, bats and all manner of other animals. It was a brilliant talk and definitely a highlight. Lucy was on hand afterwards to sign and sell copies of her book.

 Lucy Cooke (at right) signs copies of  The Unexpected Truth About Animals . Image: Darren Naish.

Lucy Cooke (at right) signs copies of The Unexpected Truth About Animals. Image: Darren Naish.

Following Lucy was Katrina van Grouw on another book-themed talk, this one devoted to her fantastic Unnatural Selection (van Grouw 2017). Selective breeding is very much a perfect illustration of evolution in action, and Katrina took us through remarkable examples from the world of pigeons, poultry, dogs, pigs, cats and other animals. The hamster gag was possibly inspired by a similar fish-themed gag employed by that Tetrapod Zoology guy. Katrina was also selling and signing her book. A brief initial take on Katrina’s book has already appeared here at Tet Zoo.

 Katrina van Grouw and her fabulous book  Unnatural Selection  at TetZooCon 2018. Image: Xane/Michael Lesniowski.

Katrina van Grouw and her fabulous book Unnatural Selection at TetZooCon 2018. Image: Xane/Michael Lesniowski.

The Palaeoart Workshop – led by John Conway – featured talks from Luis Rey and Mark Witton in addition to an art-making event in which participants were invited to depict prehistoric animals in an unfamiliar style. Bob Nicholls was also in attendance. Unusual and often attractive artwork that resulted from the workshop was up on the walls for the duration – we sure do have a lot of skilled arty-types among us, but then we already know this from previous TetZooCons. Luis and Mark also had art on sale at the event, and Mark’s brand-new book The Palaeoartist’s Handbook (Witton 2018) (which I hope to see properly some time soon… hint hint) was the object of much discussion and interest at the meeting. I should also mention that Joschua Knüppe – he of palaeostream and much else new palaeoart-themed goodness online – was in attendance and showing people original art he’d brought with him. I was carrier for one of the few printed copies of Joschua’s new #Palaeostream: Sketches of Prehistoric Life book (Knüppe 2018), which is wonderful.

 Bob Nicholls of Palaeocreations was selling prints, including those featuring the cover and concept art for Naish & Barrett’s  Dinosaurs: How They Lived and Evolved . Image: Xane/Michael Lesniowski.

Bob Nicholls of Palaeocreations was selling prints, including those featuring the cover and concept art for Naish & Barrett’s Dinosaurs: How They Lived and Evolved. Image: Xane/Michael Lesniowski.

In order to pack in more stuff this year, we opted to have the palaeoart workshop running as a parallel stream to the rest of the conference. Not ideal, and it means that many of us – myself included – didn’t get to go as we stayed in the main hall for the other talks, but there you go. A very good run-down of what happened can be found in Marc Vincent’s article at LITC, if you’re interested.

 A selection of palaeoartists present at TetZooCon 2018 (and this isn’t all of them). Left to right: Luis Rey, John Conway, Bob Nicholls, Steve White, Mark Witton and Rebecca Groom. Image: Georgia Witton-Maclean.

A selection of palaeoartists present at TetZooCon 2018 (and this isn’t all of them). Left to right: Luis Rey, John Conway, Bob Nicholls, Steve White, Mark Witton and Rebecca Groom. Image: Georgia Witton-Maclean.

And what was happening in the main hall at the same time? PhD student and herper extraordinaire (it says so in October’s BBC Wildlife) Steve Allain gave a really interesting (albeit slightly scary and depressing) review of snake fungal disease. Steve will be back at TetZooCon in the near future to discuss his on-going work on introduced British midwife toads, mark my words.

Ian Redmond. We were then on to my favourite talk of the meeting (no offence intended to our other very excellent speakers): Ian Redmond’s ‘The Reluctant Conservationist, 40 Years On: From Gorilla Parasites and Poachers to Virtual Safaris’. You’ll be familiar with Ian and his work if you know anything about primates, elephants, rhinos, or conservation in general. I first got to know of Ian due to his association with the late Dian Fossey in the Virunga Mountains of Rwanda, and maybe you did too; he was also on hand to advise David Attenborough and his team during the 1978 filming of that iconic scene with the mountain gorillas. Ian was also involved with the making of the 1988 movie Gorillas in the Mist.

 Ian Redmond talks about gorillas, Dian Fossey and conservation at TetZooCon 2018. Image: Xane/Michael Lesniowski.

Ian Redmond talks about gorillas, Dian Fossey and conservation at TetZooCon 2018. Image: Xane/Michael Lesniowski.

The talk itself was fantastic. Ian had brought with him his original field notes and read aloud the section relevant to what occurred 40 years ago on the same date. As it happened, it was a particularly eventful day, since Ian and his colleagues discovered, confronted and apprehended a group of poachers setting snares. He also discussed his work with the famous mining elephants of Mount Elgon and his encounters with forest elephants in general, his first-hand inter-species friendships with individual gorillas, and his on-going work on the preservation of tropical forests, a vital resource as goes the health of the entire planet and the function of our atmosphere and water cycle. Teaching local children the value of their gorillas and other wildlife is an essential part of current work. A small excerpt of Ian’s talk was filmed (very poorly) on my phone and streamed live: I’m hoping that we have the entire talk as I very much want to see it again. You can find out more about Ian’s VR work at www.vEcotourism.org, documentaries on www.ecostreamz.com and follow him on Twitter at @4apes.

 Ian Redmond, Darren Naish and Archie the Elephant (follow him on twitter  @ArchietheEle ) at TetZooCon 2018. Image: Darren Naish.

Ian Redmond, Darren Naish and Archie the Elephant (follow him on twitter @ArchietheEle) at TetZooCon 2018. Image: Darren Naish.

Mark O’Shea and forensic historical herpetology. Ian’s talk was followed by one from another celebrity of the zoological world: herpetologist, author, explorer and conservationist Mark O’Shea. Mark’s talk was on forensic historical herpetology: on cases in which he (working with colleague Hinrich Kaiser) worked hard to track down the true geographic origins of worm-eating snake specimens that didn’t otherwise seem right given other knowledge of the group’s distribution, taxonomy and biogeography. Fascinating stuff.

 Mark O’Shea at TetZooCon 2018: Mark certainly won the prize for longest talk title. Image: Darren Naish.

Mark O’Shea at TetZooCon 2018: Mark certainly won the prize for longest talk title. Image: Darren Naish.

Mark also brought posters (pressures of running a conference meant I never got to have a proper look at them) and stock of his brand-new The Book of Snakes (O’Shea 2018). I purchased a signed copy, and it’s an amazing piece of work. I admire the format, design and fact that he’s consistently said interesting things about the 600 featured species. Yes, the photos are life-sized but this means (obviously) that – in many cases – it’s only the animal’s head and adjacent loop of its body that’s shown at full size, not the whole snake.

  Mark O’Shea’s 2018  The Book of Snakes  , a must-have for those interested in reptiles. Image: Darren Naish.

Mark O’Shea’s 2018 The Book of Snakes, a must-have for those interested in reptiles. Image: Darren Naish.

For the first time – yeah, there were a lot of firsts at this one – we finished the evening with both a drinks reception (which didn’t go exactly to plan but was still a thing) and a conference meal. The latter was popular enough (as in, a sufficient number of people wanted to come along) that we could potentially have booked another 20 or more places and filled them. An alternative meal was arranged by Beth Windle and places there were filled as well. I can’t remember what happened after the meal, but I know we got back home. So… so far so good, we’d survived the first day, and it had worked pretty well.

Bird evolution session. And so to Sunday. Due to a stupid mistake that neither John nor I caught in time, our schedule had things kicking off at 9am, which won’t happen again. The first several talks of the day were devoted entirely to bird evolution: both deep-time, phylogenetic and palaeontological stuff as well as evolution in the human era. General themes of the subject had of course been outlined the day before in the roundtable session. All of the bird talks were really strong.

 Robyn Womack’s cover slide. Image: Georgia Witton-Maclean.

Robyn Womack’s cover slide. Image: Georgia Witton-Maclean.

Robyn Womack reviewed what we know about biological clocks in birds. Her research is on how birds are adapting (or not) to light pollution, the results showing that some birds are undergoing a modification in their timetables and behaviours while others are not. Albert Chen gave us an overview of crown-bird evolution: he covered bird survival and extinction across the KPg event, the shape of the neornithine tree and competing models on how crown-birds might have evolved. His formative role in TetZoo Time and special guest article at TetZoo (ver 3) were mentioned in the introduction…. and then there were the memes. I don’t want to spoil the surprises, but they were good. Ok, I have to give one away: an enantiornithine says “Mr Vegavis, I don’t feel so good”, and turns to ash.

 TetZooCon 2018 was the Albert Chen TetZooCon, and he is quite literally wearing the t-shirt. Image: Darren Naish.

TetZooCon 2018 was the Albert Chen TetZooCon, and he is quite literally wearing the t-shirt. Image: Darren Naish.

Caitlin Kight followed with ‘Sonic Doom’, another talk on the effects of anthropogenic disturbance on birds. A substantial but pretty obscure literature shows that noise pollution can result in a huge number of biological consequences for birds, and there can also be unexpected knock-on environmental effects due to how and where birds feed and forage. Caitlin also bought along copies of her very nice 2015 book Flamingo (Kight 2015); I got her to sign mine. Caitlin was followed by Hanneke Meijer, who discussed the fossil birds of southeast Asia. It’s not just giant marabou storks on Flores; she also spoke about vultures, the mysterious absence of chickens from the prehistoric sediments of the region and much else besides.

 Hanneke Meijer talks about the giant marabou storks and other birds that lived alongside the hobbits (and other mammals) of Flores in the past. The artwork is by Simon Roy. Image: Darren Naish.

Hanneke Meijer talks about the giant marabou storks and other birds that lived alongside the hobbits (and other mammals) of Flores in the past. The artwork is by Simon Roy. Image: Darren Naish.

The final bird talk – ‘What a Boring Duck: Why Southern Hemisphere Ducks Are So Dull’ – was by Glyn Young of the Durrell Wildlife Conservation Trust. Long-time TetZoo readers will know how much I love ducks (here’s the proof). A traditional view that the Northern mallard Anas platyrhynchos is the Best Duck and that all other ducks find it irresistibly magnetic (like, sexually) – promoted by Konrad Lorenz and those who followed him – is just not accurate, as established by Glyn’s research on Meller’s duck A. melleri.

 It’s thanks to Glyn Young that I have a great interest in Meller’s duck (at left is a captive individual at Bristol Zoo, biting my finger), so it was great to have Glyn himself speak about this species (and others) at TetZooCon 2018. Images: Darren Naish, Xane/Michael Lesniowski.

It’s thanks to Glyn Young that I have a great interest in Meller’s duck (at left is a captive individual at Bristol Zoo, biting my finger), so it was great to have Glyn himself speak about this species (and others) at TetZooCon 2018. Images: Darren Naish, Xane/Michael Lesniowski.

Aron Ra. Yes, I said ARON RA. Aron was at TetZooCon to discuss his new, incredibly ambitious and extremely worthy Phylogeny Explorer Project, an interactive, searchable database that is essentially an annotated, navigable, online tree of life – exactly the sort of thing the internet needs. Aron’s team of backers and colleagues were also at the meeting: the first time they’ve all been physically together in the same place, I believe. Aron discussed previous efforts to provide searchable versions of the tree of life to online public use (like the Tree of Life web project and Mikko’s Phylogeny Archive) and also explained why such a project is worthy. It was a great talk. Aron was also selling copies of his book Foundational Falsehoods of Creationism (Ra 2016).

 Aron Ra at TetZooCon 2018: at left, with his 2016 book; at right, with Naish. Images: Georgia Witton-Maclean, Darren Naish.

Aron Ra at TetZooCon 2018: at left, with his 2016 book; at right, with Naish. Images: Georgia Witton-Maclean, Darren Naish.

A Speculative Biology Discussion. The very first TetZooCon – the one of 2014 – included a talk from me on speculative zoology (you can see it here), so it only seemed fitting that the fifth of our meetings revisit the subject, this time in more ambitious fashion. Accordingly, we had an on-stage discussion featuring me, Dougal Dixon and Gert van Dijk. You’ll know who Dougal is (if not, see my interview with him here and my report on the After Man launch event of September); Gert is the author and creator of the Furaha website and the SpecBio-themed blog Furahan Biology and Allied Matters.

 From left to right: Gert van Dijk, Darren Naish and Dougal Dixon on stage during the SpecBio discussion at TetZooCon 2018. Dougal Dixon illustrations relevant to (but not included within)  After Man  are on the screen behind us. Image: Georgia Witton-Maclean.

From left to right: Gert van Dijk, Darren Naish and Dougal Dixon on stage during the SpecBio discussion at TetZooCon 2018. Dougal Dixon illustrations relevant to (but not included within) After Man are on the screen behind us. Image: Georgia Witton-Maclean.

After discussing SpecBio in general – what it is, what its appeal is, whether there’s a ‘community’ and so on – we were treated to a discussion of various of the speculative possibilities explored by Gert for Furaha. Think cloakfishes, tetropters, rusps and spidrids. Dougal followed with a discussion of his Green World project. At one point (while discussing the inherent popularity of SpecBio in general), Gert asked for a show of hands from those who had, at some point, invented a creature of their own. I took a photo…

 A view of the audience, taken from the stage. Note that most people have their hands up. Image: Darren Naish.

A view of the audience, taken from the stage. Note that most people have their hands up. Image: Darren Naish.

As mentioned above, this section of TetZooCon was enhanced by Dougal’s bringing along of various original pieces of art, imagery and content relevant to After Man, The Future Is Wild and Green World. As discussed in a previous article, some of this was on show at our recent event at Conway Hall, but only some of it. Dougal was also selling and signing copies of the 2018 edition of After Man (Dixon 2018).

 Dougal Dixon with copies of the  2018 Breakdown Press edition of  After Man  … and note the exclusive DVDs as well. Image: Georgia Witton-Maclean.

Dougal Dixon with copies of the 2018 Breakdown Press edition of After Man… and note the exclusive DVDs as well. Image: Georgia Witton-Maclean.

Dinosaurs in the Wild, the quiz, the fieldtrip. The very last talk of the day was next: it was mine on Dinosaurs in the Wild, the immersive, interactive, travelling exhibit that treated visitors to a view of life in the Cretaceous where time-travel has been invented. My aim here was to discuss the backstory to the whole thing and explain why we decided to depict the animals we did, and what decisions we had to make when reconstructing them. A short movie (and one hilarious joke) were, sadly, ruined by the fact that – for reasons beyond our control and unrelated to the function of the conference laptop – sound was no longer functioning at the venue, so I had to improvise and do the audio for the movie myself. Among other things, the talk covered tyrannosaur facial tissue, terrestrial stalking azhdarchids, cuddly mosasaurs, nose balloons in ornithischians and much else. Behind-the-scenes discussions relevant to Dinosaurs in the Wild continue, and we hope to have news at some point.

 Just some of the prizes available to quiz winners this year. Thanks to those who made the many kind donations. Image: Xane/Michael Lesniowski.

Just some of the prizes available to quiz winners this year. Thanks to those who made the many kind donations. Image: Xane/Michael Lesniowski.

Sunday also finished with a quiz. 30 questions on all manner of things relevant to the TetZooniverse, and with many of the questions relevant to issues covered in the talks of the conference. First place was tied by Albert Chen and Lars Dietz, with Albert winning a tie-breaker question (on genome size in axolotls). Our selection of amazing prizes were very generously provided by Everything Dinosaur, the team at Dinosaurs in the Wild, Katrina van Grouw and Crowood Press (publishers of Mark Witton’s The Palaeoartist’s Handbook). The quiz is never easy, but even so there are always people who get scores in the 20s, and well done and thanks to all who played along and enjoyed it.

 Well done Albert, the 2018 quiz winner. He chose the Fauna Figures bichir and Dinosaurs in the Wild  Dakotaraptor . Image: Georgia Witton-Maclean.

Well done Albert, the 2018 quiz winner. He chose the Fauna Figures bichir and Dinosaurs in the Wild Dakotaraptor. Image: Georgia Witton-Maclean.

After an evening spent in the pub… again, we got back to base at some point, and thanks to those who bought me beers… there was one thing left to do on Monday, and this was to lead the post-TetZooCon fieldtrip to Crystal palace. There was a small charge for this, the money raised being donated to the Friends of Crystal Palace Dinosaurs where it will assist with renovation work on the models. I could talk a lot about the stuff we looked at and spoke about but an article dedicated to the Crystal Palace models and to another recent event is due to appear here soon.

 The post-TetZooCon fieldtrip to Crystal Palace. It isn’t coincidental that we stopped to take the group photo at the  Megaloceros  models. Image: Will Naish.

The post-TetZooCon fieldtrip to Crystal Palace. It isn’t coincidental that we stopped to take the group photo at the Megaloceros models. Image: Will Naish.

Until next time. And that about draws things to a close. Running TetZooCon this year was extremely stressful – way more so than in previous years – and I now understand why people stop running conferences and conventions. But I don’t want to complain, because none of the problems really affected what happened and it all ran pretty smoothly in the end. We had a great crowd, a strong turnout, and a lot of positive feedback. The vendors and merchants did pretty well, book signings and sales mostly went really well (I shifted all copies of my Dinosaurs: How They Lived and Evolved, the second edition), and the talks were outstandingly good.

 Just some of the books I got signed by their authors at TetZooCon 2018. Image: Darren Naish.

Just some of the books I got signed by their authors at TetZooCon 2018. Image: Darren Naish.

It only remains for me to thank everyone who helped: huge thanks to all of our speakers and presenters, to Jenny, Will and Tilly for help, to our moderators Beth, Dani and Georgia, to the other Georgia for photography, to Xane and everyone else who took photos, shared material online and tweeted (sooo much tweeting!), to Luis, Mark, Bob and everyone else involved in the palaeoart workshop, to The Venue staff for assistance, and to everyone who attended, came on the fieldtrip and purchased stuff.

The fifth TetZooCon has happened. Now to start planning for the sixth.

For previous TetZoo articles on TetZooCon, see…

Refs - -

Cooke, L. 2018. The Unexpected Truth About Animals. Penguin Random House, London.

Dixon, D. 2018. After Man: A Zoology of the Future. Breakdown Press, London.

Kight, K. 2015. Flamingo. Reaktion Books, London.

Knüppe, J. 2018. #Palaeostream: Sketches of Prehistoric Life. Studio 252MYA.

O’Shea, M. 2018. The Book of Snakes. Ivy Books, London.

Ra, A. 2016. Foundational Falsehoods of Creationism. Pitchstone Publishing, Durham, North Carolina.

Van Grouw, K. 2017. Unnatural Selection. Princeton University Press, Princeton and Oxford.

Witton, M. 2018. The Palaeoartist’s Handbook. The Crowood Press, Marlborough.

Young, H. G. & Rhymer, J. M. 1998. Meller’s duck: a threatened species receives recognition at last. Biodiversity and Conservation 7, 1313-1323.

Reminiscing on Tetrapood Zoology: Book One…

Happy birthday, Book One

Tet-Zoo-Book-One-is-8-final-cover-1000-px-tiny-Sept-2018-Darren-Naish-Tetrapod-Zoology.jpg

There’s a huge quantity of stuff I want to – indeed, need to – blog about. But I can’t. Too much work and not enough time. But something interesting just happened and I feel it’s worthy of mention here. In September 2010, Tetrapod Zoology Book One (Naish 2010) was published (on or around the 27th, to be precise), and thank you facebook for reminding me that eight years have passed since then (yup, the maths checks out). This is painful to hear, because my intention was always to churn out additional volumes in the series every few years: the idea being that I would, by now, be four or five or more books into the series. But no. I despair over how little time I have to do the things I want, and how much there is that I want to do and still haven’t done. It’s on my mind all the time. Why couldn’t I be born rich? Huh.

 If anything might be considered emblematic of  Book One  it’s giant killer eagles. Golden eagles can and do kill domestic cattle calves. Image: Darren Naish.

If anything might be considered emblematic of Book One it’s giant killer eagles. Golden eagles can and do kill domestic cattle calves. Image: Darren Naish.

Anyway… Tetrapod Zoology Book One was well received at the time of publishing; I know of one published review (Pihlstrom 2011), and the few I’ve seen on blogs and on amazon are fair. The book feels something like a random assortment of essays on diverse tetrapod groups but at least it compiles articles (albeit not all of them) that appeared on Tet Zoo ver 1 back in 2006. Giant killer eagles, British big cats, Indian Ocean giant tortoises, the Ichthyosaur Wars, olms, azhdarchids, eagles owls in Britain, the discovery of the Kipunji, bird-eating bats and more get coverage. Steve Backshall very kindly wrote the foreword.

 A most amusing montage depicting assorted humans who assisted in some way with stuff that led to the production of the book. Images: Darren Naish, Neil Phillips, (c) University of Portsmouth, (c) Steve Backshall

A most amusing montage depicting assorted humans who assisted in some way with stuff that led to the production of the book. Images: Darren Naish, Neil Phillips, (c) University of Portsmouth, (c) Steve Backshall

The book’s existence is owed to Karl Shuker who emphasised to me in a bar one time how getting (quality, ahem) material written for a blog into actual print is worthwhile, possibly a necessity. I’m inclined to agree. Not everyone thinks this way in the age of the internet and digital storage, but I do and I remain attached to books as physical objects. I literally do not remember the ebooks I own but cannot say the same of the dead-tree objects that I associate with the physical spaces where I’ve interacted with them. Wow, that sounded weird. I later discussed the idea with Jon Downes and ended up having the book published through his CFZ Press, though I misunderstood the finances of the deal we made because I’m a freakin’ idiot. Whatever.

 No no  NO  — you do  NOT  own too many book already! BUY MORE!!! Digital books don’t work, I tell you. Image: Darren Naish.

No no NO — you do NOT own too many book already! BUY MORE!!! Digital books don’t work, I tell you. Image: Darren Naish.

A printing error (though you could describe it in another way if you wished) led to the very first batch of this book being called Tetrapood Zoology: Book One on the spine. That initial batch (of around 300 books, I think) sold out and are now much sought after – I’ve retained one copy and I know other people who won’t sell theirs because they like the typo so much. The second printing is technically a second edition (it even says so inside the book); two editions within the space of a year, woo-hoo!

 Spot the cryptic typo. Image: Dallas Krentzel.

Spot the cryptic typo. Image: Dallas Krentzel.

The excellent cover art was produced by my friend and colleague Memo Kösemen. I don’t think I’ve previously shared the fact that we went through several iterations before deciding on the final version, and here are two of them.

 Art by the brilliant C. M. K ö semen. Incidentally - and wholly coincidentally - Kevin Schreck’s movie  Tangent Realms: The World of C. M. Kösemen  premiered today in New York!

Art by the brilliant C. M. Kösemen. Incidentally - and wholly coincidentally - Kevin Schreck’s movie Tangent Realms: The World of C. M. Kösemen premiered today in New York!

That’ll do. The plan to publish follow-up volumes remains very much alive, but there are several major projects in the way before anything happens there, and there are things in the way stopping those things from happening, and yet other things too that stop those things from happening as well. I’ll get it all done one day. Or maybe I won’t. It sure feels that way at the moment. Gah.

Thank you to those who support my research and writing at patreon.

Refs - -

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

Pihlstrom, H. 2011. Book review: Tetrapod zoology book one. Historical Biology 23, 439-440.

Reasons to Attend TetZooCon 2018

The end of September is approaching, which means we’re getting worryingly close to the start of October and hence to the TetZoo-themed event of the year: TetZooCon, this year happening on Saturday 6th and Sunday 7th October (at The Venue, Malet Street, London).

 Our 2018 banner. It incorporates illustrations produced by participants in the palaeoart workshop of 2017 and looks pretty neat.

Our 2018 banner. It incorporates illustrations produced by participants in the palaeoart workshop of 2017 and looks pretty neat.

If you want to come along but haven’t yet bought a ticket, you’d better hurry up and do so, since time’s nearly up and we have to stop selling soon. Go here if you’re interested. This article is essentially a last call, and also a reminder of how much awesome stuff we have happening this year. It’s by far the biggest TetZooCon yet. So…

 My god, we have some amazing merchandise on sale this year (read on for more). Once again, Rebecca Groom will be selling her palaeoplushies — last year, the WHOLE LOT sold out before lunch. Image: (c) Rebecca Groom/palaeoplushies.

My god, we have some amazing merchandise on sale this year (read on for more). Once again, Rebecca Groom will be selling her palaeoplushies — last year, the WHOLE LOT sold out before lunch. Image: (c) Rebecca Groom/palaeoplushies.

Aron Ra – best known for his work as an atheist activist and his countering of creationists and other anti-science types – is joining us to talk about his Phylogeny Explorer Project. Aron has a vast international following and it’s a real coup to get him as a speaker.

 Two of many TetZooCon 2018 speakers. Left: Ian Redmond; right: Aron Ra. Images: (c) Ian Redmond, (c) Aron Ra.

Two of many TetZooCon 2018 speakers. Left: Ian Redmond; right: Aron Ra. Images: (c) Ian Redmond, (c) Aron Ra.

Ian Redmond – conservationist, primate and elephant expert, famous for his association with Dian Fossey, David Attenborough and the Rwandan gorillas – is speaking about his contributions to conservation and education. His talk is titled ‘The Reluctant Conservationist, 40 Years On: From Gorilla Parasites and Poachers to Virtual Safaris’.

We have an entire section of talks devoted to bird evolution – covering fossil history and phylogeny, the distribution and diversity of modern birds, and how birds are adapting to the human world – as well as a roundtable discussion on the same subject. Speakers and panellists are Robyn Womack, Albert Chen, Caitlin Kight, Hanneke Meijer and Glyn Young.

 This image has no special relevance to any of the TetZooCon bird talks, but here it is anyway. It’s a (now somewhat dated) bird phylogeny, produced for my 2014 paper on bird palaeobehaviour ( available here ). Image: Darren Naish.

This image has no special relevance to any of the TetZooCon bird talks, but here it is anyway. It’s a (now somewhat dated) bird phylogeny, produced for my 2014 paper on bird palaeobehaviour (available here). Image: Darren Naish.

Katrina van Grouw is talking about her amazing new book Unnatural Selection and the work behind it; Katrina will also be selling and signing copies of the book, and copies of her previous work The Unfeathered Bird too. Unnatural Selection is about the themes and patterns of evolution as revealed through the variation we’ve discovered via the selective breeding of domestic animals, not about domestic animals per se, and it’s a phenomenal tour de force that I cannot recommend highly enough. Some previous comments on the book can be found here.

 Katrina van Grouw (with duck), and the cover of her amazing 2018 book  Unnatural Selection  (to be reviewed here, hopefully soon). Images: (c) Katrina van Grouw.

Katrina van Grouw (with duck), and the cover of her amazing 2018 book Unnatural Selection (to be reviewed here, hopefully soon). Images: (c) Katrina van Grouw.

On Sunday, we have an on-stage discussion about Speculative Biology involving Gert van Dijk (of Furahan Biology and Allied Matters) and Dougal Dixon (After Man, The New Dinosaurs, Green World, The Future is Wild etc). Both will be discussing their own projects, their thoughts on SpecBio in general, and the past, present and future of the movement. Dougal will also be signing copies of the new edition of After Man and is also bringing along archive material relevant to some of his projects: I hope it will include some of the pieces brought to the recent After Man event at Conway Hall, but there will be other stuff too…

 A SpecBio montage relevant to TetZooCon 2018. At left: the Vortex from Dougal Dixon’s 1981  After Man . At right: Gert van Dijk, here photographed at the LonCon 72nd World Science Fiction Convention in 2014. Images: Dixon 1981, Darren Naish.

A SpecBio montage relevant to TetZooCon 2018. At left: the Vortex from Dougal Dixon’s 1981 After Man. At right: Gert van Dijk, here photographed at the LonCon 72nd World Science Fiction Convention in 2014. Images: Dixon 1981, Darren Naish.

World famous distinguished herpetologist and author Mark O’Shea joins us to talk about a detective story concerning the snakes of New Guinea and is also selling and signing copies of his brand-new, 656-page The Book of Snakes. Mark is also showing a short movie about reptiles. I’ve long hoped to have Mark as a TetZooCon speaker so am thrilled to have him with us this year.

 We have Professor Mark O’Shea! Mark will be selling and signing his new book. Images: (c) Mark O’Shea.

We have Professor Mark O’Shea! Mark will be selling and signing his new book. Images: (c) Mark O’Shea.

Film-maker, zoologist and author Lucy Cooke is also with us, and will be speaking about her neat 2018 book The Truth About Animals (originally titled The Unexpected Truth About Animals), which she’ll also be selling and signing. I must remember to ask her if she any copies of her previous book, A Little Book of Sloth, because who doesn’t like books on sloths?

 Lucy Cooke and equine friends, and the cover of her 2018 book  The Truth About Animals  (which I’m planning to review here later this year). Images: (c) Lucy Cooke, Darren Naish.

Lucy Cooke and equine friends, and the cover of her 2018 book The Truth About Animals (which I’m planning to review here later this year). Images: (c) Lucy Cooke, Darren Naish.

As per usual, John Conway is leading our Palaeoart Workshop: an interactive event in which – no doubt – something great is planned, I’m sure. The workshop occurs in parallel to some of the talks in a separate room. John is joined by Bob Nicholls, Mark Witton, Luis Rey and Steve White; Mark, Luis and Steve will be giving talks on (variously) the future of palaeoart (Mark), bringing dinosaurs back to life (Luis) and dinosaurs as portrayed in comics (Steve). There will also be palaeoart-themed book signings and a chance to meet the artists and see their latest projects.

 The 2018 palaeoart workshop is going to be outstanding. We have Bob Nicholls (though he won’t be bringing the lifesized pliosaur head with him, alas); Mark Witton will be selling  The Palaeoartist’s Handbook . I’ve been lucky enough to see a copy and…. it’s one of the most spectacular books I’ve ever seen, I love it. Images: (c) Bob Nicholls, Mark Witton.

The 2018 palaeoart workshop is going to be outstanding. We have Bob Nicholls (though he won’t be bringing the lifesized pliosaur head with him, alas); Mark Witton will be selling The Palaeoartist’s Handbook. I’ve been lucky enough to see a copy and…. it’s one of the most spectacular books I’ve ever seen, I love it. Images: (c) Bob Nicholls, Mark Witton.

While that’s not everything, I think that’ll do. For the first time, we’re having a drinks reception and conference meal (sold out, I’m afraid). There are additional talks on whales, extinct elephants, snakes, music in wildlife documentaries and – oh – the Dinosaurs in the Wild project (from me). There will also be a quiz (with numerous amazing prizes, as usual), additional book signings, and a substantial amount of merchandise on sale. I’m also intending to lead a post-conference fieldtrip to the Crystal Palace dinosaurs on the Monday morning.

 An amazing display of merchandise will be on sale at TetZooCon this year, be sure to bring £££. These dromaeosaur-themed mugs will be there, and are by the inimitable Jed Taylor, what a star. Image: Jed Taylor.

An amazing display of merchandise will be on sale at TetZooCon this year, be sure to bring £££. These dromaeosaur-themed mugs will be there, and are by the inimitable Jed Taylor, what a star. Image: Jed Taylor.

And that, as they say, is that. Again: tickets and more information can be found here. I look forward to seeing many of you there. Watch #TetZooCon for tweeting. All I have to do now is find time to finish getting things ready.

For articles on previous TetZooCons, see…

The Dougal Dixon After Man Event of September 2018

“Speculative biology, or speculative evolution, is a term that refers to a very hypothetical field of science that makes predictions and hypotheses on the evolution of life in a wide variety of scenarios and is also a form of fiction to an extent. It uses scientific principles and laws and applies them to a "what if" question” — the Speculative Evolution Wiki

 Dixon 1981; Dixon 2018.

Dixon 1981; Dixon 2018.

On September 11th 2018, I had the extraordinary privilege of appearing on stage with author, artist, editor, model-maker and visionary Dougal Dixon to discuss his famous book of 1981 After Man: A Zoology of the Future (Dixon 1981). We were joined by more than 215 interested members of the public, effectively filling the venue (Conway Hall in London).

 At Conway Hall once again, such a noble venue. Image: Will Naish.

At Conway Hall once again, such a noble venue. Image: Will Naish.

As you might have gathered if you’re a regular Tet Zoo reader, After Man has just been republished, and our on-stage event – hosted by New Lands London, and arranged by Scott Wood – also served as the launch for the new, 2018 edition (Dixon 2018). This was on sale and available for signing at the meeting.

As you can see from these photos, Dougal brought along a treasure trove of material relevant to the genesis of After Man, including his original sketches, text, draft spreads and the original page plan for the entire work. The latter consisted of large card mock-ups with small, rectangular vignettes depicting the planned look for each page. Dougal explained how he took these documents along to two – yes, just two – publishers and immediately got the green light from both. Evidently, he knew exactly what he was doing.

 Dougal on stage, showing the original page plan for  After Man . Small vignettes, showing the planned look of all the pages, are arranged in sequence. Image: Darren Naish.

Dougal on stage, showing the original page plan for After Man. Small vignettes, showing the planned look of all the pages, are arranged in sequence. Image: Darren Naish.

The original sketches are excellent and a testament to Dougal’s skill and planning. The quality of these illustrations also leave you wondering why it isn’t Dougal’s art that we see in the final book, and I impressed upon him during our discussion how fantastic it would be to one day see these unpublished illustrations in another book: a ‘The Making of After Man’ or something along those lines. We’ll see.

 Foreign language translations of  After Man , a Vortex and Raboon model, and relevant magazine issues (like the October 1981 ish of  BBC Wildlife ). (c) Dougal Dixon. Image: Darren Naish.

Foreign language translations of After Man, a Vortex and Raboon model, and relevant magazine issues (like the October 1981 ish of BBC Wildlife). (c) Dougal Dixon. Image: Darren Naish.

Several of the creatures pictured on the draft spreads were obvious prototypes of versions that made it into the final book but others were evidently abandoned at some point. A few of those ‘prototypes’ showed how the original animals had a different look relative to the published descendants: the bone-cracking Ghole Pallidogale nudicollum, to take one example, looked a lot more like a big mongoose in its original guise than is obvious in the book.

 Original text and original draft double-page spread for  After Man , showing creatures inhabiting tropical grasslands. You’ll recognise some (but not all!) of the creatures as the prototypes of versions that made it to final publication. (c) Dougal Dixon. Image: Darren Naish.

Original text and original draft double-page spread for After Man, showing creatures inhabiting tropical grasslands. You’ll recognise some (but not all!) of the creatures as the prototypes of versions that made it to final publication. (c) Dougal Dixon. Image: Darren Naish.

Dougal also brought foreign-language editions of After Man and a few models of After Man’s creatures with him, including a resin model of the Vortex Balenornis vivipara (a reasonable number were made, but it may be that this is the only one still in existence) and a wonderfully detailed Desert leaper Aquator adepsicautus. Alas, the Night stalker Manambulus perhorridus model pictured on the dustjacket of Dixon 1981 – the model I most wanted to see – is not in Dougal’s possession so was a no-show.

 Desert leaper model. For the handful of you that haven’t read  After Man , the Desert leaper is a giant, desert-dwelling muroid rodent (in cases more than 3 m long) that undergoes significant fluctuation in fat deposition (and hence mass) according to season. (c) Dougal Dixon. Image: Darren Naish.

Desert leaper model. For the handful of you that haven’t read After Man, the Desert leaper is a giant, desert-dwelling muroid rodent (in cases more than 3 m long) that undergoes significant fluctuation in fat deposition (and hence mass) according to season. (c) Dougal Dixon. Image: Darren Naish.

As goes our actual discussion, we covered the backstory to After Man (some of which will be familiar if you know the interview I published at ver 3 back in 2014), the response from critics and reviewers and the many overseas trips Dougal got to enjoy as a consequence of the book’s success, the substantial interest from Japanese markets and the Japanese stop-motion and animated movies (we watched a short segment from the stop-motion movie, copies of which were given away on DVD to people buying the book), the various efforts by studios in Hollywood to get an After Man movie off the ground, and the connection between After Man and Dougal’s more recent project Green World (thus far only published in Japanese).

 Original sketches, by Dougal, of creatures illustrated for  After Man . The animals were then re-illustrated by various other artists. (c) Dougal Dixon. Image: Darren Naish.

Original sketches, by Dougal, of creatures illustrated for After Man. The animals were then re-illustrated by various other artists. (c) Dougal Dixon. Image: Darren Naish.

We finished with a Q&A session and audience participation. Questions included the ‘new look’ Night stalker (yup… I shall say no more), Dougal’s thoughts on the future of humanity, how and which fossil species had influenced the creatures of After Man, and what might be different in After Man if Dougal were to write the book today.

 The Vortex model that Dougal brought along. It’s about 60 cm long. (c) Dougal Dixon. Image: Darren Naish.

The Vortex model that Dougal brought along. It’s about 60 cm long. (c) Dougal Dixon. Image: Darren Naish.

For a lifelong fan of After Man and Dougal’s connected writings, this event was an absolute thrill and I’m tremendously happy to have been involved. And judging by our audience’s response, it was enjoyed by everyone who attended too: thanks so much to everyone who came along and participated.

Two final things are worth saying. Firstly, I was asked innumerable times whether the event was going to be recorded. Alas, I was simply unable to organise this or even remember it given all the other stuff I had to worry about, though I think (and hope) than an audio recording exists. Secondly, this is not the only Dougal Dixon-themed event of 2018! Dougal is also on stage at this year’s TetZooCon when he will be joined by Gert van Dijk (of Furahan Biology and Allied Matters) in a discussion on speculative biology. TetZooCon happens on Oct 6th and 7th in London and tickets are still available. Dougal will also be bringing archive material to that meeting as well!

 Darren Naish (l) and Dougal Dixon (r) on stage at Conway Hall, September 2018. Image: Will Naish.

Darren Naish (l) and Dougal Dixon (r) on stage at Conway Hall, September 2018. Image: Will Naish.

My thanks to Dougal for being such a brilliant person to talk to and for all the material he brought along, to Scott and everyone else at Conway Hall and New Lands for organising things and setting it all up, to the Breakdown Press people for the book selling, and to our brilliant audience for their interest, enthusiasm and participation.

For previous articles on speculative biology, see…

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 - -

Dixon, D. 1981. After Man: A Zoology of the Future. Granada, London.

Dixon, D. 2018. After Man: A Zoology of the Future. Breakdown Press, London.

The Life Appearance of the Giant Deer Megaloceros

Eurasian Pleistocene megafauna are among the most familiar and oft-depicted of prehistoric animals. And among these grand, charismatic and imposing animals is the giant deer Megaloceros giganteus, an Ice Age giant that occurred from Ireland and Iberia in the west to southern Siberia in the east. It persisted beyond the end of the Pleistocene, surviving into the Early Holocene on the Isle of Man (Gonzalez et al. 2000) and western Siberia (Stuart et al. 2004)*. It is often erroneously termed the Irish elk, though it certainly wasn’t restricted to Ireland, nor should it really be termed an ‘elk’ (ugh… we’ll avoid that whole hornet’s nest for the time being). It’s been termed the Shelk by others [UPDATE: but see comments!!]. It could be 1.8 m tall at the shoulder and weigh somewhere around 600 kg, the antlers spanning 3.5 m in cases and weighing 35-45 kg (Geist 1999).

 A very conventional, traditional image of  Megaloceros giganteus : it's depicted looking like a giant red deer, basically. Males and females are not that different in size, but males are often shown as maned. Most interest in this deer has, of course, concerned the spectacularly antlered males. This image is from Hutchinson's  Extinct Monsters  (published several times over the 1890s). Image:  Hutchinson (1892) .

A very conventional, traditional image of Megaloceros giganteus: it's depicted looking like a giant red deer, basically. Males and females are not that different in size, but males are often shown as maned. Most interest in this deer has, of course, concerned the spectacularly antlered males. This image is from Hutchinson's Extinct Monsters (published several times over the 1890s). Image: Hutchinson (1892).

* In a previous edit of this article, I said that M. giganteus also survived into the Holocene in central Europe, as demonstrated by Immel et al. (2015). I missed the fact that this research concerns specimens dated to the Upper Pleistocene, not the Holocene. Furthermore, I’ve also been told that the Isle of Man data proved incorrectly dated. Am chasing confirmation on this.

While big, M. giganteus was not the biggest deer ever, since it seems that the extinct, moose-like Cervalces latifrons was even bigger. I promise to talk more about that species when I get round to discussing moose and kin at length. And while the antlers of M. giganteus were obviously very big, they weren’t especially big relative to its body size: proportionally, they were about similar in size to those of large Fallow deer Dama dama, and well exceeded in proportional size by the antlers of reindeer and caribou.

 A fine  Megaloceros  skull on show at London's Grant Museum. I seem to recall hearing or reading - possibly in one of Stephen J. Gould's papers - that this is one of the largest specimens in existence. Image: Darren Naish.

A fine Megaloceros skull on show at London's Grant Museum. I seem to recall hearing or reading - possibly in one of Stephen J. Gould's papers - that this is one of the largest specimens in existence. Image: Darren Naish.

I should add that M. giganteus was not the only Megaloceros species. Several others are known, differing in how palmate or slender and branching their antlers were, and not all were as large as M. giganteus (some were island-dwelling dwarves). There are other genera within this deer lineage (Megacerini) as well. Also of relevance to our discussion here is the position of these deer within the cervid family tree. Some experts have argued that megacerines are close to deer like the Red deer Cervus elaphus (Kuehn et al. 2005), while others point to genetic and morphological data indicating a close relationship with the Fallow deer Dama dama (Lister et al. 2005, Hughes et al. 2006, Immel et al. 2015, Mennecart et al. 2017). I have a definite preference for the latter idea, and right now it's a far better supported relationship than the alternative.

 Male  M. giganteus  skulls in the collections of the National Museum of Ireland, Dublin, examined in 2008. Yes, there is indeed a preponderance of males. Image: Darren Naish.

Male M. giganteus skulls in the collections of the National Museum of Ireland, Dublin, examined in 2008. Yes, there is indeed a preponderance of males. Image: Darren Naish.

Like most European people who’ve been lucky enough to visit museums and other such institutions, I’ve seen Megaloceros specimens on a great many occasions – there are a many of them on display. I’ve also seen and handled a reasonable number of the Irish bog specimens during time spent in Dublin. There does appear to be a preponderance of big, mature males. Maybe this reflects collecting bias (in that people were more inclined to extract the skulls and skeletons of big, prominently antlered males), but it also seems to be a valid biological signal: it has been argued that the calcium-hungry males were likely attracted to calcium-rich plants like willow at the edges of lakes and ponds, and were thus more prone to drowning, miring or falling through ice in such places than females (Geist 1999). Oh, we also know that male mammals across many species are more inclined to take stupid risks, be reckless, and even display deliberate bravado more than their female counterparts.

Here we come to the main reason for this article: what, exactly, did M. giganteus look like when alive? I’ve surely mentioned this topic on several occasions over the years here; I’m pretty sure I threatened to write about it after producing similar articles on the life appearance of the Woolly rhino and Ice Age horses. M. giganteus has been illustrated a great many times in works on prehistoric life, and the vast majority of reconstructions show it a near-monotone dark brown or reddish-brown. It’s very often depicted with a shaggy neck mane. In short, it’s usually made to look like a big, shaggy Red deer, and the tradition whereby this is done – it extends back to Zdenek Burian, Charles Knight and other founding palaeoartists – seems to me to be another of those palaeoart memes I’ve written about before. I’ve taken to calling this one the ‘Monarch of the Glen’ meme (see my palaeoart meme talk here). I will add here that we're generally talking about males of the species (since people mostly want to see depictions of specimens with those awesome antlers), though virtually all that I say below applies to females too.

Alas, this view of M. giganteus is almost certainly very wrong. Why do I say this?

 Note the many obvious external features of this male Fallow deer: a throat bulge corresponding with the larynx - an 'Adam's apple' - is obvious, and this is a boldly marked deer overall, with prominent spots (including some that have coalesced into stripes), a white rump patch, and pale ventral regions. If megacerines are close kin of  Dama  deer, we might predict a similar ancestral condition for  Megaloceros  and its relatives. Image: Dave Hone.

Note the many obvious external features of this male Fallow deer: a throat bulge corresponding with the larynx - an 'Adam's apple' - is obvious, and this is a boldly marked deer overall, with prominent spots (including some that have coalesced into stripes), a white rump patch, and pale ventral regions. If megacerines are close kin of Dama deer, we might predict a similar ancestral condition for Megaloceros and its relatives. Image: Dave Hone.

Firstly, if we look at the colours and patterns present across cervine deer as a whole, we see quite a bit of variation and no strong and obvious reason why a ‘Red deer look’ should be favoured. Secondly, we have that data indicating that M. giganteus is phylogenetically closer to Dama deer than to Cervus, in which case we would predict that it descended from ancestors with prominent spotting, pale flank stripes, and dark markings on the tail, all features typical of modern Dama populations. If the ‘Dama hypothesis’ is correct, there is again no reason to favour a ‘Red deer look’ for M. giganteus. Thirdly, body size, limb proportions, antler size and habitat choice all indicate that M. giganteus was an open-country (Clutton-Brock et al. 1980), cursorial specialist, and in fact the most cursorial of all deer (Geist 1999). Cursorial, open-country artiodactyls are often pale, with large white areas across the rump, legs and belly (examples include addax, some Arctic caribou and some argali). Again, no reason here to suspect that ‘Red deer look’.

And... fourthly, we have direct eyewitness data on the life appearance of this animal. Members of our own species saw it in life and drew it, seemingly to a very high degree of accuracy. What did they show?

 The famous panel at Cougnac, southwest France, showing  M. giganteus  males and females. This part of the cave is also interesting in depicting a short-horned bovid (at upper right) sometimes interpreted as a tahr. There are also ibex here too. I'm uncertain of the exact origin of the photo shown here: I took it from  Fabio Manucci's blog Agathaumus . Numerous additional photos of the same cave can be seen at  Don's Maps .

The famous panel at Cougnac, southwest France, showing M. giganteus males and females. This part of the cave is also interesting in depicting a short-horned bovid (at upper right) sometimes interpreted as a tahr. There are also ibex here too. I'm uncertain of the exact origin of the photo shown here: I took it from Fabio Manucci's blog Agathaumus. Numerous additional photos of the same cave can be seen at Don's Maps.

Virtually all cave art depicting M. giganteus shows a rounded, tall shoulder hump that’s sometimes shown as if it had a crest of raised hairs. Guthrie (2005) termed this a ‘hackle tuft’. There’s no obvious indication from the skeleton that a hump like this was present (indeed, fatty humps in mammals very often do not have an underlying skeletal correlate), so this is a neat thing that we wouldn’t know from skeletons alone. A protruding lump on the throat that seems to correspond to the larynx is also shown in images at Lascaux, Roucadour and elsewhere (Guthrie 2005). This feature is very reminiscent of Fallow deer.

 Cave art depicting  M. giganteus  is not all that numerous (most ancient depictions of deer are of reindeer or red deer), but what does exist shows several details worthy of note, here emphasised in illustrations produced by R. Dale Guthrie. The shoulder hump is a consistent feature. Image:  Guthrie (2005) .

Cave art depicting M. giganteus is not all that numerous (most ancient depictions of deer are of reindeer or red deer), but what does exist shows several details worthy of note, here emphasised in illustrations produced by R. Dale Guthrie. The shoulder hump is a consistent feature. Image: Guthrie (2005).

Some of the art provides information on pigmentation. A collar-like band is depicted encircling the neck in images from Chauvet and Cougnac, the shoulder hump is shown as being dark in images from Cougnac and elsewhere (Lister 1994), and some of the Chauvet and Roucadour images show a dark diagonal line that extends across the side of the body from the shoulder to the edge of the groin, and sometimes across the leg as far as the hock (ankle). An especially detailed image at Cougnac, partially illustrated on a stalactite, shows what looks like a dark vertical stripe descending from the shoulder hump and forming a division between the deep neck and the rest of the body. The same image also shows dark near-vertical markings around what might be a pale rump patch (Guthrie 2005).

 Other people have taken the same evidence I've discussed here and produced very similar reconstructions. This piece - which I hadn't seen until after producing my own illustrations (on which, see below) - is by Pavel Riha. Image:  Pavel Riha , CC BY-SA 3.0.

Other people have taken the same evidence I've discussed here and produced very similar reconstructions. This piece - which I hadn't seen until after producing my own illustrations (on which, see below) - is by Pavel Riha. Image: Pavel Riha, CC BY-SA 3.0.

If these details have been interpreted correctly, M. giganteus was boldly marked, with obvious dark striping across its neck, shoulders and torso, and on its rump too. R. Dale Guthrie proposed that the vertical shoulder stripe formed a boundary between a near-white neck and head region and the rest of the body, with the latter being pale just posterior to the stripe but darker across the legs, rump and flank (Guthrie 2005). I’m not absolutely convinced by the evidence from cave art for a near-white neck and head or for a white rump patch but these things are consistent with what I said above about the open-country lifestyle and cursoriality of this deer. Geist (1999) was a fan of this idea, and his reconstruction of M. giganteus – shown here – is meant to show the animal as being quite pale apart from its obvious striping and other dark markings.

  M. giganteus  as reconstructed by Valerius Geist, and shown to scale with the extant  Dama dama . Geist was (and presumably is) a strong advocate of the idea that megacerines (yes: megacerines, not 'megalocerines') are part of the same lineage as  Dama . Image:  Geist (1999) .

M. giganteus as reconstructed by Valerius Geist, and shown to scale with the extant Dama dama. Geist was (and presumably is) a strong advocate of the idea that megacerines (yes: megacerines, not 'megalocerines') are part of the same lineage as Dama. Image: Geist (1999).

Guthrie produced a very striking illustration depicting all of these details, but his drawing, as reproduced in his book (Guthrie 2005), is less than 4 cm long. Here it is (below), but note that I’ve produced a larger illustration here (scroll down) that shows the same details.

 At left, the best of the  M. giganteus  images from Cougnac in France, as re-drawn by  Guthrie (2005) . At right, Guthrie's reconstruction of the animal's life appearance. Image:  Guthrie (2005) .

At left, the best of the M. giganteus images from Cougnac in France, as re-drawn by Guthrie (2005). At right, Guthrie's reconstruction of the animal's life appearance. Image: Guthrie (2005).

And that just about brings us to a close. Over the years, I’ve been perpetually dismayed by the fact that most people illustrating this animal aren’t aware of the information I’ve discussed here – I mean, we have direct eyewitness data that should be pretty much the first thing we take account of when reconstructing this animal. Alas, the usual problem here is that the people who provide advice on reconstructions of fossil animals to artists are virtually never that interested in or knowledgeable about the life appearance of the animals concerned (sorry, palaeontologists). That’s an unfair generalisation though, and there are of course exceptions. Indeed, I should note that accurate, informed reconstructions of M. giganteus have appeared here and there over the years: the Megaloceros depicted in the Impossible Pictures TV series Walking With Beasts, for example, includes most of the features I’ve discussed here and obviously benefitted from the input of an informed consultant.

Megaloceros-appearance-2018-Megaloceros-cheat-sheet-1000-px-tiny-Sept-Darren-Naish-Tetrapod-Zoology.jpg

Anyway, my hope for the article you’re reading now is that it will inspire the current generation of palaeoartists to start illustrating Megaloceros in a way that’s more in accord with the data from prehistoric art, all of which has been out there in the literature for years now (Lister 1994, Guthrie 2005).

Megaloceros-appearance-2018-Megaloceros-Naish-black-background-1000-px-tiny-Sept-2018-Darren-Naish-Tetrapod-Zoology.jpg

I have further articles of this sort in mind and hope to get them published here eventually. On that note, here’s your reminder that I rely on your kind support at patreon, and that the more such support I receive, the more time and effort I can devote to Tet Zoo, and to my various book projects.

For previous Tet Zoo articles on Pleistocene megafauna, see...

And for articles on deer, see...

Refs - -

Clutton-Brock, T. H., Albon, S. D. & Harvey, P. H. 1980. Antlers, body size and breeding group size in the Cervidae. Nature 285, 565-567.

Geist, V. 1999. Deer of the World. Swan Hill Press, Shrewsbury.

Gonzalez, S., Kitchener, A. C. & Lister, A. M. 2000. Survival of the Irish elk into the Holocene. Nature 405, 753-754.

Guthrie, R. D. 2005. The Nature of Paleolithic Art. The University of Chicago Press, Chicago and London.

Hughes, S., Hayden, Th. J., Douady, Ch. J., Tougard, Ch., Germonpré, M., Stuart, A., Lbova, L., Garden, R. F., Hänni, C. & Say, L. 2006. Molecular phylogeny of the extinct giant deer, Megaloceros giganteus. Molecular Phylogeny and Evolution 40, 285-291.

Hutchinson, H. N. 1892. Extinct Monsters, 2nd edition. London: Chapman & Hall.

Immel, A., Drucker, D. G., Bonazzi, M., Jahnke, T. K., Münzel, S. C., Schuenemann, V. J., Herbig, A., Kind, C.-J. & Krause, J. 2015. Mitochondrial genomes of giant deers suggest their late survival in Central Europe. Scientific Reports 5: 10853.

Kuehn, R., Ludt, C. J., Schroeder, W. & Rottmann, O. 2005. Molecular phylogeny of Megaloceros giganteus - the Giant deer or just a giant red deer? Zoological Science 22, 1031-1044.

Lister, A. M. 1994. The evolution of the giant deer, Megaloceros giganteus (Blumenbach). Zoological Journal of the Linnean Society 112, 65-100.

Lister, A. M., Edwards, C. J., Nock, D. A. W., Bunce, M., van Pijlen, I. A., Bradley, D. G., Thomas, M. G. & Barnes, I. 2005. The phylogenetic position of the ‘giant deer’ Megaloceros giganteus. Nature 438, 850-853.

Mennecart, B., deMiguel, D., Bibi, F., Rössner, G. E., Métais, G., Neenan, J. M., Wang, S., Schulz, G., Müller, B. & Costeur, L. 2017. Bony labyrinth morphology clarifies the origin and evolution of deer. Scientific Reports 7: 13176.

Stuart, A. J., Kosintsev, P. A., Higham, T. F. G. & Lister, A. M. 2004. Pleistocene to Holocene extinction dynamics in giant deer and woolly mammoth. Nature 431, 684-689.

The Last Day of Dinosaurs in the Wild

On Sunday 2nd September 2018, the immersive, interactive time-travelling visitor attraction known as Dinosaurs in the Wild closed its doors for the last time. Yes, Dinosaurs in the Wild is now officially off-show, and if you didn’t get to see it before that fateful Sunday… where were you? I was determined to embark on one final tour, and of course I also needed to go to the grand send-off party and say those sad final goodbyes…

 Preparing to embark on a last ever tour of Late Maastrichtian western North America. Chrononaut Jasmine Arden-Brown introduces us to the world of Chronotex. Image: Darren Naish.

Preparing to embark on a last ever tour of Late Maastrichtian western North America. Chrononaut Jasmine Arden-Brown introduces us to the world of Chronotex. Image: Darren Naish.

As discussed at Tet Zoo several times already (all at ver 3, I hasten to add), I was scientific advisor for this grand project and thus very much feel that the look, behaviour and biology of the Late Cretaceous animals brought to life for the experience was and is ‘mine’, the MAJOR disclaimers being (1) that a whole team of people actually did the work that resulted in the vision coming to fruition, and (2) any ideas that I have about extinct animal biology or appearance or whatever involve the proverbial standing on the shoulders of giants, and the work and discoveries of a great many other people.

 As with any project of this size and scale, there's the long process whereby models and other props come together over time, and then there's the concept art, the explanatory diagrams, and so on. I've kept a record of as much of this stuff as I could. Image: Darren Naish.

As with any project of this size and scale, there's the long process whereby models and other props come together over time, and then there's the concept art, the explanatory diagrams, and so on. I've kept a record of as much of this stuff as I could. Image: Darren Naish.

With its fully – indeed, extensively – feathered dromaeosaurs, fuzzy-coated, muscular tyrannosaurs, terrestrial stalking azhdarchid pterosaurs (cough cough Witton & Naish 2008 cough cough), sleek, chunky mosasaurs, balloon-faced ankylosaurs and more (Conway et al. 2012), Dinosaurs in the Wild has – I really hope and feel – introduced a substantial chunk of the human public to a very up-to-date view of the Mesozoic world, and has thus gone some way towards undoing the damage of Jurassic World. No to the scaly, shit-brown, roaring monsters of the past, and yes to a more interesting, biologically plausible and often more surprising view of what these animals were like. Incidentally, Colin Trevorrow visited Dinosaurs in the Wild within the last few weeks, spoke to our associate live action director Cameron Wenn, and said really positive things (Colin and I spoke briefly over twitter).

 Here are two of the (normally nocturnal) Dinosaurs in the Wild animals seen in full illumination. At left, the metatherian mammal  Didelphodon ; at right, the small dromaeosaur  Acheroraptor  (it never stays still for long, hence the motion blur). Image: Darren Naish.

Here are two of the (normally nocturnal) Dinosaurs in the Wild animals seen in full illumination. At left, the metatherian mammal Didelphodon; at right, the small dromaeosaur Acheroraptor (it never stays still for long, hence the motion blur). Image: Darren Naish.

And did Dinosaurs in the Wild have an impact on the public? I don’t know if I’m allowed to release all the figures, but I will say that many thousands of people attended the experience during its 13 or so months of operation at Birmingham, Manchester and London. Our amazing actors and other staff all became worthy ambassadors of ‘new look’ Mesozoic animals and their biology, and the substantial amount of scientific content included in the show surely introduced the public to a great deal of information they haven’t seen or heard before. All results indicate that we certainly received the sort of feedback and accolade we hoped for: we scored really well as goes visitor feedback, indeed sufficiently well that Dinosaurs in the Wild can be regarded as a world class attraction. The palaeontologists and other scientists and experts who visited were all extraordinarily positive, and thanks indeed to those colleagues of mine who voiced their thoughts in public (Dean Lomax, Mark Witton, Albert Chen, Dave Hone, among others).

 Our final goodbye party was a solemn, quiet affair. Obviously. Thanks, Mike. Image: Darren Naish.

Our final goodbye party was a solemn, quiet affair. Obviously. Thanks, Mike. Image: Darren Naish.

Our venues were all great – Manchester’s Event City was certainly quite the sight to behold – but were perhaps not as centrally placed as might be ideal, though there are all kinds of factors controlling where and how a given exhibit can be located.

 A very dangerous box. Working in the Mesozoic is not all that easy. Image: Darren Naish.

A very dangerous box. Working in the Mesozoic is not all that easy. Image: Darren Naish.

Even now, and even after me writing that fairly substantial ‘behind the scenes’ article I published at Tet Zoo ver 3 in July 2018 (and here’s assuming that SciAm haven’t removed it due to an issue with image rights, ha ha ha), there’s a huge amount that could be said about the ‘making of’ this project. As some of you already know, the backstory to the world of Dinosaurs in the Wild is already written-up in an extensive document that we took to referring to as The Bible, but despite efforts I’ve had to give up on plans to get it published. I will be talking about much of the ‘behind the scenes’ stuff at TetZooCon this year (BUY TICKETS HERE), however, and will be bringing The Bible along for those interested in seeing it.

 There's so much to see through the windows that, even after multiple visits, I still haven't seen it all. In this sequence (seen while looking across the  Dakotaraptor  nesting colony), two female dromaeosaurs engage in a squabble. Image: Kerry Mulvihill.

There's so much to see through the windows that, even after multiple visits, I still haven't seen it all. In this sequence (seen while looking across the Dakotaraptor nesting colony), two female dromaeosaurs engage in a squabble. Image: Kerry Mulvihill.

Huge thanks to everyone at the event last night, and to everyone who made Dinosaurs in the Wild the success it was. Special thanks to producers Jill Bryant and Bob Deere, creative director Tim Haines, the team at Freeman Ryan, to live action directors Scott Faris and Cameron Wenn, to all the amazing people at Impossible, Milk VFX and Crawley Creatures, to every single one of our amazing actors, to our support staff, our sponsors and everyone else. And thanks also to Sam, Simon, Heather and the others who accompanied me on the same, final tour I took just yesterday.

 The temporal field generator is always on. Image: Darren Naish.

The temporal field generator is always on. Image: Darren Naish.

The Tet Zoo Guide to Mastigures

Among my favourite lizards are the Uromastyx agamids, variously termed mastigures, dabbs, dabs, dhubs, spinytails, spiny-tailed agamas, spiny-tailed lizards or thorny-tailed lizards. In the pet trade they’re often called ‘uros’. Here, I’ll be calling them mastigures.

 A large mastigure is a fine, handsome, happy-looking lizard. The dark overall colour and yellow dorsal occellations show that this captive specimen is a Moroccan mastigure  Uromastyx acanthinurus . Image: Darren Naish.

A large mastigure is a fine, handsome, happy-looking lizard. The dark overall colour and yellow dorsal occellations show that this captive specimen is a Moroccan mastigure Uromastyx acanthinurus. Image: Darren Naish.

Mid-sized for lizards (25 cm in total length is typical, though read on), they’re rather chunky, short-headed and wide-bodied with a proportionally short, broad tail that’s covered in 10 to 30 transverse, parallel rows of posterodistally projecting spines. The rows have a ring-like form and (rather confusingly) are typically called whorls. The tail is said to function as a ‘burrow blocker’ and also to be lashed from side to side when deterring would-be attackers. Enlarged, thorn-like scales are also present on the hindlimbs of some species. The head is short and deep by lizard standards and a neat feature is that the labial scales are large, serrated structures that sometimes look like external pseudoteeth.

 Head detail of a captive  U. acanthinurus . Note the pseudoteeth-like upper labial scales. The white exudate around the nostrils is pretty typical: it's salt discharge and evidence of nasal salt excretion typical for desert-dwelling lizards. Image: Darren Naish.

Head detail of a captive U. acanthinurus. Note the pseudoteeth-like upper labial scales. The white exudate around the nostrils is pretty typical: it's salt discharge and evidence of nasal salt excretion typical for desert-dwelling lizards. Image: Darren Naish.

Mastigures are extremely variable in colour, ranging from almost black to almost white dorsally; areas of yellow and even bright orange are present in some species, sometimes forming eye-like markings, distinct spots large or small, or transverse bands. The head may be much darker than the rest of the animal, and sometimes the tail is different in colouration too. Adding to this complexity is that individuals change colour according to temperature and time of day. The tail is variable in size: it's similar in length to the body in most species but is very short and broad in a few species, most notably the Omani spiny-tailed lizard or Thomas's mastigure U. thomasi.

 A captive  U. thomasi . The complex colouration - the facial banding in particular - is notable, as is the very short, broad, plump tail. This is a small mastigure with a total length of less than 15 cm. Recent surveys indicate that it is now extinct on mainland Oman - its type location - and is now unique to Masirah Island where local extinction has also occurred due to habitat destruction. There are anecdotal 2012 references to its persistence on the mainland, however. Image: Darren Naish.

A captive U. thomasi. The complex colouration - the facial banding in particular - is notable, as is the very short, broad, plump tail. This is a small mastigure with a total length of less than 15 cm. Recent surveys indicate that it is now extinct on mainland Oman - its type location - and is now unique to Masirah Island where local extinction has also occurred due to habitat destruction. There are anecdotal 2012 references to its persistence on the mainland, however. Image: Darren Naish.

The teeth are especially interesting: they’re short, low-crowned and fused to the jaw bones on their lingual (tongue) side, are largest at the back of the jaws, have crescentic shearing tips, and possess oblique wear facets that become so pronounced with age that entire teeth can be worn right down to the jaw (Cooper & Poole 1973). As you might guess, these animals do not possess regular tooth replacement of the sort we associate with reptiles (Robinson 1976). This is linked with a style of jaw movement (termed propaliny) where the lower jaw slides forwards to create a shearing bite when the jaws are closed (Throckmorton 1976). In the premaxillae, the upper central incisiforms are replaced by projecting structures that have been interpreted as bony pseudoteeth (Anderson 1999), though I don’t know if the histological work required to demonstrate this has been performed and they might be fused teeth.

 The skull of  U. aegyptia , as scanned for The Deep Scaly Project and available  here . Note that the partially fused teeth are largest posteriorly. The mandible is deep, the front of the dentary is toothless and bony pseudoteeth are present in the premaxilla. Image:  Digimorph.

The skull of U. aegyptia, as scanned for The Deep Scaly Project and available here. Note that the partially fused teeth are largest posteriorly. The mandible is deep, the front of the dentary is toothless and bony pseudoteeth are present in the premaxilla. Image: Digimorph.

Mastigures occur throughout the steppes, deserts and semi-deserts of northern Africa, the Middle East and western and central Asia. They aren’t associated with dune-fields, instead inhabiting rocky or gravel-covered regions or areas with compacted sand. They use and build burrows that are sometimes 3 m long or so, though I would expect based on data from other burrow-digging reptiles that burrows at least twice as long might exist. ‘Colonial burrows’ have been mentioned in the literature (Anderson 1999), though I don’t know if this means that many burrows were located in close proximity or if the burrows were known to contain some or many lizards.

 An Iraqi, Mesopotamian or Small-scaled spiny-tailed lizard  Saara loricata  (formerly  U. loricatus ), a mid-sized species of Iran and Iraq, as illustrated in one of Boulenger's 1885 catalogues of amphibians and reptiles kept in the collections of the British Museum. Image: Boulenger 1885.

An Iraqi, Mesopotamian or Small-scaled spiny-tailed lizard Saara loricata (formerly U. loricatus), a mid-sized species of Iran and Iraq, as illustrated in one of Boulenger's 1885 catalogues of amphibians and reptiles kept in the collections of the British Museum. Image: Boulenger 1885.

Around 15 extant species are recognised with Uromastyx, five of which have been named since 1990: U. maliensis Joger & Lambert, 1996, U. occidentalis Mateo et al., 1999 (or 1998), U. leptieni Wilms & Böhme, 2001 (or 2000…), U. alfredschmidti Wilms & Böhme, 2001 (or 2000…) and U. yemenensis Wilms & Schmitz, 2007. The total number of recognised species is a bit vague since some taxa are regarded as subspecies by some authors and as distinct species by others. An additional three Asian species have recently been removed from Uromastyx and placed in the resurrected genus Saara, first named by Gray in 1845 (Wilms et al. 2009). Saara species possess so-called intercalary scales between the spine whorls on the tail and molecular data finds them to be the sister-group to Uromastyx (Tamar et al. 2018).

 Persian or Iranian spiny-tailed lizard  Saara asmussi , as illustrated in William Blanford's paper of 1876. This species occurs in Iran, southern Afghanistan and Pakistan. The  Saara  species were included within  Uromastyx  prior to Wilms et al. (2009). Image: Blanford 1876.

Persian or Iranian spiny-tailed lizard Saara asmussi, as illustrated in William Blanford's paper of 1876. This species occurs in Iran, southern Afghanistan and Pakistan. The Saara species were included within Uromastyx prior to Wilms et al. (2009). Image: Blanford 1876.

In recent years, Uromastyx mastigures have become increasingly common in the pet trade and it’s now normal to see them on show in places that sell pet reptiles. I have seen them in the wild while on fieldwork in the Sahara, but the individuals concerned were dead and I never have seen a live one in the wild.

 A sadly deceased baby mastigure (probably  U. acanthinurus ), discovered in the Moroccan Sahara. Cause of death unknown. Note that the tail is fully developed and sports the full complement of tail spines, despite the animal's small size. Image: Darren Naish.

A sadly deceased baby mastigure (probably U. acanthinurus), discovered in the Moroccan Sahara. Cause of death unknown. Note that the tail is fully developed and sports the full complement of tail spines, despite the animal's small size. Image: Darren Naish.

Biology and behaviour. Mastigures are omnivorous, but they’re (seemingly) essentially herbivorous as adults, only occasionally eating arthropods or smaller lizards. The presence of symbiotic gut flora has been demonstrated for some species (a feature seen elsewhere in agamids in the Hydrosaurus sailfin dragons). Their lifestyle requires their taking refuge in rock crevices or burrows when they’re not feeding, foraging, basking or interacting socially, a behavioural syndrome where a compressed body shape and defensive spiny tail are advantageous, and one that has evolved convergently in other iguanians – the American chuckwallas and ctenosaurs and Madagascan oplurines – and in the Australian Egernia skinks and in some African corylids (Pianka & Vitt 2003).

 The tail of a deceased mastigure (probably  U. acanthinurus ), discovered in the Moroccan Sahara. Image: Darren Naish.

The tail of a deceased mastigure (probably U. acanthinurus), discovered in the Moroccan Sahara. Image: Darren Naish.

Herbivory in lizards works best at large size for the obvious reason of how much nutrition can be recovered (though it’s worth saying that there are many exceptions to this tendency: see Espinoza et al. 2004); it follows, then, that mastigures are relatively large compared to other agamids. I don’t know if there are any studies that do demonstrate this specifically, but the fact that most species are 25-45 cm long as adults does seem large, and the biggest species – the Egyptian or Leptien’s mastigure U. aegyptia – is positively enormous, reaching 75 cm on occasion and even more (specimens nearly 1 m long have been reported… can you imagine a mastigure this size? Amazing). It’s worth saying here that an especially large Paleogene lizard – Barbaturex from the middle Eocene of Myanmar, it perhaps reached 2 m in total – appears to be an especially close relative of Uromastyx (Head et al. 2013).

 I was curious to know what a c 90 cm mastigure would look like compared to a person. The smaller of these silhouettes reveals the answer. Not as impressive as I was hoping. The larger lizard silhouette depicts the approximate size of the Eocene taxon  Barbaturex , though we don't know that it had spiny whorls on its tail as shown in the illustration. The human figure is 1.7 m tall. Image: Darren Naish.

I was curious to know what a c 90 cm mastigure would look like compared to a person. The smaller of these silhouettes reveals the answer. Not as impressive as I was hoping. The larger lizard silhouette depicts the approximate size of the Eocene taxon Barbaturex, though we don't know that it had spiny whorls on its tail as shown in the illustration. The human figure is 1.7 m tall. Image: Darren Naish.

Mastigures are oviparous, females laying clutches of 6-20 elliptical eggs within a burrow. The hatchlings stay within the burrow for a few weeks, possibly even for months. The mother remains in attendance across this time and her burrow-guarding behaviour might be a form of parental care (directed both at the eggs and the hatchlings). Given that these lizards possess a symbiotic gut flora, the babies are presumably coprophagous. I’ve seen this stated informally but am not aware of a study that demonstrates it. Remember that tetrapods that possess a symbiotic gut flora must obtain it from their parents, and thus must eat their parent's dung early in life. Mm-mm.

 An Egyptian spiny-tailed lizard  U. aegyptia , as depicted in John Anderson's 1898 volume on the amphibians and reptiles of Egypt. Image: Anderson 1898.

An Egyptian spiny-tailed lizard U. aegyptia, as depicted in John Anderson's 1898 volume on the amphibians and reptiles of Egypt. Image: Anderson 1898.

Antiquity, taxonomy, biogeography. Having mentioned fossils, jaw fragments that appear to be from Uromastyx-like agamids (though not necessarily Uromastyx itself) are known from the Lower Eocene of Kyrgyzstan (Averianov & Danilov 1996) and hence establish an age of around 50 million years for this lineage. A number of Paleocene and Eocene lizards from Mongolia and China appear to be additional uromastycines. Rather younger, Oligocene fossils from the famous Jebel Qatrani Formation of the Fayum in Egypt’s Western Desert are sufficiently mastigure-like that they’ve been identified as ‘cf. Uromastyx’ (‘cf’ is an abbreviation of the Latin ‘confer’ and, when used in a taxonomic identification, basically means ‘we think that these fossils are so comparable to [insert taxon of interest] that they might belong to it, though we can’t be sure’). They date to the Lower Oligocene and hence are around 33 million years old (Holmes et al. 2010). There’s also a Lower Oligocene Uromastyx mastigure from France – yes, a European member of the group.

 Just one of the many uromastycine fossil jaw fragments from the Lower Eocene of Kyrgyzstan descibed by Averianov & Danilov (1996). These fossils - and others - demonstrate the antiquity of this group within Eurasia and show that it didn't arrive in the region after its Miocene collision with Africa. The scales bars = 1 mm. Image: Averianov & Danilov (1996).

Just one of the many uromastycine fossil jaw fragments from the Lower Eocene of Kyrgyzstan descibed by Averianov & Danilov (1996). These fossils - and others - demonstrate the antiquity of this group within Eurasia and show that it didn't arrive in the region after its Miocene collision with Africa. The scales bars = 1 mm. Image: Averianov & Danilov (1996).

This antiquity is in keeping with the idea – made on the basis of their highly distinctive anatomy – that mastigures are ‘distinct enough’ from other agamids to be worthy of their own ‘subfamily’: Uromastycinae. This view derives support from those studies that have found or inferred mastigures to be a distinct lineage outside the clade containing all remaining crown-agamids (e.g., Frost & Etheridge 1989, Macey et al. 2000, Schulte et al. 2003, Pyron et al. 2013), and perhaps even outside the clade that includes chameleons and conventional agamids (Honda et al. 2000, Gauthier et al. 2012). That last result would push mastigure origins into the Cretaceous given amber fossils that seem to be stem-chameleons.

 A phylogeny for agamids and their close kin, as recovered by Honda  et al . (2000). Mastigures and butterfly agamas form a clade, and both are outside the clade that includes chameleons and 'Agamidae' of tradition.   Like all of these sorts of diagrams, this was produced for my in-prep Vertebrate Fossil Record book, progress on which can be seen here.   Image: Darren Naish.

A phylogeny for agamids and their close kin, as recovered by Honda et al. (2000). Mastigures and butterfly agamas form a clade, and both are outside the clade that includes chameleons and 'Agamidae' of tradition. Like all of these sorts of diagrams, this was produced for my in-prep Vertebrate Fossil Record book, progress on which can be seen here. Image: Darren Naish.

Oh, you want Cretaceous stem-mastigures? In 2016, Apesteguía et al. (2016) described Jeddaherdan aleadonta from the Cenomanian of Morocco, and concluded that both this taxon and Gueragama sulamericana from the Upper Cretaceous of Brazil – both represented by partial lower jaws – are exactly that. Fossil evidence does, therefore, now back up the idea that these lizards were in existence before the end of the Cretaceous, and that acrodonts* (and thus iguanians more generally) had evolved at least some of their variation before the Cenozoic.

* Acrodonts (properly Acrodonta): the iguanian lizard clade that includes chameleons and agamids. They are named for their acrodont teeth: that is, those fused to the jawbones (though this condition is not fully developed across all members of the clade, and note that there are acrodont reptiles that are not part of Acrodonta).

 The Cretaceous uromastycine  Jeddaherdan aleadonta  is known from the chunk of lower jaw shown here, depicted within a silhouetted skull of  Uromastyx . The scale bar is in mm. Image: Apesteguía  et al . (2016).

The Cretaceous uromastycine Jeddaherdan aleadonta is known from the chunk of lower jaw shown here, depicted within a silhouetted skull of Uromastyx. The scale bar is in mm. Image: Apesteguía et al. (2016).

At least some studies find mastigures to form a clade with the east Asian butterfly agamas Leiolepis (e.g., Honda et al. 2000, Hugall & Lee 2004, Gauthier et al. 2012), both then being united within Leiolepidinae*. Butterfly agamas are fascinating for all sorts of reasons and I really should write about them at some point as well.

* There’s a long and complex argument over whether Leiolepidinae/Leiolepididae or Uromastycinae/Uromastycidae should win in a priority battle. Modern authors have tended to prefer the former, since it’s 1843 as opposed to 1863 for Theobald’s Uromastycidae. Anderson (1999) argued that the 1843 use of Fitzinger’s name cannot win this battle, since it was originally ‘Leiolepides’ and was not written in its ‘modern’ form by authors pre-1900.

 Mastigures and butterfly agamas have not been found to form a clade in all phylogenetic studies: in  Pyron  et al .'s (2013)  study - this cladogram depicts the topology they recovered - the two are successively closer to remaining Agamidae. Note the taxonomic names they used for the agamid lineages.   Like all of these sorts of diagrams, this was produced for my in-prep Vertebrate Fossil Record book, progress on which can be seen here.   Image: Darren Naish.

Mastigures and butterfly agamas have not been found to form a clade in all phylogenetic studies: in Pyron et al.'s (2013) study - this cladogram depicts the topology they recovered - the two are successively closer to remaining Agamidae. Note the taxonomic names they used for the agamid lineages. Like all of these sorts of diagrams, this was produced for my in-prep Vertebrate Fossil Record book, progress on which can be seen here. Image: Darren Naish.

Anyway: here I’ll say what I usually do and remind you that if these animals were mammals or birds they’d almost definitely be considered ‘distinct enough’ to warrant their own ‘family’, a decision that would require Agamidae of tradition to be split into several ‘families’ (I put these taxonomic ranks in quotes because they’re still effectively subjective). In addition to a mastigure family and butterfly agama family, there would be one for Hydrosaurus, one for the Australasian dragons (or amphibolurines), another for the Asian draconines, and so on. A few authors – most notably Scott Moody in his studies of the early 1980s – have at least separated mastigures and butterfly agamas from remaining agamids in a version of Theobald's ‘family’ Uromastycidae.

 Butterfly agamas ( Leiolepis ) do look mastigure-like in some features of the face (those tall ridges over the orbits especially), but are otherwise far slimmer, longer-limbed and without their other specialisations. The two groups may be closely related - though it still seems that they diverged during the Late Cretaceous, at least. Image:  TheReptilarium , CC BY 2.0.

Butterfly agamas (Leiolepis) do look mastigure-like in some features of the face (those tall ridges over the orbits especially), but are otherwise far slimmer, longer-limbed and without their other specialisations. The two groups may be closely related - though it still seems that they diverged during the Late Cretaceous, at least. Image: TheReptilarium, CC BY 2.0.

Macey et al. (2000) assumed an Indian origin for mastigures, in which case they’re among several tetrapod groups that followed an ‘Out of India’ dispersal route hypothesised elsewhere for ostriches and certain caecilians and frogs. But this is also contradicted by fossils, since Paleocene members of the lineage – if correctly identified and correctly dated – show that members of the lineage were living in Eurasia before India docked with Eurasia during the Eocene. The best model, therefore, might be one in which mastigures moved into Eurasia at the end of the Cretaceous.

 An Eocene map depicting the planet as of around 40 million years ago. At this point, Afro-Arabia had not docked with Eurasia. But members of the mastigure lineage were already present in Eurasia and Afro-Arabia by the time. Image: the original version was used in  Angst et al. (2013) ; this has been modified as per CC BY 2.5.

An Eocene map depicting the planet as of around 40 million years ago. At this point, Afro-Arabia had not docked with Eurasia. But members of the mastigure lineage were already present in Eurasia and Afro-Arabia by the time. Image: the original version was used in Angst et al. (2013); this has been modified as per CC BY 2.5.

Tamar et al. (2018) posited an initial, middle Miocene diversification of the Uromastyx crown-group in south-east Asia followed by Afro-Arabian invasion and diversification. But note that this only applies to crown-group Uromastyx, not to the Saara + Uromastyx clade, nor to the mastigure lineage as a whole, and thus is not inconsistent with an earlier origin and diversification elsewhere.

 Tamar  et al . (2018) found  Uromastyx  to consist of two main clades, one mostly associated with the Arabian Peninsula and the borders of the Red Sea, one with the western Sahara.  Saara  forms the sister-group to  Uromastyx . Image: Tamar  et al . (2018).

Tamar et al. (2018) found Uromastyx to consist of two main clades, one mostly associated with the Arabian Peninsula and the borders of the Red Sea, one with the western Sahara. Saara forms the sister-group to Uromastyx. Image: Tamar et al. (2018).

Your regular dose of misanthropy. Finally, all is not well as goes the future of mastigures. As you might guess given my earlier mentions of the pet trade, the sad fact is that uncontrolled, indiscriminate and often illegal collection from the wild is a threat to many populations. Many people involved in the pet reptile trade – those at the sharp end where animals are taken from the wild and smuggled to other countries – have no scruples whatsoever as goes the ethical or managed treatment of animals, and if you don’t believe me look up articles on Anson Wong, the Malaysian wildlife smuggler known as the ‘Lizard King’ (a most inappropriate moniker, given that Kings are supposed to be worthy of respect or admiration).

 I would love to see a large, spectacular mastigure in the wild. This mastigure ( U. aegyptia microlepis ), photographed in Al Anbar, Iraq, is a grand, magnificent animal. Image: U.S. Federal Government, Public Domain.

I would love to see a large, spectacular mastigure in the wild. This mastigure (U. aegyptia microlepis), photographed in Al Anbar, Iraq, is a grand, magnificent animal. Image: U.S. Federal Government, Public Domain.

Mastigures have also been much used for food, medicine and as ritual objects (a cleaned mastigure body serves as a traditional baby’s bottle in Morocco, for example), all of which is fine (in theory) when harvesting is kept to sustainable levels… but less fine when exploitation begins to outstrip supply. Those mastigures that have been studied are declining or locally extinct across their range and all species are CITES listed as of 1977. Specifically, they’re on Appendix II of CITES, which refers to species that are not necessarily in immediate danger of extinction but do nonetheless require a control in their trade.

 The large size and interesting appearance of many mastigure species - this is a captive  U. aegyptia  - has long made them appealing objects of trade and medicinal use, and as objects for the table too. Image: Darren Naish.

The large size and interesting appearance of many mastigure species - this is a captive U. aegyptia - has long made them appealing objects of trade and medicinal use, and as objects for the table too. Image: Darren Naish.

In some countries where these lizards occur it’s considered a rite of passage for young men to go out and kill as many mastigures as they can, and if you want verification for that you can find photos online where there are great piles of tens or even hundreds of dead mastigures in the backs of trucks. That’s depressing and vile behaviour. Like Anne Frank, I do think that people are essentially good but it’s difficult to maintain a rosy view of humanity when our stated aim seems to be the denuding of wild spaces of their animals.

On that depressing note, we move on.

This article took a lot of work and quickly expanded way beyond the brief ‘here’s a picture of a lizard’ article it was originally intended to be. However, I think that articles like this are useful and would like to keep doing them. Here’s your regular reminder that I require support if this is to continue. I would do more if support allowed. Thanks to those who support me at patreon already.

Iguanian lizards have now been covered quite a few times at Tet Zoo. For previous articles see...

Refs - -

Anderson, S. C. 1999. The Lizards of Iran. Society for the Study of Amphibians and Reptiles, Saint Louis.

Apesteguía, S., Daza, J. D., Simões, T. R. & Rage, J. C. 2016 The first iguanian lizard from the Mesozoic of Africa. Royal Society Open Science 3: 160462.

Averianov, A. & Danilov, I. 1996. Agamid lizards (Reptilia, Sauria, Agamidae) from the Early Eocene of Kyrgyzstan. Neues Jahrbuch fur Geologie und Paläontologie, Monatshefte 1996 (12), 739-750.

Cooper, J. S. & Poole, F. G. 1973. The dentition and dental tissues of the agamid lizard Uromastyx. Journal of Zoology 169, 85-100.

Espinoza, R. E., Wiens, J. J. & Tracy, C. R. 2004. Recurrent evolution of herbivory in small, cold-climate lizards: breaking the ecophysiological rules of reptilian herbivory. Proceedings of the National Academy of Sciences 101, 16819-16824.

Frost, D. R. & Etheridge, R. 1989. A phylogenetic analysis and taxonomy of iguanian lizards (Reptilia: Squamata). University of Kansas, Museum of Natural History, Miscellaneous Publication 81, 1-65.

Gauthier, J. A., Kearney, M., Maisano, J. A., Rieppel, O. & Behlke, D. B. 2012. Assembling the squamate tree of life: perspectives from the phenotype and the fossil record. Bulletin of the Peabody Museum of Natural History 53, 3-308.

Head, J. J. Gunnell, G. F., Holroyd, P. A., Hutchinson, J. H. & Ciochon, R. L. 2013. Giant lizards occupied herbivorous mammalian ecospace during the paleogene greenhouse in SouthEast Asia. Proceedings of the Royal Society B 20130665 http://dx.doi.org/10.1098/rspb.2013.0665

Holmes, R. B., Murray, A. M., Chatrath, P., Attia, Y. S. & Simons, E. L. 2010. Agamid lizards (Agamidae: Uromastycinae) from the Lower Oligocene of Egypt. Historical Biology 22, 215-223.

Honda, M., Ota, H., Kobayashi, M., Nabhitanhata, J., Yong, H.-S., Sengoku, S. & Hikida, T. 2000. Phylogenetic relationships of the family Agamidae (Reptilia: Iguania) inferred from mitochondrial DNA sequences. Zoological Science 17, 527-537.

Hugall, A. F. & Lee, M. S. Y. 2004. Molecular claims of Gondwanan age for Australian agamid lizards are untenable. Molecular Biology and Evolution 21, 2102-2110.

Macey, J. R., Schulte, J. A., Larson, A., Ananjeva, N. B., Wang, Y., Pethiyagoda, R., Rastegar-Pouyani, N. & Papenfuss, T. J. 2000. Evaluating trans-Tethys migration: an example using acrodont lizard phylogenetics. Systematic Biology 49, 233-256.

Pianka, E. R. & Vitt, L. J. 2003. Lizards: Windows 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 2013, 13:93 doi:10.1186/1471-2148-13-93

Robinson, P. L. 1976. How Sphenodon and Uromastyx grow their teeth and use them. In Bellairs, A. d’A. & Cox, C. B. (eds) Morphology and Biology of Reptiles. Academic Press (London), pp. 43-64.

Schulte, J.A., Valladares, J. P. & Larson, A. 2003. Phylogenetic relationships within Iguanidae inferred using molecular and morphological data and a phylogenetic taxonomy of iguanian lizards. Herpetology 59, 399-419.

Tamar, K., Metallinou, M., Wilms, T., Schmitz, A., Crochet, P.-A., Geniez, P. & Carranza, S. 2018. Evolutionary history of spiny-tailed lizards (Agamidae: Uromastyx) from the Saharo-Arabian region. Zoologica Scripta 47, 159-173.

Throckmorton, G. S. 1976. Oral food processing in two herbivorous lizards, Iguana iguana (Iguanidae) and Uromastix [sic] aegyptius [sic] (Agamidae). Journal of Morphology 148, 363-390.

Wilms, T. Böhme, W., Wagner, P., Lutzmann, N. & Schmitz, A. 2009. On the phylogeny and taxonomy of the genus Uromastyx Merrem, 1820 (Reptilia: Squamata: Agamidae: Uromastycinae) – resurrection of the genus Saara Gray, 1845. Bonner Zoologische Beiträge 56, 55-99.

Postcranial Palaeoneurology and the Lifestyles of Pterosaurs

Regular readers will know that I – with colleagues – publish fairly regularly on azhdarchoid pterosaurs, the very special pterosaur group that includes the short-faced tapejarids, the sometimes gigantic, long-jawed azhdarchids and a few groups seemingly intermediate between these two. Azhdarchoids are quite obviously the best and most interesting of the pterosaurs. And back in 2013, I and colleagues published a description and analysis of a new species from the Early Cretaceous of the Isle of Wight known only from a three-dimensional pelvis and some associated vertebrae. We called it Vectidraco daisymorrisae, its name honouring Daisy Morris, the young woman who discovered it (Naish et al. 2013).

 The  Vectidraco daisymorrisae  holotype (NHMUK PV R36621) in (A) left lateral, (B) right lateral, (C) dorsal and (D) ventral views, and - at right - shown in anatomical position as per the animal's presumed profile in life. Image: figures from  Naish  et al . (2013) .

The Vectidraco daisymorrisae holotype (NHMUK PV R36621) in (A) left lateral, (B) right lateral, (C) dorsal and (D) ventral views, and - at right - shown in anatomical position as per the animal's presumed profile in life. Image: figures from Naish et al. (2013).

Enough is known of Vectidraco for us to make some determination as goes what sort of pterosaur it is (it seems to be a tapejarid or tapejarid-like azhdarchoid), and we can also say interesting things as goes its size and degree of skeletal pneumatisation (Naish et al. 2013). It’s well preserved enough that quite a few other things can be done with it as well. Last year Rachel Frigot used it as a model in the determination of pelvic and hindlimb musculature (Frigot 2017). And, as part of her PhD work on pterosaur pneumaticity and anatomy, my colleague Liz Martin-Silverstone sought to do some neat science with it as well. This work has just been published (Martin-Silverstone et al. 2018), and that’s why we’re here today.

 My friend and colleague Dr Liz Martin-Silverstone, at work in the field (at left, Liz is finding fossils in a river in Romania) and in a museum exhibition at right (Liz is standing next to an exhibition panel all about her work. Let's not talk about that weird silhouette at upper right...). Images: Darren Naish.

My friend and colleague Dr Liz Martin-Silverstone, at work in the field (at left, Liz is finding fossils in a river in Romania) and in a museum exhibition at right (Liz is standing next to an exhibition panel all about her work. Let's not talk about that weird silhouette at upper right...). Images: Darren Naish.

Much of Liz’s work has involved CT-scanning (you can read about her own adventures here on her blog) and the relationship between pneumatisation, mass and flight. Vectidraco is at the other end of the scale from many of the pterosaurs that Liz has worked on (it was a small pterosaur with a wingspan likely less than 1 m as an adult) and was readily available, so it seemed sensible to incorporate it into her work. We scanned the specimen at its home (the Natural History Museum, London), and compared the results with those obtained from other pterosaurs we had to hand: namely, the ornithocheirids* Anhanguera and Coloborhynchus. scanned variously at Stony Brook University Hospital (thanks to Pat O’Connor for that data) and at the µ-VIS (pronounced ‘mu-vis’) X-Ray Imaging Centre at the University of Southampton (Martin-Silverstone et al. 2018). And we got pretty good results.

* Ornithocheirids: the mostly marine, long-jawed, long-winged pterodactyloid pterosaur group named for Ornithocheirus from the 'middle' Cretaceous of the UK. The group names Anhangueria and Anhangueridae refer to the same group... views differ on which taxonomic system we should adopt.

 Pelvic regions of the three pterosaurs included in our study, to scale: (A)  Vectidraco daisymorrisae  holotype NHMUK PV R36621, (B)  Anhanguera  specimen AMNH FARB 22555, (C)  Coloborhynchus robustus  specimen SMNK PAL 1133. Scale bar = 50 mm. Image: Martin-Silverstone et al. (2018).

Pelvic regions of the three pterosaurs included in our study, to scale: (A) Vectidraco daisymorrisae holotype NHMUK PV R36621, (B) Anhanguera specimen AMNH FARB 22555, (C) Coloborhynchus robustus specimen SMNK PAL 1133. Scale bar = 50 mm. Image: Martin-Silverstone et al. (2018).

The first interesting thing to note is that the work corrects, updates and augments various anatomical details I reported in the initial description of Vectidraco (Naish et al. 2013). Bony openings that I interpreted as pneumatic foramina turn out to be foramina for spinal nerves (properly termed intervertebral foramina), and convex transverse ridges present on the sides and undersides of some of the vertebrae are misidentified intervertebral junctions. Cool – it’s good to learn more. The identification of intervertebral foramina is not a big deal at all given that these structures are ubiquitous in tetrapods but it's worth bringing attention to them given that they’re virtually unmentioned elsewhere in the pterosaur literature.

 The  Vectidraco  holotype is one of those wonderful specimens that preserves a great many neat little anatomical details - look at these various pneumatic cavities on the T-shaped post-acetabular process on the posterior part of the ilium. Waitaminute.... aren't  all the specimens  like this? Image: Darren Naish.

The Vectidraco holotype is one of those wonderful specimens that preserves a great many neat little anatomical details - look at these various pneumatic cavities on the T-shaped post-acetabular process on the posterior part of the ilium. Waitaminute.... aren't all the specimens like this? Image: Darren Naish.

And if you’re wondering why I and my colleagues didn’t CT-scan the specimen the first time around and get this stuff correct on our first, 2013 attempt, it’s because CT-scanning requires money and virtually everything I do has been, and is, unfunded.

Anyway… what else could we do with the CT-scan data? Well…

During the late 1980s and 90s, Emily Giffin (later Emily Buchholtz) published several papers in which she used data from the size of the neural canal in the vertebrae of fossil tetrapods to make inferences about nerve size, the size then serving as a proxy for degree of innervation, this then serving as a guide to things like limb function and posture (Giffin 1989, 1990, 1992, 1995a, b). Her studies looked variously at non-bird theropods, extinct crocodylians and fossil pinnipeds, and she reported encouraging results. Non-bird theropods with large hands, to take one example, possessed neural canals in the corresponding part of the spine that were proportionally large, and hence suggestive of the well-developed nervous anatomy we would expect for animals that regularly used their hands in grabbing, piercing and tearing (Giffin 1995a). Her work has inspired other researchers to use the same (or similar) techniques on plesiosaurs and fossil raptors (again, here’s your helpful reminder that I will only ever use this term in the correct fashion. It applies to hawks, eagles and falcons and has done since the 1800s at least).

 In a series of really interesting papers, Emily Giffin linked neural canal size with form and function in diverse tetrapods. This graph (from Giffin 1995b) shows how birds flying and flightless differ as goes the position of the largest parts of their spinal cords. The ostrich ( Struthio ) lacks a large spinal cord section in the anterior (brachial) part of its spinal column. Image: Giffin (1995b).

In a series of really interesting papers, Emily Giffin linked neural canal size with form and function in diverse tetrapods. This graph (from Giffin 1995b) shows how birds flying and flightless differ as goes the position of the largest parts of their spinal cords. The ostrich (Struthio) lacks a large spinal cord section in the anterior (brachial) part of its spinal column. Image: Giffin (1995b).

Several caveats make this technique far from fool-proof (Giffin 1995a). With these things in mind, we wondered if data from pterosaurs might be informative as goes ideas on their ecology and lifestyle. What we found is that Vectidraco has an unusually large neural canal in its sacral region compared to Anhanguera, indicating that it therefore had a proportionally large spinal cord (and lumbosacral plexus) in its lumbosacral region. Anhanguera and Coloborhynchus both had enlarged neural canals in the area corresponding to the brachial plexus, larger than the neural canals in their sacral regions. Vectidraco’s shoulder region is entirely unknown at the moment so we couldn't make any comparison here.

 Neural canal cross-sectional area in our three pterosaur taxa: when normalised for centrum size,  Vectidraco  has a proportionally large neural canal. This composite image incorporates figures from Martin-Silverstone  et al . (2018) but was produced by the Palaeontological Association. Image: Martin-Silverstone  et al . (2018).

Neural canal cross-sectional area in our three pterosaur taxa: when normalised for centrum size, Vectidraco has a proportionally large neural canal. This composite image incorporates figures from Martin-Silverstone et al. (2018) but was produced by the Palaeontological Association. Image: Martin-Silverstone et al. (2018).

Taken together this suggests the following: evidence for a large brachial enlargement in ornithocheirids is consistent with the idea (based on their long, high-aspect wings and small hindlimbs) that they were highly aerial animals, while the large sacral neural canal in Vectidraco indicates that it was more proficient at terrestrial locomotion than Anhanguera. There are already indications from pelvic morphology, hindlimb size and so on that azhdarchoids and ornithocheirids were doing very different things in terms of ecology and behaviour (Witton & Naish 2008, 2015, Witton 2013, Naish & Witton 2017), so this matches what we might predict.

  Vectidraco  could almost certainly fly well, as shown at left. But - like many, most or all azhdarchoids - it was likely a proficient and regular terrestrial walker as well, as shown at right. Image: Mark Witton (left), Darren Naish (right).

Vectidraco could almost certainly fly well, as shown at left. But - like many, most or all azhdarchoids - it was likely a proficient and regular terrestrial walker as well, as shown at right. Image: Mark Witton (left), Darren Naish (right).

So far so good. But the complication comes from the second ornithocheirid we looked at: Coloborhynchus. Oh, here I’ll avoid the whole mess concerning the taxonomy of Anhanguera and Coloborhynchus. All I’ll do for now is say that “it’s complicated” and promise that I’ll come back to it in the near future. Anyway… the Coloborhynchus specimen we analysed is not like our Anhanguera specimen as goes the proportional size of the neural canal in its lumbosacral region -- it lacks a distinct lumbosacral enlargement but is superficially Vectidraco-like in having a larger neural canal in the relevant region (Martin-Silverstone et al. 2018). This is an unexpected result. Does it mean that some ornithocheirids were far more terrestrially capable than others, this perhaps reflecting niche differentiation or some other form of variation? Or does it mean that some ornithocheirids were far more aerially specialised than others and that the default condition was to have a larger sacral neural canal? Or does it mean that there’s something else we haven’t accounted for? (example: maybe some of these pterosaurs had enlarged neural canals due to pneumatisation in the neural canal? Yes, air sacs dorsal to the spinal cord. This is a thing). I don’t think it means that Coloborhynchus-type pterosaurs were as terrestrially proficient as Vectidraco, nor that our ideas on said terrestrial proficiency in azhdarchoids like Vectidraco are bogus, given the fact that Vectidraco appears to have a lumbosacral enlargement while the ornithocheirids do not, and that azhdarchoids like Vectidraco possess weird features indicative of terrestrial specialisation (like the giant, T-shaped postacetabular process on the ilium) lacking in ornithocheirids. But it’s clear that more data and more work is needed, as usual with these sorts of things.

 Behaviour speculatively inferred for the pterosaurs incorporated in our study. (A) A dedicated aerial lifestyle involving little terrestrial behaviour, as per  Anhanguera ; (B) reasonable terrestrial abilities in an animal otherwise very similar to its close, highly aerial relatives, as per  Coloborhynchus ; (C) proficient and regular terrestrial behaviour in an animal that routinely feeds and forages on the ground, as per  Vectidraco . Image: Darren Naish.

Behaviour speculatively inferred for the pterosaurs incorporated in our study. (A) A dedicated aerial lifestyle involving little terrestrial behaviour, as per Anhanguera; (B) reasonable terrestrial abilities in an animal otherwise very similar to its close, highly aerial relatives, as per Coloborhynchus; (C) proficient and regular terrestrial behaviour in an animal that routinely feeds and forages on the ground, as per Vectidraco. Image: Darren Naish.

It’s also important to remember that this data – this work as a whole – is complimentary to other studies. CT scan data on neural canal size provides nothing like a ‘Rosetta Stone’ on behaviour or lifestyle, and the more we learn about anatomy and form-function correlation, the less likely it seems that such things exist. We already have lots of reasons for thinking that azhdarchoids were better adapted for terrestriality and life in cluttered, inland settings than were many other pterosaurs, and likewise there are excellent reasons for thinking that ornithocheirids were aerial specialists and something like frigatebirds as goes ecology and lifestyle (though seemingly capable of swimming and perhaps diving, in contrast to the more piratical frigatebirds). So, this work is part of the dataset, part of the puzzle. And this is effectively an opening foray in this most intriguing area.

That’s where we’ll end for now. More on pterosaurs here soon!

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. The more independence I achieve, the more time I can spend producing the content you enjoy.

For previous Tet Zoo articles on pterosaurs, see...

Refs - -

Frigot, R. 2017. Pelvic musculature of Vectidraco daisymorrisae and consequences for pterosaur locomotion. In Hone, D. W. E., Witton, M. P. & Martill, D. M. (eds) New Perspectives on Pterosaur Palaeobiology. Geological Society, London, Special Publications Special Publications 455, 45-55.

Giffin, E. B. 1989. Gross spinal anatomy and limb function in living and fossil reptiles and birds. American Zoologist 29, 181A.

Giffin, E. B. 1990. Gross spinal anatomy and limb use in living and fossil reptiles. Paleobiology 16, 448-458.

Giffin, E. B. 1992. Functional implications of neural canal anatomy in recent and fossil marine carnivores. Journal of Morphology 214, 357-374.

Giffin, E. B. 1995a. Functional interpretation of spinal anatomy in living and fossil amniotes. In Thomason, J. J. (ed) Functional Morphology in Vertebrate Paleontology. Cambridge University Press, pp. 235-248.

Giffin, E. B. 1995b. Postcranial paleoneurology of the Diapsida. Journal of Zoology 235, 389-410.

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

Naish, D., Simpson, M. I. & Dyke, G. J. 2013. A new small-bodied azhdarchoid pterosaur from the Lower Cretaceous of England and its implications for pterosaur anatomy, diversity and phylogeny. PLoS ONE 8 (3): e58451.

Naish, D. & Witton, M. P. 2017. Neck biomechanics indicate that giant Transylvanian azhdarchid pterosaurs were short-necked arch predators. PeerJ 5: e2908.

Witton, M. P. 2013. Pterosaurs. Princeton University Press, Princeton & London.

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.

Could We Domesticate (Non-Bird) Dinosaurs?

Could we domesticate non-bird dinosaurs, supposing – hypothetically – that we lived in some alternate timeline? This might not seem like the sort of issue I readily tackle here at Tet Zoo (or… it might), but I’ve been inspired. My attention, you see, was drawn to an internet site where an expert attempted to answer the same question. Aaaand the answer they gave was pretty much useless, so here we are.

 A friendly pet  Parasaurolophus , as illustrated by Mike Skrepnick (and used with his permission) for a 'Would Dinosaurs Make Good Pets' project produced in conjunction with Dinosaur Provincial Park. Image: Mike Skrepnick.

A friendly pet Parasaurolophus, as illustrated by Mike Skrepnick (and used with his permission) for a 'Would Dinosaurs Make Good Pets' project produced in conjunction with Dinosaur Provincial Park. Image: Mike Skrepnick.

Ever keen to give fair credit to those who’ve gone before me, I feel duty-bound to remind you that the issue of hypothetical (non-bird) dinosaur domestication has been covered at least a few times before, mostly in speculative fiction. During both the 1980s and early 2000s (Mash 2003), Robert Mash published his whimsical How to Keep Dinosaurs whereby non-bird dinosaurs as shown acting as pets, livestock, guards and so on in a world where the extinction event never happened. James Gurney's 1992 Dinotopia (and its sequels and spin-offs) features dinosaurs and other animals that - while not technically domesticated - are often featured performing tasks akin to those associated with domesticated animals today. And the 1990s Orbis children’s magazine Dinosaurs! included a well illustrated feature on imaginary dinosaur-human interaction, it being implied that at least some of the dinosaurs were domesticate.

 During the 1990s, British artist Jim Robins produced a whole set of illustrations depicting non-bird dinosaurs in modern, human-dominated scenes. With his permission, I reproduce several of them here. Image: James Robins.

During the 1990s, British artist Jim Robins produced a whole set of illustrations depicting non-bird dinosaurs in modern, human-dominated scenes. With his permission, I reproduce several of them here. Image: James Robins.

Finally, an extremely odd book titled Who Lies Sleeping? – it’s predominantly concerned with the idea that Mesozoic dinosaurs might have evolved human-like intelligence and then caused their own premature extinction via nuclear annihilation or something – urges us to consider the possibility that the extravagant crests, horns and frills of Cretaceous ornithischians might provide compelling evidence of domestication by big-brained dinosaur overlords (Magee 1993). I see.

 An industrial  Parasaurolophus , from  Magee (1993) : "Did the hadrosaurs evolve breathing apparatus to protect themselves from atmospheric pollution", Magee asks. Ok. You might recognise the  Parasaurolophus  if you're familiar with the contents of Bakker's  Dinosaur Heresies . Image:  Magee (1993) .

An industrial Parasaurolophus, from Magee (1993): "Did the hadrosaurs evolve breathing apparatus to protect themselves from atmospheric pollution", Magee asks. Ok. You might recognise the Parasaurolophus if you're familiar with the contents of Bakker's Dinosaur Heresies. Image: Magee (1993).

First things first: what is domestication? Domestication is the creation of a population of living things, distinct in some way from their wild ancestors, that are selectively bred by humans. The main function of domestication involves use of those living things for food, labour, companionship or ornamentation. As evidenced by the extraordinary number and variety of domestic plants and animals, it’s either that we’re really good at it or that it’s really easy once you’re smart enough to wilfully shape the evolution of other living things. Or both. Note that domestication is not simply the grabbing and taming of animals from the wild.

While – when thinking of animals – we typically associate domestication with mammals and birds, don’t forget that animals of virtually all major groups can be domesticated, in theory. Witness domestic insects (Chinese silk moth Bombyx mori and Honeybee Apis mellifera) and fish (Goldfish Carassius auratus, among others) (Zeuner 1964). An argument has been made that there aren’t, technically speaking, any domesticated squamates, turtles or amphibians, but a counter-argument is that we’re actually pretty close to such animals being in existence (look at – for example – the many captive morphs of bearded dragon, some of which lack scales and could not exist outside of human care) and there’s no reason why they couldn’t exist given sufficient selective breeding. Time is one of the key factors here. We could – potentially, perhaps – have giant stripy domestic Komodo dragons by now if only we’d started a selective breeding project a few centuries ago [UPDATE: as discussed in the comments, an argument can be made that the ancient Egyptians domesticated crocodylians].

 Today there are captive lizards - forms of the Central bearded dragon  Pogona vitticeps  - that are significantly novel relative to their wild ancestors. The weirdest is the Silkback or Silkie: a lizard that lacks scales. There's some debate, but these animals are pretty close to being considered 'domestic'. Image: CC0, Pixabay.

Today there are captive lizards - forms of the Central bearded dragon Pogona vitticeps - that are significantly novel relative to their wild ancestors. The weirdest is the Silkback or Silkie: a lizard that lacks scales. There's some debate, but these animals are pretty close to being considered 'domestic'. Image: CC0, Pixabay.

So – could non-bird dinosaurs be domesticated? The short answer is surely: yes, if we had to, or wanted to, and I say this given that humans have proven themselves capable of domesticating just about any organism given sufficient time and motivation, as noted above. That’s a ‘general’ “yes”, however, by which I mean that it refers to non-bird dinosaurs in the general sense of referring to these animals as a group, not in the specific sense of having any one species in mind. For comparison: we’ve domesticated mammals (given that there are sheep and cows and dogs and so on), but that doesn’t mean that we’ve domesticated lions or aye-ayes, to my knowledge, so far.

On that note, it’s rather harder to be confident about the hypothetical domestication of a given species – let’s say, for argument’s sake, Tyrannosaurus rex – given that said hypothetical domestication would be contingent on the biology, behaviour, geographical range, physiology and ecology of that species. T. rex could, for example, be prone to some particular disease that would make its living alongside humans impossible, it could prove truly intractable and just impossible to tame or maintain in a human-controlled environment, or there could be something specific about – say – its life as a juvenile or nestling or even the precise place in which it lives that might also prevent domestication.

 Moose  Alces alces  (these are Alaskan moose) are often mentioned in discussions of domestication attempts. The commonest thing said is that they come with too much baggage to allow domestication -- but this simply might not be true at all. Image: Ryan Hagerty, in the public domain.  Original at Pixnio.

Moose Alces alces (these are Alaskan moose) are often mentioned in discussions of domestication attempts. The commonest thing said is that they come with too much baggage to allow domestication -- but this simply might not be true at all. Image: Ryan Hagerty, in the public domain. Original at Pixnio.

These unknowns are very much hypothetical, obviously, but would surely apply to some species: there are numerous instances where extant animals have proved problematic in the way I imagine. It’s been said, for example, that American moose have been dificult to keep in captivity in Europe, the implication being that some sort of disease prevents their survival in the region (Geist 1999), and then there’s the whole list of animals that have thus far proved either impossible or, at the least, very, very difficult to breed in captivity, predominantly because they don’t seem to like the conditions we provide them with (famous examples include giant pandas, cheetahs and some cranes). Maybe these animals could surely be domesticated in time, but why bother when there are so many other species that don’t come with the same baggage.

Some dinosaurs would be easy to domesticate…. perhaps. It should also be noted that non-bird dinosaurs might actually be well suited for domestication, and not difficult to domesticate at all. The rapid generational turnover, large clutch sizes, and relatively rapid growth of many species (especially small and mid-sized ones) could all make selective breeding and human exploitation desirable and relatively easy, so long as we’re talking about a human effort that extends over decades or centuries, at least. Remember that some workers posit life cycles for non-bird dinosaur where even big species (say, the hadrosaur Maiasaura peeblesorum) were sexually mature at two years of age and had exceeded 1.5 tons by their third year (Woodward et al. 2015).

 If dinosaurs like hadrosaurs (this graph shows  Maiasaura ) grew this quickly - we're talking about an animal that is well over 1000 kg within 2 years  and  capable of breeding by that time, and can be raised on a diet of easily obtained, cheap fodder and produces large clutches to boot - well, we have an animal that might be very desirable as a subject of domestication. Image: Woodward  et al . (2015).

If dinosaurs like hadrosaurs (this graph shows Maiasaura) grew this quickly - we're talking about an animal that is well over 1000 kg within 2 years and capable of breeding by that time, and can be raised on a diet of easily obtained, cheap fodder and produces large clutches to boot - well, we have an animal that might be very desirable as a subject of domestication. Image: Woodward et al. (2015).

The ability of at least some herbivorous dinosaurs to successfully consume and digest material that might be considered low quality, cheap and easy to obtain – we’re talking conifer branches, green vegetation of just about any sort and even wood – would be a major score as goes our ability to maintain these animals as captives over generations.

And the very fact that these animals lay eggs would also be potentially advantageous in a world where dinosaur domestication is possible since it would theoretically be easier to steal babies and raise them in a controlled environment away from their parents. We just don’t know if non-bird dinosaurs exhibited the same sort of imprinting on parental figures that so many birds do, but if they did we again have a feature highly advantageous to human manipulation.

Working With Dinosaurs. When discussing the issues covered so far, I’ve mostly been thinking about domestication that might produce animals bred for meat or egg production, or for use in labour. But what about the maintenance and creation of species used as companions or hunting partners? Here we come to that issue so perennial in discussions of dinosaur biology: intelligence.

 Trained or restrained combat dromaeosaurs: a familiar part of the Jurassic Park universe long pre-dating the Jurassic World movies. This panel is from a 1994 comic published in the Dark Horse Jurassic Park 'Raptors Attack' storyline.

Trained or restrained combat dromaeosaurs: a familiar part of the Jurassic Park universe long pre-dating the Jurassic World movies. This panel is from a 1994 comic published in the Dark Horse Jurassic Park 'Raptors Attack' storyline.

The intelligence – whatever we mean by that term – of non-bird dinosaurs is a subject of eternal debate and uncertainty, and it might be that we never have a good, confident handle on this issue. Intelligence is certainly a red herring when it comes to domestication as a whole. As in, it’s effectively irrelevant: if domestication involves selective breeding and the creation and maintenance of populations designed to be valuable to humans, intelligence has nothing to do with it.

Intelligence does, however, have a role to play if we’re talking about the use of domestic dinosaurs in hunting, herding or in performing other tasks that involve co-operation with a human or another animal or animals. Humans work with dogs in this way, of course, but we also have a long-standing and culturally important association with raptors (yeah, I told you it would be confusing if that word were co-opted for dromaeosaurs. It’s the word we use for hawks, eagles, falcons and such). Without going down the intelligence rabbit-hole too far, it’s conceivable that at least some non-bird theropods, and some ornithischians too, were on par with raptors, owls and other birds that can be used as hunting partners, though I happily admit that this is largely intuitive, difficult to demonstrate and isn’t all that well supported right now (e.g., Balanoff et al. 2013).

 At least some bird-like non-bird maniraptorans were plausibly similar to, or approached, various modern birds in intelligence. Were they as smart as raptors and owls? ... neither of which are that smart compared to birds more 'properly' considered intelligent. At left, a trained Striated caracara  Phalcoboenus australis  extracts sticks in order to obtain a reward. At right, a hybrid Turkmenistan x Siberian eagle owl  Bubo bubo  trained to follow commands and perform in display flights. Neither birds shown here belong to domesticated species, but they're still relevant. Images: Darren Naish.

At least some bird-like non-bird maniraptorans were plausibly similar to, or approached, various modern birds in intelligence. Were they as smart as raptors and owls? ... neither of which are that smart compared to birds more 'properly' considered intelligent. At left, a trained Striated caracara Phalcoboenus australis extracts sticks in order to obtain a reward. At right, a hybrid Turkmenistan x Siberian eagle owl Bubo bubo trained to follow commands and perform in display flights. Neither birds shown here belong to domesticated species, but they're still relevant. Images: Darren Naish.

Let’s say that we could train dromaeosaurs, troodontids or oviraptorosaurs as hunting partners. There would have to be a cut-off as goes which species were considered manageable enough or safe enough to work with – above a certain size and firepower, a predatory theropod, or even a given omnivore or herbivore, could simply be too risky to be around for any length of time. More on this in a minute.

 The effective function of a 'trained police dromaeosaur' as big as this one might not be plausible, but it sure is a cool image (even if the dromaeosaur is shown as unfeathered, tsk). Image: James Robins.

The effective function of a 'trained police dromaeosaur' as big as this one might not be plausible, but it sure is a cool image (even if the dromaeosaur is shown as unfeathered, tsk). Image: James Robins.

Matters of size. A non-trivial aspect of this discussion is dinosaur size. As per the discussion above, it would surely be plausible to raise and tame babies and youngsters of some, most or many dinosaur species that reach gargantuan size as adults. But what about when they reach that size whereby they become uncontrollable?

Elephants are tameable, as we all know, and can generally be controlled by a skilled person following an earlier phase in which the elephant is essentially forced to understand that it has no option but to obey the humans in charge (tangent: there is considerable debate as goes whether tamed elephants should be considered domesticated, since many – but not all – elephants used by people are not domestic as per the definition used earlier in this article). But every now and again a trained elephant snaps, refuses to obey the human giving it commands, and becomes a deadly force that ends in bloodshed, including that of the elephant. I’m not implying in any way that any of the dinosaurs applicable here could have been of intelligence or willpower or ingenuity or whatever on par with an elephant, but you do have to wonder if an angry, disgruntled or disobedient Triceratops or giant sauropod could be controlled or stopped should things go bad. So, what to do about dinosaurs that grow to be ten times – or more! –  the size of an elephant?

 Super-sized sauropods could be bred to be fatter, more muscular and hence higher-yielding than their wild ancestors if domesticated and used for meat production, the result being chubbier, bulkier animals. But what would this mean for the controlling and handling of such massive, powerful animals? Could we even control them at all? Image: Ethan Kocak.

Super-sized sauropods could be bred to be fatter, more muscular and hence higher-yielding than their wild ancestors if domesticated and used for meat production, the result being chubbier, bulkier animals. But what would this mean for the controlling and handling of such massive, powerful animals? Could we even control them at all? Image: Ethan Kocak.

  • Solution 1: cull or release said animals when they reach whatever size has been found to be at the unmanageable threshold. Releasing them might be unworkable given that their familiarity with humans could then make them a nuisance or disruptive problem to whatever society was using them.
  • Solution 2: via selective breeding, produce dwarf forms that do not get to that threshold. Though… why bother when there are non-giant species of similar form?
  • Solution 3: invent and employ technology that forces the animal in question to comply, or prevents them from disobeying. We might speculate that devices like shock collars, neural transmitters or instant chemical suppressants would not act swiftly enough to stop any dinosaur giant from inflicting problematic damage… unless it were instantly fatal. I’m reminded here of the method used by Hasdrubal of the Carthaginian Republic – Hannibal’s brother – to deal with an armoured war elephant should it do an about-turn on the battlefield and start charging through the lines of its own side: supposedly, the elephant’s riders were armed with a mallet and spike, and would drive the spike into the elephant’s spine should it need to be disabled (Spinage 1994). I don’t know how easy this would be, but there we go. Solution 3 would surely involve practices rightly deemed unethical.
 Armoured war elephants have been used by several cultures. This image originally appeared in Friedrich Arnold Brokhaus's atlas, published sometime before 1850. Image: Karl Gröning;  Elephants: A Cultural and Natural History .

Armoured war elephants have been used by several cultures. This image originally appeared in Friedrich Arnold Brokhaus's atlas, published sometime before 1850. Image: Karl Gröning; Elephants: A Cultural and Natural History.

I should also add that the emergency disabling or killing of a giant dinosaur would create all kinds of practical and logistic issues given the size of the carcass. The swift and efficient removal of a multi-ton carcass would be a necessity if the disobedient dinosaur in question succumbed in a settled area, else we could well be talking about a serious public health issue: there are the swarms of scavenging insects, enough fluidic runoff to influence local water sources, noxious smells not generally considered fun to be around by humans, and potentially the attraction of scavengers… maybe lots of them.

Domestication is ‘good’, domestication is ‘bad’. As is the case for some modern domesticate animals, becoming domesticated could be a short-term ‘wise move’ for the species concerned given human protection and propagation. Conversely, non-domesticate wild relatives of the species concerned would be at a disadvantage as their habitat was captured or managed for the domestic one, and perhaps as the risk of competition, hybridisation and aggression from said wild relatives would encourage humans to persecute and even destroy them. Exactly such a thing is said to have affected tarpans – wild relatives and near-ancestors of the domestic horse – given that tarpan stallions were apparently able to out-compete domestic stallions in mating battles and thus introduce non-desirable traits into the next generation.

A logical outcome of non-bird dinosaur domestication, then, might be ‘monocultures’ where a handful of species are super-abundant, the landscape has been changed to best suit the domestic species concerned, and wild relatives are rare, marginalised, or forced into extinction.

 Small theropods, and maybe other dinosaurs too, might make good pets, though they may prove incompatible with smaller pets of different species, as implied here. This is another of the illustrations produced by Jim Robins described above (and used here with permission). Image: James Robins.

Small theropods, and maybe other dinosaurs too, might make good pets, though they may prove incompatible with smaller pets of different species, as implied here. This is another of the illustrations produced by Jim Robins described above (and used here with permission). Image: James Robins.

That which might be possible. An over-riding pattern of domestication has been and is our ability to find variable anatomical traits in animals, and to induce, via often skilled selective breeding, the elaboration or modification of that trait, often at reasonable speed across the generations. I should add that this is the issue discussed and illustrated at length in Katrina van Grouw’s 2018 book Unnatural Selection (van Grouw 2018). We might not know anything particularly useful about potential intraspecific variation within non-bird dinosaur species, but we at least have some indication of what could be possible given the features that proved variable in their real word, natural evolution.

With that in mind, let’s speculate and have fun considering some of the things that might be possible in a world where non-bird dinosaurs have been domesticated, and where humans have become good at selectively breeding them. The showiness and elaborate ornamentation of ceratopsians and other ornithischians could mean that whichever species have been domesticated have been selected for ever more elaborate, ever showier structures.

Alternatively, the dangerous nature of horned and armoured ornithischians could encourage the development of unarmoured, less destructive, more benign forms that are positively plain and bland relative to their ancestors.

 A bland, domesticated ceratopsid with reduced cranial structures relative to its wild ancestor. Image: Darren Naish.

A bland, domesticated ceratopsid with reduced cranial structures relative to its wild ancestor. Image: Darren Naish.

Fighting dromaeosaurs, troodontids or small ceratopsians – equipped with more lethal armament and tougher hides and external coverings than their wild ancestors – could be developed in a society that did not prevent or make illegal the maintenance of such animals, a history that could lead to the creation of incredibly dangerous, near-uncontrollable strains or individuals. Maybe such animals could form the breeding stock of dinosaurs designed for urban pacification, crime-fighting and military application, though the last of these idea is now sadly and woefully unoriginal given the several appearances of trained or semi-trained ‘combat velociraptors’ in the Jurassic Park and Jurassic World franchises. Picture a trained, burly dromaeosaur with body and limb armour, prosthetic pseudoteeth, and a tight, perpetually reinforced bond with the lone human handler who has lived with it its whole life.

 A girl and her ornithomimid. Greg Paul said of theropods in   Predatory Dinosaurs of the World   that "Their stiff, perhaps feathery bodies were not what one would care to have sleep at the foot of the bed" ( Paul 1988 , p. 19), but maybe he was wrong. This is another of Mike Skrepnick's illustrations from his 'Would Dinosaurs Make Good Pets' project. Image: Mike Skrepnick.

A girl and her ornithomimid. Greg Paul said of theropods in Predatory Dinosaurs of the World that "Their stiff, perhaps feathery bodies were not what one would care to have sleep at the foot of the bed" (Paul 1988, p. 19), but maybe he was wrong. This is another of Mike Skrepnick's illustrations from his 'Would Dinosaurs Make Good Pets' project. Image: Mike Skrepnick.

At the other end of the scale, the cute appeal of at least some non-bird dinosaurs – various of the short-faced maniraptorans and small ornithischians – could result in ubercute, cuddly, dwarf forms or super-showy ornamental forms with spangly, iridescent exteriors and hypertrophied tails. There might be a subculture whereby keepers of fancy maniraptorans have bred radically surreal, immaculately groomed and bejewelled creatures deliberately made to mimic objects of heraldry or fiction, like dragons, griffins or aliens from sci-fi stories.

 I give you captively bred domestic theropods created by Ethan Kocak: a short-faced, miniature tyranno-pug, and a plush, poodle-like maniraptoran with luxuriant plumage. Image: Ethan Kocak. He's a  New York Times  bestselling artist, dontchaknow.

I give you captively bred domestic theropods created by Ethan Kocak: a short-faced, miniature tyranno-pug, and a plush, poodle-like maniraptoran with luxuriant plumage. Image: Ethan Kocak. He's a New York Times bestselling artist, dontchaknow.

Here's to the future of domestic dinosaurs. My overall conclusion – I think it’s pretty obvious by now – is that non-bird dinosaurs of several, many or most sorts could indeed be domesticated, and might in fact be domesticated quite easily. We might maintain species of many sorts as beasts of burden or for food; predators and nimble omnivores and herbivores could perhaps be used in stock control or as partners in hunting; and those animals we might consider attractive or visually or vocally interesting might be bred, modified and managed as show animals or companions. It goes without saying that all of this is nothing more than an exercise in speculative fiction, and that none of it has any real relevance as goes our relationship with those animals we really do live alongside.

But wouldn’t it make a cool book?

Tetrapod Zoology is now an independent blog. I can devote more time to it and produce more content the more support I receive. Support me at patreon and you also get to see the more than 550 in-prep illustrations being produced for my various projects.

Refs - -

Balanoff, A. M., Bever, G. S., Rowe, T. B. & Norell, M. A. 2013. Evolutionary origins of the avian brain. Nature 501, 93-96.

Geist, V. 1999. Deer of the World. Swan Hill Press, Shrewsbury.

Magee, M. 1993. Who Lies Sleeping: the Dinosaur Heritage and the Extinction of Man. AskWhy! Publications, Frome.

Mash, R. 2003. How to Keep Dinosaurs. Weidenfeld & Nicolson, London.

Paul, G. S. 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.

Spinage, C. A. 1994. Elephants. T & A D Poyser, London.

van Grouw, K. 2018. Unnatural Selection. Princeton University Press, Princeton and Oxford.

Woodward, H. N., E. A., Freedman Fowler, Farlow, J. O. & Horner, J. R. 2015. Maiasaura, a model organism for extinct vertebrate population biology: a large sample statistical assessment of growth dynamics and survivorship. Paleobiology 41, 503-527.

Zeuner, F. E. 1964. A History of Domesticated Animals. Hutchinson of London, London.

A 1996 Letter from Dr Phil Currie

I think I might get into the habit of posting short articles here: putting longer pieces together takes a while and is often difficult to do in view of other work (hint goddam hint). Let me know what you think about this. Anyway...

  Phil Currie's  The Flying Dinosaurs  (Currie 1991) . The artwork is innovative and often really interesting, even though the coelurosaurian dinosaurs are mostly shown as un-feathered. The book includes pterosaurs... which is weird but in keeping with the 'evolution of flight' subtitle. Image: Darren Naish

Phil Currie's The Flying Dinosaurs (Currie 1991). The artwork is innovative and often really interesting, even though the coelurosaurian dinosaurs are mostly shown as un-feathered. The book includes pterosaurs... which is weird but in keeping with the 'evolution of flight' subtitle. Image: Darren Naish

I used to write a lot of letters. I mean: physical, printed letters, on paper. Some of them resulted in interesting things, and others didn't. During the mid-1990s, I obtained and read Philip Currie's 1991 book The Flying Dinosaurs, richly illustrated by Jan Sovak and showing many animals brand-new at the time. And it has some weird stuff in it that had me curious (I won't start discussing that "weird stuff", as it will take me a while to find it in the book, explain it, and put it into context). I managed to obtain Dr Currie's postal address, and wrote to him. And I was thrilled to get a response! Here it is...

Philip-Currie-letter-1996-Aug-2018-Darren-Naish-Tetrapod-Zoology.jpg

Please understand that I am not, in any way, sharing this to shame Dr Currie - heavens, no. I merely opted to share it because it's (for me) an interesting piece of personal history. And it also shows what the situation was - and/or is - like for many busy academics. Today, I know this pain all too well. Modern correspondence, of course, has mostly changed from paper letters to emails. I'm not as famous as Dr Currie, but even I get so much correspondence that I either have to deliberately ignore some of it, or put it to one side such that I can 'deal with it later', only for 'later' to become 'never' as other things destroy those various other plans. I would like to remind others of this when they start sending reminder messages about the responses they'd like. Sorry: there comes a point in life when it is IMPOSSIBLE to keep up with correspondence.

Anyway, I was later to meet Phil Currie on one or two occasions and talk with him about a whole bunch of stuff. Here he is (standing) at a London-based conference that happened in May 2008...

 Image: Darren Naish.

Image: Darren Naish.

New Dinosaur Books, Part 2: Ben Garrod’s ‘So You Think You Know About… Dinosaurs’ Series

Today we press on with my brief(ish) reviews of recently(ish) published books on Mesozoic dinosaurs – I have quite a backlog – and this time it’s…. the So You Think You Know About… Dinosaurs books, by Ben Garrod. Ben is a qualified zoologist and TV presenter, perhaps best known for the BBC 2014 series Secrets of Bones. I should also add that Ben spoke at the 2017 TetZooCon, so now you know he’s awesome.

 Ben Garrod's new dinosaur books - another three are due to appear very soon. Image: Darren Naish.

Ben Garrod's new dinosaur books - another three are due to appear very soon. Image: Darren Naish.

Anyway, Dr Garrod has now made a foray into the World of Mesozoic Dinosaurs, and a pretty respectable foray it is. These small, compact books – they remind me of the Horrible Histories volumes a little – provide a huge quantity of information on the dinosaurs they cover, do so in a fun, attractive way, and are very much on-the-ball as goes the current state of our knowledge. They are very much science advocacy tools that don’t shy away from presenting and discussing such topics as complex as sexual selection and evolutionary arms races. The dinosaurian nature of birds is emphasised – dinosaurs can’t be extinct so long as around 10,000 of them are currently alive – and the books are good at conveying the phylogenetic position of the relevant dinosaurs and where they fit within geological time.

Black and white cartoons, diagrams and other images are on virtually every page. If you move in the same social media spheres that I do (I’m @TetZoo on Twitter and Instagram) – surprise! – the cartoons are by Ethan Kocak, and illustrations and diagrams by Gabriel Ugueto and Scott Hartman also appear within.

 Excellent illustrations by Gabriel Ugueto appear in each of the books. Hey, Gabriel and I actually worked together on a large poster about tyrannosaurs included in a magazine -- it was published early in 2018 but I still haven't seen it because the publishers never sent me a copy and no longer have any to provide. Huh. Image: Gabriel Ugueto/Ben Garrod/Zephyr.

Excellent illustrations by Gabriel Ugueto appear in each of the books. Hey, Gabriel and I actually worked together on a large poster about tyrannosaurs included in a magazine -- it was published early in 2018 but I still haven't seen it because the publishers never sent me a copy and no longer have any to provide. Huh. Image: Gabriel Ugueto/Ben Garrod/Zephyr.

Regarding what I said a moment ago about these books reflecting ‘the current state of our knowledge’, I’m especially pleased to see inflatable nose balloons featured in the Triceratops volume (for more on this issue see this Tet Zoo ver 3 article from 2016). The Diplodocus volume has a brief discussion of the whole trunks issue (covered here at Tet Zoo ver 3).

 Ceratopsian nose balloons go mainstream (images by Ethan Kocak). I can't take credit for this.  Or can I . Images: Ethan Kocak/Ben Garrod/Zephyr.

Ceratopsian nose balloons go mainstream (images by Ethan Kocak). I can't take credit for this. Or can I. Images: Ethan Kocak/Ben Garrod/Zephyr.

Quibbles: two or three maniraptoran silhouettes are shown as un-feathered, and I will forgive the Tyrannosaurus book for using the full binomial throughout even though the dinosaurs of the other books are only ever mentioned by their generic names. T. rex exceptionalism, we call it.

 It's nothing to with Ben Garrod's books, but I thought I'd include another image of ceratopsian nose balloons for good measure. This brilliant piece is by J. W. Kirby and the original can be seen  here at KirbyniferousRegret's deviantart page . Image: J. W. Kirby.

It's nothing to with Ben Garrod's books, but I thought I'd include another image of ceratopsian nose balloons for good measure. This brilliant piece is by J. W. Kirby and the original can be seen here at KirbyniferousRegret's deviantart page. Image: J. W. Kirby.

I really like these books and strongly recommend them for young readers (I suppose the target audience is kids between 6 and 16, or so). Ben is big on ‘embracing your inner geek’, and on encouraging young people to be proud of their curiosity and interest in science and nature, and I’m sure this is something that everyone reading this wants to see promoted and celebrated too. Three additional volumes – on Spinosaurus, Velociraptor and Stegosaurus – are finished and due to appear in print very soon (so I understand).

Ben Garrod, 2018. So You Think You Know About… Tyrannosaurus rex? Zephyr, London. ISBN 9781786697844. Hardback, 106 pp. Here at amazon. Here at amazon.co.uk. Here from the publishers.

Ben Garrod, 2018. So You Think You Know About… Diplodocus? Zephyr, London. ISBN 9781786697868. Hardback, 112 pp. Here at amazon. Here at amazon.co.uk. Here from the publishers.

Ben Garrod, 2018. So You Think You Know About… Triceratops? Zephyr, London. ISBN 9781786697882. Hardback, 108 pp. Here at amazon. Here at amazon.co.uk. Here from the publishers.

Once these book reviews are out of the way, get set for some novel dinosaur-themed content here. Here's your regular reminder that this blog relies on support via patreon, thank you to those providing support already.

For previous Tet Zoo book reviews on dinosaurs (I've now taken to adding dates to articles of the past; I find this useful)...

New Dinosaur Books, Part 1: Barrett on Stegosaurs

I’ve always reviewed books here at Tet Zoo, and here at ver 4 I fully intend to condition this fine tradition. Over the next few articles I’ll be discussing new (or newish) books on Mesozoic dinosaurs. And we start with something very special – a whole book devoted to stegosaurs. This might be a first (UPDATE: I just remembered that the late Beverly Halstead wrote a semi-fictional book on the life history of Stegosaurus).

 Cover of Barrett (2017)... perhaps the only published volume fully devoted to stegosaurs and stegosaurs alone.

Cover of Barrett (2017)... perhaps the only published volume fully devoted to stegosaurs and stegosaurs alone.

Stegosaurus: An Extraordinary Specimen and the Secrets it Reveals by Paul M. Barrett tells the story of ‘Sophie’ (initially ‘Sarah’, and technically NHMUK PV R36730), the excellent, complete Stegosaurus specimen discovered and excavated at Shell, Wyoming in 2003/4. You’ll have seen this outstanding specimen mounted on display if you’ve visited London’s Natural History Museum. Already Sophie has been the focus of a whole bunch of quality technical publications by the author and his colleagues (Brassey et al. 2015, Maidment et al. 2015, Lautenschlager et al. 2016).

 A look inside. This is an info-packed book with great graphics and loads of photos and diagrams. 

A look inside. This is an info-packed book with great graphics and loads of photos and diagrams. 

 Sophie the  Stegosaurus  - or NHMUK PV R36730, if you prefer - in person at the Natural History Museum, London. I'm sure many of us have an unreasonable number of photos of this amazing specimen. Image: Darren Naish.

Sophie the Stegosaurus - or NHMUK PV R36730, if you prefer - in person at the Natural History Museum, London. I'm sure many of us have an unreasonable number of photos of this amazing specimen. Image: Darren Naish.

The book does so much more than tell the story of Sophie: it’s also a very good review of what we know (or think we know) about stegosaur biology. The text is dense, packed with scientific content, and highly readable. Excellent images appear throughout, ranging from photos of the specimen and CT renders generated during research to life reconstructions and maps. I initially assumed – no offence intended to the author – that this would be a text-light book written for kids, with more space than text on its pages. That’s not the case at all: there’s a ton of information here and anyone interested in dinosaurs should get this book. Older kids with an interest in science or dinosaurs will enjoy it, and adults will too. Some of the taxonomic conclusions discussed in the review of stegosaur diversity – that Hesperosaurus is synonymous with Stegosaurus and Miragaia is synonymous with Dacentrurus – have been overturned in work published since the book saw print (Raven & Maidment 2017).

 A reminder that the stegosaurs of your parents or grandparents - or those of a recent terrible movie franchise - are not in keeping with the way these animals more likely looked. These drawings are old, excuse the GSP tail muscles. Image: Darren Naish.

A reminder that the stegosaurs of your parents or grandparents - or those of a recent terrible movie franchise - are not in keeping with the way these animals more likely looked. These drawings are old, excuse the GSP tail muscles. Image: Darren Naish.

In short: highly recommended for everyone, and dead cheap as well.

Disclaimer: the author and I previously worked together on another dinosaur book published by The Natural History Museum, namely Dinosaurs: How They Lived and Evolved (Naish & Barrett 2016). As some of you know, a second edition of that book – including numerous tweaks and updates – appears in print very soon, so I’ll be talking about it then.

Paul M. Barrett, 2017. Stegosaurus: An Extraordinary Specimen and the Secrets it Reveals. Natural History Museum, London. ISBN 9780565093884. Hardback, 108 pp. Here at amazon. Here at amazon.co.uk. Here from the publishers.

Stegosaurs have been covered quite a few times at Tet Zoo previously. See…

Things are going very well here at ver 4: I’m keeping an eye on the hits counter and the comments, and things are good. Many thanks to everyone who’s helping to make it work. Here’s your reminder that the more support I receive at patreon, the more time I can spend writing and publish the stuff you like to read. Thanks to those who support this endeavour already. As a Tet Zoo patron you get to see stuff coming together behind-the-scenes: there are already over 550 in-prep illustrations and pieces of text there.

Refs - -

Brassey, C. A., Maidment, S. C. R. & Barrett, P. M. 2015 Body mass estimates of an exceptionally complete Stegosaurus (Ornithischia: Thyreophora): comparing volumetric and linear bivariate mass estimation methods. Biology Letters 11: 20140984.

Lautenschlager, S., Brassey, C. A., Button, D. J. & Barrett, P. M. 2016. Decoupled form and function in disparate herbivorous dinosaur clades. Scientific Reports 6, 26495.

Naish, D. & Barrett, P. M. 2016. Dinosaurs: How They Lived and Evolved. The Natural History Museum, London.

Maidment, S. C. R., Brassey, C. & Barrett, P. M. 2015. The postcranial skeleton of an exceptionally complete individual of the plated dinosaur Stegosaurus stenops (Dinosauria: Thyreophora) from the Upper Jurassic Morrison Formation of Wyoming, U.S.A. PLoS ONE 10 (10): e0138352.

 Raven, T. J. & Maidment, S. C. R. 2017. A new phylogeny of Stegosauria (Dinosauria, Ornithischia). Paleontology 60, 401-408.