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.

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.

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.

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).

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 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).

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).

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).

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).

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…

• Paul Brinkman’s The Second Jurassic Dinosaur Rush, November 2011

• That Brontosaurus Thing, April 2015

• New Dinosaur Books, Part 1: Barrett on Stegosaurs, August 2018

• New Dinosaur Books, Part 2: Ben Garrod’s ‘So You Think You Know About… Dinosaurs’ Series, August 2018

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.

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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.

Caption: at left, an 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.

Caption: 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).

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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).

Caption: 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.

Caption: 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!

Caption: 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 reasons 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…

• Surreal caecilians part I: tentacles and protrusible eyes, January 2008

• Surreal caecilians part II: pass mum’s skin, hold the mayo, January 2008

• Close up to Andrias, despite the smell and the teeth, December 2010

• Welcome to the Squamozoic!, April 2013

• If Bigfoot Were Real, June 2016

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. Gigantopithecus. Scientific 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).

Caption: 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.

Caption: 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).

Caption: 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).

Caption: 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.

Caption: 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).

Caption: 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.

Caption: 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.

Caption: 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…

Caption: 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.

Caption: 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…

• Multiple new species of large, living mammal (part I), June 2007

• Because you can never have too many tapirs, September 2009

• The biggest tapir, September 2009

• Tapir attacks past, present, but hopefully not future, August 2013

• A new living species of large mammal: hello, Tapirus kabomani!, December 2013

• World Tapir Day, 2015, April 2015

• On World Tapir Day, a Quick Look at (Part of) Tapir History, April 2016

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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 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.

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 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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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…

• The events of TetZooCon 2014, July 2014

• TetZooCon 2015 Is On, July 2015

• The Events of TetZooCon 2015, November 2015

• Coming Soon: TetZooCon 2016, September 2016

• The Day After TetZooCon, October 2016

• The Fourth TetZooCon, September 2017

• The TetZooCon of 2017, October 2017

• Reasons to Attend TetZooCon 2018, September 2018

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…

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.

Caption: 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.

Caption: 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.

Caption: 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!

Caption: 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.

Caption: 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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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 events of TetZooCon 2014, July 2014

• TetZooCon 2015 Is On, July 2015

• The Events of TetZooCon 2015, November 2015

• Coming Soon: TetZooCon 2016, September 2016

• The Day After TetZooCon, October 2016

• The Fourth TetZooCon, September 2017

• The TetZooCon of 2017, October 2017

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

Caption: 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).

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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).

Caption: 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.

Caption: 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!

Caption: 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…

• Speculative Zoology: Wedel throws down the gauntlet, February 2007

• Oh no, not another giant predatory flightless bat from the future, March 2007

• Come back Lank, (nearly) all is forgiven, September 2008

• The Tet Zoo guide to the creatures of Avatar, January 2010

• The science of Godzilla, 2010, November 2010

• The anatomy of Zilla, the TriStar ‘Godzilla’, November 2010

• Giant flightless bats from the future, November 2010

• Welcome to the Squamozoic!, April 2013

• Of After Man, The New Dinosaurs and Greenworld: an interview with Dougal Dixon, April 2014

• The LonCon3 Speculative Biology event, August 2014

• Speculative Zoology at Tet Zoo, The Story So Far, May 2015

• Summer 2018 at Tet Zoo Towers, July 2018

• Speculative Zoology, a Discussion, July 2018

• Could We Domesticate (Non-Bird) Dinosaurs?, August 2018

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).

Caption: 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.

Caption: 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.

Caption: 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?

Caption: 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?

Caption: 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.

Caption: 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).

Caption: 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.

Caption: 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.

Caption: 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.

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).

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...

• Tet Zoo picture of the day # 3 (Elasmotherium), May 2007

• The remarkable life appearance of the Woolly rhino, November 2013

• Spots, Stripes and Spreading Hooves in the Horses of the Ice Age, February 2015

And for articles on deer, see...

• Oh deer oh deer oh deer, October 2006

• Deer oh deer, this joke gets worse every time I use it, November 2007

• Britain: wildlife theme-park, December 2007

• At the 56th SVPCA - hello Dublin! (Megaloceros), September 2008

• The plasticity of deer, October 2008

• Pouches, pockets and sacs in the heads, necks and chests of mammals, part IV: reindeer and a whole slew of others, October 2010

• The seemingly endless weirdosity of the Milu, October 2011

• Eld's deer: endangered, persisting in fragmented populations, and morphologically weird... but it wasn't always so, April 2012

• South America's Many Remarkable Deer, November 2014

• Confrontational behaviour and bipedality in deer, December 2014

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,

Caption: 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.

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…

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.

Caption: 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).

Caption: 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).

Caption: 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.

Caption: 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.

Caption: 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.

Caption: the temporal field generator is always on. Image: Darren Naish.

Is this really the very end of Dinosaurs in the Wild? Will it re-appear elsewhere at some future date? As you might guess, there are certainly plans that such a thing might happen. Right now, I can say no more. News will be reported here, and on relevant social media streams (I'm at @TetZoo, and check @DinosInTheWild too).

This blog, my research and many of my projects are made possible thanks to your kind support at patreon. The more support I receive, the more able I become to spend time producing the content you like to see: the Dinosaurs in the Wild book discussed here, for example, could be published if only I were able to devote the time to it.

Refs - -

Conway, J., Kosemen, C. M., Naish, D. & Hartman, S. 2012. All Yesterdays: Unique and Speculative Views of Dinosaurs and Other Prehistoric Animals. Irregular Books.

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

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.

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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 within 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).

Caption: 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.

Caption: 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).

Caption: 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).

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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).

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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.

Caption: 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).

Caption: 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.

Caption: 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...

• Harduns and toad-heads; a tale of arenicoly and over-looked convergence, December 2006

• Ermentrude the liolaemine, February 2008

• ‘Cryptic intermediates’ and the evolution of chameleons, June 2008

• Tell me something new about basilisks, puh-lease, January 2009

• Amazing social life of the Green iguana, September 2012

• The Squamozoic actually happened (kind of): giant herbivorous lizards in the Paleogene, June 2013

• The enormous liolaemine radiation: paradoxical herbivory, viviparity, evolutionary cul-de-sacs and the impending mass extinction, June 2013

• Leiosaurus: big heads, bold patterns, October 2013

• Grassland earless dragons, January 2014

• Australia, land of dragons (by which I mean: agamids) (part I), January 2014

• Australia, land of dragons (part II), February 2014

• What's With All These New Chameleon Names?, Part 1, February 2016

• By the Horns of Trioceros, the Casque of Calumma, the Brood of Bradypodion--Chameleons, Part 2, February 2016

• Palleon, Archaius, Kinyongia, Nadzikambia--The Last Chameleons, Part 3, March 2016

• The Tropidurine Treerunners, December 2017

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).

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.

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).

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.

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).

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).

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).

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.

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...

• Terrestrial stalking azhdarchids, the paper, May 2008

• Shemhazai and other flightless pterosaurs, September 2008

• Come back Lank, (nearly) all is forgiven, September 2008

• Giant pterosaurs invade London, Summer 2010, July 2010

• Pterosaurs, err, indoors (the Summer 2010 exhibition), July 2010

• Getting scansoriopterygids, terrestrial-stalking azhdarchids, sauropod pneumaticity and the word palaeontography into a kid’s book, July 2010

• The Cretaceous birds and pterosaurs of Cornet: part II, the pterosaurs, September 2010

• Milton Keynes: where giant pterosaurs go to die, May 2011

• What does it feel like to get bitten by a ground hornbill, I hear you ask?, June 2011

• A new azhdarchid pterosaur: the view from Europe becomes ever more interesting, Jan 2013

• Daisy’s Isle of Wight Dragon and why China has what Europe does not, March 2013

• In Rio for the 2013 International Symposium on Pterosaurs, May 2013

• Quetzalcoatlus: the evil, pin-headed, toothy nightmare monster that wants to eat your soul, August 2013

• Were azhdarchid pterosaurs really terrestrial stalkers? The evidence says yes, yes they (probably) were, December 2013

• Some Azhdarchid Pterosaurs Were Robust-Necked Top-Tier Predators, February 2017

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.

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

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...

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.

Seated at the table, we see Darren Tanke (left, near Phil), Carmen Naranjo-Marquez (far left), Luis Rey (far right) and C. M. Kosemen (closest to us, right).

Next: probably more dinosaur-themed book reviews...

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.

Ref - -

Currie, P. J. 1991. The Flying Dinosaurs. Red Deer College Press, Red Deer, Alberta.

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.

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.

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.

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.

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)...

• Long and Schouten's Feathered Dinosaurs, a review, January 2008
• Luis Chiappe's Glorified Dinosaurs: The Origin and Early Evolution of Birds, January 2011
• Greg Paul's Dinosaurs: A Field Guide, February 2012
• New Dinosaur Books, Part 1: Barrett on Stegosaurs, August 2018

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The podcast feed has moved to http://tetzoo.com/podcast/?format=rss, please subscribe to that URL in your podcatchers to continue getting episodes!

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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.

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. 

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.

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…

• A most atypical stegosaur, November 2007
• Stegosaur Wars: the SJG stegosaur special, part I, December 2010
• Heinrich’s digital Kentrosaurus: the SJG stegosaur special, part II, January 2011
• Life as a stegosaur: the SJG stegosaur special, part III, January 2011
• Dinosaurs and their exaggerated structures : species recognition aids, or sexual display devices?, April 2013
• The Stegosaurus Plate Controversy, July 2016
• Stegosaurs and the Species Recognition Hypothesis, August 2016

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.

Bigfoot’s Genitals: What Do We Know?

Ok, I will fully admit upfront that the article you’re about to read (or: about to stop reading, depending on your preference) is included here both because it’s linked to something in the news, and because it’s sensational and perhaps amusing.

Caption: the offending tweet. What we're all asking is: how anatomically accurate might this depiction be? Spoiler: not accurate at all (err.. assuming that Bigfoot even exists, a minor detail). Image: Twitter.

Without going into all the details, US Republican candidate Denver Riggleman (great name) has been outed as an alleged fan of Bigfoot erotic fiction… yeah, this is a thing, come on you knew that don't pretend you didn't… though I do wonder whether this is some sort of ploy to downplay said candidate’s more worrying association with white supremacists [UPDATE: Loren Coleman tells me that Mr Riggleman has a long-term, serious interest in Bigfootery]. Bigfoot erotic fiction? The fact that – like it or not – Bigfoot is regarded as light-relief and harmless hokum means that any link between Bigfoot and sex is immediately treated as a joke. News stories associated with Riggleman’s niche interests have featured an illustration of a male Bigfoot, its seemingly substantial sexual organ safely censored by a long and subtle black box (see if you can spot it in the image above). And this leads us to the ultimate question: what have people honestly, seriously, thought about the genital anatomy of Bigfoot? Inspired by a twitter exchange with Russ Dobler and Kyle Marian, I thought this was as good an opportunity as any to summarise what’s on the record.

First things first: does this mean that I consider Bigfoot a real, genuine, flesh and blood, undiscovered primate? Let’s just say for now that I’m really sceptical of this notion (for reasons discussed here, and in my 2017 book Hunting Monsters). I’m not a knee-jerk sceptic though, and long to have my mind changed…

Caption: I've only had one up-close encounter with a Bigfoot myself (it happened in California); I didn't have opportunity at the time to do any checking as goes any details of anatomy, but here's the proof. Image: [safely anonymous source]/Darren Naish.

One of the main aims of my writing is to summarise or convey what others have said before me and, like it or not, a bunch of authors who’ve written about Bigfoot as a real biological entity have indeed considered the subject of its genital anatomy. After all, if you regard Bigfoot as a real animal it’s perfectly valid to consider all kinds of aspects of its biology, behaviour and evolution, even if we’re doing little more than speculating.

Caption: if Bigfoot is real, just remember that it's probably the most terrifying animal in existence. Seriously. I tried to modify my drawing (available on merchandise!) so that the animal appears to be in the dark. I failed. Image: Darren Naish.

So, if you look at the Bigfoot literature you’ll see a fair few mentions of male genital anatomy, these most typically being references to “a small penis and scrotum” (Krantz 1999, p. 155). When a penis is mentioned, it’s virtually always flaccid… though there are exceptions. The infamous ‘Kong’ account (in which the semi-anonymous ‘Jan Klement’ described his long-time association during the 1970s with an animal that regularly visited his property) includes an event whereby ‘Kong’ exhibited a tumid penis and interacted sexually with a cow.

Then there’s the ‘Redwoods’ incident. In 1995, an alleged Bigfoot was filmed (from a vehicle) in Jedediah Smith Redwoods State Park, California. And what appears to be a slender, tapering, mobile, sickle-shaped penis flexes upwards during the footage. Some writers have even interpreted this as a possible threat display (Meldrum & Greenwell 1998). This piece of footage is known generally as the ‘Redwoods Footage’, but the fact that it was filmed by a crew working for the Playboy company – and hence is also often called the ‘Playboy Footage’ – hasn’t exactly helped its credibility.

Caption: at left, we see a rather grainy still from the 'Redwoods footage'; at right is an interpretation (drawn by Peter Visscher, based on an initial interpretation produced by Jeff Meldrum). The penis isn't visible in this part of the footage. Image: Meldrum & Greenwell (1998)/BBC Wildlife.

A reasonably large, erect penis has also been described in an observation of a Central Asian bar-manu, one of several creatures sometimes suggested to be Eurasian relatives or versions of North America’s Bigfoot. And then there’s the whole world of speculation concerning whether ancient depictions of satyrs, wildmen and so on with erect penises are relevant to such sightings and their validity. I discuss this issue further in Hunting Monsters (Naish 2017).

Caption: herpetologist and cryptozoologist Jordi Magraner drew this obviously male bar-manu (a crypto-hominid reported from Pakistan) as described by a witness. The account was published by Michel Raynal (2001). Image: Raynal (2001).

Loren Coleman included a whole chapter on sex and genital anatomy in his 2003 Bigfoot! The True Story of Apes in America (Coleman 2003). Indeed, he noted at the start of this chapter that the general absence of writings on Bigfoot sexual behaviour and anatomy seems weird given the importance and significance of these things to primates. “Bigfoot are most frequently shown without any male genitalia even though a penis might be part of a witness description”, he wrote (Coleman 2003, p. 185).

Caption: Loren Coleman's 2003 Bigfoot! includes a whole chapter on ideas and observations about sexual behaviour and anatomy. If Bigfoot isn't real, maybe this stuff is fatuous. Then again, even if Bigfoot isn't real, our attitude to this stuff might still tell us something. And if Bigfoot is real? Well... Image: Simon & Schuster.

A few other eyewitness encounters have described a penis. Albert Ostman – the Canadian logger and construction worker supposedly kidnapped by a Bigfoot family during the 1920s – described the penis of the large male boss of the group as short (around two inches long) and “hooded with skin”, or – according to researcher John Green following his communication with Ostman – “like an inverted funnel, which sounds horse-like” (Coleman 2003, p. 191). I should add for the record that I don’t believe Ostman’s encounter ever happened, strange as that might seem. And the other accounts of which I’m aware refer to an organ proportionally smaller than that of an average human – there aren’t (so far as I’m aware) reports of giant, swinging structures of impressive length or girth, excuse the mental image.

The general impression we get from this limited information (assuming here that Bigfoot is real, of course) is somewhat contradictory but we can couch what we know within the social systems and breeding strategies of other primates (e.g., Harcourt & Gardiner 1994). Because there’s no indication of giant testes (like those of chimps), we might infer that sperm competition is not an evolutionary driver for Bigfoot, and thus that they presumably have a monogamous or near-monogamous mating system. This is in keeping with the general idea that Bigfoot lives in small bands, perhaps involving a mating pair and their offspring. The apparently proportionally small size of the penis could be seen as being consistent with this, since it might indicate that the genitals are not used in social display and intimidation, as they are in humans and (to a degree) chimps and bonobos. On the other hand, a proportionally small penis could also be inconsistent here, given the hypothesis that large penises in humans are supposedly linked with monogamy/near-monogamy (note: supposedly. I must avoid discussing the argument over human sexual behaviour and mating systems). The contradiction comes from the Redwoods footage, though, since this seemingly reported a relatively large penis used as a display, err, tool.

I will finish this discussion on male genitals by noting that there’s been some serious discussions within Bigfoot research circles as goes such details of anatomy as the presence or otherwise of the baculum. You can read about that discussion, if you wish, in this 2010 article by Loren Coleman.

Caption: the creature in the Patterson-Gimlin footage of 1967 – now affectionately known as 'Patty' within the Bigfoot research community – seemingly has breasts comparable to those of some humans. Is it coincidental that Roger Patterson was very familiar with William Roe's female Bigfoot of the 1950s (see below)? Or is this consistent with the ostensible biological reality of this animal? Because images of the Patterson-Gimlin film are copyright protected (like virtually all images of Bigfoot), I made this myself and it's available for use. Image: Darren Naish.

So that’s enough of that. Everything you’ve read so far is focused on males of this ostensible species. What about females? I think it would be fair to assume that very little – perhaps nothing – has been written about female anatomy. Bigfoot is, after all, mostly imagined as a quintessentially male creature linked to masculinity and the manly male humans that pursue it. But female Bigfoots have been reported too, and indeed among the most influential and historically significant of all Bigfoot accounts ever – William Roe’s Canadian story of the mid-1950s and Roger Patterson and Bob Gimlin’s famous/infamous encounter of 1967 – describe obvious females.

Caption: late in the 1950s, William Roe reported his encounter (supposedly from earlier in the decade) with another obviously female Bigfoot in Canada. This drawing was produced by Roe's daughter, under his direction. I can't be the only one who thinks the anatomy here is a bit... gravity-defying. This is not the original drawing, but a re-drawing produced by John Conway for our 2013 book The Cryptozoologicon: Volume One. Image: John Conway.

Breasts have been written about and commented on a great many times in discussions of this creature. A few reports have described elongate or ‘droopy’ breasts but those most often discussed – those of ‘Patty’ and Roe’s creature – were rotund and even engorged in appearance (Bindernagel 1998). Indeed, those working to demonstrate the reality of the Patterson-Gimlin film have gone to some trouble to argue that ‘Patty’s’ breasts can’t be fake because they exhibit movement and flexibility consistent with genuine breasts, and inconsistent with any of the available prosthetic alternatives. William Munns has a whole section on this issue in his book When Roger Met Patty (Munns 2014). Human models were filmed and photographed to show, via comparison, how the ‘Patty’ breasts perform in a consistent and realistic manner.

I could go on – as I’ve said, a lot has been said about Bigfoot breasts in the literature (there’s also been a whole discussion on how realistic the concept of furry breasts might be) – but I think that’ll do. Moving elsewhere as goes females, why do we never hear about female genitalia (I’m here excluding breasts from the definition of ‘genitalia’), in contrast to all those mentions of Bigfoot junk? Maybe it’s true that those who’ve considered Bigfoot as a biological entity have been biased by patriarchy. I don’t doubt that this is true but it should also be noted that observations and recollections concerning the finer details of female genital anatomy are few and far between, as they might be for logical reasons (I mean: all the goddam hair). Anyway, a few mentions are out there. A very few.

In the aforementioned discussion provided by Loren Coleman, Loren refers to the rarity of discussions of female genital anatomy and notes that he (at the time) was only aware of one. In the November of 1968, hunter John Thomas is said to have encountered two Bigfoots sleeping out in the open. They were in the posture that – curiously enough – has been described on several occasions by witnesses; that is, with the limbs partially folded beneath the body and the back facing upwards. Anyway, one of these animals had breasts in addition to a “swelling” in its genital region that it kept rubbing (Coleman 2003, p. 198). Without more information it’s difficult to know the story here. Presumably we’re talking about a modest swelling of the sort seen in various primates (including gibbons and gorillas among hominoids), and not the pronounced and striking structures seen in chimps, bonobos and various monkeys.

Caption: several accounts describe crypto-hominids sleeping in this very unusual posture, or at least in postures like it. This drawing was apparently produced by a Soviet zoologist called Khlakhlov during the early 1900s and depicts an Almas – an Asian crypto-hominid – encountered in the Dzungarian region. The drawing is reproduced in Myra Shackley's 1983 book. Image: Shackley (1983).

A second case concerns what’s said to be the imprint of a Bigfoot butt, left in sandy soil in Walla Walla, Washington. The print was cast by Paul Freeman – an individual with a controversial track record in the world of Bigfoot evidence, shall we say – and passed to Jeff Meldrum who described it in his 2006 book. The print reveals well demarcated, muscular buttocks but also what appears to be the suggestion of labia (Meldrum 2006). With a bit of imagination, they look – from the cast of the impression – to be about similar in form and proportion to those of a human.

Needless to say, there isn’t – to my knowledge – any discussion out there as goes any other aspects of female genital anatomy in this alleged creature. Not only is eyewitness data on such details unreported (so far as I know), but any sensible idea we might have on what’s predicted or assumed is dependent on whatever phylogenetic affinity we prefer for the species: Bigfoot has, variously, been suggested to be close to gibbons, orangutans, hominines and even hominins and humans by those who endorse its existence. There's an argument for platyrrhine status out there as well.

Caption: non-human primates of many sorts have diverse and remarkable genitals. Variously platyrrhines – spider monkeys are the most famous – have enormous clitorides. This is a Colombian or Black-headed spider monkey Ateles fusciceps rufiventris using a stick as a scratching tool. Image: Darren Naish.

That about brings us to a close on this niche yet worthy subject. As usual with arcane topics like this, it’s never true that “nobody ever talks about ---- [insert weird niche subject]”: on the contrary, quite a few writers have. Clearly, if Bigfoot is real, we have scant data to work with, and a conclusion must be that Bigfoot genitalia are not especially noticeable in average encounters with humans. And if Bigfoot isn't real, but is a sociocultural phenomenon of some sort, the fact that so few encounters discuss its genitalia in detail might be consistent with the near-irrelevance of such features to an entity of this sort. This might be inconsistent with erotic fan fiction – sorry, Mr Riggleman – but there it is.

For previous Tet Zoo articles on Bigfoot and other cryptozoological matters, see…

• Cryptozoology at the Zoological Society of London. Cryptozoology: time to come in from the cold? Or, Cryptozoology: avoid at all costs?, July 2011 (ruined by hosters: no longer includes pictures)

• The Cryptozoologicon (Volume I): here, at last, December 2013 (ruined by hosters: no longer includes pictures)

• Is Cryptozoology Good or Bad for Science? (review of Loxton & Prothero 2013), September 2014 (ruined by hosters: no longer includes pictures)

• My New Book Hunting Monsters: Cryptozoology and the Reality Behind the Myths, February 2016 (ruined by hosters: no longer includes pictures)

• If Bigfoot Were Real, July 2016

And here's your reminder that the persistence and success of Tetrapod Zoology now depends entirely on funding via patreon. Thanks to those who support me! The more secure my funding, the more time I can and will spend generating content for Tet Zoo.

Refs - -

Bindernagel, J. A. 1998. North America’s Great Ape: the Sasquatch. Beachcomber Books, Courtenay, B.C.

Coleman, L. 2003. Bigfoot! The True Story of Apes in America. Paraview, New York.

Conway, J., Kosemen, C. M. & Naish, D. 2013. Cryptozoologicon Volume I. Irregular Books.

Harcourt, A. & Gardiner, J. 1994. Sexual selection and genital anatomy of male primates. Proceedings of the Royal Society, London B 255, 47-53.

Krantz, G. S. 1999. Bigfoot Sasquatch Evidence. Hancock House, Surrey, B.C. & Blaine, WA.

Meldrum, D. J. 2006. Sasquatch: Legend Meets Science. Tom Doherty Associates, New York.

Meldrum, J. & Greenwell, R. 1998. Bigfoot: take two. BBC Wildlife 16 (9), 68-71.

Munns, W. 2014. When Roger Met Patty. William Munns.

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

Raynal, M. 2001. ‘Jordi Magraner’s field research on the bar-manu: evidence for the authenticity of Heuvelmans’ Homo pongoides.’ In Heinselman, C. (ed) Hominology Special Number 1. Craig Heinselman (Francestown, New Hampshire), unpaginated.

Shackley, M. 1983. Wildmen: Yeti, Sasquatch and the Neanderthal Enigma. Thames and Hudson, London.