Friday, 25 July 2014

"Think Batman x Iron Man": how pterosaurs are inspiring the next generation of aircraft

Admit it, whatever you drive to work seems a little less adequate now.

Pterosaurophile Mike Habib was recently featured in a Scientific American article about the utility of pterosaur research. Let's face it, as cool as pterosaurs are, it can be hard to justify research into them when the world is faced with real problems like climate change, overpopulation, an enormous biodiversity crisis and Michael Bay movies. But Mike's interest in pterosaurs principally concerns biomechanics, quantifying the mechanical properties of pterosaur anatomy and seeing what it was capable of, and this sometimes allows transference of their evolutionary solutions to our own technological problems. Among other things, pterosaur biomechanics might be applied to some big projects: developing unmanned vehicles - including some which may explore other planets - and developing wind-stable fabrics. The latter may not sound very exciting, but wind-resistant fabrics are essential in all sorts of extreme activities, from exploring remote corners of the world (think tents), lightweight aircraft (parachutes, hang gliders, etc.) and extreme sports (wingsuits).

But that's small fry compared to one idea mentioned in the article. As part of an international team - including myself - pterosaurs may be launching air travel in a whole new direction. The manner in which pterosaurs took off - so called quadrupedal launch - offers a solution to a problem faced thousands of times around the globe each day: launching aircraft into the air as effectively as possible. As we all know, three lines of evidence point to pterosaurs launching quadrupedally, with most effort coming from their forelimbs. 1) animals launch using from their 'default' gait, and pterosaurs were quadrupeds; 2) pterosaur forelimbs are much more developed than their hindlimbs, whereas the opposite is true in hindlimb launchers and, 3) above a certain size, pterosaur hindlimb bones would actually fail in launch (Habib 2008, 2013; Witton and Habib 2010). These point to a powerful, quadrupdal launch mechanic which permitted even the largest, 200-250kg pterosaurs to take to the skies from a standing start, while birds - with their hindlimb launches - are seemingly capped at 70-80kg.

It's not only large birds which look enviously on pterosaurs. Most of our own aircraft require runways for takeoff. Vehicles which can take off without runways, like helicopters, are constrained to large size because of their power requirements and required wingspans. All aircraft launches require lots of fuel, and lots of space. It's unsurprising, then, that quad-launching giant pterosaurs have attracted the attention of engineers, as they clearly evolved a method of launch which is not only space and fuel-efficient, but also incredibly powerful. Practical results are undoubtedly years away, but the notion of a small, solo-pilot aircraft being capable of quad-launch and powered flight is realistic enough that we're seeking money for a project to test the waters. The concept we have in mind resembles a suit more than a plane - as Mike put it on Twitter, "think Batman x Iron Man" - alluding to concepts of the craft being controlled by a person strapped within the chassis, sort of like wearing a multi-million dollar pterosaur costume.

A visual history of pterosaur-inspired flying machines. 1, Ernst Stromer, 1913, basic glider model of Rhamphorhynchus wing membranes; 2, Hankin and Watson, 1914, a model based on their pioneering studies of Pteranodon flight (Hankin and Watson 1914); 3, Erich von Holst, 1957, a rubber band powered, wing flapping Rhamphorhynchus glider; 4, Cherrie Bramwell and George Whitfield, 1974, 7m wingspan Pteranodon glider based on their seminal 1974 paper; 5, Bramwell and Whitfield, 1984, half scale 4.5m wingspan Pteranodon made for the BBC; 6, Paul MacReady, 1984-85, 5 m span Quetzalcoatlus remote controlled, computer balanced glider (see MacCready 1985); 7, Margot Gerritsen, 2005, scaled Anhanguera with fully articulated wings built for National Geographic; 8, Matt Wilkinson, Rodger Highfield, and Vivian Bock, 2007, wind tunnel model of Anhanguera used to test Wilkinson’s hypotheses on pteroid orientation, 9, PteroFlight, our new project looking into pterosaur wing performance and its applications. Image compiled by Iain McCreary, used with permission.

What might such a thing look like? Sadly, it's not going to look like the thing at the top of this post. What you've got there is food for thought rendered by someone who's aircraft design skills boils down to watching science fiction movies, and who's engineering protocols are determined by Cool Points. It takes the idea of a 'pterosaur exoskeleton' to an extreme definition, right down to the limb proportions, wing folding and ability to walk about on all fours. Undeniably cool looking, just not very practical. But technologies and ideas taken to an extreme in this painting actually do exist. Augmentation of human frames with robotic exoskeletons is an intensive area of research and already employed to aid physically disabled people, as well as boosting the carrying strength of ground troops. Computers capable of flying deliberately unstable and responsive aircraft -manned or unmanned - are widely utilised. Large, controllable pterosaur-inspired vehicles with moving, adaptable wings have been researched for 100 years (above) and achieved flight (albeit not launch) on numerous occasions, with recent models featuring automatic computer control. The basic elements of this project - essentially a computer-supported, pterosaur-inspired lightweight flying exoskeleton - are at the far end of known technological spectra, not fantasy and hokum.

Of course, we're not going to see pterosaur-inspired suits catapulting people skywards tomorrow. Some serious research and developmental work is required before we see anything like a working concept or even - if we're honest - if it's possible at all. At this stage, however, this ultimate application of pterosaur research is not being ruled out. In other words, keep watching the skies - and check out Mike's Scientific American feature for more details.


  • Bramwell, C. D., & Whitfield, G. R. (1974). Biomechanics of PteranodonPhilosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 503-581.
  • Habib, M. B. (2008). Comparative evidence for quadrupedal launch in pterosaurs. Zitteliana, 159-166.
  • Habib, M. (2013). Constraining the air giants: limits on size in flying animals as an example of constraint-based biomechanical theories of form. Biological Theory8(3), 245-252.
  • Hankin, E. H., & Watson, D. M. S. (1914). On the flight of pterodactyls. Aeronautical journal, 324-335.
  • MacCready Jr, P. B. (1985). The great pterodactyl project. Engineering and Science49(2), 18-24.
  • Witton, M. P., & Habib, M. B. (2010). On the size and flight diversity of giant pterosaurs, the use of birds as pterosaur analogues and comments on pterosaur flightlessness. PloS one5(11), e13982.

Thursday, 17 July 2014

TetZooCon 2014: the event the palaeozoological blogosphere deserves

Last Saturday hosted an event which might, in future years, be considered a strange experiment. Set up in typical convention manner with attendance fees, invited talks and interactive audience activities, its unique selling point was its inspiration: a (largely) technical science blog which covers obscure animals in as much, often more, detail than you'll find in any textbook or scientific paper, as well as arcane topics such as speculative biology, natural history art memes, and cryptozoology. I'm talking, of course, of TetZooCon 2014.

NB. Like a chump, I didn't take a single photograph the entire day, so you'll have to make do with a very bland blog post.

Held at the London Wetland Centre on the 12th of July, TetZooCon 2014 was the latest expansion of the 'TetZooVerse', an internet enterprise founded on three incarnations of the Tetrapod Zoology blog and, more recently, a podcast, two internet comics and on-demand merchandise. The brainchild of Darren Naish and (more recently) John Conway, it's undoubtedly one of the longest running and most successful science outreach exercises on the internet, and notable for covering complex narratives and scientific problems in the world of tetrapod studies. TetZoo fans thus comprise not only casual internet surfers but also researchers and practising scientists. Few other blogs can boast such appeal and far reach, making TetZoo one of the few internet enterprises which might manage the tricky move from the 'free' virtual world to one of admission fees, travel expenses and conference overheads.

In many respects, TetZooCon almost felt like watching a live version of the blog as different talks - essentially 'live blog posts' - covered an array of TetZoo-relevant topics. Unlike TetZoo, the floor did not solely belong to Darren and John, but shared by a host of excellent speakers. I'm not going to cover the talks in detail here because others have already done so, but the topics included speculative zoology, amphibian conservation, wildlife photography, vertebrate palaeontology, crytozoology and mythical animals. Regular readers will know that I was among the invited speakers and covered changing perceptions of azhdarchid pterosaurs. As with other elements of the TetZooverse, these talks meandered from pure science (sauropod neck length) to almost humanist topics (mermaids, the cryptozoological leanings of Shakespeare). Many struck ground between these extremes, noting the interplay between science and culture and how they've influenced each other - for better and worse. Arguably, providing a platform for such talks and the diversity of topics was TetZooCon's greatest success. I've not been to a conference where talk topics varied so considerably and, in contrast to conferences with homogenous themes, there was no chance for getting subject-weary here. The talks were presented at pitch-perfect semi-technical level, assuming that the audience was intelligent and would have some prior knowledge of the broader subjects at hand (e.g. there were no, or only very brief, explanations for what things like Orang Pendek, sauropods or azhdarchid pterosaurs are), while also appreciating the room was not full of experts. It helped, of course, that the speakers and presentations were excellent. I definitely walked away with a greater education than I walked in with.

Other events included a palaeoart workshop, where attendees - led by palaeoartists John Conway, Bob Nicholls and, er, me - attempted to restore the life appearance of the historic 'Mantell Piece' Mantellisaurus fossil, and a TetZoo-themed quiz. The former was of interest for not only palaeoart aficionados, but also anyone wanting to know how fossils are interpreted. There was discussion over bone identification, how many individuals were represented by the specimen, how we could deduce the affinities of the animal and so-on, and we all compared images and notes at the end. The work of the lead artists was beamed onto the screen behind us so audience members could not only see what we were sketching, but engage in discussion with us about specifics of the fossil. Suffice to say (cheating ne'er-do-wells aside who recognised the specimen and simply drew an ornithopod), there was virtually no reconstruction consensus. Tours of the wetland centre and the obligatory pub dinner followed, while merchandise - including prints, 'official' TetZoo products and the much lauded Palaeoplushies - was on sale all day.

Was the event a success? As a speaker and delegate, my opinion is an unreserved 'yes'. There were enough delegates to generate that 'real' conference feel, the day was varied and interesting, and it was a lot of fun to be part of. With strict scheduling, custom 'palaeoart cams', delegation packs and almost flawless audiovisual performance (except for my own talk!), the day was pulled off with the sort of professionalism you'd expect from a long running conference rather than a first-time event. Most importantly, the day felt fresh and different from other conferences. As millions of TetZoo readers and listeners attest, there is a large audience for the 'offshoots' of zoological science such as palaeoart, speculative zoology and so-on, but few venues exist to chat about these topics outside of the internet. TetZooCon is a welcome plug in that gap.

Of course, whether we'll see a second TetZooCon depends on the transformation of online (and free) participation in the TetZooverse to financial and time commitments from potential delegates. This point is really why I wanted to write this short post. Turnout for this first event was good and, happily, conference overheads were recouped. At the same time, I don't think Darren and John slept in beds of gold leaves that evening. Events like these live and die on the whims of potential delegates so, if you were 50:50 about attending this time around and decided against it, rest assured that it was a blast and you won't want to miss out again. If the event passed you by entirely, but you like the idea of an annual celebration of the palaeontological and zoological blogospheres, then you'll also want to get on board next time around. This is a conference with lots of potential and, with enough support, it could become one of the most accessible, unique and interesting fixtures of the conference calendar. If it happens, I'll be booking my place for TetZooCon 2015 as soon as I can next year. If that's not high recommendation, I don't know what is.

Monday, 30 June 2014

Azhdarchid pterosaurs vs. the world

Azhdarchids: also available in flying. Depicted animal here is based on Quetzalcoatlus sp., but no taxon in particular.
In just a few weeks the world will stop for TetZooCon, a one day convention of all things we associate with the famous Tetrapod Zoology blog and podcast - tetrapods real and scientifically-speculative from wondrous, charismatic fossil reptiles to deceptively interesting small, brown herpetofauna. If you’re reading this, you’re slap bang in the middle of the TetZoo demographic and I guarantee* you’ll have a good time. Tickets are available until Friday 4th July, and it’s all going down one week later - Saturday 12th. Get your place while you can, or forever live with the shame.

*Guarantee not guaranteed.

In a surprise move, my contribution to TetZooCon features pterosaurs. Specifically, I’m looking at the way one group of pterosaurs has made major ripples in the palaeontology pond in recent years - and not just scientifically. The changing face of pterosaur science is certainly interesting, but an equally intriguing, rarely told story exists on the popular face of flying reptile research. To whet your appetite, here’s an 'extended abstract' of my TetZooCon talk, giving some insight into what we'll be talking about in a couple of weeks.


Pterosaurs are not unfamiliar characters in popular culture. They have been mainstays of science fiction literature since at least 1874 (Jules Verne’s Journey to the Centre of the Earth), made the jump to the silver screen in 1925 (The Lost World), and since starred in uncountable stories of time-travel, lost worlds and Jurassic Park-inspired de-extinction fiction. Neither are they strangers to public education, from being part of Benjamin Waterhouse Hawkins’ 1854 Crystal Palace menagerie to modern appearances in £multi-million documentaries. The hundreds of years of popular pterosaurs have rarely showed much adherence to flying reptile science: their appearances, behaviours and lifestyles mostly reflect a shorebird-like 'pterosaur archetype' rather than the specific anatomy and habits of a once-living taxon.

One of the many curious things about early azhdarchid reconstructions is the head nubbin - a short, posteriorly-directed conical(?) crest on the back of the head. There is nothing like this known from any azhdarchid. Far from being an early mistake, reconstructions with head nubbins persist until at least 2000. Slide from my TetZooCon talk.
But all that is changing. One pterosaur lineage has overshadowed ‘generic’ popular pterosaurs to bring aspects of ‘real’ pterosaurology to the masses, and has even stolen some limelight from dinosaur celebrities in the process. Media as disparate as documentaries and comic books show these animals in (basically) anatomically correct forms, with accurate but atypical postures, and behaviours which are far removed from the idea of pterosaurs being ancient seabird analogues. The animals in question are, of course, the long-necked, toothless and gigantic flying reptiles, Azhdarchidae.

In some respects, the recent surge of azhdarchid pop-culture uptake is a bit strange. It is not, for instance, that azhdarchids are a newly discovered group. Far from it, their fossils were found by at least the latest 1930s or early 1940s; good remains were apparent the 1970s, and the concept of Azhdarchidae was formalised in the early 1980s. They’re not new to popular culture either, having hung around its periphery since the 1970s to be wheeled out as 'Largest Flying Animals Ever' on occasion. These early popular azhdarchids showed little uniformity in their reconstruction - maybe even less than other pterosaurs at that time. Most bore little resemblance to actual azhdarchid fossils, either anatomically of functionally (above): hugely elongate wings, longirostrine skulls with snub-noses, pin-heads, toothed jaws, short-necks, long necks with swan-like flexibility were all rendered in artwork from 1970 - 1990. Art produced in the 1980s - 2D work by Greg Paul, Paul MacCready’s 1985 glider (below), and a (largely sculpted) azhdarchid skeleton mounted by the Texas Memorial Museum - were probably the first works to strike close to reality, but they’re still a bit short of the mark. John Sibbick’s better known and more influential 1991 snub-nosed Quetzalcoatlus was a step back from these more accurate works, accidentally making a chimeric azhdarchid from at least two Javelina Formation azhdarchids (this ‘snub nose’ almost certainly belongs to an unnamed, short-skulled azhdarchid from the same horizon as Quetzalcoatlus).

The famous 1985 'QN' pterosaur, a half-size gliding Quetzalcoatlus northropi and friends, including its designer, the late Paul MacCready (right of middle, in the tie). The model flew successfully multiple times and isn't a bad rendition of an azhdarchid, although many assumptions made in its construction conflict with modern pterosaurology. From MacCready (1985).
Why all the confusion over azhdarchid appearance? Most azhdarchid material known until the late 1990s was either too scrappy to inform artists about life appearance, while the more complete material (the small Quetzalcoatlus species) was infamously not published (still isn't!). The world at large was therefore not able to appreciate azhdarchid anatomy, so any artwork of them required more guesswork than usual. In some cases, entire 'reconstructions' were products of imagination. Not aiding the murky early phase of azhdarchid palaeoart was the transforming nature of pterosaur science which, in the 1980s and 90s, saw much of what we thought we knew about these animals turned on its head. Thus, artists who wanted answers to simple questions like standing postures, wing membrane attachment and so forth weren't always presented with straight answers. 1997 saw a potential change for the better when Unwin and Lü (1997) reclassified the Chinese Maastrichtian ‘nyctosaurid’ Zhejiangopterus linhaiensis as an azhdarchid, but few paid attention to this obscure species when reconstructing 90s azhdarchids, and artwork continued to remain of variable accuracy. The azhdarchid fossil record has not improved fantastically since 1997, only expanding via isolated, scrappy bits and pieces. Their sudden popularity and uniformity of reconstruction has nothing to do with a significantly improved azhdarchid fossil record, then.

Azhdarchids: over 40 Megafonzies of cool!

So, if azhdarchids aren’t new, and they’ve not sent a burst of insightful fossil material our way, why are they now so popular? Perhaps recent reappraisals of their appearance and behaviour have more influence here than anything else. Reconsideration of azhdarchid mass estimates (e.g. Paul 2002; Witton 2008; Henderson 2010; Sato et al. 2010), re-interpretations of lifestyles (Hwang et al. 2002; Witton and Naish 2008, 2013; Carroll et al. 2013) and flight characteristics (Habib 2008, 2013; Witton and Habib 2010) have recast azhdarchids from billboards of flighted animal size to muscular, terrestrially-competent predators and powerful fliers which were also giant. This has seen azhdarchids landing ‘major roles’ in palaeo pop media. In the last five years, erect-limbed, terrestrially stalking and quad-launching azhdarchids featuring in the BBC documentary Planet Dinosaur, Atlantic Productions’ Flying Monsters 3D, the 20th Century Fox film Walking with Dinosaurs 3D, the 20,000AD comic series Flesh and recent comics of Teenage Mutant Ninja Turtles, been made into at least two figurines by CollectA, in the upcoming, Steam-released multiplayer game The Stomping Land, in Nathan Carroll's wearable pedagogic puppet form and even a rap. This uptake of the same pterosaur lineage is all the more surprising when you consider the diversity of influences and goals of these projects, as well as the near-infinite sea of fossil species which could take their place. More remarkably, these depictions of azhdarchids aren’t anatomically bad or variable, either: they have large, pointy heads with posteriorly placed crests, long necks, short wing fingers and long limbs. It seems azhdarchids have genuinely penetrated the pop-palaeo zeitgeist.

Azhdarchids in recent comic books, including Teenage Mutant Ninja Turtles and the fabulous 20,000AD series Flesh. Note the far right panel - terrestrially stalked TO DEATH!!! Awesome stuff - should really cover Flesh in more detail here some time. Slide from my TetZooCon talk.

The times, they are becoming very different

I think there's several points of interest here. Firstly, we seem to be witnessing a relative rarity within palaeo pop culture: the rapid indoctrination of a new lineage into the canon. What takes a fossil species from an occasional extra and bit-part player to relative superstardom in the space of a few years? There must be aspects of ‘new’ azhdarchids which have suddenly made them marketable and appealing, and very quickly following a spate of new research. Before anyone mentions it, I can vouch for azhdarchid uptake not being overtly pushed by the scientists involved in reinventing them. I've acted as a consultant for three of the projects listed above because ‘new’ azhdarchids were sought after by the media producers, and not as a generic pterosaur expert who pushed his own ideas. The other media, as far as I'm aware, just moved forward with these ideas on their own. 

There is doubtless a myriad of factors making azhdarchids popular - good publicity, a sudden glut of tv and movie interest in prehistoric animals etc. - but I suspect the most important factor is that science accidentally gave azhdarchids a more appealing ‘character’. When we restore fossil animals in art and science we cannot help but impose certain ‘character traits’ into them, and, as with fictitious characters, those with traits we consider desirable are more likely to be popular. 'New' azhdarchids embody everything which is classically cool: they’re original; imagined as assertive, confident animals of great skill and energy; undoubtedly stylish and unusual to behold; instantly recognisable, and their large size creates a cool sense of indifference -metaphorically and physically bigger than us and our problems. Plus, they have that edge of danger: big, predatory species which harvest smaller ones for their own use: bad guys and anti-heroes are always cooler than the good guys. In short, it’s not surprising that ‘new’ azhdarchids are popular because they embody the same characteristics as most iconic literary monsters. The traits outlined above could easily apply to H. G. Wells’ Martian tripods (below) or the Star Wars mechanical walkers. Prior to their reinvention, azhdarchids didn’t - and couldn’t - have this appeal, as their appearance was ill-defined, their lifestyles too poorly constrained (skim-feeding? sediment probing? aerial hawking? aquatic pursuit predation? wading?), and much of our science pointed to rather ineffective, flimsy animals. This not only prevented crystallisation of an appealing and memorable palaeo pop ‘character’, but also didn't give them much popular clout.  

The uptake of azhdarchids in pop-culture may reflect their recent recasting as stylish, dominating predators of small, defenceless animals, a formula known to strike a nerve with the public - ask H. G. Wells. Do our palaeo pop-culture icons attain iconic status because, like some literary ones, they simply evoke cool characteristics and styles? 
Perhaps more importantly, azhdarchids have - for what seems like the first time - persuaded popular culture to widely depict actual pterosaurs rather than an anonymous set of wing membranes and toothy jaws. That's pretty neat, as it means we're starting to break the notion that different pterosaurs are just minor variants on the same basic animal. I wonder if this applies to pterosaur science too, as research into azhdarchid lifestyles and habits is providing compelling evidence of palaeoecological variation within the group: shorebird, fish-eating habits applied almost universally to Pterosauria just doesn't work for these guys. With it being increasingly obvious that azhdarchids were doing their own thing, it's easier to start seeing other pterosaurs as potentially having distinctive lifestyles as well. Azhdarchids may be the thin end of the wedge in this respect for both popular and scientific circles.

Of course, no-one can predict how long our current interest in azhdarchids will last, nor what will happen to hypotheses concerning terrestrial stalking, quad-launch and so on. My gut feeling is that these ideas will stand up to scrutiny, but we can never predict what the fossil record or new studies will tell us. Whatever happens, these ideas and the animals they concern have gone some way to superseding generic ‘pterodactyls’ in palaeontological culture, replacing them with a more accurate and detailed appreciation of pterosaur diversity. But what next for azhdarchids? What advances in azhdarchid science are on the horizon? How might these impact their portrayal in popular culture? For that, you’ll have to attend TetZooCon and my talk. Tickets!


  • Carroll, N. R., Poust, A. W. & Varricchio, D. J. (2013). A third azhdarchid pterosaur from the Two Medicine Formation (Campanian) of Montana. In: Sayão, J. M., Costa, F. R., Bantim, R. A. M. And Kellner, A. W. A. International Symposium on Pterosaurs, Rio Ptero 2013, Short Communications. Universidad Federal do Rio de Janeiro: pp 40-42. 
  • Habib, M. B. (2008). Comparative evidence for quadrupedal launch in pterosaurs. Zitteliana, 159-166.
  • Habib, M. (2013). Constraining the air giants: limits on size in flying animals as an example of constraint-based biomechanical theories of form. Biological Theory, 8(3), 245-252.
  • Henderson, D. M. (2010). Pterosaur body mass estimates from three-dimensional mathematical slicing. Journal of Vertebrate Paleontology, 30(3), 768-785.
  • Hwang, K. G., Huh, M., Lockley, M. G., Unwin, D. M., & Wright, J. L. (2002). New pterosaur tracks (Pteraichnidae) from the Late Cretaceous Uhangri Formation, southwestern Korea. Geological Magazine, 139(04), 421-435.
  • MacCready Jr, P. B. (1985). The great pterodactyl project. Engineering and Science, 49(2), 18-24.
  • Paul, G. S. (2002). Dinosaurs of the air: the evolution and loss of flight in dinosaurs and birds. JHU Press.
  • Sato, K., Sakamoto, K. Q., Watanuki, Y., Takahashi, A., Katsumata, N., Bost, C. A., & Weimerskirch, H. (2009). Scaling of soaring seabirds and implications for flight abilities of giant pterosaurs. PloS one, 4(4), e5400.
  • Unwin, D. M., & Lü, J. C. (1997). On Zhejiangopterus and the relationships of pterodactyloid pterosaurs. Historical Biology, 12(3-4), 199-210.
  • Witton, M. P. (2008). A new approach to determining pterosaur body mass and its implications for pterosaur flight. Zitteliana, 143-158.
  • Witton, M. P., & Habib, M. B. (2010). On the size and flight diversity of giant pterosaurs, the use of birds as pterosaur analogues and comments on pterosaur flightlessness. PloS one, 5(11), e13982.
  • 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. (2013). Azhdarchid pterosaurs: water-trawling pelican mimics or "terrestrial stalkers"?. Acta Palaeontologica Polonica doi: http://dx. doi. org/10.4202/app, 5.

Thursday, 26 June 2014

Azhdarchid pterosaurs - invading TetZooCon this July!

Mountains of respect to whoever can name the sources and years of each of these silhouettes. If your surname is 'Naish', you're not allowed to play. Title slide from my TetZooCon talk.

What are you doing on the 12th of July this year? If you're in London and you fancy a day of tetrapod-themed fun - and who doesn't? - you could do a lot worse than attending TetZooCon, a whole day inspired by the famous Tetrapod Zoology blog and its podcast. TetZoo really needs no introduction to anyone reading this, being renowned for mixing semi-technical zoological discussions of Recent and fossil tetrapods with speculative biology, fictitious creatures, cryptozoolgy and, well, whatever else takes the fancy of hosts Dazza Nash and Johnny Conners.

The full TetZooCon timetable of events is appropriately broad and covers dinosaurs, cryptozoology, conservation, speculative biology, primatology, wildlife watching and two regular stalwarts topics of this blog - pterosaurs and palaeoart. I'm taking reigns for the pterosaur aspect of the morning (you can get a sense of what I'll be talking about above - more details on this soon) as well as taking part in an interactive palaeoart event with two real artists - Bob Nicholls and John Conway. This promises to be great fun, and allows audience members an unusually good insight into palaeoart processes. I'll be selling prints of my work alongside other artists and merchandisers - the Palaeoplushies will be in town! - so bring your pennies for products you won't find anywhere else. And if that doesn't already convince you that there's something for everyone, the venue is none other than the London Wetland Centre, and delegates are free to wander around it all day.

In all, it sounds like it's going to be a terrific day, and I'm pretty stoked to be invited along. If this post tickles your fancy, and especially you'd like events like this to become regular fixtures in your calendar, support the event by spreading the word and grabbing some tickets. Looking forward to seeing you there!

Monday, 16 June 2014

Darwinopterus vs. Bat Out of Hell

Darwinopterus robustodens and his pal, a European robin (Erithacus rubecula). When not posing for artwork, they drive around in a van solving mysteries.
Wukongopterid pterosaurs have been on my mind this week thanks to a near-complete manuscript about them. This required* rendering everyone's favourite wukongopterid Darwinopterus - results above (note that this is D. robustodens, not the more familiar modularis). In the final version he's joined by something else, but you'll have to wait to just what that is. Wukongopteridae is the group of recently discovered Chinese pterosaurs which bridge early pterosaur and pterodactyloid-grades of pterosaur morphology, famously combining head and neck characteristics of the latter with the bodies of the former. They're best known for Darwinopterus modularis, but there's actually now nine(!) taxa known from the same horizon in north-east China, all morphologically very similar and almost certainly oversplit.

*By 'required', what I really mean is that I'm compelled to construct papers with extraneous artwork in them somewhere, because I'm a sucker for punishment.

Illustrations of Darwinopterus and I go way back. I was asked to do press images for two early publications on it: the initial description and assessment of its unusually 'modular' evolution (Lü et al., 2010), and the equally fantastic discovery of its sexual dimorphism and an egg-mother association (Lü et al., 2011a). Below is the first of these images, published in 2009.

Ah, 2009. Shrink wrapping, relatively light integuments and very low soft-tissue crests were still in fashion. Fun fact: Darwinopterus was referred to as 'Frank' before it was given a binomial, a nod to it being a Frankenstein's Creature-like mash of body parts.
Looking back on Darwinopterus 2009, I'm not enormously happy with it. This in itself isn't that unusual. Artists often look back with dissatisfaction with older works, but this has an additional reason for dissatisfaction: I never really agreed with the notion of Darwinopterus as an aerial-hawker of flying tetrapods, and I think this comes across in its execution. I outlined my basic concerns with this idea in Pterosaurs (Witton 2013):
Given that wukongopterids have provided an insight into macroevolutionary processes, filled a gap in pterosaur phylogeny, and present a very unique pterosaur bauplan, expectations may be high that their proposed foraging strategies will also be rather amazing. Fittingly, some have proposed that wukongopterids were pterosaur top guns, their newly evolved long necks and oversize heads being used to prey upon dinosaurs, other pterosaurs, and even gliding mammals in midair (Lü et al. 2010). Such acts would be rather remarkable because, with even a generous mass estimate, the biggest Tiaojishan wukongopterids would not weigh much over 300 g (extrapolating data from Witton 2008), which is about the same as a modern feral pigeon.
At that size, tackling squirrel- sized mammals or crow-sized dinosaurs on the wing would be a feat earning praise from even the hardiest modern raptors, and wukongopterid skeletons would have to be brimming with offensive weaponry for this purpose. Vertebrate-hawking birds are renowned for their talons, incredibly strong feet, robust skulls, and powerful beaks (e.g., Hertel 1995; Fowler et al. 2009), while bats that subdue large vertebrates in fl“ight are also armed with formidable teeth and powerful jaws (Ibáñez et al. 2001). These adaptations provide the means to immobilize their prey quickly and efficiently, and are obvious advantages for animals grappling with large prey while in “flight. Vertebrate hawkers are also powerful fliers that can chase down their quarry and, once immobilized, carry the prey to a safe spot to eat. Pterosaurian equivalents would therefore require equally powerful “flight musculature to permit the same tasks. Unfortunately for the Darwinopterus raptor hypothesis, wukongopterids do not possess any of these requirements. None of their appendages bear the chunky digits and talons ideal for subduing large aerial prey items, and their long, comparatively delicate skulls and unimpressive teeth are ill suited to this task. Nor, for that matter, do they have the expanded shoulder regions indicative of the powerful “flight muscles needed to chase and eventually carry their prey. With this in mind, raptorial pursuits look doubtful for wukongopterids.
Witton (2013), p. 141-142.
There's a lot more that could have been said about this, but you get the idea: Darwinopterus and chums were small (jackdaw-sized - see image at the top of the post), delicately built animals for which aerial predation seems counter-intuitive and unlikely. I'm not the only person saying this, either: Lü et al. (2011b) and Sullivan et al. (2014) also raise points against the aerial hawking idea. We could go so far as to to label wukongopterid aerial hawking as another example of an 'extreme' palaeoecology based on cherry picked characteristics rather than considering a full suite of functional data.

So, on reflection, we probably started on the wrong foot but, hey, what can you do? Working as an artist is quite different to being involved in research: ultimately, you're a guy with a paintbrush being told to illustrate someone else's idea, even if you don't necessarily agree with it (also see Csotonyi and White 2014). With the notion of aerial predation being raised in the paper, specifically as a possible explanation for the development of pterodactyloid head and neck features Lü et al. (2010), it was an obvious choice for the image. But we didn't help matters by walking into some of the cheesiest, silliest things stereotypes of palaeoart, primarily because we were trying to make a small, fairly inoffensive animal look like a skydiving, dinosaur-eating-badass. We tried several different takes on this, varying aerial and terrestrial prey, and compositions which upped the voracity, such as this:

You'll need to make your own whooshing jet fighter noises and 'pew pew' laser sounds.
Eventually death-from-above was decided over death from sideways. But a key issue to tackle was that that the proposed prey for Darwinopterus was the same size as the predator itself (Sullivan et al. 2014 go into this more), and it's obvious that Darwinopterus is not a fighting, wrestling creature. To make the image work at all we had to play liberally with animal sizes: the maniraptoran (loosely based on Anchiornis in the final version) is tiny, and the Darwinopterus is a huge, ferocious juggernaut. I'm not the only artist who used this trick: the disappointing pterosaur documentary Flying Monsters 3D also had their Darwinopterus plundering undersized theropods. After we tinkered with reality, we then started piling on the cheese: the maniraptoran was made more reptilian-looking, it's head twisted to stare into the Maw of Destiny and, of course, it's mouth open to 'NOOOOOO!!!' it's impending fate. The result is 'Meat Loaf palaeoart', the sort which resembles the worst kind of rock album covers more than nature.

Think I'm being daft comparing palaeoart to cheesy hard rock album covers? How many depictions of extinct animals have the exact same pose as the titular bat here? See links below for more examples. From Wikipedia.

The result isn't awful, but I think I've had more successful collaborations with the same authors, just because this doesn't feel realistic at all. From the science to the composition, the whole thing just seems 'forced'. Don't get me wrong: I'm sure Darwinopterus could be terrifying instruments of mortality to the right prey (probably terrestrial invertebrates according to Lü et al. 2011b and Witton 2013), and I'm not belittling the drama or viciousness which can occur in the lives of small animals. But life - thank God - doesn't look like the cover of a Meat Loaf album. There are some fossil animals that can - just about - pull off Meat Loaf palaeoart, but it's just not possible to turn small, fluffy, unthreatening animals into monsters. When we do, they're more liable to look goofy than impressive. Despite this, we do it all the time. I understand why we do - it's exciting and marketable, and feeds into expectation that Deep Time as a monster-filled fantasy realm - but it also reinforces the perception that palaeoart is unsophisticated and aimed at children. But there may be another way.

Palaeontologists: next time you have a new, diminutive animal you want illustrated, forget the monster: play the cute card. The chubby little Darwinopterus at the top of this post intuitively seems closer to the reality of this animal than Darwinopterus 2009: AwesomeoSuperKiller. It's proportions, fluffiness and posture are all accurate, and it only looks cute because, well, it probably was: it's a small fuzzball with an oversize head and mischievous grin. If the Internet's obsession with cats and baby sloths has taught us anything, it's that cute sells better than violence. Indeed, this image is one of the most popular things I've ever put on Facebook, and (at time of writing) it's was posted less than a day ago. The great thing about the cute card is that everyone wins: the artwork should promote research just as well or better than it's Meat Loaf variant, because it appeals to wider demographics. The artist gets to render an animal in a more realistic light without jumping through hoops to monsterise it, and the world gets a new picture to 'coo' at. A few folks might even start to appreciate extinct animals in ways they never could when they're always shown screaming and fighting.

And if nothing else, it will mean we'll never have to discuss Meat Loaf album covers here again.

May God have mercy on us all. (Wikipedia)


  • Csotonyi, J. & White, S. (2014). The Paleoart of Julius Csotonyi: Dinosaurs, Sabre Tooths and Beyond. Titan Books, London.
  • Fowler, D. W., Freedman, E. A., & Scannella, J. B. (2009). Predatory functional morphology in raptors: interdigital variation in talon size is related to prey restraint and immobilisation technique. PloS one, 4(11), e7999.
  • Hertel, F. (1995). Ecomorphological indicators of feeding behavior in recent and fossil raptors. Auk, 112(4), 890-903.
  • Ibáñez, C., Juste, J., García-Mudarra, J. L., & Agirre-Mendi, P. T. (2001). Bat predation on nocturnally migrating birds. Proceedings of the National Academy of Sciences, 98(17), 9700-9702.
  • Lü, J., Unwin, D. M., Jin, X., Liu, Y., & Ji, Q. (2010). Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proceedings of the Royal Society B: Biological Sciences, 277(1680), 383-389.
  • Lü, J., Unwin, D. M., Deeming, D. C., Jin, X., Liu, Y., & Ji, Q. (2011a). An egg-adult association, gender, and reproduction in pterosaurs. Science, 331(6015), 321-324.
  • Lü, J., Xu, L., Chang, H., & Zhang, X. (2011b). A new darwinopterid pterosaur from the Middle Jurassic of western Liaoning, northeastern China and its ecological implications. Acta Geologica Sinica‐English Edition, 85(3), 507-514.
  • Sullivan, C., Wang, Y., Hone, D. W., Wang, Y., Xu, X., & Zhang, F. (2014). The vertebrates of the Jurassic Daohugou Biota of northeastern China. Journal of Vertebrate Paleontology, 34(2), 243-280.
  • Witton, M. P. (2008). A new approach to determining pterosaur body mass and its implications for pterosaur flight. Zitteliana, 143-158.
  • Witton, M. P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.

Friday, 30 May 2014

Book review: The Paleoart of Julius Csotonyi, Csotonyi and White 2014

From Titan Books.
The Paleoart of Julius Csotonyi: Dinosaurs, Sabretooths and Beyond (Csotonyi and White 2014) is another palaeoart-focused book from Titan Books, who brought us the acclaimed Dinosaur Art: the World’s Greatest Paleoart two years ago (White 2012). Anyone familiar with this book will immediately note the very similar format and high production quality in this recent Titan release. Although slightly smaller than Dinosaur Art, this will - again - leave readers wondering how the extremely affordable price (£25) covers production costs. Csotonyi was, of course, one of the artists featured in Dinosaur Art but, this time, is entirely running the show. Fans of his work will have little doubt that he can carry an entire book by himself. For the last decade Csotonyi has been establishing himself as one of the world’s premier palaeoartists, illustrating countless press releases, books, articles and museum walls with intricate paintings or digitally-manipulated photograph composites. His work is in such demand that he is one of the few individuals globally who can make a living out of palaeoart, a status which is testament to the quality of his work.

Before we get into the review itself, I want to stress how much of a milestone this book is. Palaeoart and palaeoartists suffer a PR problem where artists are considered unimportant and interchangeable: individuals who are secondary to the scientists pushing palaeontology forward and the audience who – often superficially – experience their work. Titan Books showed that palaeoart could be tackled more seriously and respectfully with Dinosaur Art and are cementing this idea in dedicating a whole book to a leading palaeoartists. Csotonyi's position as a working palaeoartist with major publisher support is rather exclusive, but exactly the sort of treatment palaeoart needs. I hope that Csotonyi’s solo album sells well enough to kickstart a series of books featuring other artists. Intentionally or not, Julius’ artwork is a good place to start this hypothetical series: aesthetically pleasing, extremely high quality, and blending traditional palaeoart approaches with some more complex and radical compositions. As a means to test the market for these sort of books, Julius is one of the strongest candidates currently available.

As an industy, palaeoart needs all the help it can get, starting with this logo.
Right, big-picture stuff out of the way now: what of the book itself? At its most basic level, The Paleoart of Julius Csotonyi is effectively an expanded version of his chapter from Dinosaur Art, juxtaposing imagery alongside an interview about Csotonyi’s art, influences and background. The interview, confined to the first 23 of the 156 pages, features intelligent questions and the interesting responses from the artist. Csotonyi’s passion for art and science are clear even before his images are displayed in earnest, as is the amount of work required to produce the large, ultra-high-quality imagery he is famous for. He leaves no doubt that many personal sacrifices are required to work as one of the world’s leading palaeoartists. This section also contains rarely-seen early works and non-palaeontological artwork, including some dedicated to astronomy. Some of the interview responses and other text features words which may be unfamiliar to lay audiences, but a glossary is provided to help readers navigate these terms.

The real meat of the book is relatively text-light so as to provide maximum space for Csotonyi’s art – large format is the only way to appreciate the detail it contains. The art is roughly arranged in chronostratigraphic order, with Palaeoazoic, Mesozoic and Cainozoic subjects separated into different chapters. As usual within palaeoart, the bulk of the artworks depict Mesozoic dinosaurs, and theropods are particularly well represented. Each piece is accompanied by brief details of the composition and commissioners, and some featuring additional comments from scientists about the subject animals. These comments mostly complement Csotonyi’s talents or spin yarns about research associated with the depicted species and, I guess, are designed to boost the scientific content of the book. I do feel a trick has been missed here because none pass particular comment on the decisions made when reconstructing the animals. Seeing as a lot of Csotonyi's art is produced alongside consulting scientists, I’d like to know what input they had. Even the most tightly constrained reconstructions of a fossil animal requires a lot educated guesswork and speculation about palaeobiology and life appearance and - in my own experience at least - not all of this is left to the artist. After all, this is a primarily a book about scientific art, and it seems that these comments could be more insightful than discussions about fossil localities, chance discoveries, or another complement for Julius' artwork (meant with all due respect, of course, but we know he's good. That's why we bought the book!).

A busy day in Permian Texas. Photo composite by Julius Csotonyi, from The Palaeoart of Julius Csotonyi. Image from here.
Csotonyi himself gives some works longer explanations about the processes involved in the reconstruction. These often highlight works with unusual compositions or viewpoints (such as the excellent ‘fish-eye’ sauropod view) and describes the way each piece was executed, often with alongside draft versions. These provide some insights into his process and will doubtlessly be useful to budding artists. My personal take-home message from these is the exhaustive consideration and research required to understand not only fossil animals, but to also reproduce realistic landscapes and lighting, particularly when odd perspectives and water are involved.

On the art itself: Csotonyi’s images are created using a range of media, including traditional and digital painting, sketches and – most commonly – digital photographic manipulation. I’m going to come clean here and admit that I’m not enormously fond of photographic manipulation. Many such works often fall into palaeoart’s own variant of the ‘uncanny valley’ or, all too often, present oddly-proportioned, strangely posed creatures which have little in common with their known anatomy. Julius’ photo composites are easily among the best, if not the best, attempts at photo-realistic 2D palaeoart out there however, and present reasonably reconstructed animals at either photo-realistic quality, or within inches of it. Some images, particularly the more ambitious, crowded scenes (fans of ‘a busy day in deep time’-type images are well served here) do bear niggles which jar the illusion, such as animals appearing too sharply defined against the background. To a certain extent, this is unavoidable: photomanipulation is incredibly difficult to pull off even remotely well, and even Csotonyi’s lesser successes are still amazing efforts. There are no overused photographic elements, no blurred skin textures, no cloning of animals to create herds of the same individual. When the photomanipulation does work well – and it frequently does – the effects are nothing short of stunning (e.g. below). The image of the resting Edaphosaurus on page 33 could easily be mistaken for a genuine, beautifully shot photograph. As with Dinosaur Art, some panoramic scenes unfold to show enormous vistas stuffed with detail. Many of these fold-outs allow those of us with empty pockets our first detailed look at the many murals Julius has created for North American and Australian museums.

Photo composite Acrotholus audeti and Neurankylus lithographicus  by Julius Csotonyi, from The Palaeoart of Julius Csotonyi. Image from here.
My favourite images in the book are digital paintings (below), such as the dancing Guanlong, the mothering polycotylid and the ceratopsid portrait gallery on pages 102-103. Not only do these show the trademark Csotonyi attention to detail but they’re wonderfully lit and composed: they feel more ‘of a scene’ than the photo composites. A neat touch is that alternative versions of well-known paintings are sometimes included. I actually prefer the near greyscale version of the Acheroraptor press release image on page 43 to the original, its dusky palate and the removal of the mammal from the hero animal’s mouth creating an entirely different tone to the more familiar version.

Digitally painted Brachiosaurus by Julius Csotonyi, from The Palaeoart of Julius Csotonyi. Image from here.
Of course, scientific accuracy is also essential for palaeoartworks to be considered successful. In this respect, the book also delivers. Thoroughly modern reconstructions of fossil subjects are presented: extensively feathered maniraptorans, diverse integuments in other dinosaurs, correctly orientated limbs and so on. In light of All Yesterdays (Conway et al. 2013), Csotonyi’s approach to extinct animal reconstruction may be considered conservative - there are no outlandish, speculative audacities here in terms of appearance or behaviour. Most of the depicted animal interactions are predatory, and the soft-tissues of the reconstructed species are not especially elaborate. Perhaps this is because nearly all of the artworks were commissioned by researchers and museums, clients who tend to favour safer, more conservative palaeoartworks. We should not lose sight of how progressive even ‘conservative’ modern palaeoart actually is. Many sights now familiar to us would have been considered heretical just a few years ago: Csotonyi shows several tyrannosaurs with variable amounts of feathering in the book with little fanfare, for instance. For dinosaurs at least, it’s becoming harder to produce wholly shocking palaeoart without unreasonably bending palaeontological science or speculating wildly. While Csotonyi’s book may lack the accessory frills, wattles and elaborate behaviours of some modern palaeoart, it acts as a fantastic milestone for how far palaeontology and palaeoart has moved in recent years. Moreover, I do not want to give the impression that the images are not interesting or novel: fishing Dimetrodon (above), Polycotylus nuzzling its offspring to the water surface to breathe and reptiles swimming between the dredging fronds of rafting crinoids are just some thought-provoking Csotonyian innovations.

The unkillable skim-feeding hypothesis lives on. Art by Julius Csotonyi, from The Palaeoart of Julius Csotonyi. Image from here.
I do have a few issues with some science behind the artwork. I’m told that an unfortunate misunderstanding resulted in the extensive discussion of Rhamphorhynchus skim-feeding on pages 136-139 (Hone, pers. comm. 2014; above. See the comment from Julius below for the sull story). This was, in fact, meant to reflect dip-feeding or surface-gleaning. Folks who keep up with pterosaur research will know that skim-feeding habits in flying reptiles has been looked into several times, consistently found problematic (e.g. Chatterjee and Templin 2004; Humphries et al. 2007; Witton and Naish 2008, 2013), and widely publicised. It’s a surprise and a shame, then, that this idea made it into the book without someone noticing, and particularly so because the science elsewhere is pretty tight. I also wonder if some of the photo composite crocodyliforms are shown with entirely accurate scute patterns, as most seem to have been taken from modern crocodylians – many Mesozoic crocs had very different, often simpler scute morphologies. And while we’re moaning, I do wonder if some more complex images would have benefited from small ‘key’ illustrations demonstrating the position of each animal. This is not only because the animals can be hard to spot in the complex, detailed scenes on offer (this is not meant as a slight – remember that many of the more complex images are intended to be hundreds of times larger on museum walls), but because linking a list of unfamiliar names to specific creatures can be difficult. Individuals intimately familiar with genera of all major vertebrate groups should be OK (they exist, honest), but I suspect they will only make up a fraction of this books audience.

These are only minor issues in the grand scheme of things, however. The intelligence and quality of The Paleoart of Julius Csotonyi makes it essential for anyone interested in palaeoart, as well as more general aficionados of palaeontology, natural history, or natural history art. I have no doubt that palaeoartists will be keeping a close eye on its success, and hoping that it presents the first of a wave of similar tomes from Titan Books. That’s all to come, though: for the time being, The Paleoart of Julius Csotonyi cements Csotonyi’s status as a world leader among the current crop of palaeoartists, and this book will only further his success.


  • Chatterjee, S., & Templin, R. J. (2004). Posture, locomotion, and paleoecology of pterosaurs (Vol. 376). Geological Society of America.
  • Csotonyi, J. & White, S. (2014). The Paleoart of Julius Csotonyi: Dinosaurs, Sabre Tooths and Beyond. Titan Books, London.
  • Humphries, S., Bonser, R. H., Witton, M. P., & Martill, D. M. (2007). Did pterosaurs feed by skimming? Physical modelling and anatomical evaluation of an unusual feeding method. PLoS biology, 5(8), e204.
  • White, S. (2012). Dinosaur Art: the World’s Greatest Paleoart. Titan Books, 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. 2013. Azhdarchid pterosaurs: water-trawling pelican mimics or" terrestrial stalkers"? Acta Palaeontologica Polonica. (In press).

Saturday, 24 May 2014

Godzilla and MUTOs vs. birds and newts: reasons why the creatures in Godzilla (2014) are (sort of) plausible

Inside every 29 year and 11 month year-old man is a 5 year old who only watches monster movies so he can produce little doodles of them afterwards.
After much hype, Godzilla (2014) has finally stomped into cinemas around the world. It seems to have been a divisive monster movie, with some taking its slow pacing and focus on human characters as a breath of fresh air in a stale genre, and others wishing it had the wham-bam-thank-you-mam approach of Pacific Rim. Personally, I'm in the former camp, and am especially glad the new Godzilla franchise has not started with two hours of giant monster wrestling. While Godzilla (2014) is not flawless, it takes a lot of risks and feels like a modernised 'classic' monster movie, and not the focus-grouped, merchandise-led fanboy service it could have been. I should add that I'm not especially keen on the last 59 years of Godzilla movies, much preferring the genuine drama and symbolism of the original 1954 movie to the decades of monster fights which followed. Because Godzilla (2014) is tonally closer to the original than the later escapades, I'm not surprised that people just wanting two hours of skyscraper-sized wrestling are feeling a bit disappointed. Anyway, there's a lot we could say about this, but that's not quite what I want to write about  here.

A big part of updating Godzilla has been the overhaul of Big G himself, as well as bringing new creatures to this franchise. These are exciting times for monster film fans because, for only the second time in its 60 year history, one of the most famous movie monsters of all is in for a complete redesign (we all know what happened the first time) and, of course, is now free of the constraint of 'suitmation' effects - the fancy term applied to the 'guy-in-latex-suit' approach Godzilla movies are (in?)famous for. A fun fact about the original 1954 Godzilla is that the now-inconic chap-in-suit technology was a compromise, the studio initially desiring stop-motion effects like those of American-made films King Kong or The Beast from 20,000 Fathoms. Toho, the studio which created Godzilla, had neither the money or expertise to execute stop motion effectively, so opted for using costumes instead. Despite using their second choice, the Japanese trailer for Godzilla triumphantly declares their special effects superior to anything the American movie machine was making at the time! And it certainly worked out for Toho: after a difficult teething process, the 'suitmation' approach worked well for the franchise and wasn't tinkered with outside of the much maligned 1998 Tristar Godzilla.

For 2014, Godzilla is a fully digital and, as we all know, relatively faithful to the original designs. It has, however, been altered in ways which would be difficult to execute if we were still watching a man in a suit. A lot of these changes, as well as the design of Godzilla's adversaries, were pretty neat because they tie into what we know about animal biology, scaling and functionality, and I get the impression that the guys behind this latest Godzilla - Legendary Pictures - put a lot of effort into making half-sensible creatures which biologists, biomechanicists and functional anatomists can be relatively happy with. And yes, yes yes: there's a buttload of stuff which is clearly nonsense: there's no way these animals could be the size they are, or firing beams of nuclear fire from their throats and so forth. But that's just par for the course for a Godzilla movie, and I'm not going to jump on boring old bandwagon of highlighting how impossible the whole lot is. What's far more interesting, and what I want to focus on here, is how Legendary built their animals around standard movie monster tenets to produce creatures which are not only intriguing and cool-looking, but also chime with real animal biology and functionality.

Twinkle toes

Godzilla's stubby, new-look foot, as seen from an airport terminal. Image from the Godzilla movie forum.

One of the most striking aspects of Godzilla’s redesign is its short, stubby feet instead of the long, plantigrade feet we’re familiar with from the suitmation costumes (above). This is easily one of the best changes to make in the entire redesign. Not only does it move away from Godzilla looking like he’s wearing a pair of comedy slippers, but it makes a lot of sense from a functional perspective. Long, plantigrade feet of bipedal animals (like our own, as well as those of chaps in Godzilla suits) lift our entire body weight with every step, our metatarsals and ankles both supporting and propelling us forward. This works fine for lightweight animals like ourselves, but comparable foot anatomy in a skyscraper-sized animal would need to be immense to avoid buckling under thousands of tonnes of bending force. The superficially sauropod-, tortoise- or elephant-like foot of the new Godzilla negates this risk however, creating a more columnar distal limb structure which is not employed in lift and propulsion but instead ideal for supporting terrific weight. This has several effects on limb structure and locomotion. The foot musculature (anchored to the shin) is reduced, keeping the weight of the limb down overall, thus making movement more efficient. However, it also decreases the foot mobility, limits stride length and overall gracility. Some of these effects are countered, however, by the elongate thigh region common to all Godzilla designs, which can swing the shortened distal-leg over great distances with every step. This increased length of the proximal limb region and shortening of distal is a trend we commonly associate with larger animals, even in species which are quite active and sprightly, and descended from cursorial ancestors (e.g. rhinos). Godzilla’s enormous thighs also fit with observations that proximal hindlimb musculature becomes relatively huge in larger bipeds - see the pelves of large bipedal dinosaurs for examples - providing the powerhouses necessary to move its huge bulk around. The result is a limb which isn’t going to win Godzilla any prizes for sprinting but, more importantly, is much more suited to the animal itself than the older designs. And let’s face it, when you can take strides measured in hundreds of metres, moving relatively quickly isn’t a problem.


Gills ahoy, on the lower portion of Godzilla's neck. Image from Detroit News.

A clear departure from all other Godzilla movies is that new version can breathe underwater. They’re easy to miss, but look closely and you can spot a series of fish-like gills on the side of its neck. At certain points in the movie, you can see them opening and closing as the big chap respires. Apparently, these were added not only because Big G spends a lot of its time swimming from location to location in the movie, but also to explain how an animal as large as Godzilla could keep a relatively low profile for so long. Air breathing animals of such size would surely be better known to the world at large if they were routinely surfacing to take breaths. These are all that's left of fish-inspired designs for the new Godzilla, which also featured fins instead of rows of bony plates; fish-like scales instead of reptilian skin and so forth.

Gills on a big, adult tetrapod sound a bit crazy, and I will admit - the fish-like (bony?) gills of the new Godzilla are difficult to rationalise entirely among real animals. If we’re willing to stretch belief a bit (I assume we are, what with a fictitious 100 m tall reptile being the subject of discussion here), we can find saving grace from early tetrapods and their descendants, modern amphibians. The latter have internal and external gills in at least their larval stages with most species losing them as they become adults. These gills are supported by a series of bones behind the skull which lead to openings in the body wall which allow swallowed water to escape - these are known known as gill clefts. Some amphibians - specifically select caudatans (salamanders, axylotyls, newts etc. - see below) - famously retain gill structures into adulthood, including their elaborate external gills. We know from exquisitely preserved fossils that such gills were present in the ancestors of all modern tetrapods, some of which also retained these gills in adulthood, and others which at least maintained their gill clefts (Schoch 2009). It seems likely, then, that the possession of gills, and possibly retention of gill structures into adult life* is the ancestral condition for all early tetrapods - including, presumably - Godzilla. Amniotes, which also presumably includes Godzilla, have obviously abandoned these structures and co-opted the gill apparatus for other purposes.

An axolotl with its external gills. © Kevin Schafer /, linked from ARKive.

Could Godzilla or other creatures ever get their gills back? Well, maybe. For a long time it was thought that evolution was irreversible ('Dollo's Law of Irreversibility') and, under this concept, loss of gills would be a one way street: Godzilla would have to evolve an entirely new set of gills, which is entirely possible. However, it's now appreciated that Dollo's Law is not really a law at all, and that organisms frequently do reverse their evolution, including backtracking on complex aspects of their anatomy and life history: digits, dentitions, wings, larval stages, reproductive strategies are all examples of things which have been lost and then recovered in animals. What once seemed like a law is likely more statistical improbability: a lineage can evolve in many ways at any point in time, and the likelihood that it will directly reverse along a familiar path is relatively low. However, in theory, there's no real reason why this shouldn't happen, so long as the selection pressures are correct. It would take some hefty tinkering with genes and mountains of heterochrony, and it's difficult to think of conditions which would promote the development of early-stage gill anatomy, but I guess it's not impossible for an amphibious creature to regain its ancestral gill condition under the right circumstances. My guess is that they would look more like the gills of caudatans than they would the scale-like fishy apparatus of Newzilla, but, let's face it, big feathery gills would look incredibly silly on a city-smashing monster.

See? Less "Godzilla, King of the Monsters", more "Preszilla, Queen of the Desert". Both are scary, but in very different ways.

Has Godzilla put on weight? No, it's just in it's head

One of the key comments made about the new Godzilla design is that it’s a bit of a heffer compared to other versions. This is debatable (they’ve always been pretty chunky if you ask me), but Legendary’s Godzilla design probably appears fatter than the rest because of its proportionally small head. There are good design reasons for this move: the smaller head gives a natural taper towards the top of the animal, distorting its perceived perspective and size, and it helps avoid the comically oversized, googly-eyed heads of previous versions. It's also in keeping with the natural world. Animals show a disproportionate reduction in skull length with respect to body mass - that is to say, larger animals generally have proportionally smaller heads than smaller ones. Note that this applies to carnivores as well as herbivores (Van Valkenburgh 1990; Christansen 1999). Godzilla represents a real extreme of animal gigantism, so it makes sense that its skull and head is going to carry this allometric trend to an unprecedented limit. What’s more, whatever Godzilla actually is (the new movie, thankfully, doesn’t really concern itself with this), it’s clearly on the reptile branch of the animal tree. Reptile cranial musculature is generally less developed than that of mammals, so their skulls and heads are relatively smaller at a given body mass than equivalently sized mammals (Christansen 1999). Again, this fits neatly with the small cranium of our new-look Godzilla. Of course, making the head look even smaller are the big legs and counterbalancing tail but, as mentioned above, this is also to be expected. Bottom line: far from being fat, 2014's Godzilla is just showing the extremes of proportion that we would expect if an animal ever grew to the ridiculous sizes we see on screen, and my feeling is that we're looking at a far more 'realistically' proportioned version that we've seen before.

"I'm not fat, I just have a head of predictably small proportion based on typical animal scaling allometries which distorts your overall perception of my size!" Whatever, buddy. Image from Godzilla Movies

MUTO: Finally, a convincing giant, flying movie creature

Moving away from the main attraction: Godzilla vies for attention in the 2014 movie with another species of giant critter, termed ‘MUTO’ (below). The design for the MUTO(s) is very cool: gigantic, eight limbed, insect-ungulate-pterosaur beasts which defy easy classification. We’re shown pronounced dimorphism in adult MUTOs, the females being larger and more robust than the males, and devoting all limb sets to either terrestrial locomotion or gripping. The males, however, allocate one set of limbs to flight, bearing a spectacular pair of enormous membranous wings which recall monowing plane designs of the early 20th century (this isn't a coincidence: male MUTO wings are actually based on stealth bombers). These wings are largely unfoldable and situated on the dorsal surface of the torso so, when grounded, they extend behind the MUTO like a huge cape. Despite being the smallest creature in the film, male MUTO is still huge, so there’s little risk of it clipping its flowing wings on any structures when locomoting terrestrially.

MUTO in flight, from the Godzilla trailer.
There’s a lot to like about the male MUTO’s wing anatomy and flight mechanics. Firstly, the wings of this animal are simply enormous, as they should be for a creature of its size. Not only do they dwarf the body of the animal, but we could measure their span in metric villages. All too often, flying movie creatures are equipped with wings far too small for their body size, but - running off intuition here - these look ‘right’ for its size. As demonstrated by the birds below, wings increase in length and area disproportionately with body size because of standard rules of scaling: any linear size increases equates to a cubed increase in mass, which gives more for gravity to pull against and greater wing area requirements to achieve lift. If we are ever to expect a several-hundred tonne flying creature to become become and stay airborne, it would need significant wingage: MUTO delivers on this front.

A rule of thumb for designing flying animals: as mass increases, wing length increases faster. Demonstrated here by various bird planforms from Rayner (1988).
But it’s not just wing size that’s cool about the male MUTO: it’s flapping, or lack thereof, was also neat. Flapping amplitude - the degree of movement of a full wing stoke - decreases with wingspan in all flying creatures. The larger an animal is, the less movement is required in its wings to produce an effective flap cycle. A familiar example of this is the wing motions of flapping geese compared to pigeons: both flap their wings nearly constantly while flying, but the former move their wings around the horizontal, while pigeons swing them in great arcs. All too often, giant flying movie creatures (including several giant pterosaurs) show whopping huge flapping amplitudes well beyond necessity for flight and, probably, aerial stability. Godzilla’s male MUTO moves his wings just a little, however: he definitely flaps, but his wings move just a few degrees around the horizontal, not in huge arcs (watch the clip, below, for an example. Skip to 0:24 if you're a particularly impatient type). He also largely soars, as would be expected for any uber-large flier: flapping those wings would take a lot of effort, so soaring is the way to go. It would be loads of fun to get some basic wing size and mass data for this guy to work out some basic flight parameters - cruising speed, travelling distance etc. Given that giant pterosaurs seem capable of cruising at 100kph (Witton and Habib 2010), and yet are mere pipsqueeks at 10 m wingspans compared to the male MUTO, I reckon we’re looking at cruising speeds of several hundred kph. And that’s if he’s not in a hurry.

Finally, we get to see the MUTO launch a couple of times in the film, and each time we see something reminiscent of quad-launching. Pterosaur fans will know this trick as the best hypothesis on the table for explaining how flying reptiles became airborne, and a core factor explaining their attainment of gigantic proportions (Habib 2008). The kimeatics of MUTOs takeoff aren't quite the same - which is to be expected seeing as the limbs of pterosaurs are quite different to those of MUTOs - but it's neat to see the same basic components - a standing start, the rocking back crouch, forward thrust, upward push and leaping phase - incorporated into MUTOs takeoff. These actions are all performed with the four walking limbs rather than the wings themselves, as in pterosaurs and bats, but this also chimes well: MUTOs would need all the power they could to become airborne, so using every available limb is logical. Note that insects are an unlikely model for MUTO takeoff because, while seemingly comparable with MUTO because of their similar limb and wing configuration, at least some insects only use one pair of limbs (the mesothoracic) to launch (Trimarchi and Schneiderman 1995). This is probably because insects enjoy much better power/weight ratios than large vertebrates/skycraper-sized-monsters. 2-4 mm long fruit flies can jump up to 15x their own body lengths - about 30 mm - using their middle limbs only (Zumstein et al. 2004). This sets them apart from bats and pterosaurs, which use all their limbs in launching, and makes these the best candidates for MUTO launch models. It would be neat to know how much attention the animators and designers have been paying to bat and pterosaur research or if they came up with this launch strategy on their own. My experience of explaining quad-launch is that a lot of folks find it counter-intuitive at first, probably because we don't see many animals routinely taking off in this way, so I do wonder if the similarity between these takeoff strategies reflects research into animal takeoff.

And that's it for now. If you want to know more about how the animals of Godzilla (2014) were constructed, you should check out the concept art book which accompanies the film: Godzilla: the Art of Destruction (Cotta Vaz 2014). Closer to home, more fun with famous fictitious monsters can be found here, and for more on movie creatures - specifically those of special effects legend Ray Harryhausen, check out this.


  • Christiansen, P. (1999). On the head size of sauropodomorph dinosaurs: implications for ecology and physiology. Historical Biology, 13(4), 269-297.
  • Cotta Vaz, M. (2014). Godzilla: the Art of Destruction. Titan Books.
  • Habib, M. B. (2008). Comparative evidence for quadrupedal launch in pterosaurs. Zitteliana, 159-166.
  • Rayner, J. M. (1988). Form and function in avian flight. In Current ornithology (pp. 1-66). Springer US.
  • Schoch, R. R. (2009). Evolution of life cycles in early amphibians. Annual Review of Earth and Planetary Sciences, 37, 135-162.
  • Trimarchi, J. R., & Schneiderman, A. M. (1995). Initiation of flight in the unrestrained fly, Drosophila melanogaster. Journal of Zoology, 235(2), 211-222.
  • Van Valkenburgh, B. (1990). Skeletal and dental predictors of body mass in carnivores. In: Damuth, J. and MacFadden, B. (eds). Body size in mammalian paleobiology: estimation and biological implications, 18, 1-205.
  • Witton, M. P., & Habib, M. B. (2010). On the size and flight diversity of giant pterosaurs, the use of birds as pterosaur analogues and comments on pterosaur flightlessness. PLoS One, 5(11), e13982.
  • Zumstein, N., Forman, O., Nongthomba, U., Sparrow, J. C., & Elliott, C. J. (2004). Distance and force production during jumping in wild-type and mutant Drosophila melanogaster. Journal of experimental biology, 207(20), 3515-3522.