9 things you may not know about giant azhdarchid pterosaurs

The 2015 version of the giraffe vs. azhdarchid vs. person image, now in it's fifth iteration (see the general history of these images through the years: 2006; 2007; 2009). The giraffe is a big bull Masai individual, standing a healthy 5.6 m tall, close to the maximum known Masai giraffe height. The pterosaur is a 10 m wingspan Arambourgiania philadelphiae (for reasons I cannot go into now, it is not wise to consider the appearance of giant azhdarchid taxa interchangeable any more: this should not be considered Hatzegopteryx thambema or Quetzalcoatlus northropi). The Disaknowlegement provides the human touch. These characters will receive some additional company soon.

The splendid beasts that are giant azhdarchid pterosaurs have occupied my thoughts a lot of late, mostly thanks to three upcoming talks I'll be delivering about them at upcoming conferences and society meetings. Preparing that number of talks in a short space of time has given me a whole new interpretation of the term 'death by PowerPoint' so, to take a break from animating slides and producing diagrams, here's a quick run down of 10 factoids you may not know about giant azhdarchid pterosaurs (the likes of Quetzalcoatlus northropi, Hatzegopteryx thambema and Arambourgiania philadelphiae - as if they need introduction) and their smaller relatives. Even if these facts are familiar, please feel free to enjoy the new bits of artwork accompanying the post. Those of you really into the ever evolving depiction of these pterosaurs really should also pay a visit to this recent Tetrapod Zoology post: I'm clearly not the only one with giant pterosaurs on the brain.

They nearly weren't called 'azhdarchids'
'Azhdarchidae' is a terrific name. It's short but mysterious, relatively easy to spell, and PR friendly enough that even the British tabloid The Sun has used the term on at least two (I think) occasions. The name 'Azhdarchinae' was coined by the late Lev Alexandrovich Nesov in 1984 from the Uzbek word 'azhdarkho', a name for a mythical dragon, and also the nomenclatural basis for the medium-sized Uzbek azhdarchid Azhdarcho lancicollis. Nesov's name encompassed all three azhdarchid genera known at that time: Azhdarcho, Titanopteryx (now known as Armabourgiania) and Quetzalcoatlus. Almost simultaneously, however, the exact same set of taxa was being roped into another group by Kevin Padian, which he termed Titanopterygiidae after, obviously, Titanopteryx. Nesov's 'Azhdarchinae' pipped the far-less elegant Titanopterygiidae to the publishing punch by a matter of months, and took nomenclatural priority for the group. Padian elevated Azhdarchinae to 'family' level in a short note in 1986, giving us our now familiar term, 'Azhdarchidae'.

Lev Alexandrovich Nesov holds the fossil cervical vertebra, notarium and jaw tip of the azhdarchid Azhdarcho. Image from Unwin (2005).

Tiny bodies
Despite their giraffian proportions, giant azhdarchid torso were relatively tiny. Witton and Habib (2010) noted that, like many pterodactyloid pterosaurs, their torsos were probably only a third or so longer than their humeri, suggesting a shoulder-hip length of about 65-75 cm for an animal with a 10 m wingspan. That's a torso length not much larger than your own, although they were considerably more stocky and swamped with muscle. Azhdarchid shoulders, in particular, are well endowed with attachment sites for flight muscles, as are (for pterosaurs) their pelves and hindquarters.

Giant azhdarchids did not suffer from flight power shortages
Many internet commenters often roll out the idea that giant azhdarchids would struggle to take off from the ground, even allowing for new ideas like quadrupedal launching. These folks need to get out of their armchairs, however, and check out some classic work on animal flight and giant pterosaur takeoff. James Marden's 1994 work on animal takeoff found some surprisingly consistent scaling trends among animal flight power and takeoff ability, allowing us to predict the muscle power of even long extinct fliers like Meganeura, Archaeopteryx and a 10 m span azhdarchid. The resulting aerobic power output of azhdarchid flight muscles - all 60 kg of them (a fairly safe bet for a 250 kg azhdarchid given what we know of animal flight muscle fractions among modern fliers) - is a bit rubbish, only 4.52 N/kg of body weight. Animals need to be generating 9.8 N/kg to fight gravity, so this would seemingly ground our giants. Bear in mind, however, that swans, albatross, vultures and turkeys also have aerobic power outputs of around 4.5 N/kg from their flight muscles, and they can fly just fine. The secret to their takeoff lies in the great power of anaerobic muscle contraction, which provides twice the power achieved under aerobic regimes. Using anaerobic power, giant azhdarchid power outputs are 10.098 N/kg of body weight, a value surpassing the 9.8 N/kg and matching the anaerobic power outputs of a 10 kg swan or 1 kg vulture (see graph, below). In terms of power availability, then, giant azhdarchids would not have struggled to launch any more than a large bird, so all these suggestions about poor takeoff ability and whatnot can be put to bed.

Scaling of flight performance with body size under anaerobic power output. The dashed line is the minimum lift needed to overcome gravity. Anaerobic power is 225W/kg, the upper limit of avian anaerobic output. From Marden (1994).

An unsurpassed 80 million years of evolutionary history, and growing
Azhdarchids are undeniably best known from Upper Cretaceous rocks, but they also have a patchy and sometimes controversial Lower Cretaceous record. Recently, Gareth Dyke and colleagues (2011) demonstrated that the group were probably present at the very base of the Cretaceous, in Berriasian (c. 140-145 Ma) deposits of Romania. Given that azhdarchids are definitely present at the final stage of the Cretaceous, this gives the group a stratigraphic record spanning the entire Cretaceous: 80 million years in total. This is longer than any other pterosaur group. Two cervical vertebrae from the Late Jurassic of Africa may extend their temporal range another 5 million years, although the affinity of these specimens remains controversial.

A much improved skeletal reconstruction of the small azhdarchid Zhejiangotperus linhaeiensis over my oft-reproduced effort from Witton and Naish 2008. Note the use of pacing strides, a gait indicated by pterosaur trackways but seldom seen in pterosaur palaeoart.

More than just long necks
When we describe azhdarchids, we often use two qualifiers: 'toothless' and 'long-necked'. In fact, these pterosaurs are brimming with characterising features (above). Their rostra are particularly elongate compared to all other pterosaurs, their orbits are depressed well into the lower half of their skulls, their wing metacarpals and femora are atypically long, and their extremities are short and robust. Their mid-series cervical vertebrae are famously simplified into almost tube-like structures, and their humeri are deceptively derived from the pterodactyloid norm. The wing fingers of azhdarchids occupy a relatively small 47% of their wing lengths, a value only approximated by one other pterosaur group, the closely related thalassodromids. Artists, take note: grounded azhdarchids should not be reconstructed with their folded wing fingers stretching skywards over their backs: they couldn't reach that far.

But no, seriously, the long necks
The cervical vertebrae of giant azhdarchids are poorly known, with only a few specimens (and even fewer good ones) being recovered to date. These rare fossils do, however, clearly indicate substantial neck proportions in at least animals like Arambourgiania. The holotype cervical V of this animal is around 660 mm long, and is missing an estimated 100 mm from its posterior end. Steel et al. (1997) scaled this vertebra isometrically with relatively complete neck skeleton material from the 4.7 m wingspan azhdarchid Quetzalcoatlus sp. to predict a whopping 3071 mm length for cervicals III-IX in Arambourgiania. The use of isometry here is questionable (Witton and Habib 2010), but is defensible given the amount of azhdarchid neck material available to these authors in the mid-nineties. Ongoing work I'm involved with (which will hopefully be published before we're too much older) has attempted to apply allometry to calculations of giant azhdarchid neck lengths. The results are a little more conservative than the 3 m offered here, but we're still landing in the "seriously long neck" camp. Whether azhdarchids will retain the title of absolutely longest necks outside of Sauropoda (Taylor and Wedel 2013) remains to be seen however: I suspect they may ultimately just be pipped by the weirdo protorosaur Tanystropheus. Dammit.

The 'Big Necks Which Don't Belong to Sauropods Competition', won by the giant azhdarchid Arambourgiania. From Taylor and Wedel (2013).

Finally, some data on neck arthrology
The necks of azhdarchids are not just famous for their size, but are also renowned for their rather inflexible joints. These widely discussed features have been the bane of many azhdarchid lifestyle interpretations (see Witton and Naish 2008 for a review), but actual quantification of their arthrological range has been lacking until recently. This is, in part, because a complete 3D azhdarchid cervical series has been elusive for many years, but Alex Averianov (2013) recently produced a composite digital neck skeleton for Azhdarcho to figure out their range of motion. The results were more-or-less what we all expected: very limited range in the mid-series, with most of the mobility limited to the extremes. A surprising amount (but still fairly restricted) range of motion was afforded at the neck base, however. As may be expected, this study is very welcome to those of us interested in the biomechanics and functional anatomy of these animals, and I'm glad to see it.

Averianov's (2013) reconstructed neck arthrology of Azhdarcho lancicollis. That's one stiff neck.

Incidentally, some folks have asked me what I think of Averianov's suggestion that azhdarchids weren't what Darren Naish and I termed 'terrestrial stalkers' in our 2008 paper (generalised terrestrial foragers which spent much of their time wandering over Cretaceous plains in search of small animals and rich plant material like modern storks and ground hornbills). I won't say much now, but Darren and I don't agree with the alleged 'flaws' put forward against our hypothesis, and especially do not agree with the proposed 'aerial scoop feeding' counter hypothesis. Our formal reply has just been through peer review, and we hope to complete the minor tweaks needed to get it ready for publication very soon.

Swimming piscivores and aerial hawking: genuinely suggested azhdarchid lifestyles
It's well known that most recent 'serious' proposals of azhdarchid lifestyles are things like skim-feeding, terrestrial stalking and wading, but many other, frankly outlandish palaeoecological hypotheses have been thrown at azhdarchids over the years. Lev Nesov perhaps takes home the prize for the most bizarre ideas, proposing in his 1984 paper that azhdarchids could swim to find food (both along the surface and by diving) and pursue 'poorly flying' vertebrates through the air. In the same paper, he also advocates skim-feeding as a probably azhdarchid lifestyle. I remain unsure which part of azhdarchid anatomy indicated to Nesov that these animals had superhero-like abilities to acquire food.

Sauropods give a giant azhdarchid the evils. Seems they don't like being buzzed at close range. 

The awesomopower of giant pterosaur flight
Although azhdarchids are frequently discussed for their natty terrestrial capability nowadays, it's important to remember than any substantial travelling they had to do was probably performed in the air. Computations of the flight abilities of giant azhdarchids have returned seriously impressive results (Witton and Habib 2010). As mentioned above, azhdarchids likely employed anaerobic power for strenuous flight activities like takeoff and perhaps flapping bursts, and likely relied mostly on thermal soaring and flap-gliding like modern raptors to remain airborne for long periods. Their minimum sink and best glide speeds are steady cruises at 16.3 - 24.9 m/s (58.7 - 89.4 kph) but, if they were in a hurry (such as looking for a source of uplift), speeds of up to 48.3m/s (173 kph) were possible for short durations. We estimated that azhdarchids had about 90 - 120 seconds of anaerobic burst power before tiring, meaning these animals could go from a standing start to - literally - several kilometres away in the space of a few minutes. Yowsers. What's more, the size and bodily resources available to such large creatures permitted tremendous flight times: up to 16,000 km of travelling without resting or foraging were likely possible. That's the equivalent of an animal flying from London to Vegas non-stop, realising it forgot its passport, and then flying home again without touching the ground.

And that's your lot for now, folks. If you want to know more about azhdarchids, be sure to check out my book for a whole chapter about them, which is something like the second biggest entry in the entire thing. Things may go quiet over the next few weeks while I'm away at various conferences, but posting will resume when I get back.

References

• Averianov, A. O. (2013). Reconstruction of the neck of Azhdarcho lancicollis and lifestyle of azhdarchids (Pterosauria, Azhdarchidae). Paleontological Journal, 47(2), 203-209.
• Dyke, G. J., Benton, M. J., Posmosanu, E., & Naish, D. (2011). Early Cretaceous (Berriasian) birds and pterosaurs from the Cornet bauxite mine, Romania. Palaeontology, 54(1), 79-95.
• Marden, J. H. (1994). From damselflies to pterosaurs: how burst and sustainable flight performance scale with size. American Journal of Physiology-Regulatory, Integrative and Comparative Physiology, 266(4), R1077-R1084.
• Nesov, L. A. (1984). Pterosaurs and birds of the Late Cretaceous of Central Asia. Paläontologische Zeitschrift, 1, 47-57.
• Padian, K. (1984). A large pterodactyloid pterosaur from the Two Medicine Formation (Campanian) of Montana. Journal of Vertebrate Paleontology, 4(4), 516-524.
• Padian, K. (1986). A taxonomic note on two pterodactyloid families. Journal of Vertebrate Paleontology, 6(3), 289-289.
• Steel, L., Martill, D. M., Kirk, J. R. J., Anders, A., Loveridge, R. F., Frey, E. & Martin, J. G. (1997). Arambourgiania philidelphiae: giant wings in small halls. The Geological Curator, 6, 305-313.
• Taylor, M. P., & Wedel, M. J. (2013). Why sauropods had long necks; and why giraffes have short necks. PeerJ, 1, e36.
• Unwin, D. M. (2005). The pterosaurs from deep time. Pi Press, New York.
• 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.

Pterosaurs invades the newpapers, this weekend!

Good news, everyone! If you've not already made the plunge and bought Pterosaurs, you can enjoy a teaser article in this Sunday's (11/08/13) Observer to sample it's flavour, available both online and in dead-tree format. The piece covers mostly familiar ground for those with an ear for the jungle drums of pterosaur research, but may serve as a useful taster for current pterosaur science for anyone else. Perhaps of wider interest will be the showcasing of artwork taken straight from the book itself, including several pieces which are yet to be seen elsewhere. If this is the final impetus you need to buy the book, there's promise of a discounted price and free shipping from The Guardian Bookshop. I'll pop a link to the article into this post when it becomes available. (UPDATE: The article has now been posted: find it here).

While I'm stroking my PR cat, of further note is that Pterosaurs was reviewed in New Scientist by Jeff Hecht earlier this week, concluding that "[the] explanations and diagrams shed vivid light on the most intriguing creatures that ever flew, and in richer detail than even [Wellnhofer's 1991 pterosaur encyclopedia] could have managed just two decades ago." The article also features images from the University of Portsmouth Southbank pterosaur display, including our late, 10 m wingspan model of a flying azhdarchid. RIP, BigQuetz.

If this isn't quite enough of a pterosaurian fix for you, don't forget about the Pterosaurs talk being held at the Natural History Museum, London, this September.

Childhood dinomania: the greatest of all palaeontological mysteries?

Sinornithoides youngi, a long-legged, gracile troodontid from China. But why are he and his contemporaries so darned popular, and particularly with children?

Perhaps the greatest question that surrounds prehistoric animals is nothing to do with their palaeobiology or evolution at all. An enormous question, and one that perhaps continues to defy satisfactory answer, concerns their endless cultural appeal. Why are these long-dead animals so darned fascinating to us? Internationally, vast sums of money are spent on research into them, the care of their fossils and the educating of others about them. We put their remains on display in vast, elaborate museums, write no-end of books and articles on them. But why? Why are we do dedicated and passionate about these animals, to the point of near obsession in some individuals? I can understand that a few components of palaeontology have clear rewards. Palaeontologists specialising in invertebrates and microfossils are essential components of any team hoping to find hydrocarbon reserves, giving their research obvious application and financial implications. Such studies may also shed light on rates and mechanisms of evolution, which has bearing for the conservation and preservation of our modern biota. Most fossil vertebrate lineages, however, are of little use for, well anything. They may be interesting, but the scant nature of their fossil record doesn't allow their use in any applied studies. Our interest in them is purely academic. Knowledge for the sake of knowledge, I guess we could say.

Minor interests
Of course, interest in fossil vertebrates is not confined to academics and, in fact, the largest audiences for prehistorically-themed topics is under 10 years old. Of all fossil animals, the appeal of Mesozoic reptiles to young individuals is particularly well known, and encouraged by adults for good reason. Mesozoic reptiles introduce children to important concepts of science and the natural world, provide wonderful material to teach mathematics, literary and drawing skills, and, unlike many other things kids are interested in, they represent reality. Learned information about Mesozoic reptiles are learned facts about things that actually happened, not some silliness about Pokémon, ThunderCats, or... blast, who the devil do children like?... Or Morgan Freeman.

Young interests in palaeontology aren't really questioned, they're just accepted and ran with. Discussions as to why young people are so interested in Mesozoic reptiles aren't uncommon, but they're of secondary concern to nurturing childhood interests in the topic. As Dave Hone wrote about this topic at his Lost Worlds, " I won't pretend to know why, but kids really do love dinosaurs and the important thing is that they do." There's certainly nothing wrong with this attitude, but the why of this question has been on my mind of late. My second cousin is as dinosaur-obsessed as any young boy should be and I'll be spending the day with him next week. I'll also be gaining a nephew before too long. Between these two small members of my family clan, I'm expecting to have to play the cool, 'dinosaur'-researching relative for a while yet. All of which makes me wonder why, why why are small people so interested in these animals?

Proof that I liked 'safe monsters' as much as anyone when I was small. What's a 'safe monster'? Read on! (image by me, age 7[ish]. My younger self bore amazing powers of prediction for my own future with the pterosaur attacking dinosaur).

Standard responses
The most common explanation I've heard to this question is stressed in this article and others like it. Mesozoic reptiles are monstrous, and kids like monsters. They like these even more however, because they're long dead, and therefore 'safe'. Unlike real monsters, like the bogeyman, things that live under the bed and recent Discovery Communications documentaries, Mesozoic reptiles can't hurt us any more. Under this logic, extinction is the key agent here. Kids like the security that extinction offers between themselves and the monsters they're reading about.

Jurassic Park author Michael Crichton offered a completely different explanation, linking dinosaurs with authority figures, like parents. He wrote in his 1993 novel:

"...he mused on what it was about Dinosaurs that appealed to kids. He decided that dinosaurs represented a sort of symbolic authority to kids, a sort of surrogate parent. Just like a parent, they were simultaneously frightening yet accessible, and they presented an authority figure they could love. He also thought that children found satisfaction in saying the names of the animals, as that represented a sort of power of the vanished giants, showing a form of control."

Here, it would seem, it's not extinction at all that's key: it's the perception of authority and accessibility that Mesozoic animals seemingly offer children, and their own desire to master and control their expression. Other common explanations include an escapist quality to learning about the distant past, being able to express our childhood selves through acting out dinosaur fantasies, and because dinosaurs are strange, and yet real beings.

I've got to admit that I cannot really reconcile any of these explanations with what I know about being interested in Mesozoic animals, either as an adult or a child. They - particularly the first two suggestions - seem to complicated, too 'psychological'. I cannot ever remember associating Tyrannosaurus with my parents, or disliking other monstrous creatures because they weren't long extinct. I don't think the 'distance' between myself and dinosaurs, or any other monsters, really mattered. The fact that Mesozoic animals once existed was kinda cool I guess, but clearly not a deal-clincher: I was interested in plenty of make-believe things when I was small. And while kids are undoubtedly irrational sometimes, I don't think their grasp of what is a tangible, 'real' threat and slightly scary but fantastic beast isn't as blurred as the above explanations suggest. I note that many of the suggested points are rather anthropocentric, explaining that our childhood selves are interested in these animals because they reflect our own lives somehow, but that also doesn't seem right. My childhood interest in dinosaurs and the like seemed more innocent than that: I just wanted to know more about them and play within their universe. These ideas don't even seem like explanations which, in hindsight, chime with a deeply buried feeling associated with my childhood obsession with all things Mesozoic. Conversations with friends and colleagues suggest these explanations are similarly unfamiliar to them.

This makes me wonder if we're thinking about this the wrong way. We seem to expect that the appeal of Mesozoic reptiles to children is a unique trait, an X-factor, something inherently mystical about these animals which mean most children will be under their spell at some point. There may be, but I wonder if we're over-thinking this. Perhaps there is no unique factor behind the popularity of Mesozoic reptiles with young humans, and they're popular with kids for the same reasons that a lot of things are. Maybe the reason children like Mesozoic reptiles is very simple: they're just really cool.

The Anatomy of Cool
Let's run with this idea for just a moment. Mesozoic reptiles certainly tick all the boxes for Cool Things That Kids Like. Starting with the most obvious: they look awesome. Innumerable cartoons and comics featuring appealing characters and creatures are testament to the power awesome-looking beings have over children. The muscular bodies, dynamic postures, horns, frills, teeth and claws of many Mesozoic reptiles are clear signs of badassery, and kids of all ages respond positively to that. Perhaps the consistent choice of favourite dinosaurs in youngsters reflects this. Although most children's dinosaur books introduce a wide selection of species, it's the most anatomically extreme and charismatic species that are picked out by generation after generation as Top Dino. Triceratops, Tyrannosaurus, Baryonyx, Velociraptor, Ankylosaurus, Brachiosaurus and so forth are consistent favourites. Some kids - especially cootie-ridden girls, because they're rubbish and smelly - might like prefer cuter, baby versions of dinosaurs,  but they still pick babies of the most immediately interesting taxa. By contrast, no kid has ever said that their favourite dinosaur is Iguanodon or Hypsilophodon, because they're freakin' boring to a sub-10 year old. This is despite them being among the 'safest' dinosaurs, bearing no real offensive equipment and having no interest in eating children. Kids dig awesome, even if it's a little scary, and their favourite dinosaurs are full of it.

Do kids like Sinornithoides? I don't know that they do, but it would definitely score Cute Points when they realised that the holotype was found in a Mei-like sleeping posture. Of course, Sinornithoides was described by Russell and Dong way back in 1993, including discussion of its sleeping posture, meaning it pre-dates the announcement of sleepy Mei by over a decade. This fact seems mostly overlooked nowadays, however.

Mesozoic animals are also immediately characterisable. A cursory glance at a menagerie of Mesozoic animals reveals which ones are 'good' - the plant and fish eaters - and which are 'bad' - the carnivores. Universes designed with young people in mind go to great lengths to give their characters similarly recognisable traits of good and bad. They also, as with Mesozoic reptiles, make their characters wear their lifestyles on their sleeves. It's immediately clear that they spend their time doing interesting things because their appearance (clothing, physical characteristics, objects they carry) consistently reflects their habits. When do we see warrior characters in children's shows put their weapons down, or adventurers leave their backpacks and hats behind? Never, because it's part of who they are. The same is true of Mesozoic reptiles: their lifestyles are clear from their anatomy, and their habits are obviously interesting. As with invented universes, this allows even young children to have a fairly immediate, if very basic understanding of the dynamics of the Mesozoic world, and that makes it fun to play with and think about. I've written before about how some Mesozoic creatures even come with pre-conceived ideas of 'character': the frills and horns of some dinosaurs recall the armaments of knights, the powerful jaws and teeth of tyrannosaurs make them obvious threats, and so on. These perceived anthropomorphisms may tie into the choice of favourite species among children, perhaps reflecting elements of wish-fulfilment and reflection of individual  personalities, but the same applies to their selection of a favourite Transformer or mutated ninja turtle.

What about complicated Latin and Greek names? Surely they must have some unique appeal? It's perhaps no coincidence that many favourite Mesozoic animals are also those with the coolest names. Animals with undoubtedly disastrous (Futalognkosaurus) or boring (the infinite numbers of Placename-osaurus we now have) names are unlikely candidates for being any child's favourite. The strong, weighty names of Deinonynchus, Pteranodon, Plesiosaurus and Diplodocus are where it's at. Really, they aren't actually too different or more difficult to say than invented names of child-approved fantasy universes. A of extinct animal names are no trickier or less familiar to children than the names of Star Wars or Lord of the Rings characters, for instance. There may be no more psychological significance to a child saying the word 'Gallimimus' than there is them saying 'Legolas' or 'Dagobah'.

The stats and factoids associated with Mesozoic reptiles are perhaps also factors in childhood palaeo cool. Any juvenile palaeo nut worth their salt knows the size, mass, biogeography, geological period, lineage, and diet of a hundred extinct species. Our brains are sponges for that kind of stuff when we're small, but not only for Mesozoic animals. Kids get obsessive about all manner of data, hence the success of all these newfangled Japanese card playing games with weird animals and, before them, things like Top Trumps, complex board and video games and RPGs. It seems that, if children like a topic, and the information is there to be learned, they'll take it in whether it has a dinosaur stamped on it or not.

Sinornithoides again, acting as end-of-post wallpaper

We've now also created a rich array of Mesozoic reptile merchandise for children to enjoy - toys, games, books, films and TV shows and so on - which, again, mirrors the development of universes invented for child consumption. These are food in the purest form for the imaginations of small children, enhancing their ability to play out their own interpretations of the Mesozoic in the same way that the merchandise of invented franchises allows kids to play within other universes. Unlike many franchises aimed at children, however, Mesozoic reptiles rarely disappear from fashion, and their merchandise is always easy to obtain. Indeed, I wonder if the perpetual availability of Mesozoic reptile merchandise and media has made it almost certain that child interests in these animals will never go away. Given the ripeness of childhood minds for the awesomeness of Mesozoic reptiles and the associated financial gain from making Mesozoic merchandise, we may be creating a self-fulfilling prophecy that children will always be introduced to and inspired to learn more about these animals.

The discussion at the end

With all these things considered, I really wonder if Mesozoic reptiles have, or indeed need a mysterious 'X-factor' to explain their appeal. I don't think it's been an intended goal of palaeontologists or merchandisers, but these two contrasting industries have created a window into the Mesozoic that children can enjoy on many levels, developing a world which couldn't be more child-friendly if someone designed it. The many parallels we see between childhood palaeo culture and industries designing universes to interest children are surely a reflection of this. Cool, identifiable creatures with interesting lives, awesome names and stats, and a wealth of merchandise. That description could describe how children will interpret palaeontology, or it could describe the way they'll interpret Doctor Who.

As a final point to chew on, I think it's interesting that we don't really feel a need to explain the childhood appeal of superheroes, spaceships and giant robots by means of an X-factor', but we do for Mesozoic reptiles. Adults just accept that kids find these more anthropocentric topics inherently awesome and interesting, and that's good enough. Why doesn't that work for palaeontological topics? Is it a little worrying that we think like this? That the raw appeal of the natural world, which kids seem to intuitively grasp as interesting and awesome, isn't a strong enough draw on it's own, and requires rationalising into a more a anthropocentric model to explain it's childhood appeal? Maybe there's something to be learned from that. General knowledge and understanding of the natural world is critically poor, biological education is consistently being attacked by anti-scientific groups, and media groups increasingly think that the natural world needs sexing up with human interaction and made-up science. Maybe if we just remembered that it's OK to find the natural world fascinating and awesome because it is, and that we don't need to make ourselves the centre of everything, these issues wouldn't be anywhere near as big and worrying as they are.

Reference

• Russell, D. A., and Dong, Z. M. 1993. A nearly complete skeleton of a new troodontid dinosaur from the Early Cretaceous of the Ordos Basin, Inner Mongolia, People's Republic of China. Canadian Journal of Earth Sciences, 30, 2163-2173.

Pterosaurs: The Natural History Museum Talk, September 2013

HyPtA D does the NHM logo. What's a HyPtA D? You need to buy Pterosaurs to find out, or attend the Pterosaurs NHM event in September. HyPta D image from Witton (2013); NHM logo borrowed from here.

Those of you with long-term memories may recall that, this September 10th (2013), the critically-acclaimed tome Pterosaurs: Natural History, Evolution, Anatomy is being celebrated with a public lecture at none other than the Natural History Museum, London. Details of the event have now crystalised and been made public over at the NHM's website. The talk will take place at 19:00-20:00 in the Neil Chalmers Seminar Room and cover all things pterosaurian: our changing understanding of these animals since their discovery, current ideas on many aspects of their palaeobiology, lots of palaeoart and photographs of specimens and, no doubt, some over-the-top PowerPoint animations which I'll spend hours tweaking for no obvious benefit. I'll do my best to keep to time (I have an excellent track record for punchy, concise talks, honest) so I can field questions at the end of the talk.

Note that the event is for NHM Members Only, will cost £5.50 per ticket, and booking is required. Details about membership with the NHM can be found here, and further information on the event and NHM memberships are available from your telephone, at +44(0)20 7942 5792.

I'm already looking forward to this, and hope to see many of you there.

UPDATE: I've just realised that the NHM's Lorna Steel, a pterosaur worker herself, is giving a behind the scenes tour of the NHM's extensive pterosaur collections on the same day as this talk (10/09/13) for NHM members, also for a mere £5.50. It seems that, if pterosaurs are your thing and you're an NHM member, you'll really want to be in London on the 10th of September.

Reference

• Witton, M. P. 2013. Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.

Engaging my Disacknowledgement

Tupandactylus imperator, a rat, and the Disacknowledgement. Not in that order. From Witton (2013).

Weirdest thing happened today: I finished two planned bits of work ahead of time (I know. I'm scared too!). That doesn't mean I'm off the hook work-wise. I was away last week (hence the lack of a fresh post) and catching up is making for a very hectic few days, but the fact that two bits of work were finished bang on time for an afternoon tea break means I've got time to slip in a quick bit of blogging without the usual guilt trip. We'll have to keep it brief, though, and discuss the rather straightforward image shown above.

Rather obviously, this image is a play on those most standard of palaeontological artworks, the extinct-creature-human-scale diagram, in this case showing the flamboyantly crested tapejarid pterosaur Tupandactylus imperator and a fully-fleshed Homo sapiens instead of a silhouette. Those who've read my book, Pterosaurs, may recognise this image from page 221. Note that the Tupandactylus soft-tissue crest is convex along its posterior margin instead of concave, as it's often been depicted. New fossils, such as those mentioned in Pinheiro et al. (2011), suggest that this 'fuller' crest is more likely than the concave crest indicated by the T. imperator holotype (Campos and Kellner 1997). The mandible shape also follows the specimen described by Pinheiro et al. (2011) rather than, as in most depictions of this animal, a generalised tapejarid mandible based on closely related species. Believe it or don't, virtually all that's known for certain of T. imperator is skulls, with only two specimens preserving mandible remains. Tapejarid skeletons without skulls are known from the Tupandactylus-bearing Crato Formation, and some of them likely represent bits of T. imperator itself, but we can't be certain of this until skeletons with associated cranial remains are recovered. Until then, we'll have to be satisfied with the revelation that T. imperator has chin big enough to scare even Kurt Russell, and wait for further discoveries. Be sure to check out this image of Tupandactylus navigans, another Crato pterosaur with a penchant for elaborate headgear, for more information on this genus.

But enough about that
What makes this picture more unusual than many like it is that it depicts an actual, real-life person who I happen to know very well, and anyone who's read Pterosaurs will also be familiar with. Here's where she's mentioned at length, from the 'Acknowledgements' page. She got her own section and everything.

Disacknowledgement

Finally, although customary in book acknowledgements to honor those who help steer projects to completion, it seems unfair to not mention the tremendous negative impact on this project made by Georgia Maclean-Henry. As the single most destructive force against this work, she took my attention from this project so frequently that we ended up moving in with each other halfway through the writing process and have ended up making some sort of home together. She continues to distract me from all kinds of work to this very day and, frankly, I could not be happier about it.

So yes, here she is. The Disacknowledgement herself. The 'single most destructive force against this work', and many others. I chose this picture to post now because today is the 2nd anniversary of the aforementioned moving in with my Disacknowledgement. I'm not normally one for noting or making a big deal of such things (as the Disacknowledgement knows all too well), this ties in with a more important recent event: the Disacknowledgement agreed to marry me when I asked her last week, which is all very exciting and I'm rather over the Moon about (before anyone asks, there were no knees involved, but there was a breccia, not to mention a kick-ass grey seal sighting moments before. He was huge!). Seeing as this image puts me in a good (and probably fairly unique) position to broadcast this happy fact while clinging to some sort of palaeontological relevancy, there seemed no better way to spend this brief teabreak. Speaking of which, said break ended quite a few minutes ago: best get on with other things, including hoping that the Disacknowledgement doesn't mind being referred to as 'the Disacknowledgement' all the time. Sorry dear. 

References

• Campos, D. A. and Kellner, A. W. A. 1997. Short note on the first occurrence of Tapejaridae in the Crato Member (Aptian), Santana Formation, Araripe Basin, Northeast Brazil. Anais-Academia Brasileira de Ciencias, 69, 83-88.
• Pinheiro, F. L., Fortier, D. C., Schultz, C. L., De Andrade, J. A. F. and Bantim, R. A. 2011. New information on the pterosaur Tupandactylus imperator, with comments on the relationships of Tapejaridae. Acta Palaeontologica Polonica, 56, 567-580.
• Witton, M. P. 2013. Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.

The Golden Age of Palaeoart?

Polacanthus foxii, a Lower Cretaceous ankylosaur from the Wealden of southern Britain, trying to shake some tiny birds off his nose as he strolls around a knoll of horsetails. Prints of this image are available.

It's a bad time for many industries, communities and institutions at present. Economies have crashed around the world, money for scientific research and artistic projects is at an all time low, education and human rights are being challenged by extreme social movements and the global biosphere is in crisis. To make matters worse, there's not even anything good on TV to take our minds off it all. But all is not lost. At least one institution, and likely one that folks reading this will care about a lot, is thriving. Indeed, it may be a veritable Golden Age for this industry, creatively at least, and there's no sign this boom period is going to end soon. I'm talking, of course, about palaeoart.

The recent history of palaeoart has been pretty interesting. We've seen the publication of several new books which are, for various reasons, likely to be considered seminal palaeoartistic works in years to come. New philosophies to palaeoart have emerged, and, all the while, the internet palaeoart community continues to grow and evolve. It's not only these developments which have helped palaeoart blossom particularly brightly in recent years, however. The advancement of new technologies, as well changes to academic publishing models, have also help usher in this wave of palaeoartistic splendidness. Some of these developments may sound a little humdrum, and some may be so obvious that we've not really sat to think about them before, but they may well make today one of the best times ever to be practising the art of reconstructing extinct organisms.

The Future, now!, and what it means for palaeoart
We'll start at the most fundamental end of palaeoart procedure: acquiring reference material. This has long been a problem for palaeoartists. Because palaeoart has an obvious firm rooting in palaeontological science, no self-respecting palaeoartist begins their work without developing an understanding of the anatomy of their target species. This typically involves working from scientific texts which have been, classically, hidden away behind publisher paywalls, unavailable to anyone without access to a university library, an online subscription to their services, or unwilling to part with obscenely-large sums of money for a single pdf download. There have always been ways around this - emailing authors, asking connected friends for photocopies or pdfs - but there's still an initial barrier to hurdle before work is even started.

The sign of better times. From Wikimedia Commons.

Happily, these walls are starting to crumble. The Open Access movement, which makes scientific literature available free to everyone, everywhere, has taken the academic world by storm, and even the greediest publishers are being forced to make their archived material more readily available. The battle for entirely free scientific literature is not yet over, but its victories are already making the life of palaeoartists far easier: high-quality reference material is increasingly often only a click away. It's not just the availability of literature which is a boon, either. New, online-only journals have thrown aside pesky limitations on figure numbers and image quality, permitting publication of numerous, top-quality illustrations for reference. Sometimes, we even get spinny-rotatey movies of CT scans to work from, showing anatomies in almost any view we could desire. As a bonus, the digital format of this information makes it easy to zoom up to tiny details which would be lost in print publications, allows for easily compiling our own anatomical reference sheets, and makes tracing anatomies for skeletal reconstructions infinitely easier than it's been in the past.

It's not just scientific papers that are an increasingly accessible resource, either. Dedicated websites committed to archiving the anatomical information craved by palaeoartists are becoming more plentiful. Sites like Digimorph, the Witmerlab pages, and the Bird Skull Collection are full of detailed graphics of animal anatomy of both extinct and modern species, and they're entirely free to access. Even Wikipedia is doing a fairly good job at providing references for palaeoartists. The excellent image of the sacral shield of Polacanthus, below, was sourced from the Wikipedia entry for that species for use the reconstructions adorning this post, for instance, and there's a lot more where that came from. Add this the increasing wealth of information scientists are putting online through blogs, podcasts, media releases and Youtube channels, and palaeoartists are on considerably better ground for finding reference material than we were even a decade ago.

 Woodward's 1881 depiction of the Polacanthus foxii sacral shield. From Wikipedia.

By palaeoartists, for palaeoartists
It's not only the scientific community that's feeding the blooming palaeoart scene. Several authors, many of them world-class palaeoartists themselves, have recently been producing easily-accessed works which are almost solely designed as references for fellow palaeoartists. I'm referring, of course, to the digital and printed works of the likes of Gregory Paul, Scott Hartman, Jaime Headden, Mike Hanson, and others. These chaps have published extensive libraries of anatomical orthographics for numerous fossil species, made suggestions about the most appropriate ways to reconstruct extinct anatomies, and generally helped artists transition fossil bones to restored animals significantly in recent years. (Note that a lot of Paul's guidance in this area pre-dates the Internet, and has mostly been published in books and magazines. Pdf versions of these articles, including his seminal 'rigorous guide' to dinosaur reconstruction from 1987, are available from his website). Matthew Martyniuk has also been contributing to this area recently, his excellent guide to Mesozoic birds (Martyniuk 2012)* and website detailing approaches to reconstructing avialans and their immediate ancestors. Elsewhere, I made a stab at providing a few helpful pointers to reconstructing pterosaurs in my own book, with some text and graphics specifically targeted at palaeoartists (Witton 2013). Even non-artists, like the guys behind Tetrapod Zoology and SV:POW!, are getting in on the act, regularly commenting on the possible life appearance of their extinct subjects and providing constructive criticism on certain bits of palaeoart.

The books, imagery and accessible discussions of these individuals will almost certainly inform palaeoartists for years to come. In particular, Paul's (2010) Field Guide to Dinosaurs is probably the most comprehensive collection of dinosaur skeletal reconstructions in a single volume, ever, and is of obvious utility for the high numbers of palaeoartists interested in Mesozoic dinosaurs. Pushing Paul (2010) as a key reference is, of course, complicated by Paul's insistence that his orthographics are not to be used as reference material by other artists. I must admit to finding this request a little odd and I suspect it's mostly falling on deaf ears. Still, however you want to use it, Paul's book, and other recently published resources like it, are invaluable resources for palaeoartists, and make it easier than ever for everyone to get basic anatomical data for their subject matter.

Taking it to strange new places
Is there any reason to think our conveyance of palaeontological subjects has improved in recent years? This is very much a matter of opinion, but I think it has, for some species at least. I've got to stress that this isn't because modern artists are more talented than their forebears. It's because modern palaeoartists seem to have finally got a handle on the rather dramatic new ideas and data that have coming our way for the last few decades, which has helped make newer depictions of certain species a little more convincing than those that came before. For instance, we've been bombarded with the discoveries of feathers on numerous type of non-avian dinosaurs, new considerations of soft-tissue masses (e.g. pterosaur limb musculature, dinosaur caudofemoralis mass) and novel ideas on the locomotory habits and postures of many species. That's a lot to take in and regurgitate into an image. It's not only changing scientific opinions which have altered the way we restore the ancient world, either. Radical shifts in palaeoartistic culture have also occurred, such as the well documented (including here) move away from overly conservative and 'shrink-wrapped' depictions of fossil species. It's been a changeable and dynamic few decades for palaeoartists, and I think it's understandable that it's taken a little time to work out what all this new information and philosophical changes equate to in artwork.

Perceptions of pterosaurs have changed a lot in recent years. The thin, lanky azhdarchid on the left didn't look odd in 2006, but looks positively anorexic compared to more recent reconstructions (right, from Witton 2013).

The upshot of this, in my opinion at least, is reconstructions of several species - including well-known taxa - have only started to appear in the last few years. This particularly applies to animals like maniraptoran dinosaurs and pterosaurs, species which have been classically portrayed rather outlandishly because of their unusual anatomy and some outlandish approaches to their depiction (Slasher Poses, I'm looking at you). I'm astounded at the convincingness of some of Matt Martyniuk's reconstructions in his Field Guide to Mesozoic Birds, for instance. His oviraptorosaurs look like elegant creatures painted from life, instead of the incredibly goofy forms we're more familiar with. The same applies to Emily Wolloughby's bouncy, eutherian-chasing Velociraptor, John Conway's famous squatting Therizinosaurus and many others. These animals make a lot more sense to look at now now, as do a lot of other restored ancient creatures. My feeling, at least, is that these very recent interpretations of these animals are closer to reality than anything we've seen before, and that can only be a good thing.

As an added bonus, an almost postmodern approach to palaeoart has emerged (crystallised, of course, in All Yesterdays [Conway et al. 2012]), which helps avoid the many tropes and clichéd artwork we see again and again. The result is that modern palaeoart may not only be more accurate to life, but is more interesting to look at. I almost get the feeling that there's greater confidence in palaeoart than there used to be. Because we know more about the animals, more about the most convincing ways to depict them, and more about our own collective foibles, palaeoart seems to be moving confidently into new and interesting areas. Again, I stress that I'm not downplaying the quality or importance of other artists and previous work - as with any artform, palaeoart is a product of its history - but it feels that a lot of modern palaeoart is trying hard to break into new territory. And that's great.

Being seen and heard
Of course, all of this work would be for nothing if it weren't seen. As a child living in the pre-Internet age, the only way I'd see new palaeoart was though purchasing new books, or sometimes in magazines and newspapers. New bits of artwork were rare, important events. The situation nowadays couldn't be more different. Most palaeoartists have online galleries or blogs (or both), and they post new artwork to Twitter, Tumblr, and Facebook. We don't just see new press release artwork because it's been a slow news day: if a new bit of artwork is released, there's a good chance we'll see it. What's more, the artists themselves are no longer simply represented by squiggled names in the corner of an image: social media allows them to introduce and comment on their work. While most of this is happening online, it does, on occasion, still happen in print: see Dinosaur Art: The World's Greatest Palaeoart (White 2012), and Douglas Henderson's new book detailing his early work and inspirations (here) for examples. I think this is a terrific change from simply waiting for new books to be released. There's a much greater sense of community, greater familiarity with each other's work, and very liberal exchange of ideas.

A closer look at the Polacanthus from up top. He's nothing to do with anything here, just livening up the post.

The Internet has also revealed how much terrific artwork is being done by up-and-coming artists. The only palaeoart we routinely saw in the 90s was that of a few masters: Henderson, Hallett, Sibbick, Paul and so forth. Nowadays, we know of have a much broader range of individuals producing art at very high levels. The growing number of palaeoartists seems to have ruffled a few feathers among some established profressionals who have found a lot of their work being outsourced to others (see this thread on the Dinosaur Mailing List) but, on reflection, I think growth in the palaeoart community is inevitable. Greater availability of palaeontological information, the rise of the online palaeontological community, the increasing availability of palaeoartistic guidance, the affordability of digital painting setups and the ease of uploading content online are all contributing factors here. This does mean greater dispersal of the available work among more individuals, but I'm not sure there's much we can do about that.

Besides, this greater range of artists has to be a good thing, and not just because it we get to see more palaeoart. It also allows for a wider range of styles and media than we're used to. Most palaeoart I remember from my childhood was attempting to rival photographs in detail, but a lot of modern work offers more stylised - more artistic, even - interpretations of ancient life. This goes hand-in-hand with a range of media being used. Modern palaeoart is not only being produced by traditional painting and sculpture, but also by compositing photographs, 3D digital modelling, digital painting, vector art and combinations thereof. It's great seeing how these diverse media bring ancient worlds to life in different hands, and makes working within the palaeoart community a generally more interesting and inspirational place.

On top of that, the palaeoart community itself is proving to be an active and interesting one, with events like the open source Life Galleries of ArtEvolved (sadly, these galleries are coming to an end soon after a 4 year run), palaeoart competitions at Love in the Time of Chasmosaurs and from the All Yesterdays chaps, and even pooling resources to generate it's own independent products. And this doesn't even scratch the surface of palaeoart coverage and events in social media, not the mention the palaeoartist meetings and gatherings that now frequently happen at conferences. We only have to look to the rapid uptake and discussion of All Yesterdays as a good example of how lively and active the online palaeoart community is. How successful would that project have been without the online palaeoart community?

So there you have it
And that's why, all told, I think we're living in a 'Golden Age' of palaeoart. There's more of it, the artwork we have is generally better informed because of improved access to reference material and guidance from experts, we've (finally) got a a handle on ways to reconstruct some stranger ancient species and, to boot, developed a wide, friendly community of excellent artists. It's a good time to be a palaeoartist, then, and long may that continue.

References

• Conway, J., Kosemen, C. M. and Naish, D. 2012. All Yesterdays: Unique and Speculative Views of Dinosaurs and Other Prehistoric Animals. Irregular Books.
• Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs. Pan Aves.
• Paul, G. S. 1987. The science and art of restoring the life appearance of dinosaurs and their relatives – a rigorous how-to guide. In Czerkas, S. J. and Olson, E. C. (eds) Dinosaurs Past and Present Vol. II. Natural History Museum of Los Angeles County/University of Washington Press, pp. 4-49.
• Paul, G. S. 2010. The Princeton field guide to dinosaurs. Princeton University Press.
• White, S. 2012. Dinosaur Art: the World’s Greatest Paleoart. Titan Books, London.
• Witton, M. P. 2013. Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.

Something R'lyeh different

Detail of "Mark Witton's greasy, flowing mass", for Matt Wedel's Collect Call of Cthulhu. 

If my Facebook and Twitter feed tell me anything, it's that folks with interests in palaeontology are also very interested in science fiction. Chaps who liberally discuss bird origins and tyrannosaur feeding palaeobiology are likely to also discuss how many tribbles you can stuff into a Jefferies tube or which Imperial Walker would be best for car-pooling to work. Given that a lot of modern science fiction teeters under the weight of ideas borrowed from early science fiction authors, I'm going to take a wild guess that a lot of palaeofiends have, at one time or another, traced these sources back to one of the cornerstones of modern sci-fi: the nightmarish, slimy and tentacle-rich mythos of H. P. Lovecraft.

Matt Wedel, best known in palaeontological circles as a crusader of sauropod vertebrae, certainly has. Over at Echo Station 5-7 he's been musing on the nature of Lovecraft's seminal creation, Cthulhu, and inviting submissions of artwork rendering his hideous visage. A keen emphasis has been made on entrants avoiding the traditional representation of Cthulhu as a guy with squid head through either style or biological interpretations, which I agree with 100%. Lovecraft's descriptions of Cthulhu are undeniably vague, but his suggestion of a greasy, dripping, and ambiguous being is a million miles away from the beefy, green anthropoids we see striding around all too often. Indeed, even Lovecraft's own sketches of Cthulhu seem quite wide of his text:

Cthulhu takes five in this sketch by H. P. Lovecraft himself. Borrowed from the Lovecraft EZine.

Given that I'm not adverse to dipping my toes in Cthulhu mythos myself, I couldn't resist joining the fun. I've had an old, half-finished painting of Cthulhu lying dormant, dreaming, on my hard drive for well over two years, so saw Matt's open invitation as the impetus to wake him up, finish him off, and send him in. The results can be seen in part at the top of this post, but you need to head to Echo Station 5-7 to see the rest. Because I was apparently cursed to write at length about pretty much anything I create in a past life, there's also a typically lengthy explanation for the decisions I made in the rendering process.

If Lovecraftian things are up your street, you also owe it to yourself to read the comments from regular palaeoblog contributors on what they think makes a good Cthulhu, here, here and here. They're not only great demonstrations of how diverse the interpretations of vaguely-understood literary beings can be, but offer some great speculative biology because of the informed scientific backgrounds of each commenter. The discussions at Matt's blog have also spawned an interesting essay on the ecological strategies used by Lovecraft's beings, based on vernal pools, penned by Heteromeles at Putting the Life Back in Science Fiction.

All good stuff, and I've thoroughly enjoyed being part of it. Regular programming - art of old dead things - will be resumed shortly.