Postcranial Palaeoneurology and the Lifestyles of Pterosaurs

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

My technical research and my writing here at the blog continues with your kind support via patreon. Many thanks to those who assist my projects. Please consider assisting if you can. The more independence I achieve, the more time I can spend producing the content you enjoy.

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

• Terrestrial stalking azhdarchids, the paper, May 2008

• Shemhazai and other flightless pterosaurs, September 2008

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

• Giant pterosaurs invade London, Summer 2010, July 2010

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

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

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

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

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

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

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

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

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

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

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

Refs - -

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

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

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

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

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

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

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

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

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

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

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

Witton, M. P. & Naish, D. 2015. Azhdarchid pterosaurs: water-trawling pelican mimics or “terrestrial stalkers”? Acta Palaeontologica Polonica 60, 651-660.

A 1996 Letter from Dr Phil Currie

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

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

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

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

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

Image: Darren Naish.

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

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

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

Ref - -

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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New Dinosaur Books, Part 1: Barrett on Stegosaurs

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

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

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

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

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

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

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

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

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

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

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

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

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

Refs - -

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

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

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

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

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

Bigfoot’s Genitals: What Do We Know?

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

• Cryptozoology at the Zoological Society of London. Cryptozoology: time to come in from the cold? Or, Cryptozoology: avoid at all costs?

• The Cryptozoologicon (Volume I): here, at last

• Is Cryptozoology Good or Bad for Science? (review of Loxton & Prothero 2013)

• My New Book Hunting Monsters: Cryptozoology and the Reality Behind the Myths

• If Bigfoot Were Real

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

Refs - -

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

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

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

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

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

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

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

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

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

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

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

Welcome to Tetrapod Zoology ver 4

Hello and welcome to the very first article of Tet Zoo ver 4, and big thanks if you're here because you followed the link at the final ver 3 article. Yes, it’s happened – I could no longer stay at Scientific American and have had to move and go independent. More on that in a minute.

A really old banner I prepared for Tet Zoo ver 2 and used there for a while. It features various items specially relevant at the time (May 2011), much of it involving Wealden theropods (on which more soon). Image: Darren Naish.

After numerous discussions and deliberations, I’ve decided that the best option for the continuation of Tet Zoo is to host it here, at tetzoo.com, and thus to have it sharing the same platform as the podcast. Given that blog articles are very swiftly going to outnumber podcast episodes (there are only 69 of the latter at the time of writing), it won’t be long before this site becomes obvious as the home of the blog, not just the home of the podcast. And if, so far, you’ve only listened to the podcast and never read the blog, well, now maybe you’ll become better acquainted with the blog. I should note that this site will likely get a redesign in time so that it looks more blog-themed.

A Rhacodactylus gecko climbing on a copy of Tetrapod Zoology Book One, of course. One day I'll be able to publish the follow-up volumes to that book. Image: Ethan Kocak.

If you’re new here, what do I cover? Tetrapod Zoology features longish and semi-technical articles – reviews, discussions, musings and such – devoted to tetrapods of all sorts (amphibians, reptiles, birds, mammals, living and extinct, and all of their extinct relatives). Popular topics include dinosaurs, obscure evolutionary models, interesting and little-known aspects of animal behaviour, conservation biology and cryptozoology. I also write about relevant books and toys when the time is right. I mostly avoid writing about ‘newsy‘ stories, only because they get covered at a million other places and writing about them typically feels unoriginal and predictable. Having said that, I will cover newsy things when I feel I have stuff to stay that isn’t being covered elsewhere.

I approve of fan-art. I'm not sure if this counts (it's from a Christmas card for the Centre for Fortean Zoology, designed by Mark North), but it's a worthy addition. I'm playing the role of the 'Darrenadon' in an on-stage version of The Lost World. Image: Mark North.

Community-relevant things – announcements relevant to our annual TetZooCon and other appropriate conferences and meetings – also get covered here. On that note, as a denizen of the TetZooniverse (such as it’s known), do consider coming along to TetZooCon if you can, it’s great. The 2018 meeting is happening on the 6th and 7th of October at The Venue, Malet Street, London, and is by far the biggest so far. Go here for more information on the meeting, and on those of previous years.

Tapirs. Tet Zoo loves tapirs. Image: Darren Naish.

A Quick History of Tet Zoo. For those who don’t know, Tetrapod Zoology started life on January 21st 2006 when, while unable to sleep at 1am (I was in the final throes of writing my PhD thesis at the time), I got up to eat biscuits and start a blog. I was inspired by the blog-writing of one of my longest-standing friends, but I’ve opted to keep secret who that is in order to minimise the impact on his ego. Tetrapod Zoology was launched at blogspot, the first article ever being a hello and the second being devoted to the topic of megafaunal predation in eagles. The site went from strength to strength, quickly building up a good community and a large readership.

Screengrab of part of the first ever 'proper' Tet Zoo article, the killer eagles one of January 2006. That weird photo of me holding a bottle was taken at a conference held at the Natural History Museum, London; I think by Richard Forrest.

In fact, such was its success that – within that first year – it had come to the attention of those running the biggest and best scientific blogging collective of the time: ScienceBlogs (Sb from hereon). I was invited to join, and Tet Zoo ver 2 launched there on January 23rd 2007, things immediately kicking off with a discussion of blood-feeding in oxpeckers. I was now being PAID TO BLOG: Sb ran a system whereby you were paid according to number of monthly hits, there being incremental tiers corresponding to payment. Despite being in the top 5 most-hit Sb blogs, we’re talking about monthly payments of $150. Better than a poke in the eye, but not exactly enough to pay the bills.

An ominous portent of things to come -- my beloved ScienceBlogs mug suddenly made a cracking noise one day and split neatly in half. Read down to see what this was, err, portenting... is that a word? Image: Darren Naish.

ScienceBlogs was a great group to be part of. Constant encouragement and support from Bora Zivkovic, our benevolent blogfather, and a healthy backchannel discussion group involving a large number of the Sb bloggers, created a good, healthy community. My personal situation at the time meant I could blog pretty frequently, and a (mostly) well designed, user-friendly comment system resulted in long, excellent comment threads often containing more insight and information than the articles they were appended to. I will admit to being continually miffed by the fact that the site’s two highest-hitting bloggers – PZ Myers and Greg Laden (neither of whom I have anything against personally) – seemed to blog randomly about all manner of things that were either not really scientific or were outside their area of expertise, but such it is and always will be. No offence guys.

A montage depicting 'things relevant to Tet Zoo' as of July 2011, and used in the article launching Tet Zoo ver 3. 2011 is such a long time ago...

But it was not to last. A few events – most notably PepsiGate (in which the Sb overlords opted to run a commercial blog written by Pepsi alongside the other blogs) – resulted in meltdown during the latter part of 2010, and bloggers jumped ship. New collectives were created, and Bora founded a new science blog hub at Scientific American (SciAm from hereon). I was invited, and Tet Zoo ver 3 launched there on July 5th 2011. Awesome – a continuing stint of paid blogging (now we’re talking about $200 per month, eventually upped to $500). My contract at SciAm has required the publication of four articles per month, which is doable but often difficult given how many other things I now have going on (freelancing, consultancy, authoring, editing, research). But, here we are at another new location, so my time at SciAm was not to last. Why?

Screengrab of part of the very first article at Tet Zoo ver 3, the SciAm years (2011-2018).

Why leave SciAm? Scientific American is an excellent brand that I respect very much, and I always thought it was good for Tet Zoo to be associated with it. Ok, there are minor quibbles I have with the style of the site: I never liked the formatting, what with the space for articles being a narrow vertical strip surrounded (on a monitor screen) by vast borders of white nothingness, the thick horizontal bars at the top of the screen (sometimes accounting for about a quarter of the display (again, on a monitor)), and the limiting of images to 600 pixels in width and low-res…

The third (and final) part of the Tet Zoo 12th birthday review - published at Tet Zoo ver 3 - is currently offline because of image permission problems (read on). The article will have to be published here, at ver 4. Here's one of the photos from that article. It shows the High Tatras, Poland. Image: Darren Naish.

But it’s two things in particular that have forced me to leave. The first concerns commenting. SciAm, very obviously, is not interested in hosting comment sections. After months of discussion it was obvious that they were never going to get rid of an awkward (and often semi- or non-functional) login system. By merely including a login, you’re losing virtually all of your potential commenters, many of whom will only comment when it’s quick and easy to do so. I asked repeatedly about this. And I found, to my surprise, that virtually all other SciAm bloggers had or have no interest in commenting either. I realised eventually that this is because those people – no offence intended – are not bloggers at all, but science writers who only want to write about newsworthy stories, not engage with their readers or build a community. I feel bad for saying this, but there it is. Over time, I felt more and more like an odd-one-out on the site. Anyway... hopefully, we will once more have a fully functional, user-friendly commenting system here. Feel free to put it to the test  - we may even aim to get back to the 23 comments game. To regular readers: the rules of the past will apply here. Trolls and haters are not tolerated, charlatans and cranks are called out or muted, and I have ultimate discretion over what gets published.

Seabirds: yet another group of tetrapods I've never covered sufficiently at Tet Zoo (despite 12 years of operation). Let's correct that. Here, a Herring gull Larus argentatus protests at the nearby presence of a Northern fulmar Fulmarus glacialis. Image: Darren Naish.

The second issue concerns image use. In my early years of blogging, I made many mistakes and did many things that are totally unacceptable today. Within recent years, I’ve been following the rules of fair use and creative commons licences quite strictly and appropriately. But a change in policy at SciAm – forced by the larger company to which SciAm belongs (Springer Nature) – means that they’ve become unreasonably restrictive on image use, a consequence being that I was having images removed from my articles despite complying with fair use and CC licencing. Ultimately, entire articles were being removed from Tet Zoo due to such issues (they’ll be republished here at ver 4). It became obvious at this point that my style of blogging – which is contingent on the use of a large number of images, many of which are of the sort that were proving a problem at SciAm (book and magazine covers, and images from inside them) – is just not compliant with the SciAm way.

There are currently over 550 in-progress illustrations at the Tet Zoo patreon. If you support me you get to see how things are coming along behind-the-scenes.

I had to make the decision to leave. To say goodbye to those monthly earnings. In order to cover for the loss in earnings this would entail, I set a goal at patreon. But, in the end, I had to leave before that goal was reached. So here we are. A huge thanks to those who support me at patreon – this is now an independent blog dependant entirely on benefactor funding – and I hope that if you’re not a patron you’ll consider becoming one if you think it’s worth it.

On the subject of finances, I’m thinking seriously about setting up some sort of Tet Zoo Trust, Fund or Foundation – I’m tired of using my own personal money on research and travel and would also like to contribute to the research of my colleagues and associates. More on this in future. I need a new Tet Zoo logo, here's the old one (prepared back in the days of ver 2 but then abandoned)...

So – what next? As I’ve said as per previous moves, things are basically going to carry on as normal as goes what Tet Zoo does. I have, even after 12 years of blogging, a long list of things I want to write about here, many of which have never been covered before. So, let’s get to it…