A Multi-Species Nesting Assemblage in the Late Cretaceous of Europe

You don’t have to be an expert on the fossil birds of the Mesozoic Era to know that our knowledge of these animals has increased exponentially in recent years. An extraordinary number of new species have been described, we’ve learnt a great deal about their anatomy thanks to spectacular new fossils – some of which are even preserved in amber – and we’ve gained insights into their behaviour and ecology thanks to discoveries made about their teeth, feathers, stomach contents, phylogeny and the environments in which they lived.

Enantiornithines — here are just a few of them. In-prep montage from my   in-prep textbook  . Image: Darren Naish.

Enantiornithines — here are just a few of them. In-prep montage from my in-prep textbook. Image: Darren Naish.

Among the most important and species-rich of Mesozoic bird groups are the enantiornithines, or ‘opposite birds’, so named because a few aspects of their skeletal anatomy (the way their shoulder and chest bones fit together in particular) are unusual relative to those of modern birds. Enantiornithines are known from rocks worldwide and were present from the start of the Cretaceous until its close 66 million years ago. They might even have evolved in the Jurassic, in which case older members of the group await discovery.

Among the many interesting things discovered recently about enantiornithines is that at least some of them were colonial nesters. In a Naturwissenschaften paper published in 2012, Gareth Dyke, Mátyás Vremir, Gary Kaiser and myself reported a remarkable fossil assemblage: a big, lens-shaped mass of calcareous mudstone (about 80 cm long, 50 cm wide and 20 cm deep) packed with literally thousands of enantiornithine eggshell fragments. A few enantiornithine bones were present as well, but eggshell fragments form 70-80% of the entire mass (Dyke et al. 2012).

Different chunks of the (originally lens-shaped) Oarda de Jos eggshell assemblage. You can see abundant eggshell fragments (a, b) as well as crushed but complete eggs (the image at bottom). Scale bars = 1 cm. Image: Vremir, Dyke  et al .

Different chunks of the (originally lens-shaped) Oarda de Jos eggshell assemblage. You can see abundant eggshell fragments (a, b) as well as crushed but complete eggs (the image at bottom). Scale bars = 1 cm. Image: Vremir, Dyke et al.

This eggshell mass was found at Oarda de Jos near Sebeş, Transylvania, western Romania and comes from the latest Cretaceous Sebeş Formation (Dyke et al. 2012). Pleurodire turtles, azhdarchid pterosaurs and such dinosaurs as ornithopods and the peculiar Balaur bondoc (which is probably a flightless member of the bird lineage, not a dromaeosaur as originally proposed: see Cau et al. 2015) also come from the Sebeş Formation. I wrote about the discovery and initial interpretation of the eggshell assemblage at TetZoo ver 3, though (sigh) it now appears without any of its images.

The Romanian maniraptoran theropod  Balaur bondoc , originally published as a dromaeosaurid, has a number of features which indicate that an avialan position might be more likely, and this is the position it has occupied in several phylogenetic analyses. If it really is an avialan, it has to be interpreted as a big, secondarily flightless member of the group. We explored this idea in   Cau  et al . (2015)  . Image: Emily Willoughby.

The Romanian maniraptoran theropod Balaur bondoc, originally published as a dromaeosaurid, has a number of features which indicate that an avialan position might be more likely, and this is the position it has occupied in several phylogenetic analyses. If it really is an avialan, it has to be interpreted as a big, secondarily flightless member of the group. We explored this idea in Cau et al. (2015). Image: Emily Willoughby.

As described in our 2012 paper, the assemblage is a fairly big deal, since it means that colonial nesting was practised by at least some enantiornithine species, is not unique to crown-birds, and evolved in birds more stem-ward than other colonially nesting birds (Naish 2014).

But as revealed in our new paper, published in Scientific Communications this past week and led by Mariela Soledad Fernández, things turn out to be a bit more complex than we originally thought. Rather than consisting of enantiornithine eggshell fragments and bones alone, the assemblage actually contains eggshell fragments (and probably bones) of several different, additional animal groups. Namely, gekkotan lizards, crocodylomorphs, and a bird different from the enantiornithine otherwise so well represented in the assemblage (Fernández et al. 2019).

Examined under microscopes (A-B show thin-sections viewed with a standard light microscope; C-D views from the SEM), eggshell fragments look like this. These images show the crocodylomorph eggshell in the assemblage. Image:   Fernández  et al . (2019)  .

Examined under microscopes (A-B show thin-sections viewed with a standard light microscope; C-D views from the SEM), eggshell fragments look like this. These images show the crocodylomorph eggshell in the assemblage. Image: Fernández et al. (2019).

It should be noted to start with that enantiornithine remains dominate the assemblage by far, around 70% of the sampled eggshell fragments belonging to that group (and presumably to the same one species). Sadly, the gekkotan and crocodylomorph eggshell fragments aren’t informative enough to tell us anything particularly interesting about the species concerned, other than that their remains are present. The Oarda de Jos crocodylomorph eggshell is different in thickness and microscopic surface texture from crocodylomorph eggshell fragments reported from the Upper Cretaceous of the USA and Brazil and is most similar to fossil eggs from the Eocene of Colorado, called Krokolites wilsoni (Hirsch 1985). But we don’t have a good handle on what sort of crocodylomorph we’re talking about. After the enantiornithine, its eggs are the most abundant in the sample, forming about 28% of the assemblage. Those artistic reconstructions previously created for the location are thus in error: they really should have at least a few crocodylomorphs in view.

Remember this scene? Produced in 2012 by   Julio Lacerda  , it depicts the possible appearance of the enantiornithine nesting colony we infer for the locality. Maybe some of the colony did look like this. But it now seems that a few crocodylomorphs and the odd lizard were in the immediate area as well. Image:   Julio Lacerda  . UPDATE: this is a horribly lo-res version of the image, I aim to publish a better one in time.

Remember this scene? Produced in 2012 by Julio Lacerda, it depicts the possible appearance of the enantiornithine nesting colony we infer for the locality. Maybe some of the colony did look like this. But it now seems that a few crocodylomorphs and the odd lizard were in the immediate area as well. Image: Julio Lacerda. UPDATE: this is a horribly lo-res version of the image, I aim to publish a better one in time.

The gekkotan eggshell pieces – they have an eggshell morphology termed ‘geckoid’ – have features in common with the eggs of modern geckos, and hence were presumably produced by crown-geckos (Fernández et al. 2019). Beyond that, we can say no more. Less than 1% of the eggshell in the assemblage comes from this lizard, so we’re not saying that its eggs are abundant in the sample.

‘Geckoid eggshell’ in the Oarda de Jos assemblage, as seen via SEM. The images (note the different scales) show (A) a distinct two-layered structure and (B) numerous tiny holes in the prisms of the second later. Image:   Fernández  et al . (2019)  .

‘Geckoid eggshell’ in the Oarda de Jos assemblage, as seen via SEM. The images (note the different scales) show (A) a distinct two-layered structure and (B) numerous tiny holes in the prisms of the second later. Image: Fernández et al. (2019).

Similarly, the second bird in the assemblage appears (from microscopic details of eggshell anatomy) similar to crown-birds – more so than to enantiornithines – but cannot be matched with any specific bird group and might represent something new (Fernández et al. 2019). So, we don’t know exactly what sort of bird we’re dealing with. Again, less than 1% of the eggshell in the assemblage is of this type, so it’s rare in our sample.

Piecing all of this together, what does it all mean? If our enantiornithine eggshell assemblage doesn’t involve enantiornithines alone, but also includes the remains of a second bird, a lizard and a crocodylomorph… do we have an example here of a multi-species nesting colony, perhaps one involving so-called parasitic nesting or even communal or cooperative nesting?

Several examples of this sort of thing are known for the modern world. They involve turtles laying their eggs at the edges of crocodylian nests, South American geckos which lay their eggs within the nests of cormorants and gulls, and seemingly harmonious nesting associations involving rheas, tinamous and penguins. Maybe the behaviours listed here occurred in Maastrichtian Romania. Perhaps the gekkotan lizard concerned was cheekily laying its (hard-shelled, perhaps sticky-shelled) eggs at the edges of enantiornithine nests, and maybe the mystery bird and crocodylomorph were non-threatening enough to be tolerated, their nests perhaps being close to those of the abundant enantiornithines (Fernández et al. 2019).

Opportunistic, parasitic, co-operative and harmonious nesting associations exist in the modern world; here’s an example where raptor nests (in this case, that of an Osprey  Pandion haliaetus  pair) invite the association of passerines, herons and others. Cases like this could well have existed in the Cretaceous. Image: D’Ami  et al . (1969).

Opportunistic, parasitic, co-operative and harmonious nesting associations exist in the modern world; here’s an example where raptor nests (in this case, that of an Osprey Pandion haliaetus pair) invite the association of passerines, herons and others. Cases like this could well have existed in the Cretaceous. Image: D’Ami et al. (1969).

It may also be that the association we report is not quite as interesting as just described, but more to do with geological and hydrodynamic processes. That is, that the eggshell fragments and bones concerned became associated due to their presence in the same general floodplain area, their remains becoming mixed together by the actions of floodwater and not being all that informative as goes behaviour and ecology. Even if this is true, however, we can at least say that these animals were nesting in the same area and environment, and seemingly in proximity. That alone is interesting, and there are indications from elsewhere in the Cretaceous fossil record that it might have been a fairly regular occurrence.

Our paper is open access (OA) and available here; it’s one of several technical papers I hope to see published in the year. On that note, here’s a reminder that I’m not a salaried academic researcher and that any contribution I make to the technical literature is done in my own time. Thanks to those who support me via pledges at patreon, and please consider doing so if you don’t already.

For previous TetZoo articles relevant to Mesozoic birds, the Late Cretaceous animals of Romania and other relevant issues, see…

While compiling this list I’ve discovered that essentially ALL of the TetZoo articles on enantiornithines are hosted at sites (ScienceBlogs and SciAm) that have stripped them of their original images. I must therefore make some effort to republish them here, with their pictures. Great, more stuff to do.

Refs - -

Cau, A., Brougham, T. & Naish, D. 2015. The phylogenetic affinities of the bizarre Late Cretaceous Romanian theropod Balaur bondoc (Dinosauria, Maniraptora): dromaeosaurid or flightless bird? PeerJ 3: e1032.

D’Ami, R. D., Invernici, F. & Quochi, G. 1969. Animals of Lake and Marsh. Casa Editrice AMZ, Milan.

Dyke, G., Vremir, M., Kaiser, G. & Naish, D. 2012. A drowned Mesozoic bird breeding colony from the Late Cretaceous of Transylvania. Naturwissenschaften 99, 435-442.

Fernández, M. S., Wang, X., Vremir, M., Lauren, C., Naish, D., Kaiser, G. & Dyke, G. 2019. A mixed vertebrate eggshell assemblage from the Transylvanian Late Cretaceous. Scientific Reports 9, 1944.

Hirsch, K. F. 1985. Fossil crocodilian eggs from the Eocene of Colorado. Journal of Paleontology 59, 531-542.

Naish, D. 2014. The fossil record of bird behaviour. Journal of Zoology 292, 268-280.

TetZoo Bookshelf, February 2019, Part 1

There are a lot of books that require review at TetZoo. Let’s see how many I can get through right now…

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Michael J. Everhart’s Oceans of Kansas: A Natural History of the Western Interior Sea, Second Edition

Those of us interested in Mesozoic animals frequently lament the lack of any good, especially useful books that review our knowledge of Mesozoic marine life. But it’s not all bad, since we at least have this wonderful, weighty, very detailed volume. Oceans of Kansas is by far the best there is, and it’s a must-have. Ok, the limitation is that it’s devoted, of course, to Late Cretaceous North American seas alone – specifically, to the Western Interior Sea that once extended north-south across the whole of North America – but this is no fault of the book but its main feature. Indeed, Oceans of Kansas is successful enough that this is its second edition, and it’s newly expanded, revised and augmented, so much so that those owning the first, 2005 edition should consider buying this one too.

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Everhart devotes chapters to sharks, bony fishes, turtles, the various plesiosaur groups, mosasaurs, pterosaurs and dinosaurs as well as invertebrates, the history of our discovery of the Western Interior Sea and more. At 427 pages and about 3.5 cm in thickness, this book contains a huge amount of information. It feels like the compilation of a dedicated person’s life’s work, and I hope the author doesn’t mind me saying that.

Excellent colour photos and numerous diagrams and other illustrations appear throughout. Many of these images are rare or novel and there’s much here not available elsewhere. A colour plate section features art by the late, great Dan Varner, a friend and correspondent I admired and liked a great deal. Dan’s paintings are looking a bit dated now in view of recently acquired information on mosasaur anatomy and we’d all love to know what Dan would have done with them had he the chance (he died in 2012). Regardless, they’re fantastic pieces of work.

One of Dan Varner’s greatest illustrations (in my opinion): the shark  Cretoxyrhina  takes out a  Tylosaurus . This and many other Varner pieces feature in  Oceans of Kansas . Image: (c)   Dan Varner/Oceans of Kansas  .

One of Dan Varner’s greatest illustrations (in my opinion): the shark Cretoxyrhina takes out a Tylosaurus. This and many other Varner pieces feature in Oceans of Kansas. Image: (c) Dan Varner/Oceans of Kansas.

Everhart’s text combines historical review with discussions of what we know, or think we know. There are also numerous personal anecdotes about how various specimens were discovered, recovered and interpreted as well as discussions on how relevant ideas on Cretaceous marine life have developed or changed during the time in which the author has been involved. As he explains, some ideas (those on mosasaur skin and physiology, for example) have undergone substantial revision since he published the book’s first edition in 2005. It’s this personal touch that I enjoy the most.

I say again that Oceans of Kansas is absolutely a must-have if you’re interested in Mesozoic marine life. In my work as a vertebrate palaeontologist and researcher I’ve consulted it hundreds of times, and – like all very good books – it’s not just a great source of information but also a joy to look at.

Michael J. Everhart. 2017. Oceans of Kansas: A Natural History of the Western Interior Sea, Second Edition. Indiana University Press (Bloomington, Indiana), pp. 427. ISBN 978-0-253-02632-3. Hardback, index, refs. Here at amazon, here at amazon.co.uk, here from the publishers.

Katrina van Grouw’s Unnatural Selection

Much of the fandom that developed around Katrina van Grouw’s 2013 The Unfeathered Bird did so because of its spectacular, beautiful illustrations (van Grouw 2013). Van Grouw’s newest book – 2018’s Unnatural Selection – has a similar format and design and is also spectacularly well illustrated; it is, however, an important work of literature, not of illustration or art alone. I’ve already said quite a fair bit about this book – I wrote a long article on its contents, published before it was out – back in May 2018 at ver 2, so will be brief here.

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Divided into four main sections (Origin, Inheritance, Variation and Selection), Unnatural Selection describes and celebrates selective breeding and its results. We’ve bred animals with all manner of anatomical novelties not present in their wild ancestors, and – as Van Grouw explains right at the start of the book – have ended up with domestic hybrids and variants that are so modified that ancestry has been obscured, or are hybrid mish-mashes of more than one species.

Unnatural Selection  showcases an extraordinary number of unusual domestic animals (these are all domestic pigeon breeds), but the main theme is the evolutionary plasticity, and potential, that human selective breeding has created in these species. Image:   van Grouw (2018)  .

Unnatural Selection showcases an extraordinary number of unusual domestic animals (these are all domestic pigeon breeds), but the main theme is the evolutionary plasticity, and potential, that human selective breeding has created in these species. Image: van Grouw (2018).

There’s a huge quantity of information here on the diversity of domestic animals and the backstories to them. Van Grouw talks about specific breeders and what they aimed to do with particular animal breeds, as well as the (sometimes erroneous) ideas on why certain breeds are the way they are. But… this isn’t really what the book is about. The main thrust here, instead, is to highlight how selective breeding has enhanced and modified the variation and plasticity present ancestrally in the animals concerned, and on how selective breeding is super-rapid, human-controlled evolution, echoing, mirroring or replicating that which has occurred outside of human influence.

Geese (wild and domesticate) from    Unnatural Selection   . This panel combines Katrina’s illustrations with Natee Himmapaan’s amazing skills in writing and labelling. Credit: Katrina van Grouw/  van Grouw (2018)  .

Geese (wild and domesticate) from Unnatural Selection. This panel combines Katrina’s illustrations with Natee Himmapaan’s amazing skills in writing and labelling. Credit: Katrina van Grouw/van Grouw (2018).

If you’re interested in the diversity of domestic animals, in evolution, or in anatomy, you absolutely must get hold of this remarkable book, and all the more so and if you enjoyed The Unfeathered Bird.

Katrina van Grouw. 2018. Unnatural Selection. Princeton University Press (Princeton and Oxford), pp. 284. ISBN 978-0-691-15706-1. Hardback, index, refs. Here at amazon, here at amazon.co.uk, here from the publishers.

 Jonathan Losos’s Improbable Destinies: Fate, Chance, and the Future of Evolution

Why certain things have happened in evolutionary history, why certain other things haven’t happened, and what things might be like had events took a different turn has, as you’ll know, been a popular area of discussion in this neck of the woods since whenever. Broadly speaking, I have SpecBio in mind: big-brained dinosaurs, an Earth ruled by giant lizards, that sort of thing. If this subject interests you, you absolutely have to read Jonathan Losos’s Improbable Destinies. Inspired (in part) by Gould’s argument in Wonderful Life that evolutionary history is and was dependent on contingency and not destiny, and with something of a herpetological bias linked to his specialisation as a student of anoles, Losos explores the Park Grass Experiment, guppy and stickleback variation, fruit fly studies, disease biology and more.

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My own biases mean that I especially enjoyed his take on dinosauroids (“… to my surprise … The dinosauroid hypothesis was alive and thriving in cyberspace”; p. 323), both the Dale Russell version and the Kosemen/Roy one (yes, there is an illustration), and I think I recall providing him with some of the relevant literature (Naish 2008). The same chapter also includes a discussion of Perry the Platypus and platypusoids (yes, really), E. O. Wilson’s thoughts on extraterrestrials, and, ugh, Simon Conway Morris’s take on the whole thing.

Marlin Peterson’s re-drawing of one of the Kosmen/Roy dinosauroids, from Jonathan Losos’s  Improbable Destinies . Image:   Losos (2017)  .

Marlin Peterson’s re-drawing of one of the Kosmen/Roy dinosauroids, from Jonathan Losos’s Improbable Destinies. Image: Losos (2017).

Losos’s writing is fun and eloquent and the book is really well designed. References and detailed footnotes are included, and the black and white illustrations by Marlin Peterson are really nice. If you enjoy reading this blog, you will like this book. UPDATE: hilariously, I just discovered that I reviewed this book before, here at TetZoo ver 2 in December 2017. I covered exactly the same points because my brain only works one way.

Jonathan B. Losos. 2017. Improbable Destinies: Fate, Chance, and the Future of Evolution. Riverhead Books (New York), pp. 368. ISBN 978-0-399-18492-5. Hardback, index, refs. Here at amazon, here at amazon.co.uk, here from the publishers.

Laufer et al.’s Audubon’s Last Wilderness Journey: The Viviparous Quadrupeds of North America

Most people reading this, especially those interested in the portrayal of animals in art, will be familiar with the several works of naturalist, ornithologist and artist John James Audubon (1785-1851). Audubon is most famous for his grand, lavish The Birds of America, published between 1827 and 1839 and involving years of work, massive investment and hundreds of illustrations. But he should be just as well known for his equally momentous and lavish The Viviparous Quadrupeds of North America.

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This new book – including contributions from nine authors, and edited by Charles T. Butler – reproduces all 150 prints from that work, includes a great deal of material that essentially serves as ‘the making of The Viviparous Quadrupeds of North America’ (actually, this is the exact title of Ron Tyler’s article within the volume), and also features sections of text on the history of conservation and natural history in North America, and on how human-wildlife interactions have changed since Audubon’s time. Footnotes cite the relevant correspondence and literature. These ‘introductory’ sections take us all the way to p. 84, so this is a substantial volume (31 x 26 cm, 280 pp).

As goes the illustrations, the quality of reproduction is outstandingly good. The colours are deep and vibrant, and the level of detail that shows up in magnified sections is impressive. Aubudon was really good at drawing hairs, and my god his skunks, foxes, bears and others clearly involved a lot of work and are quite something to examine in detail. The plates are reproduced in entirety, but magnified sections also included – for some of them – as well, and it was these that I enjoyed looking at the most.

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The volume ends with a timeline that explains every step in the creation of The Viviparous Quadrupeds, including reproductions of correspondence and colour maps of Audubon’s journey.

This is a grand, beautiful book that will be appreciated by anyone interested in mammals or natural history more broadly. Those especially interested in Audubon and his work will appreciate the biographical and background information, and those who don’t own previous reproductions of The Viviparous Quadrupeds (I imagine that that’s most of us) will enjoy the excellent reproductions of his art.

Marilyn Laufer, Ron Tyler, Charles T. Butler, Dennis Harper, Daniel Patterson, Sarah Zohdy, Robert A. Gitzen, James B. Armstrong & Christopher A. Lepczyk. 2018. Audubon’s Last Wilderness Journey: The Viviparous Quadrupeds of North America. Jule Collins Smith Museum of Fine Art, Auburn University/D. Giles, London. pp. 280. ISBN 978-1-911282-10-5. Hardback, index, refs. Here at amazon, here at amazon.co.uk (where title is spelt wrong.. I have told amazon), here from the publishers.

John Reilly’s The Ascent of Birds

It has always been a great paradox to me why books on the evolutionary history of birds are so incredibly rare. There is – as you’ll know if you’re at all familiar with books on natural history – a wonderful yet maddeningly frustrating never-ending glut of excellent, glossy, beautiful, horridly expensive books on birds, year on year on year, yet bird evolution just never gets covered. Regular readers will know what I think of Alan Feduccia’s idiosyncratic, biased and utterly misleading writings.

It is a major problem that there has long been a total shortage of reliable books on bird evolution. Alan Feduccia’s books - two of which are shown here - are unreliable and misleading. Images:  Yale University Press (left);   Yale University Press (right) .

It is a major problem that there has long been a total shortage of reliable books on bird evolution. Alan Feduccia’s books - two of which are shown here - are unreliable and misleading. Images: Yale University Press (left); Yale University Press (right).

John Reilly’s volume substantially alleviates that gap in the market (yes, I do know about Gerald Mayr’s books), and it’s really good. Written very much in the format of Dawkins’s The Ancestor’s Tale (which I confess to not liking all that much, ask me if you want to know why), The Ascent of Birds is a very dense, very well informed, mostly up to date tour of the bird family tree, its structure following molecular phylogenies like those of Hackett et al., Jarvis et al., Prum et al. and so on: specifically, he follows the topology published by Jetz et al. (2012). I guess he used Jetz et al. (2012) specifically because it was the most up to date at the time of writing (various of its conclusions have been replaced by those more recent studies). My own (now dated) review of bird evolution (Naish 2012) is cited, but I think only as a source of information on passerine genitalia. Ok.

The excellent cover art - featuring paintings by Jon Fjeldså (whose illustrations appear in several of his papers) - depicts the cladogram published by Jetz  et al . (2012). Image:   Pelagic Publishing  .

The excellent cover art - featuring paintings by Jon Fjeldså (whose illustrations appear in several of his papers) - depicts the cladogram published by Jetz et al. (2012). Image: Pelagic Publishing.

Chapters discuss lineages in the order of their branching in the Jetz et al. cladogram.. broadly speaking (there are actually a few deviations). Each combines something on the general history of the lineage before focusing on a story specific to one, some or all members of said lineage. The chapter on albatrosses, for example, examines models of albatross biogeography before discussing dynamic soaring behaviour and olfactory biology while that on corvids mostly focuses on cognitive skills and their evolution. We thus get a broad-brush picture on the group’s evolution before zooming in on something more specific but still relevant to the group’s story as a whole. This works really well, and Reilly focuses on things that are – in my opinion – more interesting than Dawkins did.

References are provided throughout, most chapters contain a diagram or two, and a colour plate section includes images of fossils and living birds relevant to the main prose.

There are a lot of passerine lineages: this simplified cladogram features the major lineages only. It is fitting that about half of Reilly’s  The Ascent of Birds  is devoted to this one group. Image: Darren Naish,   from my textbook project  .

There are a lot of passerine lineages: this simplified cladogram features the major lineages only. It is fitting that about half of Reilly’s The Ascent of Birds is devoted to this one group. Image: Darren Naish, from my textbook project.

Long-time readers will know that I once invested considerable time in a book that, similarly, took its readers on a tour of the modern, molecular avian family tree. It was less prose-based than Reilly’s book, and with more focus on images, and was ultimately abandoned due to lack of interest from publishers (part of the story is discussed here, though the article concerned is now lacking all of its accompanying images). Reilly’s book does what my book should have done, but in a far more detail-oriented fashion, meaning that we finally have a good volume presenting the vast amount of modern work done on bird evolution to those interested. This is a notable achievement and has been well executed.

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Screengrabs from the aborted Naish ‘The Bird Family Tree’ book. Yes, I should pick it up and get it published. No, I can’t do that due to a list of other projects that prevent such action. Images: Darren Naish.

Screengrabs from the aborted Naish ‘The Bird Family Tree’ book. Yes, I should pick it up and get it published. No, I can’t do that due to a list of other projects that prevent such action. Images: Darren Naish.

John Reilly. 2018. The Ascent of Birds: How Modern Science is Revealing Their Story. Pelagic Press (Exeter), pp. 340. ISBN 978-1-78427-169-5. Hardback, index, refs. Here at amazon, here at amazon.co.uk, here from the publishers.

Philippe Geniez’s Snakes of Europe, North Africa and the Middle East: A Photographic Guide

Originated published in French, and translated into English by Tony Williams, Geniez’s Snakes is a compact, attractive, well designed and pleasingly thick volume, definitely one to get if you’re a snake specialist. An introductory section covers snake natural history, biology, habitat, the effects of snakebite and classification; the rest of the book – which goes through the species on a family by family basis – covers ‘identification’, ‘range’ as well as diet, reproduction and so on for each species (where available) and is especially good on intraspecific variation and proposed subspecies. Geniez has gone to some trouble to use the newest ideas on taxonomy and phylogeny, the various boxed-out sections providing discussion on genera and their contents. So, you’re going to meet Myriopholis, Spalerosophis, Platyceps and others, if you haven’t already.

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There are already a few (mostly very good) fieldguides to the reptiles of the European field guide region – this typically being taken to include northern Africa and at least part of the Middle East too – so what makes this book different as goes which taxa it includes? Well, there are quite a few species from outside the region usually covered, including a few surprises – too many to discuss here.  I will say that they include the Brahminy worm snake Indotyphlops braminus (yeah, not Ramphotyphlops anymore), California kingsnake Lampropeltis californiae (yup: introduced to the Canary Islands and now abundant there, and also known from Spain), and the as-yet-unnamed Israeli whip snake.

Pages from   Geniez (2018)  . It’s a great-looking book. Image:   weboryx.com twitter account.

Pages from Geniez (2018). It’s a great-looking book. Image: weboryx.com twitter account.

This is not a ‘fieldguide’ in the sense of having plates depicting related species in close proximity. Instead, photos (taken by a long list of different photographers) are distributed throughout the text. But they’re outstandingly good and make this a very colourful, attractive book. Strongly recommended for those of herpetological inclination!

Philippe Geniez. 2018. Snakes of Europe, North Africa and the Middle East: A Photographic Guide. Princeton University Press (Princeton and Oxford), pp. 379. ISBN 978-0-691-17239-2. Softback, index, refs. Here at amazon, here at amazon.co.uk, here from the publishers.

 For other TetZoo book reviews and articles relevant to the topics covered here, see…

Refs - -

Geniez. P. 2018. Snakes of Europe, North Africa and the Middle East: A Photographic Guide. Princeton University Press, Princeton and Oxford.

Jetz, W., Thomas, G. H., Joy, J. B., Hartmann, K. & Mooers, A. O. 2012. The global diversity of birds in space and time. Nature 491, 444-448.

Losos, J. B. 2017. Improbable Destinies: Fate, Chance, and the Future of Evolution. Riverhead Books, New York.

Naish, D. 2008. Intelligent dinosaurs. Fortean Times 239, 52-53.

Naish, D. 2012. Birds. In Brett-Surman, M. K., Holtz, T. R. & Farlow, J. O. (eds) The Complete Dinosaur (Second Edition). Indiana University Press (Bloomington & Indianapolis), pp. 379-423.

van Grouw, K. 2013. The Unfeathered Bird. Princeton University Press, Princeton and Oxford.

Tetrapod Zoology Is A Teenager Now

Yet again a year has passed and yet again it’s time to review events relevant to the TetZooniverse, so here we are at TetZoo’s 13th birthday (January 21st 2006 being the date on which the blog first came into being). As per usual, I’m going to discuss, comment on or mention all those TetZooniverse things that I deem worthy of discussion, comment or mention. And, as usual, I implore you not to read this if the thought of me writing about my own achievements and adventures sounds arrogant or worth a miss. Still here? Ok, we proceed…

Once again I’m going to begin by reminding you of how it all began, back in 2006… you can see that very first article (should you want to)   here  . Images: Darren Naish.

Once again I’m going to begin by reminding you of how it all began, back in 2006… you can see that very first article (should you want to) here. Images: Darren Naish.

Our story begins in late January 2018. Birds (owls and rollers, among others) were still keeping me busy as goes The Big Book, and academic work on ichthyosaurs, pterosaurs and sea monsters (of the cryptozoological sort) was underway. It’s a reminder (as if another were needed) of how slowly it takes for academic projects to come together to note that none of those projects have yet seen print, more than a year later. Fossil bats – covered only because I suddenly suffered bat guilt – got a little coverage at TetZoo during February. I visited Hill Head, West Wittering and Swanwick to see grebes, divers, waders and gulls. The conditions were mostly cold and windy but clear and sunny, which is great for looking at water- and seabirds.

There are so many great places to see water- and seabirds around the coasts of the UK. One of my favourite haunts at the moment is Swanwick in Hampshire. Teals at left; Brent geese at right. Image: Darren Naish.

There are so many great places to see water- and seabirds around the coasts of the UK. One of my favourite haunts at the moment is Swanwick in Hampshire. Teals at left; Brent geese at right. Image: Darren Naish.

Dinosaurs in the Wild (DITW) opened for its final, London-based run at the start of February. If you’re a regular reader you’ll already be familiar with this interactive, science-based visitor attraction so I won’t be saying much more about it today. If you don’t know what I’m talking about and are curious, see Dinosaurs in the Wild: An Inside View at TetZoo ver 3 and The Last Day of Dinosaurs in the Wild here at ver 4. I did various promotional events for DITW from February to June and met science journalist and author John Pickrell at the exhibition during February. Also in February, I wasted time submitting a grant application which was ultimately unsuccessful – more on that later.

Scenes from Dinosaurs in the Wild. At left, your humble author with John Pickrell (and friend: John is the one not holding the plush tyrannosaur). At right: myself, Siouxsie the  Dakotaraptor , and Chris Packham. Images: Darren Naish, Megan McCubbin.

Scenes from Dinosaurs in the Wild. At left, your humble author with John Pickrell (and friend: John is the one not holding the plush tyrannosaur). At right: myself, Siouxsie the Dakotaraptor, and Chris Packham. Images: Darren Naish, Megan McCubbin.

A TV thing filmed at Colchester Zoo during October 2017 appeared on BBC4, titled ‘The Incredible Science of Temperature’ and presented by Helen Czerski. I was filmed discussing, with Helen, what influence global temperatures have had on the distribution of ectothermic reptiles (qualifier because not all non-bird reptiles are ectothermic), and on the physiology and behaviour of Komodo dragons. It felt, at the time, like a total disaster but I’m pleased to say that the parts of my performance included in the show weren’t too bad (the relevant episode is online here). Fighting blue tits, Leguatia, and an article about Mark Witton’s palaeoart appeared at TetZoo during this part of the year. Remember that TetZoo was still hosted at SciAm at this point.

In March I assisted with a local beach clean event. I don’t talk about these to brag about my public service (heavens, no), but to help maintain awareness of plastic pollution and the problems facing marine environments today, and also to remind those of you reading this that you can help by getting involved with local environmental groups.

Beach cleaning is always such a joy (am being ironic). Here’s a haul after just a few hour’s work. Image: Darren Naish.

Beach cleaning is always such a joy (am being ironic). Here’s a haul after just a few hour’s work. Image: Darren Naish.

March was also the month in which Paul Barrett and I compiled our suggested list of corrections and updates for the second edition of Dinosaurs: How They Lived and Evolved. Again, I don’t need to say much about this here since the story behind the second edition was recently covered here in, err, The Second Edition of Dinosaurs: How They Lived and Evolved. Thanks again to those who provided help in compiling corrections, to those who’ve said positive things about the book and to those who’ve purchased it and helped spread the word about it.

Milo the Neotropical giant toad - at right - now lives at Tet Zoo Towers, but his original home was the BBC Natural History Unit, at BBC Bristol. Like Milo, I spent time at BBC Bristol during 2018. Why? Well… we shall see. Images: Darren Naish.

Milo the Neotropical giant toad - at right - now lives at Tet Zoo Towers, but his original home was the BBC Natural History Unit, at BBC Bristol. Like Milo, I spent time at BBC Bristol during 2018. Why? Well… we shall see. Images: Darren Naish.

I engaged in an online interview with the brilliant Aron Ra who I’ve long been familiar with but had not previously spoken to. We discussed TetZooCon, the Tetrapod Zoology project, my books, serial mis-educator David Peters, and more. The interview – part of Aron’s RaMen series – is online here. I also used our virtual meeting as an excuse to invite Aron as a speaker to TetZooCon 2018 since, by virtue of good timing, he was going to be in the UK at exactly the right time. More on TetZooCon later. Oh: I joined instagram in March, wholly because I read an article by a scientist bemoaning scientists who join instagram. My review of Jonathan Kane et al.’s most worthy book God’s Word or Human Reason? appeared at TetZoo in late March, as did a short review of everything columbiform. By now, the substantial passerine section was being done for The Big Book which surely meant that completion of the enormous bird section was in sight, right?

Work on The Big Book (my in-prep textbook  The Vertebrate Fossil Record ) mostly involved birds across 2018. In cases, as here, I had to revamp some old illustrations. This image shows old (at right) and new images of  Aepyornis … and surely I won’t need to revamp  Aepyornis  AGAIN, right? Some of you will know what I’m getting at. Image: Darren Naish.

Work on The Big Book (my in-prep textbook The Vertebrate Fossil Record) mostly involved birds across 2018. In cases, as here, I had to revamp some old illustrations. This image shows old (at right) and new images of Aepyornis… and surely I won’t need to revamp Aepyornis AGAIN, right? Some of you will know what I’m getting at. Image: Darren Naish.

Sunday Brunch, publishers, duckbills, sexual selection. And so to April. As – once again – part of the whole promotional Dinosaurs in the Wild thing, I appeared on the TV show Sunday Brunch… which I’ve never watched but is, by all accounts, pretty popular. It went well and I got to talk about our current knowledge of Mesozoic dinosaurs a fair bit as well as promote Dinosaurs in the Wild, but what might have been the most memorable section of the interview was at the end where presenter Tim Lovejoy asked what the future might hold in store as goes the evolution of life on Earth. Ever the optimist, I told them about the impending mass extinction and the death of biological diversity at the hands of humanity, and that was that.

A view from the set of  Sunday Brunch , April 2018. Ash, Chrissie and a hatchling  Dakotaraptor  at left. Image: Darren Naish.

A view from the set of Sunday Brunch, April 2018. Ash, Chrissie and a hatchling Dakotaraptor at left. Image: Darren Naish.

Chrissie, Darren and the  Dakotaraptor  hatchling, live on Sunday Brunch on April 8th 2018. Image: (c)  Sunday Brunch /Channel 4.

Chrissie, Darren and the Dakotaraptor hatchling, live on Sunday Brunch on April 8th 2018. Image: (c) Sunday Brunch/Channel 4.

In an effort to get the planned Dinosaurs in the Wild book off the ground, I physically dragged myself from publisher to publisher. So many meetings, and all in vain. While at William Collins, however, I did get to see the complete set of the New Naturalist series and, via publisher Myles Archibald, got to hear the backstory to that Too Big to Walk book… the one by ol’ BJ Ford in which he asserts his uncanny ability to finally get dinosaurs right (they were all aquatic, you see). More on that shortly. I also had a meeting about another planned cryptozoology-based TV series. I can basically guarantee having two or three such meetings about the same sort of project every year. They never go anywhere. Yet.

The Collins New Naturalist books on display at the Collins office in London. Most of us own at least a few of these volumes. Image: Darren Naish.

The Collins New Naturalist books on display at the Collins office in London. Most of us own at least a few of these volumes. Image: Darren Naish.

My article on why duckbilled dinosaurs really shouldn’t be called duckbilled dinosaurs appeared in April. It includes a significant gaff and a follow-up article is due to be published here within the next several weeks. The second part of of my review of the Indiana University Press 2014 hadrosaurs volume was also published in April: the first part of my review is one of those articles that SciAm removed due to perceived problems with image use so isn’t currently online. I’ll republish it here at ver 4, eventually.

Attendees of the Sexual Selection meeting at Chicheley Hall, Buckinghamshire. Image: Rob Knell.

Attendees of the Sexual Selection meeting at Chicheley Hall, Buckinghamshire. Image: Rob Knell.

I attended the Royal Society Sexual Selection meeting at Chicheley Hall in Buckinghamshire in early May and chaired one of the sessions. A TetZoo article about the meeting can be found here at ver 3.

I went to Tintagel, north Cornwall, in May 2018. Here’s a scene of the coastline there, with seabirds. Image: Darren Naish.

I went to Tintagel, north Cornwall, in May 2018. Here’s a scene of the coastline there, with seabirds. Image: Darren Naish.

Naish v Ford. The biggest event of May was the debate with Brian Ford at Conway Hall, hinted at above. It seems that this was actually the official launch for Mr Ford’s book – it’s hard to tell as I didn’t see anybody buy a copy – but it was more like a structured debate, the two of us presenting opposing viewpoints on the palaeobiology of Mesozoic dinosaurs before responding to questions from the floor. Events were reported on twitter via #FordvNaish, and my talk can be seen here online.

At left: the cover slide for the event. At right: protesters outside Conway Hall, as imagined by Gareth Monger. Well done if you recognise the reference. Images: Darren Naish, (c) Gareth Monger.

At left: the cover slide for the event. At right: protesters outside Conway Hall, as imagined by Gareth Monger. Well done if you recognise the reference. Images: Darren Naish, (c) Gareth Monger.

I’ve written about Brian Ford and his ideas at least twice before: both in print for an invited article in Laboratory News (Naish 2012) and at TetZoo ver 3. Needless to say, I think he’s flat-out wrong and also guilty of conspiracy mongering, self-aggrandisement (he has literally compared himself to Galileo) and crankery (“nobody’s smart but me!”), among other things. Evaluating the talks is difficult due to the fact that I’m just about never happy with the presentations I give; in any case, John Conway and I already dissected the event at some length in an episode of the podcast. Having said that, Brian asserted that everyone was getting dinosaurs wrong, and assert is basically all he did. I would think that the mostly negative responses that have been directed toward Mr Ford’s ideas would have been sufficient to make him realise that he could very well be wrong as goes his take on dinosaurs, but… nope… he’s still at it (at least, judging from comments he’s recently been making at ResearchGate). I don’t want to give him more publicity than I already have, but if I need to respond to him again, I will.

Darren Naish (left) and Brian Ford (right). Image: Luke Muscutt.

Darren Naish (left) and Brian Ford (right). Image: Luke Muscutt.

Podcats, Jurassic Park at 25, lego at Metro. Having mentioned Conway, we released a few episodes of the podcast – the Tet Zoo podcats or Tetrapodcats – throughout the first half of the year, but we’re still nowhere near having anything like a regular schedule. Mostly this is because John is incredibly lazy, and unproductive to boot. An added factor is that I ‘accidentally’ cleaved through a buried telephone cable in the garden with a spade one day, but I’m sure that that’s coincidental. The new cover for Dinosaurs How They Lived and Evolved, produced by the excellent Bob Nicholls, was released in May – sooner than expected. More on that below.

In May 2018, I (with my kids Will and Emma) attended the Portsmouth Comic Con, and here are some scenes. Not really relevant to TetZoo, but what the hell. Images: Darren Naish.

In May 2018, I (with my kids Will and Emma) attended the Portsmouth Comic Con, and here are some scenes. Not really relevant to TetZoo, but what the hell. Images: Darren Naish.

Hunting Monsters – the cryptozoology book that I’ve now mentioned here on some number of occasions (Naish 2017) – was mentioned on the TV show Jeopardy in early June, wow. Thanks to whoever it was who brought my attention to this. June 2018 was also the time at which Jurassic Park – a movie you might have seen or heard about – celebrated its 25th birthday. A lot of the fan response to the original Jurassic Park – we see a lot of it in vertebrate palaeontology, in fact it comes around every few years – is, my apologies for seeming rude here, mostly vacuous and doesn’t come from a position all that well informed as goes the backstory and making-of the movie. Make of that what you will; whatever, my three Jurassic Park articles are here, here and here, and my liberal use of Jurassic Park toys reflects my best efforts to circumvent the rules regarding image use at SciAm. For all that, they still objected so much to my use of a photo of an issue of National Geographic that they removed it. So I created my own version of the cover.

Hunting Monsters  appears on  Jeopardy . Image: (c)  Jeopardy , NBC/American Broadcasting Company.

Hunting Monsters appears on Jeopardy. Image: (c) Jeopardy, NBC/American Broadcasting Company.

Another bit of Dinosaurs in the Wild promotion happened in late June when – together with palaeontologist Alessandro Chiarenza and actor Ross Cooper – I got to sit in a studio with Natasha Salmon and build Jurassic World lego, all the while talking about dinosaurs. This was for the Metro newspaper, and what fun it was. I even got to keep some of the lego. I also had an article on dinosaurs – specifically on how our perception of their appearance has changed – appear in the Independent Online in mid July. It’s here.

Playing with  Jurassic World: Fallen Kingdom  lego at the Metro offices. I’m not entirely sure why Natasha is making the face she is. Image: (c) Metro.

Playing with Jurassic World: Fallen Kingdom lego at the Metro offices. I’m not entirely sure why Natasha is making the face she is. Image: (c) Metro.

Eotyrannus 2018. During late June I participated in the 2018 BioTweeps conference (an online, Twitter-based conference in which attendees present their ‘talks’ via a series of threaded tweets), my presentation being ‘Eotyrannus and the History of Tyrant Dinosaurs’ (abstract here). As an invited presenter I got to tweet as much as I wanted to in the allotted 30 minute slot. It was a very rewarding experience (with a massive, global reach, I might add), and my thanks to Dani Rabaiotti and Anthony Caravaggi for getting me involved. You can see my presented tweets here (with an addendum here).

Naish at BioTweeps 2018. Image: Darren Naish.

Naish at BioTweeps 2018. Image: Darren Naish.

Having mentioned Eotyrannus… as some of you know, I am (still) committed to publishing the monograph on this dinosaur, and while I failed to get the damned thing into print in 2018, I promise that this will happen in 2019. It’s not for want of trying: substantial progress was made on the post-review version of the manuscript throughout the year, it’s just that I was never able to commit sufficient time to getting it finished. Silly old me for leaving professional scientific research and becoming a freelancer. My co-author Andrea Cau continues to be a most worthy collaborator.

The  Eotyrannus  holotype, in its storage boxes, at Dinosaur Isle, Sandown, Isle of Wight… as it looked in 2012. One day my work on this dinosaur will be published. Image: Darren Naish.

The Eotyrannus holotype, in its storage boxes, at Dinosaur Isle, Sandown, Isle of Wight… as it looked in 2012. One day my work on this dinosaur will be published. Image: Darren Naish.

I mentioned earlier that I was unsuccessful in obtaining a grant – that’s no big deal, most grant applications in science are unsuccessful. Said grant was an attempt to win the money I require to publish an open-access version of the Eotyrannus monograph. On the advice of colleagues, I tried a different approach and set up a gofundme campaign to raise the money aaaand succeeded in raising it (and a bit extra) in about 5 hours. WOW.

Eotyrannus  at gofundme — SUCCESS. Image: gofundme.

Eotyrannus at gofundme — SUCCESS. Image: gofundme.

Speaking of long-running projects that never seem to reach completion, I and colleagues put what are supposed to be the final touches to a PhyloCode Companion Volume article on theropods during the summer. I’m also part of the team that produced the volume’s sauropod and sauropodomorph articles. But they have a somewhat different fate from the theropod article, that’s all I’ll say for now.

This image has to go somewhere and here is as good a place as any. It’s not made by me, but by naughtyhippo ( if that is her real name ).   Available on merchandise here.   Image: Darren Naish.

This image has to go somewhere and here is as good a place as any. It’s not made by me, but by naughtyhippo (if that is her real name). Available on merchandise here. Image: Darren Naish.

Goodbye SciAm. July was a significant month as goes TetZooniverse things as, on July 25th, I reached a funding goal at patreon. Regular TetZoo readers will know (my god, how many times have l mentioned it?) that I was not happy with the way things were going at SciAm. Their total clamping down on image use and removal of various of my articles was pretty much the last straw: blogging there was proving more trouble that it was worth, despite the major kudos of being attached to the SciAm brand. Achieving the relevant goal meant that I could leave (as a freelancer, I’m dependent on earning from my writing: it is my primary income) and set up shop as an independent blogger once more (harking back to the days of TetZo ver 1, 2006). Despite my unhappiness with SciAm, walking away was not a decision I took lightly. The very last TetZoo ver 3 was published on July 31st, and here we are at ver 4, at the same website at the podcast. Thanks, as ever, to the patreon supporters who made this possible.

Leaving SciAm means that I’m now entirely dependent on   support at Patreon  . What a gamble. As of right now, the site needs updating. Image: Darren Naish.

Leaving SciAm means that I’m now entirely dependent on support at Patreon. What a gamble. As of right now, the site needs updating. Image: Darren Naish.

August began with the taking on of an exciting job in the museums world. It involves an exhibition relevant to our interests, but it won’t go live until 2020 and I won’t be talking about it until then, sorry. I’ll give you a clue and say that it has meant spending time in Cornwall, one of my favourite parts of the UK.  Also during August, I spoke to New Scientist about megalodon in view of a scientific documentary called Meg. I’m not, technically, an expert on megatooth sharks but I have written about them several times: there’s a section in Cryptozoologicon Volume I (Conway et al. 2013) and another – as yet unpublished, of course – in the chondrichthyan section of The Vertebrate Fossil Record.

From time spent in Hay-on-Wye. This is Hay Bluff. Image: Darren Naish.

From time spent in Hay-on-Wye. This is Hay Bluff. Image: Darren Naish.

Articles on the potential domestication potential of non-bird dinosaurs and mastigures (or spiny-tailed agamas) appeared at TetZoo. Together with the family, I visited Hay-on-Wye, the fabled Town of Books of the Welsh borders (and filming location for certain iconic scenes in An American Werewolf in London). There aren’t half as many book shops in Hay-on-Wye as there used to be, but there are enough to make a visit very much worthwhile if you’re a bibliophile. I made a great number of amazing discoveries – beware, there’s a dedicated natural history bookshop (C. Arden) – and am really happy that I finally took the time to go there.

Don’t go to Hay-on-Wye, it will be bad. Image: Darren Naish.

Don’t go to Hay-on-Wye, it will be bad. Image: Darren Naish.

As I mention every now and again, I continue to remain involved in technical scientific research as and when I can, and late August saw the publication of my then-newest paper: a study of pelvic neural anatomy in pterosaurs and what the relevant details might mean for ecomorphology, co-authored with Liz Martin-Silverstone and Dan Sykes (Martin-Silverstone et al. 2018). This was the second 2018 paper that included an analysis of Vectidraco – a small Cretaceous azhdarchoid pterosaur I and colleagues described in 2013 (Naish et al. 2013) – the other being Rachel Frigot’s study of pelvic musculature included within the Geological Society special volume New Perspectives on Pterosaur Palaeobiology (Frigot 2018). A discussion on the new paper was provided here.

At left, an illustration of the  Vectidraco  pelvis, some key features labelled. At right: a pterosaur montage produced for the TetZoo article on pterosaur palaeoneurology. Images: Darren Naish.

At left, an illustration of the Vectidraco pelvis, some key features labelled. At right: a pterosaur montage produced for the TetZoo article on pterosaur palaeoneurology. Images: Darren Naish.

September 2nd was the very final day of Dinosaurs in the Wild. I went on a final tour, attended the staff party, and said all those sad, final goodbyes. All things must come to an end, but hope remains that DITW will be able to open its doors again one day.

Me (at left) with Tim Haines - of Impossible Pictures, Walking With Dinosaurs, Primeval and so on - doing some Dinosaurs in the Wild promotion for radio during May 2018. Image: Darren Naish.

Me (at left) with Tim Haines - of Impossible Pictures, Walking With Dinosaurs, Primeval and so on - doing some Dinosaurs in the Wild promotion for radio during May 2018. Image: Darren Naish.

Dougal Dixon, Crystal Palace, DHLTE: 2nd ed. Early September also brought the devastating news of the calamitous fire at Rio’s National Museum, a blow for human knowledge as a whole but especially for our Brazilian friends and colleagues. While at the museum in 2013, I took some reasonable number of photos, so have been submitting them to Brazilian colleagues. In happier news, September also saw the Dougal Dixon event at London’s Conway Hall (covered here at TetZoo) in which Dougal and I discussed his work and projects in front of a pretty substantial crowd. Original art, models and more were in attendance – it was great.

At Conway Hall, for the  After Man -themed event with Dougal Dixon. Images: Darren Naish.

At Conway Hall, for the After Man-themed event with Dougal Dixon. Images: Darren Naish.

Just a few days later, and I was speaking at another London-based event, this time the Crystal Palace Dinosaurs Days event, part of the Heritage Open Days weekend occurring across the UK. Other speakers were in attendance too, and we had special up-close access to the models. I’m surely repeating myself, as all of this was covered in the December 2018 article Up Close and Personal With the Crystal Palace Dinosaurs. As you may know, part of the reason for the publicity push surrounding the models is that funding is desperately needed for their care and upkeep, and September 2018 saw the launch of a crowd-funded project to get a bridge to the models constructed (said bridge then allowing the access that can otherwise only be obtained on rare occasion). The project reached its target (which was over £70,000) in December, which is excellent and exhilarating news.

Special access to the islands at Crystal Palace, what a treat. Here, I’m talking about what the Victorians thought about ichthyosaurs. For more, see   Up Close and Personal With the Crystal Palace Dinosaurs  . Image: Emma Wain.

Special access to the islands at Crystal Palace, what a treat. Here, I’m talking about what the Victorians thought about ichthyosaurs. For more, see Up Close and Personal With the Crystal Palace Dinosaurs. Image: Emma Wain.

The second edition of Dinosaurs: How They Lived and Evolved – my Natural History Museum book with Paul Barrett (Naish & Barrett 2018) – saw print in September. Again, thoughts on this have already appeared at TetZoo so I’ll avoid repeating things here. A review appeared in The Economist, which is kind of a big deal. Summer 2018 also saw the publication of the Russian edition of the book. It has the text of the second edition but the cover of the first.

Screengrab from the  Economist ’s review of the second edition of   Dinosaurs: How They Lived and Evolved  . The lead quote comes from Bob Nicholls, creator of the fuzzy  Tianyulong  we see at left. Image: (c) Economist.

Screengrab from the Economist’s review of the second edition of Dinosaurs: How They Lived and Evolved. The lead quote comes from Bob Nicholls, creator of the fuzzy Tianyulong we see at left. Image: (c) Economist.

TetZooCon 2018! And so to October, and the TetZoo-themed event of the year, by which I mean TetZooCon. As covered here on the blog, 2018’s TetZooCon was the first two day TetZooCon ever, and by far the biggest and (I think) the best. Speakers and presenters included Ian Redmond, Mark O’Shea, Aron Ra, Dougal Dixon and Gert van Dijk (on stage for a SpecBio discussion), Katrina van Grouw and myself. John Conway led a palaeoart workshop that occurred as a parallel session to some of the talks and I led a post-con fieldtrip to Crystal Palace the day after. A MonsterTalk episode with Blake Smith – done as a piece of TetZooCon promotion but mostly revolving around Jurassic Park – was released in early October (it’s here; episode 172).

Scenes from TetZooCon 2018. At left, Albert Chen discusses TetZoo. At right: a box of Dinosaurs in the Wild plushies. Images: Darren Naish.

Scenes from TetZooCon 2018. At left, Albert Chen discusses TetZoo. At right: a box of Dinosaurs in the Wild plushies. Images: Darren Naish.

Hey, I had to turn down a free, all-expenses-paid birdwatching trip to Cadiz in Spain because it clashed with TetZooCon, talk about bad luck. The success of TetZooCon 2018 means that a 2019 event is just about guaranteed (or, it is now that John and I are back on speaking terms again); it will again be a two-day event but I can’t yet say any more than that. News will be announced as and when I have it.

TetZooCon 2018 included a dedicated SpecBio discussion, and here’s Gert van Dijk talking about his own Furaha project, at said event. Image: Darren Naish.

TetZooCon 2018 included a dedicated SpecBio discussion, and here’s Gert van Dijk talking about his own Furaha project, at said event. Image: Darren Naish.

Dorling Kindersley. It was around this time in the year that I and Chris Barker put the finishing touches to a new kid’s book we’ve put together for Dorling Kindersley, titled Where on Earth? Dinosaurs and Other Prehistoric Life. I wouldn’t mention this (seeing as the book isn’t published yet), but it’s already being advertised.

Screengrabs from the in-press   Where on Earth? Dinosaurs and Other Prehistoric Life  , by Chris Barker and Darren Naish. Images: Barker & Naish/Dorling Kindersley.

Screengrabs from the in-press Where on Earth? Dinosaurs and Other Prehistoric Life, by Chris Barker and Darren Naish. Images: Barker & Naish/Dorling Kindersley.

In fact, I did a huge amount of work for Dorling Kindersley during 2018 as I and colleagues at DK worked hard to overhaul their entire prehistoric animal picture library. One volume that benefited from this overhaul – John Woodward’s The Dinosaurs Book (Woodward 2018) – was published during the year (I was consultant). It should be obvious to anyone paying attention to children’s books on dinosaurs that this overhauling has resulted in a massive improvement as goes the sorts of images the books now contain.

Castle Urquhurt, as photographed from the surface of Loch Ness (a scene from 2014, not 2018). Image: Darren Naish.

Castle Urquhurt, as photographed from the surface of Loch Ness (a scene from 2014, not 2018). Image: Darren Naish.

Loch Ness, 99% Invisible. I’ve mentioned once or twice here my communications with Professor Neil Gemmell and my role in his eDNA-based work on the biology and ecology of Loch Ness. Neil and I met up in 2017 to discuss this work (a 2018 article on the backstory to Neil’s research – one of several – can be found here), and I published two magazine articles on the subject during the year: one in a kid’s magazine in October (Naish 2018a) and another for a more adult audience in December (Naish 2018b). It’s very easy to misunderstand, or ‘mis-frame’, this project as “SCIENTIST WASTES TAX-PAYER MONEY ON LOCH NESS MONSTER NONSENSE” and, indeed, at least one author did make this misunderstanding during the year. What should be noted, I’d argue, is that this is and was a phenomenally successful science outreach programme wherein the Loch Ness Monster was used as the hook to get journalists to cover eDNA and ecological and genetic research, and as such it was absurdly successful (Naish 2018a, b).

Artwork accompanying Naish (2018b). I didn’t write that caption, since the ‘For centuries’ thing is not true. Image: (c)  BBC Focus , Naish (2018b).

Artwork accompanying Naish (2018b). I didn’t write that caption, since the ‘For centuries’ thing is not true. Image: (c) BBC Focus, Naish (2018b).

Late October also saw the release of an episode of the 99% Invisible podcast in which myself, Bob Bakker and John Conway discussed the portrayal of Mesozoic dinosaurs in art. The episode is called Welcome to Jurassic Art and can be found here. Alas, the three of us didn’t really sit around talking (though I have spoken to Bob Bakker, in person, on a few occasions). Instead, our interviews were recorded at different times, at different places. Anyway, it worked out alright.

One of the highlights of the ZSL book-themed event — Jules Howard tells us about the digital duck vagina app. Image: Darren Naish.

One of the highlights of the ZSL book-themed event — Jules Howard tells us about the digital duck vagina app. Image: Darren Naish.

The ZSL zoology books evening. Articles on New Living Animals We Want to Find, Aldrovandi’s monstrous rooster, the second edition of Dinosaurs: How They Lived and Evolved and the pouches of the Sungrebe appeared at TetZoo during November. The article on Aldrovandi’s rooster includes a notable gaff, and an update is due to appear here soon (there is not enough time to do all this stuff, argh!!). Mid-November also saw the Zoological Society of London event From Stoned Sloths to Farting Fish: Comical Tales from the Animal Kingdom, in which myself, Dani Rabaiotti, Jules Howard and Lucy Cooke discussed our adventures in the world of zoology-themed publishing. I mostly spoke about dinosaur sex.

Crew at the public engagement event of December’s Popularising Palaeontology event. L to r: Elsa Panciroli, Mark Witton, Chris Manias, Becky Wragg Sykes, Darren Naish. Image: Darren Naish.

Crew at the public engagement event of December’s Popularising Palaeontology event. L to r: Elsa Panciroli, Mark Witton, Chris Manias, Becky Wragg Sykes, Darren Naish. Image: Darren Naish.

Early in December, I stayed in Bournemouth, UK, for the Joint Scientific Meeting of the British Herpetological Society and Amphibian and Reptile Conservation, which has become one of my favourite small meetings. I spoke at last year’s event but not this one. And, later in the month, I attended the Popularising Palaeontology workshop, organised by Chris Manias and hosted by King’s College, London where I listened to some great presentations and participated in the public engagement event.

Herpetology-themed snap cards, obtained at the Joint Scientific Meeting of the British Herpetological Society and Amphibian and Reptile Conservation. Image: Darren Naish.

Herpetology-themed snap cards, obtained at the Joint Scientific Meeting of the British Herpetological Society and Amphibian and Reptile Conservation. Image: Darren Naish.

And that brings us up to January, a month mostly spent in China and away from the long list of TetZoo-related things I aim to complete in 2019. While in China (specifically, Zigong in Sichuan) I visited Panda Base (properly Chengdu Research Base of Giant Panda Breeding) and the Zigong Dinosaur Museum, among other places.

The entrance to Chengdu Research Base of Giant Panda Breeding, Sichuan, China. Image: Darren Naish.

The entrance to Chengdu Research Base of Giant Panda Breeding, Sichuan, China. Image: Darren Naish.

Thus far, the year’s only TetZoo articles have been those on Heilmann’s Proavis and the life appearance of sauropods. I so wish I could publish more – there is still so much to do. On that note, here’s this…

Achievements for 2018: the pterosaur palaeoneurology paper (Martin-Silverstone et al. 2018), Tetrapod Zoology ver 4 is launched, 2nd edition of Dinosaurs: How They Lived and Evolved sees print (as does Russian translation), Loch Ness magazine articles (Naish 2018a, b), money raised for Eotyrannus monograph, the Dorling Kindersley overhaul, biggest and best TetZooCon so far.

Failures for 2018: not finishing the Eotyrannus monograph, not finishing The Big Book, not making any progress on the Tetrapod Zoology books, Cryptozoologicon Volume 2 or the various dinosaur-themed books planned with John Conway.

Toy thylacines - did you know that there are this many? The grey 3D-printed one - and the painted version at far right (the sitting one) - were obtained at TetZooCon 2018 and were made by Rebecca Groom of palaeoplushies. Image: Darren Naish.

Toy thylacines - did you know that there are this many? The grey 3D-printed one - and the painted version at far right (the sitting one) - were obtained at TetZooCon 2018 and were made by Rebecca Groom of palaeoplushies. Image: Darren Naish.

Ok, so that’s that. How does the year’s blogging do in terms of group representation?

Miscellaneous Musings

Mammals

Lepidosaurs

Pterosaurs

Non-avialan dinosaurs

Birds

Cryptozoology

SpecBio

Tet-Zoo-13th-birthday-Tet-Zoo-13th-birthday-graph-Jan-2018-tiny-Jan-2019-Tetrapod-Zoology-Darren-Naish.jpg

So there we have it. Clearly, 2018 was dominated by non-bird dinosaurs and by articles that can only be classified as generic or miscellaneous and weren’t dedicated to any specific tetrapod group. Amphibians, croc-line archosaurs and so on received no coverage whatsoever. All in all, then, a total and epic failure of what I’ve been trying to achieve here. I figure, by now, that I should stop caring and just aim to write about whatever it is that I write about.

Oh, fish. Image: Darren Naish.

Oh, fish. Image: Darren Naish.

So — happy 13th birthday, Tetrapod Zoology. As ever, I aim to continue to publish as much as I can here: to both present new ideas, new reviews and new takes on zoological topics of interest, and to up-date and correct those things I’ve covered in the past. There is so much to do, and finding time for blogging remains perpetually difficult in view of workload and the never-ending chasing of finances. Plus there’s that textbook I so desperately want to see finished. Here’s your reminder that I’m wholly reliant on funding at patreon, and that I would be able to devote just about all of my work-time to these projects – blogging and the textbook, and the other TetZooniverse-relevant books – if more TetZoo readers were prepared to support me. Huge thanks to those who do so already. Come back soon for another thrilling instalment.

For previous Tet Zoo birthday articles, see...

Refs - -

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

Frigot, R. A. 2018. Pelvic musculature of Vectidraco daisymorrisae and consequences for pterosaur locomotion. In Hone, D. W. E., Witton, M. P. & Martill, D. M. (eds) New Perspectives on Pterosaur Palaeobiology. Geological Society, London, Special Publications 455, 45-55.

Martin-Silverstone, E., Sykes, D. & Naish, D. 2018. Does postcranial palaeoneurology provide insight into pterosaur behaviour and lifestyle? New data from the azhdarchoid Vectidraco and the ornithocheirids Coloborhynchus and Anhanguera. Palaeontology 2018, 1-14. doi: 10.1111/pala/12390

Naish, D. 2012. Palaeontology bites back… (response to Ford’s article on alleged aquatic habits of dinosaurs) Laboratory News May 2012, 31-32.

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

Naish, D. 2018a. Will we ever find Nessie? In Lipscombe-Southwell, A. (ed) BBC Focus Big Book of Mind-Blowing Answers, pp. 16-21.

Naish, D. 2018b. The genetic hunt for Nessie. BBC Focus 329, 66-71.

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

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

Woodward, J. 2018. The Dinosaurs Book. Dorling Kindersley, London.

The Life Appearance of Sauropod Dinosaurs

If you’re a regular TetZoo reader you’ll be familiar with my several articles on the life appearance of Mesozoic dinosaurs, an issue in which I have a special interest. Alas, several of these articles were published at TetZoo ver 2 – the ScienceBlogs years – and hosting issues at the site concerned mean that they’re currently appearing sans all of the many images I so lovingly uploaded. Which is a pain.

A really nice, life-sized model of a titanosaurian sauropod, on display in Romania and constructed by Brian Cooley. Image: Darren Naish.

A really nice, life-sized model of a titanosaurian sauropod, on display in Romania and constructed by Brian Cooley. Image: Darren Naish.

Partly as a consequence, and partly because the issue is on my mind due to several concurrent projects (he says, cryptically), now is a good time to talk once more about the life appearance of Mesozoic dinosaurs, and I’m going to start with sauropods. Sauropods have been covered quite a few times on TetZoo before, as you can see from the list of links at the bottom of this article.

The head, and face especially. We’ll start with the head. Sauropod skulls were proportionally small compared to the overall size of the animals, but not comically so. It’s also worth saying that their eyes – while pretty big in absolute size (based on the size of the eye socket and the sclerotic rings preserved in fossils) – are easy to over-emphasise in illustrations. If you’ve heard that sauropods might have had trunks and thought it reasonable or plausible… well, it’s a poor idea that’s neither reasonable nor plausible, and has a lot counting against it, as covered in the ver 3 article here.

Did sauropods have trunks? NO. It’s the dumbest idea ever, and every piece of evidence counts against it. Several authors have independently explored this idea, including Robert Bakker (upper right) and Bill Munns (lower right). The two images at left are from Knoll  et al .’s (2006) paper refuting the trunk idea. Images: Knoll  et al . (2006),   Bakker (1986)  , Bill Munns ( original here ).

Did sauropods have trunks? NO. It’s the dumbest idea ever, and every piece of evidence counts against it. Several authors have independently explored this idea, including Robert Bakker (upper right) and Bill Munns (lower right). The two images at left are from Knoll et al.’s (2006) paper refuting the trunk idea. Images: Knoll et al. (2006), Bakker (1986), Bill Munns (original here).

Trunks are a no, but did sauropods have ‘lips’ or ‘cheeks’? A whole article could be written on this issue. For now, I’ll summarise things by saying that sauropod skull bone texture indicates that they likely did have extra-oral tissues that mostly covered their teeth (Witton 2018), meaning that they were facially similar to lizards and kin.

A suggestion that some titanosaurs had blade-like cutting sections in the posterior sections of their jaws – so-called ‘guillotine crests’ (Apesteguía 2004) – looks unlikely given that it would require the animals concerned to do their food procurement at the sides of the jaws, rather than at the front like all other sauropods and other dinosaurs. It’s more likely that a mistake has been made here and that the jaw segments thought to support those blade-like, keratinised sections were, while sharp-edged, covered in normal lip tissue. The possibility that diplodocoids and maybe other sauropods too might have had true beaks at the font of the mouth has recently been put forward, but so far only in preliminary fashion.

Apesteguía (2004) argued that some titanosaurs - this is the rhino-sized  Bonitasaura  - had keratinised ‘guillotine crests’ on the edges of the jaws. I think that this is likely a mistake. Image: Apesteguía (2004).

Apesteguía (2004) argued that some titanosaurs - this is the rhino-sized Bonitasaura - had keratinised ‘guillotine crests’ on the edges of the jaws. I think that this is likely a mistake. Image: Apesteguía (2004).

The nostrils and nose. As is now widely known, and near-universally accepted, the external or fleshy nostrils of sauropods were almost certainly not located way up on the forehead, far from the front of the snout, as was long thought based on the retracted position of the bony nostril openings. A more ‘normal’, anterior position for the nostrils was demonstrated by Witmer (2001) who pointed to evidence from blood vessel and nerve impressions and associated cranial hollows, all of which are located on the anterior part of the snout. These indicate the most likely position of the fleshy nostril and associated blood vessel clusters.

The soft dinosaur revolution comes to Sauropod Town. At left, an image by Matt Wedel which shows why we need more soft tissue on our sauropods. At right, Larry Witmer’s (2001) depiction of the different possibilities as goes nostril position in sauropods. (a) is the most likely option based on anatomical data. Image: Mathew Wedel, Witmer (2001).

The soft dinosaur revolution comes to Sauropod Town. At left, an image by Matt Wedel which shows why we need more soft tissue on our sauropods. At right, Larry Witmer’s (2001) depiction of the different possibilities as goes nostril position in sauropods. (a) is the most likely option based on anatomical data. Image: Mathew Wedel, Witmer (2001).

There are also reasons for thinking that the giant, cavernous bony nostril openings and tall nasal bars of some macronarian sauropods supported, and were surrounded by, dome-shaped soft tissue convexities, superficially recalling the bulging nasal regions of some living monitor lizards. Part of my reason for saying this comes from the nasal anatomy of the exquisitely preserved South American titanosaur Sarmientosaurus. Here, an anteriorly projecting bony spine located along the midline and associated bar-like structures on the lateral edges of the large narial fossa – the big bony opening surrounding the bony nostril – indicate that a wide, convex mass of tissue connected the forehead with the sides and front of the snout’s upper surface (Martínez et al. 2016). Sarmientosaurus isn’t unique in this respect but is a particularly good example.

The skulls of some sauropods - this is the titanosaur  Sarmientosaurus  - indicate that there were bulbous nasal structures covering much of the snout region. Image: WitmerLab.

The skulls of some sauropods - this is the titanosaur Sarmientosaurus - indicate that there were bulbous nasal structures covering much of the snout region. Image: WitmerLab.

Indeed, skull openings in general were almost definitely not sunken in appearance or otherwise all that obvious, something that’s being said of archosaurian faces in general as artists and anatomists have learnt to take better attention of the conditions present in living animals (where cranial openings are just about never obvious in the live creature).

Accordingly, sauropod faces were seemingly ‘softer’ and more ‘padded’ than convention would have it, at least some of – perhaps all of – the nasal and forehead region being convex and fleshy, rather than shrink-wrapped and covered in thin skin alone (Witton 2018).

Were sauropod necks mostly semi-horizontal and with a limited range of motion, as argued by Martin (1987) and illustrated at left? Nope; it’s more likely that the necks were often held aloft and far more flexible, as argued by  Taylor  et al . (2009)  and depicted at right. Images: Martin (1987),  Taylor  et al . (2009) .

Were sauropod necks mostly semi-horizontal and with a limited range of motion, as argued by Martin (1987) and illustrated at left? Nope; it’s more likely that the necks were often held aloft and far more flexible, as argued by Taylor et al. (2009) and depicted at right. Images: Martin (1987), Taylor et al. (2009).

The neck. The most remarkable feature of sauropod anatomy is the neck. As goes how the neck was held and how flexible it was, several possibilities have been put forward, some workers arguing that it was held in a mostly horizontal attitude for much of the time and with only a limited range of lateral and vertical flexibility. I’m part of a group who argue for mostly elevated habitual neck poses (even in diplodocids) and a wide range of flexibility (Taylor et al. 2009). Arguments that sauropods must have been horizontal-necked do not take account of the flexibility permitted by cartilage, or – in living animals of all sorts – the ranges of motion that happen at zygapophyseal junctions and at the neck base and head-neck junctions.

There are good reasons for thinking that sauropod necks were habitually inclined upwards, as is typical for terrestrial tetrapods in general, and several fossils are actually preserved with the neck in this orientation. This montage by Greg Paul shows several of the fossil concerned. Image: Paul (1998).

There are good reasons for thinking that sauropod necks were habitually inclined upwards, as is typical for terrestrial tetrapods in general, and several fossils are actually preserved with the neck in this orientation. This montage by Greg Paul shows several of the fossil concerned. Image: Paul (1998).

It's been convention to show the sauropod neck as a featureless tubular structure, like a hose. This is mostly wrong, in part because the shapes of the vertebrae reveal a more unusual cross-sectional shape that varies from one sauropod group to the next. In some sauropods (like mamenchisaurs), the vertebrae are quite narrow and the neck would have looked laterally compressed in places, as it is (for at least some of its length) in giraffes. In diplodocoids – apatosaurines in particular – the neck is narrow close to the head but wide for much of its length, and subtriangular in cross-section, being widest across the neck’s underside. A neck that was quite narrow in its anteriormost quarter or so but was markedly wide for the rest of its length seems to have been the case in brachiosaurs and at least some titanosaurs.

The necks of some sauropods - a good example is the ultra-long-necked  Omeisaurus , photographed at Zigong Dinosaur Museum - are somewhat laterally compressed for at least part of their length. Image: Darren Naish.

The necks of some sauropods - a good example is the ultra-long-necked Omeisaurus, photographed at Zigong Dinosaur Museum - are somewhat laterally compressed for at least part of their length. Image: Darren Naish.

Indeed, the very base of the neck is remarkably broad in some sauropods – Camarasaurus is the classic example, where the neck base is not that different in width from the front of the chest – meaning that the neck would taper gradually along its length if you were looking at the animal from the front or back, or from above or below.

The neck bases of some sauropods - this is the  Camarasaurus  replica on display in London’s NHM - are shockingly broad. It would be wrong to show the neck as a narrow, hose-like object with parallel sides. Image: Darren Naish.

The neck bases of some sauropods - this is the Camarasaurus replica on display in London’s NHM - are shockingly broad. It would be wrong to show the neck as a narrow, hose-like object with parallel sides. Image: Darren Naish.

The vertebrae themselves are very complicated with large hollows on their sides, projecting neural spines on the apices and so on. In the most extreme version of the shrink-wrapping meme promoted by one or two palaeontologists and palaeoartists, sauropod necks have been depicted as if these structures should be visible in the live animal, Ely Kish’s apatosaurines from 1983 being the ultimate example. This was definitely not the case. It remains uncertain how much musculature and soft tissue surrounded the vertebrae, but it was almost certainly (based on the anatomy of living animals) enough to obscure the form of the vertebrae, their bulbous junctions perhaps being visible in sauropods of some or many sorts.

The brilliant, late Ely Kish, one of the best palaeoartists of all time, constructing a scale model of  Apatosaurus  (in preparation for a large painting). Partly on the advice of palaeontologist Dale Russell, she depicted great lateral concavities on the sides of the neck. Image: Russell (1987).

The brilliant, late Ely Kish, one of the best palaeoartists of all time, constructing a scale model of Apatosaurus (in preparation for a large painting). Partly on the advice of palaeontologist Dale Russell, she depicted great lateral concavities on the sides of the neck. Image: Russell (1987).

Necks as display banners. The unprecedented form of the sauropod neck makes it plausible – perhaps even likely – that the neck was used as a display structure. Phil Senter’s suggestion that the size and length of the neck was driven, in evolutionary terms, by its use as a display structure (Senter 2007) is not supported by evidence (Taylor et al. 2011). However, this doesn’t discount co-option of the neck in display, so it really is – while wholly speculative right now – worth taking seriously the possibility that display structures of various kinds could well have adorned sauropod necks. These could include wattles, dewlaps, spiky frills, spines, filaments, inflatable pouches or distensible flags or flaps. A few artists have explored these possibilities, most notably Brian Engh and Emiliano Troco.

As weird as it might seem, it is at least plausible that the sauropod neck was decorated with weird soft-tissue display structures, as depicted here on a diplodocid. Image: Emiliano Troco.

As weird as it might seem, it is at least plausible that the sauropod neck was decorated with weird soft-tissue display structures, as depicted here on a diplodocid. Image: Emiliano Troco.

 Hands. Sauropod hands are extremely odd, and very different from the rounded, elephant-style structures, edged with big nails and/or several claws, shown so often in artistic reconstructions and museum models. Indeed, they’re sufficiently weird and interesting that I’ve written whole articles about them before but, as I said above, these are currently lacking all of their relevant illustrations and are thus all but useless.

Sauropod hands are essentially unique. They’re weird, semi-tubular structures with pillar-like metacarpals. At left, a brachiosaur hand. At right, the hand of the turiasaur  Zby . Images: Anthony Maltese, Darren Naish.

Sauropod hands are essentially unique. They’re weird, semi-tubular structures with pillar-like metacarpals. At left, a brachiosaur hand. At right, the hand of the turiasaur Zby. Images: Anthony Maltese, Darren Naish.

The sauropod hand is essentially a semi-tubular structure formed of elongate metacarpals arranged, pillar-like, in a semi-circle. The posterior surface – corresponding to the palm – was hollow, the consequence being a semilunate area of contact with the ground. We’ve known since at least 1940 that sauropod hands had this very unusual form thanks to fossil tracks (Falkingham et al. 2014), which makes it all the weirder that people have ignored this information and merrily continued giving sauropods elephant-like hands across the decades.

Roland T. Bird’s sauropod track illustrations from the 1940s - shown here - clearly show the true, highly unusual form of the sauropod manus. Shame on those who ignored this information in the following decades. Image:  Falkingham  et al . (2014 ), CC BY 4.0.

Roland T. Bird’s sauropod track illustrations from the 1940s - shown here - clearly show the true, highly unusual form of the sauropod manus. Shame on those who ignored this information in the following decades. Image: Falkingham et al. (2014), CC BY 4.0.

Tracks also show that nails and claws were absent from the hands, except on the thumb where a pointed claw – which varied considerably in exact shape and size from one group to the next – projected inwards and slightly backwards. The thumb claw was lost within Titanosauria, meaning that at least some members of this group lacked nails and claws on their hands altogether. I will concede that poorly defined convexities corresponding to manual digits were present in at least some sauropods, but they still wouldn’t have looked like distinct digits.

Some tracks appear to show that tough semi-conical tubercles projected from the skin on the front and sides of the hand, perhaps giving part of the hand a spiky or tuberculated appearance (Milàn et al. 2005). Maybe these structures had a role in foraging, digging, display or combat, since it’s plausible that they made the hands gnarlier and harder than they would have been otherwise.

Vertical scores associated with sauropod hand prints indicate that at least some of them had tuberculate hand skin, as depicted here (at right) in this modified version of a Greg Paul illustration. Image: Milàn  et al . (2005).

Vertical scores associated with sauropod hand prints indicate that at least some of them had tuberculate hand skin, as depicted here (at right) in this modified version of a Greg Paul illustration. Image: Milàn et al. (2005).

Feet. Three large, curved claws projected anterolaterally from the inner three toes. At least some sauropods possessed four such claws. The outer two toes projected as blunt, rounded convexities in some, most or all sauropods. This is obvious from at least some Brontopodus tracks (Meyer et al. 1994). They might have had nails but were more likely lacking horny structures of any sort and essentially continuous with the rest of the foot’s outer surface. In contrast to the hand, the foot did have a massive fatty pad at its back.

Until recently it was thought that the metatarsals were held at a high angle, the consequence being a short foot shaped like that of an elephant but for the claws (Paul 1987). Data from articulated skeletons and tracks, however, show that the metatarsus was not as erect as argued, in which case the toes were longer and flatter than depicted by some artists.

Tschopp  et al . (2015)   used data from complete  Camarasaurus  hands and feet to produce the skeletal and soft-tissue models you see here (in 1 and 2), and then used these to generate tracks (3). The tracks are an exact match for real fossil tracks. Image: (c)  Tschopp  et al . (2015) .

Tschopp et al. (2015) used data from complete Camarasaurus hands and feet to produce the skeletal and soft-tissue models you see here (in 1 and 2), and then used these to generate tracks (3). The tracks are an exact match for real fossil tracks. Image: (c) Tschopp et al. (2015).

On the subject on feet and limbs, trackways also show that sauropods differed in how they placed their hands and feet, some walking with a very narrow gait (the hands and feet being placed close to the midline), others using a wide gait (where there was some short distance between the hands and feet of the left side versus those of the right), and others being intermediate. It should be remembered that even the widest-gauge sauropods did not walk with their feet all that far apart, but that the legs were almost certainly angled inwards, as is typical in living animals. For more on this issue see Scott Hartman’s article here.

Sauropods were variable in cross-sectional shape and in whether they walked with narrow-gauge or wide-gauge gaits, as depicted here by Scott Hartman. Image:   Scott Hartman’s skeletaldrawing.com

Sauropods were variable in cross-sectional shape and in whether they walked with narrow-gauge or wide-gauge gaits, as depicted here by Scott Hartman. Image: Scott Hartman’s skeletaldrawing.com

The body and tail. Sauropod bodies were variable in cross-sectional shape, length and other details. Diplodocoids, for example, were relatively narrow, deep-bodied and with a tall ridge formed from their neural spines running along the back, while titanosaurs were extremely broad across the hips and must have been just about flat across the back. The thorax may have sloped down ever so slightly in diplodocoids (in part because their forelimbs were shorter than their hindlimbs) whereas the thorax was angled upwards slightly or even markedly in some macronarians, like brachiosaurs and some titanosaurs.

Articulated skeletons show that the tail mostly projected horizontally from the pelvis (a subtle arch at the tail base is present in diplodocoids and some others, in fact), but the macronarians with the upward-sloping bodies also had a downward-sloping tail. A real curiosity is provided by the tails of some mamenchisaurs which seem to have projected upwards at an angle. This has been discussed and illustrated by Paul (2010) and more recently by Hallett & Wedel (2016), but has otherwise gone undiscussed as far as I know. It sounds so odd that surely some mistake has been made… though I really don’t think it has.

Tall neural spines, massive, wing-shaped transverse processes and other structures show that an enormous quantity of musculature would have been obvious along the proximal part of the sauropod tail. This is the tail of … ugh .. Dippy, the cast of  Diplodocus carnegii  until recently on show in London. Image: Darren Naish.

Tall neural spines, massive, wing-shaped transverse processes and other structures show that an enormous quantity of musculature would have been obvious along the proximal part of the sauropod tail. This is the tail of … ugh .. Dippy, the cast of Diplodocus carnegii until recently on show in London. Image: Darren Naish.

As is typical for non-bird dinosaurs, the musculature at the base of the tail was (so we can say from the relevant bony attachment points) evidently enormous and bulky, the proximal part of the tail likely being similar in width to the pelvis and thighs combined.

The integument. We know for definite that sauropods of all groups were scaly animals, since scaly skin impressions and actual preserved skin patches are known for diplodocids, Camarasaurus and titanosaurs at least. A few skin folds here and there were almost certainly present (say, where the limbs met the body, at the limb joints, and at mobile zones in the neck) but a fissured, wrinkled or scored elephant-like skin is a big fat no and every effort should be made to avoid it in artistic depictions.

Haestasaurus  from the English Wealden (shown at left) and a few other sauropods preserve polygonal scales that vary somewhat in size. Image: Darren Naish, Czerkas (1994).

Haestasaurus from the English Wealden (shown at left) and a few other sauropods preserve polygonal scales that vary somewhat in size. Image: Darren Naish, Czerkas (1994).

Polygonal – specifically, pentagonal, hexagonal and heptagonal – scales are known for the macronarian Haestasaurus and some diplodocids, while rounded scales possessing a papilliform texture of tiny bumps are also preserved in a diplodocid specimen (Czerkas 1994). The polygonal structures varied in size somewhat, those present in more mobile parts of the skin (say, the inner crease of the elbow) being smaller than those present in non-mobile sections. Even so, the largest scales were, at most, 60 mm across, which is not large at all on an animal more than 15 m long. This is pretty common in non-bird dinosaurs, by the way. The scales were often so small that they wouldn’t be readily visible from any distance greater than a few metres. The papilliform texture on those diplodocid scales seems to have been widespread across sauropods and means that sauropod skin would have been rough to the touch.

A diplodocid specimen from Wyoming that has scaly skin preserved also preserves tall, conical dermal spines (note: they are not horn-covered bony structures) that must have been arranged along the dorsal midline (Czerkas 1992). These are variable in height (the biggest are 18 cm tall) and seem to have formed a single row along the top of the tail’s proximal part at least. This specimen was originally implied to belong to Diplodocus but is of indeterminate identity. It might belong to Kaatedocus, since remains of this dinosaur come from the exact same quarry.

Large, laterally compressed conical and semi-conical dermal structures lined the upper surface of the tail (at least) in some diplodocids, and perhaps in other diplodocoids and sauropods too. Image: Czerkas (1994).

Large, laterally compressed conical and semi-conical dermal structures lined the upper surface of the tail (at least) in some diplodocids, and perhaps in other diplodocoids and sauropods too. Image: Czerkas (1994).

The conical and semi-conical dermal spines of some diplodocids were variable in height, breadth and shape, as illustrated here. Image: Czerkas (1994).

The conical and semi-conical dermal spines of some diplodocids were variable in height, breadth and shape, as illustrated here. Image: Czerkas (1994).

Were these structures present across all diplodocids, all diplodocoids, all neosauropods or even all sauropods, or were they exclusive to one small clade, perhaps even just to Kaatedocus itself? In the absence of further information we can’t say, but it’s appropriate to depict them on diplodocids and their close kin at least, in the absence of further information. The possibility that shorter and/or taller dermal structures of this sort were present elsewhere on other sauropods exists.

The discovery of those conical and semi-conical spines in a diplodocid led Stephen Czerkas to produce this new look for these dinosaurs. It might be correct, but it is not clear whether the structures were as extensive along the animal’s length as shown here. Image: Czerkas (1992).

The discovery of those conical and semi-conical spines in a diplodocid led Stephen Czerkas to produce this new look for these dinosaurs. It might be correct, but it is not clear whether the structures were as extensive along the animal’s length as shown here. Image: Czerkas (1992).

Horn-covered osteoderms - that is, lumps and nodules with a bony core - were arranged across the backs and flanks of some titanosaurs, specifically the members of the clade Lithostrotia. These structures were variously rounded or oval, sometimes quite flat and sometimes slightly or strongly domed. They appear to have been numerous and prominent in some of the relevant species and would have given their backs and sides a pebbly, armoured appearance.

Colours. There’s a traditional view stating that big dinosaurs were likely grey and plain because big living mammals are, as is the Komodo dragon, I suppose. I’m going to be bold here and say that there’s no reason whatsoever to take any notice of this. Sauropods were not mammals or Komodo dragons. While an argument could be made that camouflage would have been beneficial, and that certain pigments and patterns would have been helpful or necessary for physiological reasons (heat-shedding, heat retention, UV protection and so on), bold and complex patterns and bright, even vibrant colours are all consistent with the ecophysiological demands of being a sauropod, and the good colour vision and complexity of sauropod integument, and likely reliance on visual display, mean that they very likely could have been more like gigantic lizards or birds than elephants. In any case, giraffes, perenties and others show that big, terrestrial animals don’t have to be bland and grey as has often been stated.

The idea that big terrestrial animals have to be bland never was correct. Perenties  Varanus giganteus  might not be that colourful, but they help emphasise the possibility that striking patterns can be present even in very large terrestrial reptiles. Image: (c) Stephen Zozaya.

The idea that big terrestrial animals have to be bland never was correct. Perenties Varanus giganteus might not be that colourful, but they help emphasise the possibility that striking patterns can be present even in very large terrestrial reptiles. Image: (c) Stephen Zozaya.

In the absence of any direct evidence, it seems reasonable to me to reconstruct barred, striped, reticulated or dappled colour schemes, to depict bright colours on faces, necks or other areas considered relevant to display, and to imagine any colours reasonable for big, terrestrial animals associated with woodlands, parks, scrubby places, mangroves and all the other habitats frequented by this long-lived, diverse group of animals.

Want to know more about sauropod life appearance? It sometimes surprises people that there are scarcely any good books dedicated to specific Mesozoic dinosaur groups. For sauropods, the main must-have volume is   Mark Hallett and Matt Wedel’s  The Sauropod Dinosaurs   . It’s really good.    Dinosaurs Past and Present Vol II    is also worth getting, in part because it includes Greg Paul’s (now quite dated, but still useful) article on dinosaur life appearance. Images: amazon ( here  and  here ).

Want to know more about sauropod life appearance? It sometimes surprises people that there are scarcely any good books dedicated to specific Mesozoic dinosaur groups. For sauropods, the main must-have volume is Mark Hallett and Matt Wedel’s The Sauropod Dinosaurs. It’s really good. Dinosaurs Past and Present Vol II is also worth getting, in part because it includes Greg Paul’s (now quite dated, but still useful) article on dinosaur life appearance. Images: amazon (here and here).

If we really want to imagine sauropods as living animals, there is – of course – so much more to say. Body language, posture, gait, social life, anti-predator responses, feeding behaviours, digestive function, sleep and so much more are all things that need to be considered. But that’s where I’ll stop for now. I hope you found this interesting, and we’ll be looking at dinosaur life appearance again sometime soon.

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

For previous TetZoo articles on sauropods, see…

 Refs - -

Apesteguía, S. 2004. Bonitasaura salgadoi gen. et sp. nov.: a beaked sauropod from the Late Cretaceous of Patagonia. Naturwissenschaften 91, 493-497.

Bakker, R. T. 1986. The Dinosaur Heresies. New Theories Unlocking the Mystery of Dinosaurs and their Extinction. William Morrow, New York.

Czerkas, S. A. 1992. Discovery of dermal spines reveals a new look for sauropod dinosaurs. Geology 20, 1068-1070.

Czerkas, S. A. 1994. The history and interpretation of sauropod skin impressions. Gaia 10, 173-182.

Falkingham, P., Bates, K. & Farlow, J. 2014. Historical photogrammetry: Bird’s Paluxy River dinosaur chase sequence digitally reconstructed as it was prior to excavation 70 years ago. PLoS ONE 9, 4: e93247.

Hallett, M. & Wedel, M. J. 2016. The Sauropod Dinosaurs: Life in the Age of Giants. Johns Hopkins University Press, Baltimore.

Knoll, F., Galton, P. M. & López-Antoñanzas, R. 2006. Paleoneurological evidence against a proboscis in the sauropod dinosaur Diplodocus. Geobios 39, 215-221.

Martin, J. 1987. Mobility and feeding of Cetiosaurus (saurischia, sauropoda [sic]) - why the long neck? In Currie, P. J. & Koster, E. H.(eds) Fourth Symposium on Mesozoic Terrestrial Ecosystems, Short Papers. Boxtree Books (Drumheller, Alberta), pp. 154-159.

Martínez, R. D. F., Lamanna, M. C., Novas, F. E., Ridgely, R. C., Casal, G. A., Martínez, J. E., Vita, J. R. & Witmer, L. M. 2016. A basal lithostrotian titanosaur (Dinosauria: Sauropoda) with a complete skull: implications for the evolution and paleobiology of Titanosauria. PLoS ONE 11, 4: e0151661. 

Meyer, C. A., Lockley, M. G., Robinson, J. W. & dos Santos, V. F. 1994. A comparison of well-preserved sauropod tracks from the Late Jurassic of Portugal and the western United States: evidence and implications. Gaia 10, 57-64.

Milàn, J., Christiansen, P. & Mateus, O. 2005. A three-dimensionally preserved sauropod manus impression from the Upper Jurassic of Portugal: implications for sauropod manus shape and locomotor mechanics. Kaupia 14, 47-52.

Paul, G. S. 1987. The science and art of restoring the life appearance of dinosaurs and their relatives - a rigorous how-to guide. In Czerkas, S. J. & Olson, E. C. (eds) Dinosaurs Past and Present Vol. II. Natural History Museum of Los Angeles County/University of Washington Press (Seattle and London), pp. 4-49.

Paul, G.S. 1998. Terramegathermy and Cope’s rule in the land of titans. Modern Geology 23, 179-217.

Paul, G. S. 2010. Dinosaurs: A Field Guide. A & C Black, London.

Russell, D. A. 1987. Models and paintings of North American dinosaurs. In Czerkas, S. J. & Olson, E. C. (eds) Dinosaurs Past and Present, Volume I. Natural History Museum of Los Angeles County/University of Washington Press (Seattle and Washington), pp. 114-131.

Senter, P. 2007. Necks for sex: sexual selection as an explanation for sauropod dinosaur neck elongation. Journal of Zoology 271, 45-53.

Taylor, M. P., Hone, D. W. E., Wedel, M. J., & Naish, D. 2011. The long necks of sauropods did not evolve primarily through sexual selection. Journal of Zoology 285, 150-161.

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

Tschopp, E., Wings, O., Frauenfelder, T. & Brinkmann, W. 2015. Articulated bone sets of manus and pedes of Camarasaurus (Sauropoda, Dinosauria). Palaeontologia Electronica 18.2.44A: 1-65.

Witmer, L. M. 2001. Nostril position in dinosaurs and other vertebrates and its significance for nasal function. Science 293, 850-853.

Witton, M. P. 2018. The Palaeoartist’s Handbook: Recreating Prehistoric Animals in Art. The Crowood Press, Marlborough.

Heilmann, Thompson, Beebe, Tetrapteryx and the Proavian

Many people familiar with ideas on the early evolution of birds and of bird flight will know of the Proavis or proavian, a hypothetical bird ancestor illustrated and discussed by William Pycraft (1868-1942) in 1906 but made better known by Gerhard Heilmann (1859/1861-1946) during the 1910s and 20s. Versions of this creature were later illustrated in colour by Zdeněk Burian for popular books of the 1960s, 70s and 80s. In the article that you’re about to read, we’re predominantly interested in Heilmann’s take on the Proavis.

Fighting, gliding and climbing proavians, depicted in a cluttered woodland environment, by Gerhard Heilmann. This is typical of Heilmann’s many excellent scenes. He used posed plaster models of the creatures before drawing. This illustration was used in the 1916 Danish version of the book (it first appeared in one of his 1912 articles) but does not appear in the 1926 English version. As we’ll see below, the decision to exclude some images from the 1926 version might have served to make it seem more scholarly than its predecessor. Image: Heilmann (1916).

Fighting, gliding and climbing proavians, depicted in a cluttered woodland environment, by Gerhard Heilmann. This is typical of Heilmann’s many excellent scenes. He used posed plaster models of the creatures before drawing. This illustration was used in the 1916 Danish version of the book (it first appeared in one of his 1912 articles) but does not appear in the 1926 English version. As we’ll see below, the decision to exclude some images from the 1926 version might have served to make it seem more scholarly than its predecessor. Image: Heilmann (1916).

Heilmann was an artist and graphic designer by profession but his interest in bird evolution and anatomy was such that he published a series of articles on the subject for the Danish Ornithological Society between 1912 and 1916 (Nieuwland 2004, Ries 2007).

Self-portrait of Gerhard Heilmann, produced in 1902. Heilmann was an argumentative man often in conflict with “various forms of authority”, and at loggerheads with his own family   (Nieuwland 2004)  . This illustration was included in     Nieuwland (2004)  .

Self-portrait of Gerhard Heilmann, produced in 1902. Heilmann was an argumentative man often in conflict with “various forms of authority”, and at loggerheads with his own family (Nieuwland 2004). This illustration was included in Nieuwland (2004).

Compiled, these articles described Heilmann’s take on the diversity of ancient fossil birds, embryological development in birds and reptiles, and bird anatomy, with the last in the series explaining what these data meant for the appearance and lifestyle of his Proavis, an imagined animal older and anatomically more archaic than Archaeopteryx. These articles were published together in a Danish-language book Vor Nuvaerende Viden om Fuglenes Afstamning (Heilmann 1916), the title meaning Our Current Knowledge of the Descent of Birds.

Heilmann’s The Origin of Birds. This body of work was sufficiently impressive that – while Heilmann was mostly shunned in Denmark – it was well received elsewhere, and he was encouraged by scientists in Germany in particular to translate it into English. The result was the 1926 The Origin of Birds (republished in the USA in 1927), a scholarly and well-illustrated book that was well received internationally and almost immediately deemed the most authoritative work on bird origins. The 1926 volume is quite different from that of 1916 in the illustrations it includes (as we’ll see below), but also in how scientific and speculative the text is, some more fanciful sections on palaeobiology being absent from the English version.

It has been said several times that Heilmann’s work was deemed so impressive that, rather than inspire new work on the subject, it effectively bought research on bird origins to a halt. Most relevant workers (albeit not all) now considered the question of bird origins to be resolved. As noted by historian Ilja Nieuwland, things might not have gone this way had the English-speaking readers of Heilmann’s book known that he was an amateur scientist and an artist by trade (Palm 1997, Nieuwland 2004).

Heilmann’s draftmanship was superb. This illustration (fig. 140 from   Heilmann 1926  ) shows thigh feathering on the chicks of various birds, with 8 being a gliding gecko. “The animal with which we may best compare the bird-ancestor, is the Fringed gecko” [sic] (p. 197). Image:   Heilmann (1926)  .

Heilmann’s draftmanship was superb. This illustration (fig. 140 from Heilmann 1926) shows thigh feathering on the chicks of various birds, with 8 being a gliding gecko. “The animal with which we may best compare the bird-ancestor, is the Fringed gecko” [sic] (p. 197). Image: Heilmann (1926).

Heilmann thought that bird ancestors were likely quadrupedal gliders, similar superficially to living gliding geckos (Heilmann 1926, p. 197). This statement is somewhat odd in view of his endorsement elsewhere of a more cursorial view of these animals, but it seems that he imagined them adopting very different poses when on the ground versus when climbing. Heilmann also argued that birds were not dinosaurs – despite his good understanding of the compelling anatomical similarity between theropod dinosaurs and birds – but were instead the descendants of a group closely related to – and supposedly ancestral to – dinosaurs, termed ‘pseudosuchians’*. This was because of his adherence to ‘Dollo’s Law’ – the (erroneous) idea that a lost anatomical structure cannot be regained – and his mistaken contention that dinosaurs lacked clavicles. If dinosaurs couldn’t be ancestral to birds, the true ancestors must have been another group, hence Heilmann’s use of the more archaic ‘pseudosuchians’, already mooted as possible bird ancestors by Robert Broom in his 1913 description of the South African Euparkeria.

* The term pseudosuchian is today applied to the archosaur lineage that includes crocodylians and all of their extinct relatives. Given the historical baggage that comes with the term, I personally don’t think that this is a good idea at all and would prefer it if another name were used for the lineage concerned (like Crurotarsi)… but that’s an issue for another time.

Euparkeria capensis  has long been intimated as a sort of bird ancestor by those looking for such creatures outside of theropod dinosaurs. Not only is this animal only very distantly related to birds, it’s not even part of the crown-archosaur clade. Image: Taenadoman, CC BY-SA 3.0 ( original here ).

Euparkeria capensis has long been intimated as a sort of bird ancestor by those looking for such creatures outside of theropod dinosaurs. Not only is this animal only very distantly related to birds, it’s not even part of the crown-archosaur clade. Image: Taenadoman, CC BY-SA 3.0 (original here).

By combining features common to Archaeopteryx as well as to the ‘pseudosuchians’ Aetosaurus, Euparkeria, Ornithosuchus and Saltoposuchus, Heilmann (1916, 1926) described how he invented a creature that looked somewhat like a theropod but had a more archaic skull and foot, retained a fourth metacarpal, and had a smaller pelvis with far shorter pubic and ischial bones.

Heilmann’s skeletal reconstruction of Proavis, as depicted in the 1916 Danish version of the book… but not in the 1926 English version. Note the lack of long feathers on the hindlimbs. Incidentally, note also that Heilmann was partly responsible for encouraging the belief that forelimb feathers did not cover the hands during the earliest stages of bird evolution. Image: Heilmann (1916).

Heilmann’s skeletal reconstruction of Proavis, as depicted in the 1916 Danish version of the book… but not in the 1926 English version. Note the lack of long feathers on the hindlimbs. Incidentally, note also that Heilmann was partly responsible for encouraging the belief that forelimb feathers did not cover the hands during the earliest stages of bird evolution. Image: Heilmann (1916).

Heilmann, Thompson and D’Arcy Thompson grids. It’s a matter of some interest that Heilmann didn’t just guess what the imagined morphology of the proavian would be, as might be assumed given the English edition of his book. Instead, he used a technique whereby the relevant anatomical regions were mapped on to a grid and then distorted to mimic the evolutionary process. This grid-based deformation process was pioneered by D’Arcy Wentworth Thompson (1860-1948) and was explained most thoroughly in his 1917 book On Growth and Form (Thompson 1917). It was actually put forward beforehand in 1915 (Thompson 1915).

Thompson applied his Cartesian grid technique to many animal lineages, his aim being to show that mathematically predictable transformation in one or more anatomical regions could result in profound anatomical change. Here’s one of my favourite examples: how you can derive a molid sunfish ( Mola  was known to Thompson as  Orthagoriscus ) from a porcupinefish. Image: Thompson (1917).

Thompson applied his Cartesian grid technique to many animal lineages, his aim being to show that mathematically predictable transformation in one or more anatomical regions could result in profound anatomical change. Here’s one of my favourite examples: how you can derive a molid sunfish (Mola was known to Thompson as Orthagoriscus) from a porcupinefish. Image: Thompson (1917).

By placing grids on top of diagrams of related animals (say, the skull of a human and a chimp), Thompson showed how the skewing of the grid in a certain direction “would result in the sort of changes that would allow the emergence of a new species” (Naish 2017, p. 116). These grids are generally termed D’Arcy Thompson transformation grids, Cartesian transformations or Cartesian grids, and their use was quite popular in the evolutionary literature of the early 20th century. Heilmann was inspired to use the technique after Thompson wrote to him about bird evolution in 1915, and the two wrote to each other on many occasions about the technique and its application to ideas on the evolution of horses, hominids and birds. They also discussed how the resulting illustrations could be used in Thompson’s On Growth and Form (Ries 2007).

Heilmann included these Cartesian transformations in the 1916 version of his book, but they weren’t included in the 1926 English version. His take on Proavis was not, therefore, simple guesswork. Image: this montage is from Witmer (1991) but the originals are from Heilmann (1916).

Heilmann included these Cartesian transformations in the 1916 version of his book, but they weren’t included in the 1926 English version. His take on Proavis was not, therefore, simple guesswork. Image: this montage is from Witmer (1991) but the originals are from Heilmann (1916).

Heilmann’s commitment to the technique is demonstrated by the fact that the 1916 Danish edition of his book includes grids that depict the inferred evolution of the avian skull, forelimb and pelvis (I don’t own a copy of the Danish edition, but the diagrams are included in both Witmer (1991) and Ries (2007)). With Ornithosuchus and Euparkeria as ‘starting points’ and Archaeopteryx and modern birds as ‘end points’*, Heilmann used the grids to create relevant intermediates, the results allowing him to generate, piecemeal, his Proavis (Heilmann 1916). A lateral view of the hypothetical animal – showing it in a neural, non-dynamic walking pose – was included in the Danish edition (Heilmann 1916, Ries 2007).

* It should be noted that Heilmann’s Archaeopteryx was not accurate, some of its details (in the skull especially) being semi-hypothetical and informed by Heilmann’s interpretation of ‘pseudosuchians’ like Aetosaurus.

The more dynamic, climbing version of Heilmann’s Proavis skeletal reconstruction, the only version of the reconstruction included in the 1926 English version of his book. Image:   Heilmann (1926)  .

The more dynamic, climbing version of Heilmann’s Proavis skeletal reconstruction, the only version of the reconstruction included in the 1926 English version of his book. Image: Heilmann (1926).

Remarkably, none of this was included in the English edition at all, this creating the impression that Heilmann simply invented the proavian via intuition and guesswork. He didn’t: it was generated via a testable, repeatable method (albeit using flawed anatomical data). Furthermore, the fact that Heilmann’s skeletal reconstruction of Proavis included in the English edition (Heilmann 1926) is the more dynamic climbing version of the creature gives it a less scientific, more speculative, artistic air than the walking version of 1916.

Heilmann, Beebe and Tetrapteryx. When it came to feathering and other aspects of the integument, Heilmann noted his thoughts on another Proavis-type animal, this time the one invented by Charles William Beebe in 1915. Beebe (1877-1962) was an interesting person. He was a naturalist, ecologist, ornithologist, marine biologist, author and explorer, among other things. Today, he might be best known for the deep-water observations he made while in a bathysphere off the coast of Nonsuch Island off Bermuda during the 1930s.

William Beebe’s hypothetical tetrapteryx creature. A pre- Archaepteryx  glider. You’ll note that Beebe was a pretty good artist [UPDATE: this is an error. The illustrations in this publication were by Dwight Franklin, not by Beebe!]. Image: Beebe (1915).

William Beebe’s hypothetical tetrapteryx creature. A pre-Archaepteryx glider. You’ll note that Beebe was a pretty good artist [UPDATE: this is an error. The illustrations in this publication were by Dwight Franklin, not by Beebe!]. Image: Beebe (1915).

When it came to bird origins, Beebe thought that birds originated via a ‘tetrapteryx’ phase where large feathers on the proximal hindlimb were present and functioned in slowing descent during leaping and gliding (Beebe 1915). Both the fossils of Archaeopteryx and the nestlings of living bird species demonstrated the antiquity of these ‘pelvic wings’, Beebe (1915) said. Heilmann (1926) disagreed. His own examination of bird nestlings pertaining to bird lineages across the family tree resulted in his “complete disappointment, for what I found was wholly negative; there was not in any of them the slightest trace of a “pelvic wing”” (p. 194).

As for Archaeopteryx, Heilmann (1926) argued that the hindlimbs were arranged such that they could never have had an aerodynamic function, nor was there evidence for long hindlimb feathering of the sort consistent with ‘pelvic wings’. Ergo, Heilmann illustrated Archaeopteryx without long hindlimb feathering, and he didn’t think that Proavis had them either: his skeletal reconstructions (both the 1916 walking version and 1926 climbing one) lack them (the 1926 climber has short hindlimb feathers, similar in length to those elsewhere on the body) and the gliding proavians included in his various drawings clearly lack them as well. His most famous proavian figure – the semi-erect one standing in a conifer tree (shown below) – does have long feathers on the rear edge of the thigh, though they certainly aren’t as long as they should be as per Beebe’s tetrapteryx model.

The iconic tree-dwelling proavian. The iconic tree-dwelling proavian I have in mind here is one of Heilmann’s best-known and most frequently reproduced illustrations. It has what appears to be great symbolic significance if you’ve read the English version of The Origin of Birds since it’s the very last image in the entire book (Heilmann 1926, p. 199), thus appearing as a visual metaphor for an imagined phrase: “Finally, I can reconstruct the real ancestor of birds, and it looked like this”.

Heilmann’s most iconic version of his Proavis. Note the long feathers on the back of the thigh and how they’re invisible and thus not deemed important in the gliding individual in the background. Heilmann said that the long feathers close to the base of the tail would produce the required lift in this region. Image:   Heilmann (1926)  .

Heilmann’s most iconic version of his Proavis. Note the long feathers on the back of the thigh and how they’re invisible and thus not deemed important in the gliding individual in the background. Heilmann said that the long feathers close to the base of the tail would produce the required lift in this region. Image: Heilmann (1926).

I was surprised to learn that Beebe produced what looks like his own version of this illustration, and inserted it as a plate in a book published in 1915. However, the image itself is not dated 1915 and does not definitely pre-date Heilmann’s illustration. My suspicion – and that of Paul Stewart, who kindly brought my attention to the image – is that Beebe copied Heilmann’s proavian for his own personal use (the illustration was not intended for publication*). This in itself is interesting since it could mean that Beebe personally endorsed Heilmann’s view of Proavis and regarded it as a ‘descendant’ of his tetrapteryx creature… which it wasn’t, given that Heilmann rejected Beebe’s idea, as we saw earlier. Again, my thanks to Paul Stewart for sharing this image with me and permitting its use here.

* Thanks to Jonathan Kane, I’ve learnt that it was previously published in Tim Berra’s 1977 William Beebe: An Annotated Bibliography.

William Beebe seems to have produced his own take on Heilmann’s iconic proavian scene. We’re presuming that it was produced some time after Heilmann’s illustration was, but this hasn’t yet been fully confirmed. The illustration appears here courtesy of Paul Stewart.

William Beebe seems to have produced his own take on Heilmann’s iconic proavian scene. We’re presuming that it was produced some time after Heilmann’s illustration was, but this hasn’t yet been fully confirmed. The illustration appears here courtesy of Paul Stewart.

Today, the significance of Beebe’s tetrapteryx idea is a bit uncertain and, frankly, depends on who you ask. Archaeopteryx probably did have long feathers on the hindlimbs (Longrich 2006), and they perhaps had an aerodynamic role if this animal indulged in aerial locomotion (which it likely did). Furthermore, the discovery of prominent ‘hindlimb wings’ in other feathered dinosaurs (most notably the Chinese dromaeosaur Microraptor) has led some experts to note the predictive power of Beebe’s suggestion (Kane et al. 2016) and to favour the presence and importance of hindlimb feathers in the earliest birds (Longrich 2006, Chatterjee & Templin 2007, Zheng et al. 2013, Xu et al. 2014). However, it’s not at all certain that large, aerodynamically ‘functional’ hindlimb feathers were present consistently across the lineages concerned or that they had the role that Beebe imagined (O’Connor & Chang 2015).

Recent work shows that  Archaeopteryx  really did have long feathers on the proximal portions of its hindlimbs (these reconstructions are by Longrich (2006)), though they weren’t as long as the feathers Beebe imagined for his tetrapteryx stage creature. Image: Longrich (2006).

Recent work shows that Archaeopteryx really did have long feathers on the proximal portions of its hindlimbs (these reconstructions are by Longrich (2006)), though they weren’t as long as the feathers Beebe imagined for his tetrapteryx stage creature. Image: Longrich (2006).

As always, there’s more to say and this article is already longer than intended. I also wanted to talk about Pycraft’s initial concept of the Proavis, and also Burian’s depiction, since he didn’t simply copy Heilmann’s illustrations but actually produced yet another hypothetical creature. These things will have to wait to another time. Also worthy of further discussion is the point made earlier about Heilmann’s influence happening despite his status as a ‘mere’ artist, a theme that has parallels elsewhere in the history of vertebrate palaeontology and is relevant to my writings on meme perpetuation, the scientific acceptance of feathering in non-bird dinosaurs and much else besides. I aim to explore these topics (and others) in future articles.

Before Heilmann, there was Pycraft’s proavian of 1906. We’ll have to discuss this creature and its backstory another time. Image:   Pycraft (1910)  .

Before Heilmann, there was Pycraft’s proavian of 1906. We’ll have to discuss this creature and its backstory another time. Image: Pycraft (1910).

On which note, please consider supporting this blog at patreon if you don’t already do so. The more support I receive, the more time I can spend generating new content. Thanks so much.

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

Refs - -

Beebe, C. W. 1915. A tetrapteryx phase in the ancestry of birds. Zoologica 2, 38-52.

Chatterjee, S. & Templin, R. J. 2007. Biplane wing planform and flight performance of the feathered dinosaur Microraptor gui. Proceedings of the National Academy of Sciences 104, 1576-1580.

Heilmann, G. 1916. Vor Nuvaerende Viden om Fuglenes Afstamning. Unknown publisher, Copenhagen.

Heilmann, G. 1926. The Origin of Birds. Witherby, London.

Kane, J., Willoughby, E. & Keesey, T. M. 2016. God’s Word or Human Reason? An Inside Perspective on Creationism. Inkwater Press.

Longrich, N. 2006. Structure and function of hindlimb feathers in Archaeopteryx lithographica. Paleobiology 32, 417-431.

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

Nieuwland, I. J. J. 2004. Gerhard Heilmann and the artist’s eye in science, 1912-1927. www.PalArch.nl., vertebrate palaeontology 3, 2.

O’Connor, J. & Chang, H. 2015. Hindlimb feathers in paravians: primarily “wings” or ornaments? Biology Bulletin 42, 616-621.

Palm, S. 1997. The Origin of Flapping Flight in Birds. Svend Plam, Ballerop.

Pycraft, W. P. 1910. A History of Birds. Methuen & Co, London.

Ries, C. J. 2007. Creating the Proavis: bird origins in the art and science of Gerhard Heilmann 1913-1926. Archives of Natural History 34, 1-19.

Thompson, D. W. 1915. Morphology and mathematics. Transactions of the Royal Society of Edinburgh 50, 857-895.

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Xu, X., Zhou, Z., Dudley, R., Mackem, S., Chuong, C.-M., Erickson, G. M. & Varricchio, D. J. 2014. An integrative approach to understanding bird origins. Science 346 (6215), 1253293.

Zheng, X., Zhou, Z., Wang, X., Zhang, F., Zhang, X., Wang, Y., Wei, G., Wang, S. & Xu, X. 2013. Hind wings in basal birds and the evolution of leg feathers. Science 339, 1309-1312.

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.

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.

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 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.

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.

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).

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 ).

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  ).

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 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.

TetZooniverse-Christmas-2018-tiny-from-Darren-Naish.jpg

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

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.

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.

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.

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 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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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!

CP-Sept-2018-Darren-and-Iguanodon-1000-px-tiny-Dec-2018-Darren-Naish-Tetrapod-Zoology.jpg
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.

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…

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.

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.

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.

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.

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.

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.

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.

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.

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

Amphibians

Mammals

Turtles

Mesozoic marine reptiles

Pterosaurs

Non-avialan dinosaurs

Birds

Lepidosaurs

Cryptozoology

A graph...

Image: Darren Naish.

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.

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...

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).

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 ).

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.

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).

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).

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).

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 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.

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).

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).

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.

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.

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).

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.

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 ).

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.

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

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.

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 .

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.

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

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…

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

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.

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.

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.

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

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.

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

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.

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

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.

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.

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…

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 ).

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Refs - -

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

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

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

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

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

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

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

There will be books.

There will be books.

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

Books. Image: Darren Naish.

Books. Image: Darren Naish.

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

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

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

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

Brian Switek’s My Beloved Brontosaurus: A Belated Review

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

For previous TetZoo articles relevant to this one, see…

Refs - -

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

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

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

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

New Living Animals We Want to Find

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

There are all kinds of 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…

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

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

Refs - -

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

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

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

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

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

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

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

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

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

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

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

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

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

Whatever Happened to the Kabomani Tapir?

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Refs - -

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

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

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

Ruiz-García, M., Castellanos, A., Agueda Bernal, L., Navas, D., Pinedo-Castro, M. & Mark Shostell, J. 2015. Mitochondrial gene diversity of the mega-herbivorous species of the genus Tapirus (Tapiridae, Perissodactyla) in South America and some insights on their genetic conservation, systematics and the Pleistocene influence on their genetic characteristics. Advances in Genetic Research 14, 1-51.

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

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

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

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

Avocets in Flight and Phylogeny

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

The modern RSPB logo. Image: fair use.

The modern RSPB logo. Image: fair use.

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

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

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

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

Refs - -

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

  • The multi-part guide to Australian agamid lizards

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

  • Domestic horses of Africa

  • Raymond Hoser smackdown

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

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

  • On why humans and other apes are actually monkeys

  • No trunks for sauropods

  • Ornithoscelida debut

  • Piltdown man and the dualist connection

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

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

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

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

TetZooCon 2018: Best TetZooCon So Far

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

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

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

Indeed, TetZooCon 2018 was ambitious – possibly over-ambitious. In addition to a long list of talks, the event included a Palaeoart Workshop, a special session on talks devoted entirely to bird evolution, a bird evolution roundtable event, a Speculative Biology on-stage discussion, a quiz, a conference meal and drinks reception, and a post-conference fieldtrip. Abundant stalls with merchandise were there too; we also had numerous book signings and a few ‘show and tell’ events (relating to Dougal Dixon’s SpecBio projects and my in-prep textbook, among other things). For the second year running, Beth Windle brought along a TetZoo-themed cak