The Most Amazing TetZoo-Themed Discoveries of 2018

As we hurtle toward the end of the year – always a scary thing because you realise how much you didn’t get done in the year that’s passed – it’s time to look back at just a little of what happened in 2018. This article is not anything like a TetZoo review of 2018 (I’ll aim to produce something along those lines in early 2019), but, rather, a quick look at some of the year’s neatest and most exciting zoological (well, tetrapodological) discoveries. As per usual, I intended to write a whole lot more – there are so many things worthy of coverage – and what we have here is very much an abridged version of what I planned.

Animals we will meet below, a montage. Images: (c) Philippe Verbelen, (c) Kristen Grace, Florida Museum of Natural History, Graham et al. (2018), CC BY-SA 4.0.

Thanks as always to those supporting me at patreon. Time is the great constraint (and finance, of course), and the more support I have, the more time I can spend on producing blog content. Anyway, to business…

The Rote leaf warbler. New passerine bird species are still discovered on a fairly regular basis; in fact three were named in 2018*. One of these is especially remarkable. It’s a leaf warbler, or phylloscopid, endemic to Rote in the Lesser Sundas, and like most members of the group is a canopy-dwelling, insectivorous, greenish bird that gleans for prey among foliage. Leaf warblers are generally samey in profile and bill shape, so the big deal about the new Rote species – the Rote leaf warbler Phylloscopus rotiensis – is that its bill is proportionally long and curved, giving it a unique look within the group. It superficially recalls a tailorbird. Indeed, I think it’s likely that the species would be considered ‘distinct enough’ for its own genus if there weren’t compelling molecular data that nests it deeply within Phylloscopus (Ng et al. 2018).

* The others are the Cordillera Azul antbird Myrmoderus eowilsoni and the Western square-tailed drongo Dicrurus occidentalis.

A Common chiffchaff Phylloscopus collybita encountered in western Europe, a familiar Eurasian-African phylloscopid leaf warbler. Image: Darren Naish.

The story of the Rote leaf warbler’s discovery is interesting in that it’s yet another recently discovered species whose existence and novelty was suspected for a while. Colin Trainor reported leaf warblers on Rote in 2004 but never got a good look at them, Philippe Verbelen observed them in 2009 and realised how anatomically unusual they were, and it wasn’t until 2015 that a holotype specimen was procured (Ng et al. 2018). I’ve mentioned before the fact that documenting and eventually publishing a new species is rarely an instant see it catch it publish it event, but a drawn-out one that can take decades, and here we are again. Also worth noting is that the existence of a leaf warbler on Rote was not predicted based on our prior knowledge of leaf warbler distribution in view of the deep marine channel separating Rote from Timor and lack of any prior terrestrial connection. Yeah, birds can fly, but members of many groups prefer not to cross deep water channels. In this case, this did, however, happen and most likely at some point late in the Pliocene (Ng et al. 2018).

Rote leaf warbler in life, a novel member of an otherwise conservative group. Image: (c) Philippe Verbelen.

Rote has yielded other new passerines in recent years – the Rote myzomela Myzomela irianawidodoae (a honeyeater) was named in 2017 – and it’s possible that one or two others might still await discovery there.

Neanderthal cave art. Hominins don’t get covered much at TetZoo, which is weird given the amazing pace of relevant recent discoveries and the fact that they’re totally part of the remit. I mostly don’t cover them because I feel they’re sufficiently written about elsewhere in the science blogging universe, plus I tend to be preoccupied with other things. Nevertheless, I take notice, and of the many very interesting things published in 2018 was Hoffman et al.’s (2018) announcement of several different pieces of Spanish rock art, seemingly made by Neanderthals Homo neanderthalensis. The art concerned involves hand stencils, abstract lines, squares and amorphous patches of pigment, always marked in red.

Red abstract markings, discovered in several Spanish caves, are old, and in fact were seemingly made by hominins long before H. sapiens moved into Europe. The red sinuous marking and system of squares and lines near the middle of this photo are purported to have been made by Neanderthals (other images, depicting animals and present adjacent to these markings, were seemingly created more recently by H. sapiens individuals). Image: (c) P. Saura.

The main reason for the attribution of this art to Neanderthals is its age: uranium-thorium dating shows that it’s older than 64ka, which therefore makes it more than 20ka older than the time at which H. sapiens arrived in Europe (Hoffman et al. 2018). That seems compelling, and it’s consistent with a building quantity of evidence for Neanderthal cultural complexity which involves the use of shells, pigments, broken stalagmites and so on.

One of the most famous pieces of claimed Neanderthal rock art: the Gorham's Cave ‘hashtag’ from Gibraltar. Image: (c) Stewart Finlayson.

I should add here, however, that I’m slightly sceptical of the use of age as a guide to species-level identification. Why? Well, we have evidence from elsewhere in the fossil record that the range of a hominin species can be extended by around 100ka without serious issue (witness the 2017 announcement of H. sapiens remains from north Africa; a discovery which substantially increased the longevity of our species). In view of this, would a 20ka extension of H. sapiens’ presence in Europe be absolutely out of the question? Such a possibility is not supported by evidence yet, and I don’t mean to appear at all biased against Neanderthals.

A tiny Cretaceous anguimorph in amber, and other Mesozoic amber animals. As you’ll know if you follow fossil-themed news, recent years have seen the discovery of an impressive number of vertebrate fossils in Cretaceous amber, virtually all of which are from Myanmar and date to around 99 million years old. They include tiny enantiornithine birds, various feathers (most recently racquet-like ‘rachis dominated feathers’), the tiny snake Xiaophis, early members of the gecko and chameleon lineages and the small frog Electrorana. Many of these finds were published in 2018 and any one could count as an ‘amazing’ discovery.

The Barlochersaurus winhtini holotype, from Daza et al. (2018).

However, there’s one fossil in particular that I find ‘amazing’, and it hasn’t received all that much coverage. It’s the tiny (SVL* 19.1 mm!), slim-bodied anguimorph Barlochersaurus winhtini, named for a single, near-complete specimen subjected to CT-scanning (Daza et al. 2018). Remarkable images of its anatomical details are included in Daza et al.’s (2018) paper. It has short limbs, pentadactyl hands and feet and a slim, shallow, bullet-shaped skull. Phylogenetic study finds it to be somewhere close to, or within, Platynota (the clade that includes gila monsters and kin, and monitors and kin), or perhaps a shinisaurian (Daza et al. 2018). It could be a specialised dwarf form, or somehow more reflective of the ancestral bauplan for these anguimorph groups. Either way, it’s exciting and interesting. What next from Burmese amber?

* snout to vent length

Barlochersaurus in life. It’s about the size of a paperclip. Image: (c) Kristen Grace, Florida Museum of Natural History (original here).

The Reticulated Siren. Sirens are very special, long-bodied aquatic salamanders with reduced limbs and bushy external gills. They’re very weird. They can reach 95 cm in length (and some fossil species were even larger), lack hindlimbs and a pelvis, have a horny beak and pavements of crushing teeth, and eat plants in addition to gastropods, bivalves and other animal prey. A longish article on siren biology and evolution can be found here at TetZoo ver 3.

A life reconstruction of the Cretaceous siren Habrosaurus, showing features typical of the group. This animal could reach 1.5 m in total length. Image: Darren Naish (prepared for my in-prep texbook The Vertebrate Fossil Record, on which go here).

Until recently, just four living siren species were recognised. But it turns out that indications of a fifth – endemic to southern Alabama and the Florida panhandle – have been around since 1970 at least. Furthermore, they pertain to a big species, similar in size to the Great siren Siren lacertina. Known locally as the ‘leopard eel’ (a less than ideal moniker, given that there’s a real eel that already goes by this name), this animal has been published by Sean Graham and colleagues in the open-access journal PLoS ONE (Graham et al. 2018) wherein it’s formally christened the Reticulated siren S. reticulata. It reaches 60 cm in total length, has dark spots across its dorsal surface and a proportionally smaller head and longer tail than other Siren species.

A museum specimen of the species has been known since 1970 when its finder noted that it did “not conform” to descriptions of known species, and live specimens were collected by David Steen and colleagues in 2009 and 2014. Again, note that discovery and recognition was a drawn-out process. The discovery has, quite rightly, received a substantial amount of media coverage, and many interesting articles about the find are already online. Many of you will already know of David Steen due to his social media presence and Alongside Wild charity (which I’m proud to say I support via pledges at patreon).

The Reticulated siren paratype specimen, as described by Graham et al. (2018). Image: Graham et al. (2018), CC BY-SA 4.0. Original here.

The idea that a new living amphibian species 60 cm long might be discovered anew in North America in 2018 is pretty radical. I’m reminded of the 2009 TetZoo ver 2 article ‘The USA is still yielding lots of new extant tetrapod species’ (which is less fun to look at than it should be, since images aren’t currently showing at ver 2). Furthermore, Graham et al. (2018) discovered during their molecular phylogenetic work that some other siren species are not monophyletic but likely species complexes, in which case taxonomic revision is required and more new species will probably be named down the line.

And that’s where I must end things, even though there are easily another ten discoveries I’d like to write about. This is very likely the last article I’ll have time to deal with before Christmas. As I write, I’m preparing to leave for the Popularising Palaeontology conference which happens in London this week (more info here), and then there are Christmas parties and a ton of consultancy jobs to get done before the New Year. On that note, I’ll sign off with a festive message, as is tradition. Best wishes for the season, and here’s to a fruitful and action-packed 2019. Special thanks once again to those helping me out at patreon.

For previous TetZoo articles relevant to various of the subjects covered here, see…

•  The USA is still yielding lots of new extant tetrapod species, July 2009

• THE AMAZING WORLD OF SALAMANDERS, October 2013

• Chiffchaffs: a view of passerines from the peripheries (part I), August 2014

• The Biology of Sirens, June 2016

Refs - -

Daza, J. D., Bauer, A. M., Stanley, E. L., Bolet, A., Dickson, B. & Losos, J. B. 2018. An enigmatic miniaturized and attenuate whole lizard from the mid-Cretaceous amber of Myanmar. Breviora 563, 1-18.

Graham, S. P., Kline, R., Steen, D. A. & Kelehear, C. 2018. Description of an extant salamander from the Gulf Coastal Plain of North America: the Reticulated Siren, Siren reticulata. PLoS ONE 13 (12): e0207460.

Hoffman, D. L., Standish, C. D., García-Diez, M., Pettitt, P. B., Milton, J. A., Zilhão, J., Alcolea-González, J. J., Cantelejo-Duarte, P., Collado, H., de Balbín, R., Lorblanchet, M., Ramos-Muñoz, J., Weniger, G.-Ch. & Pike, A. W. G. 2018. U-Th dating of carbonate crusts reveals Neandertal origin of Iberian cave art. Science 359, 912-915.

Ng, N. S. R., Prawiradilaga, D. M., Ng, E. Y. X., Suparno, Ashari, H., Trainor, C., Verbelen, P. & Rheindt, F. E. 2018. A striking new species of leaf warbler from the Lesser Sundas as uncovered through morphology and genomics. Scientific Reports 8: 15646.

Up Close and Personal With the Crystal Palace Dinosaurs

Like many of us, I have long enjoyed looking at the Crystal Palace dinosaurs and other prehistoric animal models, created in 1854, still on show more than 160 years later, and providing a remarkable showcase of ancient life as it was imagined at the time. But I’ve only ever seen them from afar. How fantastic would it be to examine them up close? Well…

You’ve seen the Crystal Palace dinosaurs before (or images of them, anyway), but you might not have seen them up-close like this. Neither had I prior to this very special visit. Image: Darren Naish.

Way back in September 2018, I was fortunate enough to attend the Crystal Palace Dinosaur Days event, part of the Heritage Open Days weekend occurring across the UK on the weekend concerned. I gave a talk and also led a tour around the prehistoric animal models (focusing on the reptiles and amphibians alone). Adrian Lister (of mammoth and Megaloceros fame) led a tour too, Mark Witton gave a talk on ‘Palaeoart After Crystal Palace’, and much else happened besides. I also have to mention the 3D-printed models of the dinosaurs made by Perri Wheeler. How I would love for these to be commercially available: I’m sure they’d be a success. So, it was a great event; well done Ellinor Michel and everyone else involved in the Friends of Crystal Palace Dinosaurs group (follow them on Twitter at @cpdinosaurs) for putting it together.

Perri Wheeler’s brilliant models of the three Crystal Palace dinosaurs (from back to front: Megalosaurus, Hylaeosaurus, Iguanodon). As a pathological collector of model dinosaurs, I sure would like to own a set - but I also sure would like for these models (or a set very similar to them) to be commercially available. Image: Darren Naish.

The real thrill, however, was not the talks nor the presence of the amazing and sometimes spectacularly good speakers but the fact that we were awarded special, up-close access to the prehistoric animal models. A dream come true. As you’ll know if you’ve visited Crystal Palace or read about it, the models are located on islands surrounded by a snaking waterway. In other words, they aren’t readily accessible. For the duration of Dinosaurs Days, however, a temporary bridge had been erected and – like Lord Roxton striding across a felled tree to Maple White Land – we made the crossing and stepped into a bygone era.

The amazing, enormous head of the Mosasaurus. Many of the scales on the body were recently repaired as the entire skin across the body was in a poor state. More on the mosasaur below. Image: Darren Naish.

Why erect a bridge to the islands in the first place? Both so that crucial landscaping and gardening can occur, and so that the models can be examined and evaluated for repair. They’re not in the best of shape, you see, and much work needs doing. Indeed, right now there’s a major push to get funding for a permanent bridge that will allow the continual access that’s required. This project only has a few days of fundraising left and there’s some way to go before the target is reached: go here and chip in if you can. You might have heard that the Mayor of London agreed to partly fund the project… as has legendary musician and song-writer Slash, since it turns out that he’s a big fan! I should add that Slash seems to be quite the fan of science in general, his twitter account revealing a definite tendency to use his powers for good.

The standing Iguanodon was given renovation and a new paint scheme within recent years. Unfortunately, further repair work is already required. Image: Darren Naish.

The reason I’m writing this article is not just to bring attention to this push for funding, but also to discuss and illustrate various of the remarkable details I got to see thanks to this up-close encounter. Before I start, be sure to read (if you haven’t already) the August 2016 TetZoo ver 3 article on the Crystal Palace models. Thanks to the Dinosaur Days event, I should add that I’ve been able to get hold of the guide that Richard Owen wrote to accompany the exhibition, or the 2013 reprinting (Owen 2013) of this 1864 publication (Owen 1854), anyway. It provides at least some background information on why the animals look the way they do.

Head of the reclining Iguanodon. Only a privileged few have seen the head from its left side. Image: Darren Naish.

Again, relatively few people will have seen the reclining Iguanodon from this side. It’s striking how natural, realistic and well-proportioned the model looks in this view - very much like a real animal. Image: Darren Naish.

Hylaeosaurus was thought by Owen and Hawkins to be an iguana-like reptile with a “lofty serrated or jagged crest, extended along the middle of the back”, though many aspects of the reconstruction were noted as being “at present conjectural” (Owen 2013, p. 18). Image: Darren Naish.

I’ll avoid repeating here the same points I made in my 2016 article but I will repeat my primary take-homes. Namely, that it’s disingenuous and naïve to criticise the models as outdated or as inaccurate, laughable follies, as is sometimes done. They have to be seen within the context of what was known at the time, there has to some acknowledgement of the fact that scientific knowledge has improved over time, and there should also be recognition of the fact that the models are more up-to-date than, and superior in technical accuracy and craftmanship to, the vast majority of modern efforts to portray prehistoric life. In the interests of correcting a mistake made in my 2016 article I should also point out that Crystal Palace is not in Sydenham as I stated, but in Penge. With that out of the way…

The pachydermal, vaguely bear-like Megalosaurus is actually a composite of information compiled from both Jurassic and Cretaceous theropods. The tall shoulder hump was included because Owen erroneously regarded the tall-spined Altispinax (previously Becklespinax) vertebrae as belonging to the shoulder region of Megalosaurus (Naish 2010). Image: Darren Naish.

A lot of detail was added to the megalosaur’s face - some of it is superficially crocodylian-like. Note the ominous cracks at the tip of the nose and along the side of the lower jaw. Image: Darren Naish.

It was a real thrill to see the remarkably detailed appearance of the three Crystal Palace dinosaurs: Iguanodon, Megalosaurus and Hylaeosaurus. Each has a very different skin texture, the Megalosaurus being the most unusual in that it doesn’t have the tile-like scales of the other two. Instead, it’s decorated with a crazy-paving-like covering. It’s not clear what Benjamin Waterhouse Hawkins (the designer and model-maker) was trying to achieve here since this is a rather non-reptilian look. Perhaps the aim was to give the animal a fissured skin texture vaguely like that of elephants.

View of the interior of the standing Iguanodon, with and without flash. The light at the far end of the image is coming in through the Iguanodon’s mouth. Image: Darren Naish.

Holes on the undersides of the Iguanodon and Hylaeosaurus mean that their insides can be inspected. As you can see from my photos, the models look like weird gabled roofs from the inside, numerous metal struts and poles helping to provide support. The hylaeosaur’s original head was removed since its weight was causing the model’s neck to break, and was replaced with a fibreglass copy. So, peer inside the hylaeosaur from beneath and you see the translucent interior of its face.

A view of the hylaeosaur’s interior! Image: Darren Naish.

Cracks, fissures and damaged sections are visible everywhere, moss invades and covers parts of the hylaeosaur’s flanks (not good if you want the models to persist) and sections of the megalosaur’s nose look like they could fall off at any moment. Similar damage is present on some of the other models, their skin and scales flaking or cracking or looking to be in imminent danger of breaking or falling off. Some substantial (expensive) repair work has been done by the Friends of Crystal Palace Dinosaurs, but much more is required.

Looking into the mouth of the Mosasaurus. “The large pointed teeth on the jaws are very conspicuous; but, in addition to these, the gigantic reptile had teeth on a bone of the roof of the mouth (the pterygoid), like some of the modern lizards” (Owen 2013, p. 11). Image: Darren Naish.

In my previous Crystal Palace article, I discussed the fact that the models reveal a great many complex anatomical details, some of them involving details only familiar to specialists. When you see the models up close, even more such details become apparent. I’m not sure I knew that the mosasaur is equipped with accurate palatal teeth, for example. Owen specifically referred to this feature in the guidebook (Owen 2013). The temnospondyls (‘labyrinthodonts’) have big palatal teeth as well, as they should.

Anterior view of one of the temnospondyls. Check out the accurate palatal teeth. To Owen and Hawkins, this animal was Labyrinthodon salamandroides, a sort of composite based on temnospondyl bones and teeth, and inferences made from croc-line archosaur footprints, thought by Owen and those who followed his work to be made by Labyrinthodon. Image: Darren Naish.

Like the dinosaurs (except the megalosaur), the surviving pterosaurs are fantastically scaly (today, we think that pterosaurs were covered in a filamentous coat, except on their wings, the distal parts of their hindlimbs and their snouts and faces). Unfortunately, the pterosaur with folded wings has recently been damaged, its smashed snout and lower jaw meaning that you can see right inside its head. This reveals a complex internal ‘anatomy’: another reminder that the models weren’t all built to the same plan or in the same style, but that very different approaches were used for each.

The two large Crystal Palace pterosaurs represent the species known to Owen and Hawkins as Pterodactylus cuvieri (though the possibility that more than one species is represented is raised by Owen’s remarks in the accompanying guide). Unfortunately, one of the models is now badly broken. The two smaller pterosaur models are not currently on display and have had a really unfortunate history: they’ve been vandalised, broken and stolen several times. Image: Darren Naish.

Both big pterosaurs stand atop a small rocky ’cliff’. Like all the geological structures in the park, this is an installation specially created as part of the display. It looks, at first sight, to be made of nondescript grey rock. While looking at it, I began wondering about its specific composition, since we know that the other chunks of rocks in the park aren’t just random lumps of local geology, but transplanted sections of the specific geological unit the respective animal’s fossils come from.

As per usual, the model up-close - this is the Pterodactylus cuvieri posed with open wings - is a remarkable bit of craftmanship. Pterodactylus cuvieri was named for bones that have more recently been included within the genera Ornithocheirus and Anhanguera, and have most recently been awarded the new name Cimoliopterus. Image: Darren Naish.

And here’s a close-up of that detail. I absolutely adore the work here; check out all those individual scales. It seems remarkable now to think that Owen and Hawkins really imagined pterosaurs to look like this, but here’s the evidence. Image: Darren Naish.

What, then, are these pterosaurs really standing on? Mark Witton and I examined some freshly broken fragments of the cliff – the rock is chalk! This really shouldn’t have been a surprise given that the fossils these reconstructions are based on come from the English Chalk (Owen 2013), but it was great to see it confirmed. There’s even a line of dark flint nodules, just as there is in real chalk cliffs. These details are surely known to specialist researchers but were news to me.

Broken sections of the ‘pterosaur cliff’ reveal that we’re looking at chalk… which isn’t a surprise, and is exactly what we would expect, but here’s confirmation. You should be able to see a few of the dark, shiny flint nodules too. Image: Darren Naish.

Look – below – at the photo of the teleosaurs. Notice how the arrangement of scales and scutes is highly detailed, and how the animals have been given a scute arrangement that very much resembles that of living crocodylians. As it happens, the arrangement they’ve been given is dead wrong for teleosaurs but it is absolutely accurate for living crocodylians (where dorsal scute arrangement is – mostly – diagnostic to species level). What I’m saying is that I think that Hawkins looked at living Saltwater crocodiles Crocodylus porosus when designing these amazing models, since their dorsal scute pattern specifically matches this species (and, surprisingly, not gharials).

The two Teleosaurus of Crystal Palace. While compared by Owen with gharials, it’s interesting that the dorsal scute pattern they were given is very clearly based on living crocodiles. As per usual, look at the remarkable amount of well-rendered detail. Image: Darren Naish.

As usual, there’s stacks more I want to say, but time is up. I had such a great time seeing the models up close and I can’t wait to do it again. At the risk of sounding like a broken record, the Crystal Palace prehistoric animals are among the most scientifically and historically important renditions of ancient creatures ever created, and they’re amazing pieces of art, construction and craftmanship to boot. A full, thorough discussion of their ‘anatomy’, backstory, construction and history has, even today, never been published – McCarthy & Gilbert (1994) is the closest thing to it – and much remains to be compiled and discovered.

These models must be preserved for the future. On that note, don’t forget to pledge your support for the bridge project. Crystal Palace and its models will be covered here again at some point in the future, and various relevant projects will be discussed here in 2019 – watch this space!

As should be obvious from these photos, the entire area has become somewhat overgrown recently, and much maintenance is needed. The Friends of Crystal Palace Dinosaurs group are doing what they can, but help is needed. Image: Darren Naish.

For other TetZoo articles on the Crystal Palace prehistoric animals and other relevant issues, see…

• The Crystal Palace monsters, armoured tyrannosaurs and lurking sauropods: a look back at ‘Dinosaurs – A Historical Perspective’ (part I), May 2008

• The Great Dinosaur Art Event of 2012, November 2012

• All Yesterdays: Unique and Speculative Views of Dinosaurs and Other Prehistoric Animals - the book and the launch event, December 2012

• Artistic Depictions of Dinosaurs Have Undergone Two Revolutions, September 2014

• The Dinosaurs of Crystal Palace: Among the Most Accurate Renditions of Prehistoric Life Ever Made, August 2016

Refs - -

McCarthy, S. & Gilbert, M. 1994. Crystal Palace Dinosaurs: The Story of the World’s First Prehistoric Sculptures. Crystal Palace Foundation, London.

Naish, D. 2010. Pneumaticity, the early years: Wealden Supergroup dinosaurs and the hypothesis of saurischian pneumaticity. In Moody, R. T. J., Buffetaut, E., Naish, D. & Martill, D. M. (eds) Dinosaurs and Other Extinct Saurians: A Historical Perspective. Geological Society, London, Special Publications 343, pp. 229-236.

Owen, R. 1854. Geology and Inhabitants of the Ancient World. Crystal Palace Library and Euston & Evans, London.

Owen, R. 2013. Geology and Inhabitants of the Ancient World. Euston Grove Press, London.

The Much Belated Final Part of the Tetrapod Zoology 12th Birthday Event

Way, way back in January 2018 – back when TetZoo was hosted at SciAm – I published two articles on Tet Zoo’s 12th birthday (they’re here and here). Some time later – I think in late May 2018 – I finally published the third and final part. But, alas, its publication coincided with a time during which SciAm was – thanks to their new owners, the Springer Nature Group – becoming increasingly draconian as goes image use. Because I hadn’t completed the appropriate paperwork regarding use of an image (specifically, an image of me giving a talk at a conference, taken by someone more than happy to let me use said image), they pulled the entire article and it’s not online at the site right now. My plan on launching TetZoo ver 4 was therefore to eventually upload the article anew, mostly for reasons of having it published somewhere. Here it is. My apologies for posting something that’s now so, so, so far behind schedule, so much so that it’s scarcely relevant. But here we go…

2017 in the TetZooniverse: mostly Dinosaurs in the Wild and Hunting Monsters. But other stuff too. Images: Darren Naish.

Running this blog for 12 years is a pretty big deal to me, and for that reason I find it necessary to spend at least some time reviewing the year that’s passed and evaluating Tet Zoo’s performance. And so here – better later than never – we find the third and last of the 12th birthday articles. As per usual, these articles aren’t much fun if you dislike autocratic pontification or anything that might be interpreted as overt self-congratulation. Last warning.

​In which I gratuitously use Matt Baron (on the right) in the promotion of (the first edition of) Naish & Barrett’s Dinosaurs: How They Lived and Evolved. Have I ever mentioned my idea on how male Caucasian humans all look exactly alike when they grow facial hair? Image: Darren Naish.

So, to business once more. The previous article finished off with me discussing October’s TetZooCon 2017. With TetZooCon out of the way, it was back to work. I made the ‘finishing’ touches to the Eotyrannus monograph [UPDATE: HAAAA/ARGH], continued apace on the bird section of The Big Book, and spoke about Hunting Monsters (my cryptozoology book) for the Cambridge University Biological Society. I recognised a figure in the audience. It turned out to be none other than Matt ‘Ornithoscelida’ Baron. In person, he’s basically ok (I kid, I kid). White rhinos and tigers were covered at Tet Zoo at about this time.

​I’ve been drawing a whole lot of Paleogene birds for The Big Book. And the other birds as well. Image: Darren Naish.

November 2017 marked ten years since the publication of the seminal, ground-breaking, game-changing new sauropod dinosaur Xenoposeidon proneneukos (Taylor & Naish 2007). As summarised in the resultant Tet Zoo article, Xenoposeidon has not been ignored since its 2007 publication. And 2017 was an important year for this taxon given that a new paper on its phylogenetic position was proposed (Taylor 2017). My newest book – Evolution in Minutes (Naish 2017b) – appeared in November. It’s been well received and I’m really happy with it.

​It’s Evolution in Minutes, a book of bite-sized summaries of just about everything we understand about evolution. Image: Darren Naish.

On to December… and, right at the start of the month, it was time to attend (and speak at) another meeting: the Joint Scientific Meeting of ARC (Amphibian and Reptile Conservation) and the British Herpetological Society (BHS). My thoughts on the meeting were published here at Tet Zoo. I like zoology-themed conferences of all sorts, but I think herpetological ones might be best. My talk combined various ideas and bits of research on the British herpetofauna: are various of the supposedly introduced amphibians and reptiles we have in the country overlooked natives? Mostly they’re not, but the stories are interesting nonetheless.

Today, I’m the proud owner of a PalaeoPlushies thylacine - a Christmas 2017 gift from my mother-in-law, Sheila. He’s called Kid Cynoceph and I love him. Buy your own here! [UPDATE: currently sold out!] Image: Darren Naish.

​A few days later, and it was back to London for the second Popularising Palaeontology workshop, organised by Chris Manias and featuring a mix of talks from scientists, science historians and people in the museum world. My talk was on review volumes devoted to vertebrate palaeontology and on whether they’ve done a ‘fair’ job of covering the different vertebrate groups (spoiler: they totally haven’t). The talk is now online here (it’s about 20 minutes long). I will be publishing a full-length article about the talk – or, rather, its subject – here within the next few weeks [UPDATE: oops]. The talk is a sort of tie-in to The Big Book.

Preparing to present a fish-themed talk at PopPalaeo. Photo kindly provided by Jed Taylor.... whose work you absolutely have to check out if you’re interested in artistic depictions of dinosaurs (and other animals). Image: Jed Taylor. Oh yeah - this is the photo that caused this whole article to be pulled from SciAm.

​An article on Plica lizards (I sure do love the iguanians) appeared at Tet Zoo, Marilyn Munro (yup – honest) delivered the first seven volumes of Handbook to the Birds of the World to Tet Zoo Towers (only another ten volumes to go…), and Gabriel Ugueto and I worked together on a poster that still (as of late May 2018) hasn’t seen the light of day. I spent New Year (and the weeks around it) in the Tatras Mountains of far southern Poland. I went to places where there were bears, nutcrackers and assorted other neat animals… but didn’t see them (well, the bears would be hibernating, so that wasn’t a surprise).

Holy crap the High Tetras mountains are incredible. Image: Darren Naish.

​And that basically takes us up to January, which is where we bring things to a close (seeing as the actual blog birthday is January 21st). Work on a new book kicked off that month, and – with Tim Haines – I did my bit training the next batch of chrononauts for Dinosaurs in the Wild. Have I mentioned Dinosaurs in the Wild? I think I have.

​A sign that will greet you as you leave North Greenwich underground station, London. I’ll write, at length, about Dinosaurs in the Wild sometime soon.... [UPDATE: I eventually published both Dinosaurs in the Wild: An Inside View at ver 3 and The Last Day of Dinosaurs in the Wild here at ver 4]. Image: Darren Naish.

The megafaunal bias, 2017. So… standard annual procedure here is to then compile a list of the year’s articles, and to then see how things fared as goes taxonomic representation. The ultimate aim: to achieve fair balance of the different tetrapod groups. The reality: shameful bias towards charismatic megafauna. Let’s see how things turn out. Hold your breath…

Oh, I’ve used the same categories as per previous years but have combined ‘lissamphibians’ with ‘non-lissamphibian anamniotes’ given that keeping them separate now seems futile (cf Pardo et al. 2017).

Miscellaneous musings

• Animal Species Named from Photos

• Palaeoart Memes and the Unspoken Status Quo in Palaeontological Popularization

• Books That Influenced Me The Most

• Fossils We Want to Find

• An Ode to Britains Toy Animals

• The Fourth TetZooCon

• The TetZooCon of 2017

• Tet Zoo Reviews Zoos: Colchester Zoo

• Amphibian and Reptile Biology and Conservation, the 2017 Joint Scientific Meeting

• Zoology Books of 2017

• Closing Down for Christmas

Amphibians

• The Terrible Leaf Walker Frog

• The 2017 Amphibian Conservation Research Symposium

• The Microsaurs of Yore

Mammals

• The Sigmodontines

• Incredible Elephant Seals, Part 1

• Incredible Elephant Seals, Part 2

• Mountain Beaver, Boomer, Sewellel

• Darin Croft's Horned Armadillos and Rafting Monkeys

• For Rodent Week, the Gift of Jerboas

• How Many White Rhino Species Are There? The Conversation Continues

• The Tiger Subspecies Revised, 2017

Turtles

• J.Lo and Other Araripemydid Pleurodires

• Letters from the World of Turtle Evolution

• What—If Anything—Is a Geoemydid?

Mesozoic marine reptiles

• The Unique and Efficient 4-Flipper Locomotion of Plesiosaurs

• The Fall and Rise of Protoichthyosaurus

Pterosaurs

• Some Azhdarchid Pterosaurs Were Robust-Necked Top-Tier Predators

Non-avialan dinosaurs

• Ornithoscelida Rises: A New Family Tree for Dinosaurs

• Mass Survival of Multitudinous Dinosaur Lineages across the K–Pg Boundary

• Dinosaurs and the Anti-Shrink-Wrapping Revolution

• The Sensitive Face of a Big Predatory Dinosaur

• The Supracranial Sinus of the Horned Dinosaur Skull

• The "Birds Are Not Dinosaurs" Movement

• 10 Long, Happy Years of Xenoposeidon

Birds

• Secretary Birds: Weird, but Actually Not That Weird

• Raptor Raptor Raptor, Part 1

• Raptors Once More (Raptors, Part 2)

Lepidosaurs

• Predatory Actions of Amphisbaenians

• The Borioteiioids

• Stellagama the Rainbow Dragon

• The Tropidurine Treerunners

Cryptozoology

• The Soay Island Sea Monster of 1959

• The Ozenkadnook Tiger Photo Revealed as a Hoax

• A Review of Neanderthal: The Strange Saga of the Minnesota Iceman, Part 1

• A Review of Neanderthal: the Strange Saga of the Minnesota Iceman, Part 2

A graph...

​Image: Darren Naish.

And, there we have it…. surprise surprise, mammals and non-bird dinosaurs are about in the lead, and the ‘obscure’ groups I always aim to cover more (like stem-mammals and croc-line archosaurs) received no coverage at all. Cryptozoological issues were covered a bit during 2017, in part because of things connected to Hunting Monsters. I’m surprised that SpecBio received no coverage across 2017, but there we have it. Also interesting is that miscellaneous musings did pretty well, but then I find it easier these days to write ‘general’ articles where the thoughts meander across various subjects.

​I will keep drawing drawing drawing all the vertebrates until there is nothing left. Image: Darren Naish.

One final point: while I always blame myself for producing the sort of megafaunal biases you see here, the fact remains that – given the constraints of time that are now such a concern – it really is easier and quicker to generate articles on megafauna where available images are plentiful and to hand. In contrast, articles on weird, obscure animals are hard to do because getting useable images is that much harder. Would I do better, and produce more content for Tet Zoo, including on those weird, obscure animals, if only I could? Yes, I would. And should I be leaving Sci Am given that I’m now acutely aware of what a poor fit I am? [UPDATE: ummm].

And this is where we end. For previous Tet Zoo birthday articles, see...

• Happy first birthday Tetrapod Zoology (part I)

• Happy first birthday Tetrapod Zoology (part II)

• Happy second birthday Tetrapod Zoology (part I)

• Tetrapods of 2007 (happy birthday Tet Zoo part II)

• Happy THIRD birthday Tet Zoo

• Tet Zoo = 4 years old today

• 2009, a year of Tet Zooery

• Four years of Tet Zoo: to infinity... and beyond!

• It is with some dismay that I announce Tet Zoo's first hemi-decade

• Tet Zoo 5th birthday extravaganza, part II

• Happy Birthday Tetrapod Zoology: SIX YEARS of blogging

• Happy 6th Birthday, Tetrapod Zoology (part II)

• Tetrapod Zoology enters its 8th year of operation

• Today marks NINE YEARS of Tetrapod Zoology

• Tetrapod Zoology 10th-Birthday Extravaganza, Part 1: 2015 in Review

• Tetrapod Zoology 10th Birthday Extravaganza, Part II: the Rest of 2015 Reviewed

• Tetrapod Zoology 10th-Birthday Extravaganza, Part 3: Tet Zoo's Tetrapod Treatment in 2015

• Today Is Tet Zoo's 11th Birthday

• The 12th Year of Tet Zoo

• The Tet Zoo 12th-Birthday Event, Part 2

Refs - -

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

Naish, D. 2017b. Evolution in Minutes. Quercus Books, London.

Pardo, J. D., Small, B. J. & Huttenlocker, A. K. 2017. Stem caecilian from the Triassic of Colorado sheds light on the origins of Lissamphibia. Proceeding of the National Academic of Sciences, USA 114, E5389-E5395.

Taylor, M. P. 2017. Xenoposeidon is the earliest known rebbachisaurid sauropod dinosaur. PeerJ PrePrints 5: e3415.

Taylor, M. P. & Naish, D. 2007. An unusual new neosauropod dinosaur from the Lower Cretaceous Hastings Beds Group of East Sussex, England. Palaeontology 50, 1547-1564.

Pouches of the Sungrebe

Among the most obscure and poorly known of the world’s living birds are the finfoots or heliornithids, a group of duck-sized, vaguely grebe-like swimmers of the American, African and south-east Asian tropics. There are just three extant species. Heliornithids are gruiforms (part of the crane + rail clade), recent studies indicating a close relationship with the Afro-Madagascan flufftails, a group conventionally included within the rail family but increasingly thought to represent a separate lineage termed Sarothruridae (García-R. et al. 2014).

Caption: the Sungrebe is a boldly marked heliornithid that occurs from southern Mexico in the north to Bolivia and northern Argentina in the south. It is c 30 cm long, remains relatively abundant, and is associated with swamps, marshes and well-vegetated streams and rivers. Image: L. Catchick, wikipedia, CC BY 3.0 (original here).

There’s a lot to say about heliornithids – their biogeography and fossil record is confusing and fascinating – but here I want to focus on one particularly interesting aspect of their biology. Namely, that the tropical American Sungrebe Heliornis fulica (sometimes called the American finfoot) has pouches, and uses these pouches in the protection and transportation of its young. The chicks are altricial (that is, unable to walk or look after themselves) and hatch after a ridiculously short incubation period of just 10-11 days. Incubation is carried out by both parents, but it’s the male alone who has the pouches and is able to transport the chicks (Bertram 1996).

Waitaminute… pouches? In a bird? How exactly do these structures work? What do they look like? And where are they located? Books and articles that mention the pouches generally say that there are two of them and that they’re under the wings, but you don’t get much more detail than that. The primary source of detailed information on what’s going on here is Miguel Álvarez del Toro’s 1971 article, a semi-legendary paper that most interested people are aware of due to the summarised description of its contents provided by Brian Bertram in his heliornithid chapter in Handbook of the Birds of the World, Volume 3. Bertram (1996) didn’t provide photos or diagrams, basically because those that exist are not good enough or of the right style for HBW. But Álvarez del Toro (1971) did.

Caption: yes, there are published photos of Sungrebe babies inside their father’s pouches. Or.. there’s one photo, anyway. Here it is, and it comes from Álvarez del Toro’s 1971 paper. Image: Álvarez del Toro (1971).

A far older publication – one of Prince Maximilian of Wied-Neuwied’s volumes of his Beiträge zur Naturgeschichte von Brasilien – also includes discussion and description of sungrebe pouches. I haven’t seen this reference but it was cited by Álvarez del Toro (1971).

Caption: pouch configuration, with feathering removed and chick in place, in the male Sungrebe, as illustrated by Álvarez del Toro (1971). This was drawn directly from a specimen. Image: Álvarez del Toro (1971).

If – for now – we ignore feathers, the pouches are essentially shallow, oval pockets formed by muscular pleats of skin that extend along the side of the chest. The bird appears to have some muscular control over the shape and turgidity of the pleats (Álvarez del Toro (1971). A single chick can fit in each pouch, but it would likely fall out if the structure consisted of skin folds and a recessed area alone. Here, it seems, is where the feathers come in: a number of long, curved feathers grow upwards and backwards from the lower part of the side of the chest and form a feathery outer wall, this now meaning that the recess becomes a secure pouch. And let me state again that all of this is exclusive to males: females do not have any of these anatomical peculiarities.

Caption: Álvarez del Toro’s (1971) illustration of the pouch, now with the feathering in place. The feathers form a lateral wall to the pouch and keep the chick in place. Image: Álvarez del Toro (1971).

When are the pouches used? I used to think that they were only there for emergency reasons (say, nest invasion by a predator), and possibly not even used by every sungrebe father. This appears to be incorrect. Álvarez del Toro (1971) disturbed a nesting male sungrebe and then saw it both swimming and flying with “two tiny heads sticking out from the plumage of the sides under the wings”, so this might support the idea of ‘emergency’ function. But his conclusion overall was that the chicks are (somehow) placed within the pouches immediately after hatching, kept there for days as they grow their plumage and become more capable, and are fed and kept clean by the male all the while: “While carrying the young, the male reaches beneath the wing to feed them and remove the droppings” (Álvarez del Toro’s 1971, p. 86).

Caption: Heliornis, as illustrated for the HUGE bird section of the in-prep The Vertebrate Fossil Record. Yes, there are fossil heliornithids (or… claimed heliornithids, anyway). Progress on this book can be viewed here at my patreon. Image: Darren Naish.

And that, as far as I can tell, about sums up everything we know. The exact and precise details as goes what’s going on here haven’t been worked out, and a full and detailed study is still required. Presumably that will happen eventually. Are these structures definitely a novelty of Heliornis, or are they more widespread, and are there similar, related or relevant structures elsewhere in related species? It really doesn’t seem that there are, but Bertram (1996) noted that it might be difficult to say for sure given that much of our anatomical knowledge on these animals (as in, obscure gruiforms in general) comes from dried skins.

Those of you familiar with the arcane ornithology literature will know that the transportation of chicks by adults has been mooted for a reasonable range of species, including nightjars, woodcocks and other waders, various galliforms, anseriforms and others (see Ad Cameron’s illustrations below - the nightjar illustrations are almost certainly fanciful - from Perrins (1992)). Most of these cases are likely or almost definitely erroneous… and they don’t involve pouches…. but not all are. Jacanas definitely transport their babies by clasping them with their wings, but they can’t do this while flying.

Caption: ornithologists spent decades arguing over whether nightjars (these are European nightjars Caprimulgus europaeus) and Eurasian woodcocks Scolopax rusticola carry their eggs and/or chicks in flight. It now seems that woodcocks do do this (as do other waders, like some shanks), but that nightjars seemingly don’t. Image: Ad Cameron, in Perrins (1992).

I’d like to finish this article by taking you on a weird, speculative tangent in the vein of All Yesterdays (Conway et al. 2012). Regular readers of this blog will know that I often write about extinct dinosaurs, and – on occasion – about speculations pertaining to the behaviour or anatomy of extinct dinosaurs. If we were asked (as we occasionally are) how likely it might be that extinct dinosaurs – say, for example, hadrosaurs or therizinosaurs – had pouches in which they could carry and transport their young, we would typically respond by noting the total lack of evidence that might support such a possibility, and the sheer improbability of such a thing given the anatomy of the living animals that ‘bracket’ the relevant fossil animals on the family tree.  There are no pouches in crocodylians or those living birds that diverged earliest in bird history, nor are there indications of them in stem-members of these lineages.

Caption: there are no pouches here, but at least we have a speculative scene where an adult maniraptoran (presumably a dromaeosaur) is carrying its young. Image: Alex Sone (original here).

But novelty does arise – phylogenetic brackets can be violated, as I’ve said, to the amusement of some, at least once here in the past – and there’s no way we’d predict the anatomy of Heliornis if we only knew of it as a fossil. Could, then, that most bizarre novelty – a pouch for carrying babies – have arisen elsewhere in dinosaurs? To suggest such would be a grotesque and baseless novelty of the worst kind. Yup. Did I mention All Yesterdays?

My thanks to the anonymous colleague who kindly provided me with the key piece of literature that allowed the production of this article. And thanks to those who contributed to the poll at the Tetrapod Zoology facebook group and thereby forced this article to the top of the list.

Caption: put ‘A really weird bird’ into a poll … and, wow, people really like really weird birds. This is a screengrab from a facebook poll.

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 - -

Álvarez del Toro, M. 1971. On the biology of the American finfoot in southern Mexico. Living Bird 10, 79-88.

Bertram, B. C. R. 1996. Family Heliornithidae (finfoots). In del Hoyo, J., Elliott, A. & Sargatal, J. (eds) Handbook of the Birds of the World, Volume 3: Hoatzin to Auks. Lynx Edicions, Barcelona, pp. 210-217.

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

Garcia-R, J. C., Gibb, G. C. & Trewick, S. A. 2014. Deep global evolutionary radiation in birds: diversification and trait evolution in the cosmopolitan bird family Rallidae. Molecular Phylogenetics and Evolution 81, 96-108.

Perrins, C. 1992. Bird Life: An Introduction to the World of Birds. Magna Books, Liecester [sic].

Erroll Fuller’s The Passenger Pigeon

Most readers of this blog will be familiar with Erroll Fuller and the huge quantity he has published on recently extinct birds. His books typically combine a discussion on the biology, history and extinction of the species, a section on existing specimens (as in, museum mounts and skins and eggs and so on), and some good coverage of the animal’s appearances in art and popular culture.

Cover, featuring John James Audubon’s illustration from The Birds of North America. Credit: fair use.

Fuller’s 2015 The Passenger Pigeon does all this and more, and is an extremely well-illustrated, popular review of what we know of this species. The Passenger pigeon Ectopistes migratorius is/was a long-tailed, swift-flying pigeon, famous for existing in nomadic flocks of billions by the time Europeans began colonising North America. How the birds came to be so phenomenally abundant, whether this was an ephemeral or long-lived phenomenon, and how linked this was to human-caused habitat change are good questions and there are reasons for thinking that a population of billions was not a pre-Holocene condition (e.g., Mann 2011).

Juvenile, male and female Passenger pigeon, as illustrated by Louis Agassiz Fuertes in 1910. Credit: public domain, original here.

Fuller’s The Passenger Pigeon is not a gargantuan magnum opus like his The Great Auk (Fuller 1999). It’s far smaller (25 x 19 cm) and slimmer (177 pages). It includes an extraordinary number of photos, paintings and drawings, and indeed is a very attractive volume. Numerous black and white photos, mostly taken of captive birds in the early 1900s, depict the animals in life. There are photos of Martha (fabled ‘last of the passenger pigeons’) too, and reproductions of many paintings and other artworks.

The Passenger Pigeon is not an academic tome with technical citations and any extended discussion of thoughts on this bird’s evolution or biology. But it’s a good book nonetheless and I definitely recommend it for anyone interested in pigeons and/or in recently extinct birds.

The Passenger Pigeon is not as huge and impressive as Fuller’s The Great Auk book, but that’s ok. Credit: fair use.

Errol Fuller. 2015. The Passenger Pigeon. Princeton University Press (Princeton and Oxford), pp. 177. ISBN 978-0-691-16295-9. $29.95. Hardback, index, refs. Here at amazon, here at amazon.co.uk, here from the publishers.

 For previous TetZoo articles on pigeons and kin, see…

• The detachable tails of pigeons, September 2008

• Are parrots actually pigeons?, May 2009

• Giant club-winged pigeons and ninja ibises: clubs, spurs, spikes and claws on the hands of birds (part III), February 2011

• Sunbathing birds, August 2011

• The war on parasites: the pigeon’s eye view, the oviraptorosaur’s eye view, August 2012

• Footless urbanite pigeons, September 2014

• Voyeurism and Feral Pigeons, June 2015

• The Asian Dove That Ate Europe Alive, January 2017

• Pigeons and Doves 101, March 2018

Refs - -

Fuller, E. 1999. The Great Auk. Erroll Fuller, Southborough, Kent.

Mann, C. C. 2011. 1491: New Revelations of the Americas before Columbus. Vintage, New York.

The Second Edition of Naish and Barrett’s Dinosaurs: How They Lived and Evolved

Regular readers of this blog should know that 2016 saw the publication of the Natural History Museum book Dinosaurs: How They Lived and Evolved, co-authored by this blog’s humble overlord… that might be an oxymoron… and the Natural History Museum’s Paul Barrett (Naish & Barrett 2016). Dinosaurs has been well received and pretty successful in terms of sales, and so it came to pass that there was the need for a modified, softback version that included updates and corrections. Officially, the new version is a ‘fully revised and updated’ version, but it’s very literally a second edition, and that’s what I’m calling it.

Naish and Barrett, second edition - with a new cover!

First off, the production of a second edition – we’ll call it Dinosaurs 2nd ed (Naish & Barrett 2018) from hereon – allowed the correction of assorted typos and poor word choices. It never ceases to amaze me how much stuff we miss even when a given piece of text is checked, double-checked and checked again. Big thanks to fan of the book Klinsman Hinjaya for noting a number of required corrections. More importantly, said second edition also allowed us to update or modify various aspects of the science covered in the book. New data and new interpretations mean that our ideas on the biology and evolutionary history of extinct animals are constantly changing, and this was a great opportunity to get some of the relevant changes incorporated.

Some of you – especially those in possession of the first edition – are keen to know what’s different about Dinosaurs 2nd ed, so – without giving too much away – let’s take a look…

Caption: some of Emma’s drawings feature in the book. No, I’m kidding - they don’t. Or do they. Credit: Darren Naish.

A new cover. Personally, I think that Dinosaurs: How They Lived and Evolved is a pretty good book, and I hope you agree. But I never much liked the cover, and I know I’m not alone. Given that it portrays a roaring monster that’s showing us the inside of its mouth and what biiiig teeth it has, it might be construed as being contrary to the message otherwise promoted throughout the book: that non-bird dinosaurs were animals, and not the monsters of Hollywood and popular fiction.

Caption: new cover art by Bob Nicholls of Paleocreations. I own a full-sized print of this amazing piece. Damn… I own a lot of Bob Nicholls art now. Credit: Bob Nicholls.

The plan for Dinosaurs 2nd ed was thus to produce a brand-new cover that better represented modern thinking on dinosaurs. Paul and I opted to have either a feathered theropod, or an unusual ornithischian, and we ended up going with the latter for reasons I can’t recall (I think because it might look weirder and less familiar: feathered theropods are so passé, after all). Our chosen artist – Bob Nicholls – came up with a bunch of test sketches, all depicting the Chinese heterodontosaur Tianyulong in various poses and behavioural settings. We chose one, and Bob went to extraordinary trouble to get the contrast, lighting and composition right. There’s much more to it than that but… ladies and gentlemen, I give you… our new cover.

New artwork. Moving now to the insides of the book, we’ve also replaced a few other images that were used in the first edition. Bob has a few more of his excellent images in the book, we swapped out the (now defunct) diagrams on diplodocid jaw movement with a new reconstruction of a ground-feeding diplodocid (though illustrated without the keratinous beak recently proposed for this group), and Matt Martyniuk’s Anchiornis replaces a previous image with a problematic forelimb configuration. I also replaced a bunch of small images used in various of the cladograms.

Caption: some of the cladograms of Dinosaurs 2nd ed have been tweaked a little. This one depicts Theropoda, the predatory dinosaurs. Credit: Darren Naish.

Necessary coverage of the Ornithoscelida thing. Currently, one of the most talked-about issues within the study of Mesozoic dinosaurs is Matt Baron et al.’s (2017) proposal that the main branches of the dinosaur family tree be rearranged, such that sauropodomorphs are outside a theropod + ornithischian clade termed Ornithoscelida. I wrote about this proposal when it was brand-new, back in March 2017. It has been hotly contested by several teams of authors and there are already a few publications saying how reanalysis does not support Baron et al.’s model, or at least does not support it preferentially above the others that are available.

Obviously, we just had to cover this issue for Dinosaurs 2nd ed – especially so given that Paul is one of the study’s authors – and a few new paragraphs of text and new diagrams summarise the area for our readers. Our coverage of the Ornithoscelida issue resulted in various knock-on changes elsewhere in the book. New phylogenetic positions, for example, have been favoured for herrerasaurids, Eoraptor and so on.

Caption: the Baron et al. model, as depicted in the 2017 article I wrote about it for TetZoo ver 2. Credit: Darren Naish.

New taxa, new names, new phylogenetic possibilities. The world of Mesozoic dinosaur phylogeny and systematics move fast – remember that an average year right now sees the naming of between 30 and 50 new non-bird dinosaur species. We’re at a stage where phylogenetic models are never really ‘overturned’, but they certainly undergo regular tweaking, modification and augmentation. In view of this, Dinosaurs 2nd ed includes references to Stenonychosaurus, Latenivenatrix (sorry, Troodon) and halszkaraptorines, and I subtly changed the wording on the megaraptorans…

Pisanosaurus is no longer bigged-up as a possible early ornithischian given data indicating that it’s a non-dinosaur.

Revising thoughts on the origins of flight. Those familiar with discussions on both bird and flight origins within dinosaurs will know all about the ‘trees down vs ground up’ argument, and also with the contention that it might be utterly wrong to polarise things in this way. Nevertheless, there remain – for all those attempts to point to shades of grey – extremes in the debate. My take on the earliest phases of maniraptoran flight has mostly involved a weird sort of hybrid whereby the animals concerned are deemed predominantly terrestrial but also capable of climbing, their leaping, fluttering and gliding in arboreal settings being antecedent to flight. Like many people, I was originally enthused enough by WAIR (wing-assisted incline running) to think that it might be a plausible explanation of how maniraptorans first came to exploit arboreal settings and, from there, evolve flight.

Caption: Dececchi et al. (2016) showed that at least some non-bird maniraptorans do not have the right combination of anatomical features to benefit from WAIR as originally envisioned. This work affected our thinking on flight origins in Dinosaurs 2nd ed. Credit: Dececchi et al. 2016, PeerJ

The diversity of non-bird maniraptorans is such that it looks likely that these animals practised all sorts of behaviours during the long time that they were around, and thus that various different acts and adventures could have contributed to their ability to leave the ground. Having said that, recent studies indicate that at least some of the relevant animals could likely leap into flight from a ground-based start (Dececchi et al. 2016), and – at the same time – that arboreal behaviour was unlikely in such species. The possibility that flight could well have evolved without any arboreal component is interesting (and even shocking to some), and sufficiently so that we’ve alluded to it (briefly) in Dinosaurs 2nd ed.

Caption: Dinosaurs, the Russian edition. Now I know what my name looks like in Russian. Yes, the title is not the same as the English one.

And that’s it. I should also add that Dinosaurs: How They Lived and Evolved has also recently appeared in Australia (where it’s published by CSIRO), and that a Russian translation is now out as well. My thanks to everyone who’s bought this book, to those who commented on or said nice positive things about the first edition, and to everyone involved in its production and publication.

For those who haven’t purchased a copy, it’s available here from amazon, here from amazon.co.uk, and here from the publishers.

For previous articles relevant to this one, see…

• Flight of the Microraptor, November 2013

• The Integrated Maniraptoran, Part 1, May 2016

• The Integrated Maniraptoran, Part 2: Meet the Maniraptorans, May 2016

• The Integrated Maniraptoran, Part 3: Feathers Did Not Evolve in an Aerodynamic Context, May 2016

• Naish and Barrett's Dinosaurs: How They Lived and Evolved, November 2016

• Ornithoscelida Rises: A New Family Tree for Dinosaurs, March 2017

Refs - -

Baron, M. G., Norman, D. B. & Barrett, P. M. 2017. A new hypothesis of dinosaur relationships and early dinosaur evolution. Nature 543, 501-506.

Dececchi, T. A., Larsson, H. C. E. & Habib, M. B. 2016. The wings before the bird: an evaluation of flapping-based locomotory hypotheses in bird antecedents. PeerJ 4: e2159.

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

Naish, D. & Barrett, P. M. 2018. Dinosaurs: How They Lived and Evolved. The Natural History Museum, London.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

• How to prevent cannibalism in pheasants, March 2010

• The snood of the turkey, the wires and rackets of the motmot, the face of the rook, February 2011

• Chickens, 2012, April 2012

• Turkeys vs peafowl, the great debate, January 2013

• The other turkey, January 2013

•  The other peacock, January 2013

Refs - -

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

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

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

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

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

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

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

There will be books.

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.

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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.

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

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

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

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

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

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

Spot the cryptic typo. Image: Dallas Krentzel.

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

Art by the brilliant C. M. 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, and if you didn’t get to see it before that fateful Sunday… where were you? I was determined to embark on one final tour, and of course I also needed to go to the grand send-off party and say those sad final goodbyes…

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

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

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

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

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

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

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

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

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

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

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

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

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

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.