Quetzalcoatlus: the media concept vs. the science

Background: Javelina Formation forest. Mid-ground: the 4.6 m wingspan, super-famous azhdarchid pterosaur Quetzalcoatlus sp. Foreground: lunch.

As a consultant, the pterosaur I get asked about more than any other is the azhdarchid Quetzalcoatlus. In recent weeks I've been speaking to two completely independent media producers about this animal, and I think just about all my prior TV and film work has involved it somehow, even if not prominently. I suppose Quetzalcoatlus is so popular because it's not just the most famous azhdarchid pterosaur - which are now more popular than ever - but the first animal most people think of as the biggest ever flying creature.

All the pictures, museum exhibitions, sculptures and animations of Quetzalcoatlus suggest it must be a well-understood animal, but it's actually very difficult to provide consultancy on, for several related issues. The first is that the science on the animal itself is unfinished, largely unpublished, and the existing body of work is decades old. The second is that our general understanding of azhdarchid pterosaurs has moved on considerably in the last two decades, and largely without a good grounding of what Quetzalcoatlus actually is. The third, and final, issue is that most parties think Quetzalcoatlus is better understood than it really is, to the point where whole media projects are locked in around it, and consultants are asked to make calls that have little scientific backing. This third point is an important one: 'Quetzalcoatlus the media concept' is a very different beast to 'Quetzalcoatlus the scientific entity'. This can create difficulties when creating programmes, games or artwork of this animal, as the expectations for what can be achieved with Quetzalcoatlus are often beyond what scientists can provide.

In the interests of helping out those who want to use Quetzalcoatlus in their projects, I thought it might be useful to show how 'common knowledge' about Quetzalcoatlus differs from its actual, objective status within 21st century pterosaur science. An important caveat before we go further is that scientific work on Quetzalcoatus is ongoing. A full description and functional assessment is rumoured to be in the works (as, er, it has been for 40 years...) and this will change the way we view this animal tremendously. It can't fail to do so, as the data available on this animal are minimal and the publication of any new details will augment our current situation significantly. In all likelihood this post will be moot, at least in part, when this document is finally published. This piece is being written in May 2016, so please bear in mind that things might have changed if you're reading this at a later date.

The concept. Quetzalcoatlus is a giant azhdarchid pterosaur from Texas, known from substantial remains.

The science. The popular view of Quetzalcoatlus is really a conflation of two taxonomic entities, Quetzalcoatlus northropi and Quetzalcoatlus sp. Both occur in the Upper Cretaceous (Maastrichtian) Javelina Formation of southern Texas*. Q. northropi is currently the only 'officially' named species of Quetzalcoatlus, and is the reason we think this animal was so huge. It's also only known from bits and pieces of a gigantic left wing and cannot be regarded as a well known animal. Do not, if you're a TV producer or whatever, expect to film a room full of real fossils for this thing - any skeleton you see of a giant Quetzalcoatlus is almost entirely reconstruction - still impressive, but nearly all calculation and extrapolation.

*It's worth mentioning that Quetzalcoatlus has been identified outside of Texas, scraps of azhdarchid bones from both North America and Europe being allied to this genus. I suspect that most of these should not be referred to Quetzalcoatlus proper, as they're fairly 'generic' azhdarchid bones and, moreover, we have no criteria for what constitutes Quetzalcoatlus within Azhdarchidae itself (see below). I don't want to discuss these now, however, and only mention them for the sake of completeness. 

Q. sp., by contrast, is much better represented. Several incomplete skeletons are known (an example can be seen below) that collectively give a near complete picture of Q. sp. osteology. Its this material that gives us our familiar image of Quetzalcoatlus: the long neck, the oversize pointy face, the long limbs and so on. Almost all discussions about the detailed anatomy of Quetzalcoatlus pertain to this material, not the giant wing. The Q. sp. fossils are also a key source of the proportional data used in calculating the 10 m wingspan for the giant Q. northropi animal, even though Q. sp. is much smaller - the complete wing metrics of one specimen give a wingspan of 4.6 m (Unwin et al. 2000).

Q. sp. partial skeleton figured by Kellner and Langston (1996).

How the two Quetzalcoatlus species are related to each other, and other azhdarchids, remains critically unexplored and uncertain. It is no exaggeration to say that taxonomic considerations of this important, well-known genus comprise no more than a few sentences in the entirety of technical pterosaur literature. The name Q. northropi was not even erected in a descriptive paper, but as an aside in a brief comment on the likely wingspan of the giant wing specimen (Lawson 1975a). This gave us the northropi name and designated a type specimen (the big wing), and suggested that the smaller specimens were just diminutive, presumably immature versions of the giant species. However, other elements key to species creation - diagnoses, specimen inventories, supporting description or illustrations were not provided, the best alternative being a very short 1975 science paper (Lawson 1975b). Nessov (1991) provided an attempt to diagnose the genus (along with other azhdarchids) but his characters are not useful. Later, Kellner and Langston (1996) suggested that the smaller Quetzalcoatlus skeletons were not juveniles of the giant species, but a distinct species that would be named at a later date. It was this paper that created the 'Q. sp.' moniker, a place-holder for a name we'll perhaps see published in time. Alas, Q. sp. was also created using drive-by taxonomy without justification for separating sp. from northropi, or providing characters to unite these species in a distinct genus among other azhdarchids.

This might all sound like tedious detail irrelevant to reconstructions, media portrayals etc., but the upshot is that the pterosaur community is still largely in the dark about Quetzalcoatlus. We can't really comment on what makes it unique, whether all these bones from Texas should be considered one or two species, how accurate it is to scale up the smaller Quetzalcoatlus to the size of the big wing and so on: that information has not been made public yet. Some specific folks might be able to provide those details, but they are not peer-reviewed 'common knowledge'. Therefore, most researchers (including myself) can only talk about Quetzalcoatlus in terms of the few details that have been published, and fill the rest in with 'generic' details of azhdarchid pterosaur palaeobiology.

The concept. Quetzalcoatlus was like all azhdarchids: a long-necked, long-skulled creature with long limbs.

The science. As alluded to above, we have a basic idea of Quetzalcoatlus sp. appearance, even if the vast majority of it remains unpublished. The core aspects of this animal can be built from data titbits gleaned from different papers: a good description of the skulls and mandibles was provided by Kellner and Langston (1996); Witton and Naish (2008) and Steel et al. (1997) gave some details of the cervical vertebrae, and Unwin et al. (2000) provided measurements for nearly all major limb bones. From this, we can be confident that Quetzalcoatlus sp. is the long-necked, long-faced, toothless, short-winged and gracile-limbed creature we have traditionally associated with the Quetzalcoatlus name. I've attempted a skeletal reconstruction of this 4.6 m wingspan species below using these data: some of the bone anatomy is 'generic azhdarchid', but the basic proportions, skull and anterior neck vertebrae should be OK. This skeletal is the basis for the painting at the top of the post.

Quetzalcoatlus sp. skeletal, using data from Kellner and Langston (1996); Witton and Naish (2008), several other sources of azhdarchid neck data, and Unwin et al. (2000). Yes, it was that long-necked and long-legged.

A question I have much more trouble answering is what Q. northropi looked like. When we see a giant long-necked Quetzalcoatlus in TV shows or comics, we're seeing a hybrid of the two Quetzalcoatlus animals - the anatomy of sp. crossed with the size of northropi. This approach is not without merit: it's consistent with the existing taxonomy of this animal (such that it is), and other pterosaur fossils confirm that some giant azhdarchids were long-necked, perhaps Q. sp-like creatures.

On the other hand, we only have an incomplete northropi wing skeleton to work from. It's widely recognised that pterosaur wings are among the more conservative aspects of their anatomies - great for identifying pterosaurs to specific groups (we have characterised azhdarchid wings, for instance) but above a basic level of taxonomy they don't tell us much about life appearance. It wasn't always this way. When Quetzalcoatlus was found in the 1970s the smaller Q. sp. skeletons provided our only comprehensive insight into azhdarchid anatomy, and thereafter we assumed that Q. sp. typifies the group. However, azhdarchid pterosaur science has progressed considerably in the last two decades and the group can no longer be considered anatomically uniform. Their skulls can be short and broad, long and narrow, and have deep or slender lateral profiles (Witton 2013). They can have cranial crests (Kellner and Langston 1996), but they might not (Cai and Wei 1994). Their necks can be extremely long, or of more typical pterodactyloid lengths (Vremir et al. 2015). Some had stilt-like limb bones, but others had short forelimb anatomy (McGowan et al. 2001). Such variation seems present in the giants as much as their smaller cousins. I don't think we know what an 'average' azhdarchid looked like yet, and Q. sp. should be viewed as representing only one, relatively exaggerated take on azhdarchid anatomy. It historically seemed safe to make Q. sp. a giant version of this form, but that cannot be regarded as the only option today. For all we know, Q. northropi could be a short-armed, short-necked species with a truncated, deep jaw - quite the opposite of Q. sp..

A selection of azhdarchid skull and mandibles. A and B, posterior skull bits of Hatzegopteryx; C-D, Q. sp., E. Zhejiangopterus; F-G, Bakonydraco; H, TMM 42489-2, the Javelina Formation azhdarchid which isn't Quetzalcoatlus. Q. sp. must be regarded as a long-snouted, gracile form: does this also apply to Q. northropi? Image from Witton 2013.

I would be more comfortable with reconstructions of Q. northropi as a giant, scaled-up Q. sp. if we had good reason to believe the two were closely related**. As mentioned above however, necessary work on this has yet to be presented and, as evidence-led scientists, it is not unreasonable to call the situation ambiguous until more data is forthcoming. I suppose one reason we might think northropi and sp. are congeneric is because they're from the same formation. However, the remains of northropi and sp. were not associated, being found tens of kilometres apart (Lawson 1975b), and we have increasing evidence of multiple azhdarchid taxa occurring in the same geological units (e.g. McGowen et al. 2000; Godfrey and Curry 2005; Vremir et al. 2013). Where azhdarchids do coexist, they differ markedly in anatomy and overall form (Vremir et al. 2013). An additional complication is TMM 42498-2, a large, deep-jawed Javelina Formation azhdarchid which is clearly not Q. sp. (bottom panel in the image above). If this represents cranial remains of Q. northropi and not an additional Javelina species (we currently have no way of telling), northropi would look would look very different to our usual depictions.

**I'm expecting people to wonder if Quetzalcoatlus taxonomy has been looked at in detail via cladistic methods. The most complete published assessments I'm aware of are those by Brian Andres (e.g. Andres and Myers 2013) which use seven azhdarchid OTUs, including the two Quetzalcoatlus taxa. The results of such studies remain ambiguous about the affinities of Quetzalcoatlus (Q. sp. and northropi form a polytomy with Arambourgiania). I'd like to see an analysis with more azhdarchid taxa, including some of the more unusual types such as Hatzegopteryx and Montanazhdarcho.

I don't have any real answers or insight on these points. Rather, I'm getting at the fact that our 21st century understanding of azhdarchids and other flying reptiles is becoming complicated, and with our current, superficial insights into what Quetzalcoatlus is and how it's related to other azhdarchids, there may not be one 'right' way to restore northropi. We might be correct to represent it as a giant Q. sp., as per tradition, but we might not.

The concept. Quetzalcoatlus was the biggest animal to have ever flown, even larger than other giant azhdarchids

The science. As with modelling the size of most giant extinct animals, it's difficult to say what giant azhdarchid was the biggest. Q. northropi was certainly up there, recent estimates of its wingspan being in the region of 10 m and predicted body masses of 200-260 kg. But other giant pterosaurs are predicted to be about the same size (see Witton and Habib 2010 for a recent discussion)... and that's about all we can really say. Every giant azhdarchid is represented by scrappy material, so the error bars on any size estimate are large and the calculations themselves are highly sensitive. We would be foolish to use them as anything other than ballpark figures. We can say that Q. northropi was one of the biggest flying animals of all time and, along with other giant azhdarchids, it dwarfed other flying species including most pterosaurs, and all birds and bats. I appreciate some folks will find this lack of clarity frustrating, but that's just how it is: we can't say any more until we understand all the giants - not only Quetzalcoatlus - in more detail. 

The concept. The outstanding flight capabilities of giant azhdarchids allowed Quetzalcoatlus to be a continent-hopping animal that occurred in Europe as well as the USA, although the European bones were given a different name: Hatzegopteryx. Some chap called 'Witton' suggested this.

The science. Thanks, I think, to Wikipedia, I've been confronted several times about suggesting the Romanian giant azhdarchid Hatzegopteryx thambema should be synonymised with Quetzalcoatlus. I feel a bit misquoted on this. Yes, I (and colleagues) mentioned this as a qualified possibility in a 2010 abstract and conference talk (Witton et al. 2010), but as part of a broader, detailed discussion about the need to tighten up giant azhdarchid taxonomy. Specifically, we discussed the three named giant azhdarchids - Arambourgiania philadelphiae, Quetzalcoatlus northropi, and Hatzegopteryx thambema - and stressed issues with their current taxonomy. These issues - hitting some points already tackled here - included the lack of a description and diagnosis for Quetzalcoatlus; the uncertainty over the relationships between Q. northropi and Q. sp.; the scrappy nature and general incomparability of giant azhdarchid fossil remains; their representation by anatomies generally considered unnameable by pterosaur workers, and the identification of several alleged autapomorphies of giant species in other azhdarchid remains.

Holotypes of giant azhdarchids. A, Arambourgiania philadelphiae, B, Q. northropi (humerus only, the other holotype wing elements have never been published) C, the damaged proximal humerus of Hatzegopteryx thambema (humeral head and cranial remains not figured here). From Witton (2013).

Concerning Quetzalcoatlus and Hatzegopteryx, we pointed out that overlapping bits of Q. northropi and Hatzegopteryx (proximal humeri) are similar enough that they could be considered synonymous, but this says more about the use of wing bones for the Q. northropi type material than anything else. As mentioned above, wing bones aren't always that useful in detailed taxonomy. The overlapping bits of Hatzegopteryx and Q. sp. (jaw joints), however, are different enough to demonstrate they are separate taxa. We went on to say that the significance of all this isn't clear because the relationships between Q. sp and Q. northropi are not evaluable at this stage. Our take-home message, then, wasn't that Hatzegopteryx and Quetzalcoatlus are the same thing, but that the diagnoses, validity and relationships of named giant azhdarchids warrant detailed assessment in future. Since writing this abstract six years ago, ongoing work on Hatzegopteryx seems to be supporting its separation from other azhdarchids - more on that in time.

So... how can Quetzalcoatlus be used in art, film, games etc.?

It's clear by now that the way we imagine and depict Quetzalcoatlus as a media construct is very different to its status in science. My take on this animal is about as cautious and conservative as you'll find, and I suppose that's because my experience with azhdarchids in both a research and artistic capacity has frequently found Quetzalcoatlus as a tricky animal to work with. While we can't point to anything being 'wrong' with those older interpretations of Quetzalcoatlus, shifts in our understanding of azhdarchid and other pterosaur science means we can't accept those 40- or 20-year old interpretations of this animal with the same confidence as we used to. Quetzalcoatlus, as a scientific concept, needs modernisation.

Still, my overall goal here is not to be defeatist, rather to simply say what we might and might not be confident about when depicting Quetzalcoatlus. For instance, while Q. northropi is a can of palaeobiological worms, Q. sp. (or whatever it turns out to be) does offer a lot of scope for use in reconstructions and media projects. I frequently feel that these sorts of animals (i.e. the better known mid-sized or small species, not the specrapularly known giants) make interesting enough subjects for artwork and media projects. There's certainly a lot more to work with and say about them, and we can be far more confident in what is being conveyed to the public. Not everything with palaeontological content has to focus on the biggest animals!

Of course, I also appreciate that some projects just can't do without giant azhdarchids. For these, I stress that there are alternatives to Q. northropi which boast the same approximate wingspan, are known from anatomies that provide more insight into life appearance, and have a better grounding in contemporary science. These include the long-necked Jordanian form Ararmbourgiania and its Romanian contemporary, the wide-skulled, robustly-built Hatzegopteryx. Both have interesting stories of discovery and scientific development (for instance, Arambourgiania was actually the first giant azhdarchid on record, not Quetzalcoatlus; and Hatzegopteryx is so chunky that it was initially interpreted as a giant predatory dinosaur) and are palaeobiologically interesting. If these won't do - perhaps for reasons of geography - then consider that unnamed bones of giant azhdarchids are widespread, being known from North America, Europe, Africa and Asia. Although these offer fewer anatomical details than the named taxa, they can also be handled more generally because they don't need to look like a (theoretically) well understood, diagnosed and named species. This means data can be pooled from other finds into a more 'generic azhdarchid' melting pot, and that gives more wiggle room when considering appearance and form.

The take-home here, then, is that Quetzalcoatlus might be the 'best known' giant azhdarchid and the one that everyone wants to feature in their TV shows, films and games, but be forewarned that the scientific status of this animal is rather different to what popular depictions suggest. Moreover, there are alternatives which might be (at time of writing) better understood and just as interesting. If you're planning a TV show, video game or artwork of a giant azhdarchid, remember that there are choices other than the obvious.

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References

• Andres, B., & Myers, T. S. (2012). Lone star pterosaurs. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 103(3-4), 383-398.
• Cai, Z., & Wei, F. (1994). On a new pterosaur (Zhejiangopterus linhaiensis gen. et sp. nov.) from Upper Cretaceous in Linhai, Zhejiang, China. Vertebrata Pal Asiatica, 32(3), 181-194.
• Godfrey, S. J. & Currie, P. J. (2005). 16. Pterosaurs Dinosaur Provincial Park: A Spectacular Ancient Ecosystem Revealed, 1, 292.
• Kellner, A. W., & Langston Jr, W. (1996). Cranial remains of Quetzalcoatlus (Pterosauria, Azhdarchidae) from Late Cretaceous sediments of Big Bend National Park, Texas. Journal of Vertebrate Paleontology, 16(2), 222-231.
• Lawson, D. A. (1975a). Could pterosaurs fly?, Science, 188: 676-678
• Lawson, D. A. (1975b). Pterosaur from the Latest Cretaceous of West Texas. Discovery of the Largest Flying Creature. Science, 187: 947-948.
• McGowen, M.R.; Padian, K.; de Sosa, M.A.; Harmon, R.J. (2002). "Description of Montanazhdarcho minor, an azhdarchid pterosaur from the Two Medicine Formation (Campanian) of Montana". PaleoBios 22 (1): 1–9.
• Nessov, L. A. "Giant flying reptiles of the family Azhdarchidae. I. Morphology, systematics." Vestnik Leningradskogo Universiteta, Seriya 7, no. 2 (1991): 14-23.
• Unwin, D. M., Lü, J., & Bakhurina, N. N. (2000). On the systematic and stratigraphic significance of pterosaurs from the Lower Cretaceous Yixian Formation (Jehol Group) of Liaoning, China. Fossil Record, 3(1), 181-206.
• Vremir, M., Kellner, A. W., Naish, D., & Dyke, G. J. (2013). A new azhdarchid pterosaur from the Late Cretaceous of the Transylvanian Basin, Romania: implications for azhdarchid diversity and distribution. PLoS One, 8(1), e54268.
• Vremir, M., Witton, M., Naish, D., Dyke, G., Brusatte, S. L., Norell, M., & Totoianu, R. (2015). A Medium-Sized Robust-Necked Azhdarchid Pterosaur (Pterodactyloidea: Azhdarchidae) from the Maastrichtian of Pui (Hateg Basin, Transylvania, Romania). American Museum Novitates, (3827), 1-16.
• Witton, M. P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.
• 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., Martill, D. M. and Loveridge, R. F. 2010. Clipping the wings of giant pterosaurs: comments on wingspan estimations and diversity. Acta Geoscientica Sinica, 31 (1), 79-81.

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 I'll be covering in a couple of weeks.

PteroPop

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. These historic popular pterosaurs 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). They were depicted with huge variation from the 1970s-1990s: hugely elongate and narrow wings, or broad, kite-like wings? Longirostrine skulls with snub-noses or tiny pin-heads? Toothed or toothless jaws? Short necks, long necks with swan-like flexibility, or long necks stiffer than broom handles? 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) remained unpublished (and 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 just also happened to be giants. 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 indoctrination of a new lineage into the mainstream palaeontological 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 made them more marketable and appealing in short time. And 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; visually interesting and striking; instantly recognisable, as well as being gigantic: 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 - let's face it, 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 a consistent palaeo pop ‘character’, but also didn't give them much popular appeal.

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: the concept of 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!

References

• 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.

The Walking with Dinosaurs 3D trailer lands

Good news, everyone: the PR machine behind Walking with Dinosaurs 3D has provided our first detailed look at the film with its first trailer. I'm rather excited about this because it's my first real look at the film since contributing character orthographics for several (six, specifically) of its animals in early 2011, a process that involved working with a fleet of palaeontological and palaeoartist supernauts: David Krentz, Scott Hartman, Tom Holtz, Scott Sampson, Luis Chiappe, Victoria Arbour and others. Since that time, the guys at Animal Logic have been crunching the ones and zeroes required to breath life into the characters and their world, with the results posted above. There's always an air of uncertainty surrounding palaeontological film projects but, happily, this first trailer suggests WwD3D is taking good shape, and the Facebook and Twitter buzz from the contributors mentioned above suggests they're pretty happy with what they've seen, as am I. Be sure to set the trailer stream to its highest, 1080p HD quality to take in all the details.

Highlights of this first teaser, aside from some excellent zbrush creations from David Krentz and the stunning visuals and animation, include feathered dromaeosaurs, an iridescent gorgosaur and suitably snowy slopes in latest Cretaceous Alaska. I like the efforts to characterise individual animals without resorting to crazy variations in colour and form, such as the hole in the frill of the hero Pachyrhinosaurus. Presumably, this reflects a pathology that never entirely healed, and it's a cool idea. So cool, in fact, that I'm wondering why it's not more common in palaeoart. We can also see that the colour schemes of the animals are striking without being outlandishly garish, and the characters seem capable of emoting without using overblown actions akin to those of other CG dinosaurs, who seem to have gone to an acting school for 1920s silent movies. Moreover - stop press - plenty of shots even show animals not roaring and with their mouths closed. Gosh.

So, there's plenty to be happy about then, and my feeling from reading the comments of others is that I'm not alone in this view. Perhaps the only common niggle being raised is that the tyrannosaur is scaly rather than feathered, which jars with recent discoveries showing some big tyrannosauroids were covered in protofeathers (Xu et al. 2012). Obviously, it would have been extremely cool to render this in WwD3D, but the feathered tyrant Yutyrannus arrived just a little too late in the day for the design team to work feathers onto the Gorgosaurus. As hinted above, the creature designs for the WwD3D animals were being set in late 2010 and early 2011, well over a year before Yutyrannus was unleashed on the world. And we shouldn't be too upset: there is still controversial evidence that another tyrannosaur, Tyrannosaurus, was scaly, and the WwD3D Gorgosaurus still looks awesome.

But enough about dinosaurs: what about the real stars of the show, the pterosaurs? It may not be surprising that I had a hand in design and consultancy for the movie's azhdarchids, and I'm happy to say that I like what I can see thus far. Flap-gliding and full-on flapping flight are both on display, and just short of the one minute mark, we see a gang of azhdarchids surround and aggressively 'terrestrially stalk' the hero animals. If you're sad enough to have freeze-framed high-definition shots of the azhdarchids (which, er... I didn't, but a friend of mine did) you'll notice thick coats of soft pycnofibres, and that the internal anatomy of azhdarchid jaws have, for the first time, been accurately rendered on film. Indeed, the contours of the head are the best I've ever seen in a CG azhdarchid. Note the concave lateral skull facia, the tapering shape of the tomial margins... wonderful stuff. I'm very much looking forward to seeing them on screen later in the year.

So, this is all shaping up very well then, making the December 20th release date something to look forward too. It almost seems that fans of Mesozoic reptiles on film are being spoilt at present, with WwD3D following 2011's terrific (and underrated) Dinosaur Revolution/Dinotasia and the excellent BBC series Planet Dinosaur. Indeed, the BBC's dinosaur coverage is on something of a roll at present. Presumably as part of the PR campaign for the new film, its revamped Walking with Dinosaurs website has been providing good coverage of recent dinosaur topics in a way that is extremely accessible but not dumbed down (for a good example of this, see their coverage of the Torosaurus/Triceratops debate). Several well-known contributors to the palaeo blogosphere have been contributing new content to the site too, so it's definitely worth checking out if you haven't already.

And that will have to do for now: lots of work to catch up with. In the meantime, I leave you, and particularly if you're a 20th Century Fox or BBC executive, with a pitch for a WwD3D spin-off, starring the animals I'm sure will steal the entire movie. It's Box Office Gold, I tell you.

Reference

• Xu, Xing, Kebai Wang, Ke Zhang, Qingyu Ma, Lida Xing, Corwin Sullivan, Dongyu Hu, Shuqing Cheng, and Shuo Wang. 2012. A gigantic feathered dinosaur from the Lower Cretaceous of China. Nature, 484, 92-95.

"There's something in the mist!"

One of the most effective movies I've seen in recent years is the  2007 science fiction siege thriller, The Mist, which isn't to be confused with the considerably more forgettable eighties pirate horror outing, The Fog (which has one of the most long-winded and least suspenseful trailers ever). Like all good stories, the plot of The Mist is dead simple, with a mysterious, creature-filled fog mist descending on a small Maine town and trapping a small population of people within a supermarket, who proceed to bicker about the best response to their crisis, descend to the level of primitive savages, and throw cans of peas at each other. It also has one of the most killer endings of a movie I've ever seen. I won't ruin it for you here, but it's a terrific antidote for the sickly-sweet, rosy endings slapped onto films at the last minute to please test screens. 1986 Little Shop of Horrors, A.I., and the original cut of Blade Runner: I'm looking at you. And all your pansy-ending friends.

One of the best things about The Mist is that Frank Darabont, it's director, knew exactly what to do with his monsters, in that he shows as little of them as possible. Instead, most species (and there are many) are glimpsed in silhouette at the far reaches of the fog mist, so we're never really sure what they look like or what they might be capable of. This, and the handling of the bickering survivors in the supermarket, turns a very pulpy plot into an intelligent and tense movie, and I heartily recommend you track it down if you've not seen it.

The handling of the creatures in The Mist got me thinking about how bizarre, and perhaps how terrifying, the silhouettes of barely-glimpsed creatures in the Mesozoic may have looked on misty, drizzly days. It's not hard to think of a number of Mesozoic animals that would look downright weird when glimpsed through thick fog, but the outlandish proportions of giant azhdarchid pterosaurs, with their long necks and limbs, oversize heads and small bodies, make them unlikely animals at the best of times and truly strange when only seen in outline. The lightweight frames and long limbs of these animals would probably make them lithe and quick over land, and it's not hard to imagine these enormous carnivores giving many Mesozoic animals the hebbie-jebbies as they stalked silently across Cretaceous landscapes. And not just little animals, either: prey of human dimensions may also have been alarmed by a stealthy, fog-strewn azhdarchid.

With this in mind, I've been slowly adding to the painting above for the last few weeks in a very piecemeal, 5-minute burst fashion. It's a deliberately basic image with a limited colour palate and detail, which came from both a desire to attempt something a bit different with my art and enhance the dreary atmosphere. The intention was to make the pterosaur in the background looming and menacing, all but invisible in the dismal weather save for its treetop-scraping silhouette. The small azhdarchid in the foreground, who's giving its big relative an understandably wide berth, serves to add scale, as does the rotting log in the foreground. The taxa here are not meant to be any azhdarchids in particular, but this scene could take place in several places around the Late Cretaceous world. As pointed out by Matyas Vremir et al. (2013), several azhdarchid-bearing deposits yield azhdarchid species of vastly contrasting size, suggesting giants and diminutive species frequently lived alongside one another (see summary image, below, of cohabiting azhdarchids of distinct size, from Vremir et al. [2013], featuring Ron Blakey's fantastic latest Cretaceous palaeomap (Colorado Plateau Geosystems, Inc.), and my handiwork. The image features a new skull reconstruction of Quetzalcoatlus sp., which offers a sneaky peak into some of the imagery used in my book). The scant record we have of these cohabiting species indicates that at least some bore distinct jaw and skull proportions, suggesting different dietary preferences and habits which would prevent them stepping on each other's ecological toes.

Some geological units reveal evidence of two or even three sympatric azhdarchid species. Diagram produced by Mark Witton and map used with kind permission of Ron Blakey, Colorado Plateau Geosystems, Inc; from Vremir et al. (2013).

The result, hopefully, is something a little different and interesting, and I definitely see potential for more things like this in the future. That's all for now, though: got to head out to be social. Next week, or perhaps the week after, should see something fairly special in these quarters, so be sure to check back soon.

Reference

• Vremir, M., Kellner, A. W., Naish, D., & Dyke, G. J. (2013). A new azhdarchid pterosaur from the Late Cretaceous of the Transylvanian Basin, Romania: implications for azhdarchid diversity and distribution. PloS one, 8, e54268.

The Rise and Fall of Ziggy BigQuetz

Back in 2010, when I was employed in building a series of giant pterosaur models for the University of Portsmouth and Royal Society, I painted the above image of several giant azhdarchids in flight for use on our display boards and advertising work (bonus sauropods can be seen playing in the water below the pterosaurs). The azhdarchids are meant to be a fairly close match for our models, and specifically the giant, 10 m wingspan jobby we suspended between Royal Festival hall and the neighbouring buildings:

Our 2010 exhibition, with BigQuetz at the topright. Two smaller giants follow behind it, with the giant Bamofo looking on. Pterosaur worker Michael O'Sullivan can be seen in the bottom left. 

BigQuetz in all its 10 m span glory

Our giant model had a couple of goofs that I wish we could have avoided, but was far from the worst rendition of a giant pterosaur I've ever seen. Certainly, there was no doubting its enormous size, which was substantial enough to shade a gathering of people escaping the unusually intense London sun experienced in June/July 2010. Alas, impressive as it was, it's size proved to be problematic for transportation and storage purposes. For engineering reasons, the body and wings of our BigQuetz were constructed as a single, solid unit, which meant that it had to be transported on a long-base low loader and, after its display in London, finding places to store and display it proved very, very difficult.

The BigQuetz body and wing frame, with pterosaur workers for scale (I'm on the left, Dave Martill is on the right)

The upshot is that BigQuetz, after a short stint on display in the Netherlands, had to weather being stored outside the School of Earth and Environmental Sciences in the open campus of the University of Portsmouth. Dave Martill, our pterosaur model wrangler, tried repeatedly to offload the model to various institutions but, despite finding homes for the rest of our giant models, could not find anyone willing to take on the transportation costs and space requirements of BigQuetz, even if the model was otherwise theirs for the taking. We were all very aware that BigQuetz was vulnerable in being stored in this way and, indeed, the ravages of weather and occasional shunts around were begging to take a toll on its more delicate aspects. But, as is often the case, the biggest problem was protecting it from people. It was only a matter of time before someone found BigQuetz and started fooling about with it, and thus inevitable that it would be found, one day, with significant damage.

Sadly, that very day arrived at the turn of this year, when University of Portsmouth staff found the BiqQuetz head  had been smashed in by unknown individuals (sadly, I don't have any photos to show of this). It seemed that the damage was well beyond repair, and the BigQuetz story was brought to a close when the model was chopped up and disposed of. It's a terrific shame that our model should meet such an inauspicious end, and particularly stings because I personally experienced the many hours of work that were poured into its creation, often by volunteers, and know of the craftsmanship that went into its production, particularly on its aluminium frame. Plus, I find it hard to rationalise its demise being caused by anything other than a stupid stunt pulled by bored delinquents, seeing as its location on campus was not conducive to being accidentally damaged by university staff. Some consolation can be found in the fact that our other models have found homes elsewhere, and are hopefully being looked after, but it still seems a tremendous waste. Hey ho. RIP BigQuetz, we hardly knew ye. Or whatever people say about this sort of thing.