Tetrapod Zoology

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By now, it’s reasonably well known to interested people what azhdarchid pterosaurs looked like when alive. The answer: sort of like a cross between a giraffe and a stork, though with all of this being over-ridden by uniquely pterosaurian weirdness; membranous wings supported by giant fingers, a large cranial crest, plantigrade feet, and so on.

Why have azhdarchids become so familiar? The reconstruction of several life-sized skeletons and models mean that they’ve been widely exposed to the museum-going public, CG versions have recently featured on TV programmes about Mesozoic life, and a better knowledge of their appearance and biology means that they’re increasingly featured in artwork. Tetrapod Zoology has, I think it’s fair to say, done more than its fair share of azhdarchid promotion, plus there’s also a rather good paper on these animals by two chaps called Witton and Naish (Witton & Naish 2008). Mark has illustrated azhdarchids many times, and – as I hope you know – he’s working on something BIG due to be showcased very, very soon. I can’t wait. Anyway…

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I first got to know Quetzalcoatlus as a very, very different beast, however. Little known today (it seems) is that Quetzalcoatlus was depicted as a most peculiar creature in a few books from the 1970s and 80s. As you can see from the images reproduced here (the drawing above is by Guy Michel and appeared in John Gilbert’s 1979 book Dinosaurs Discovered), it was shown as being a short-headed pterosaur with a weird lump on the back of its head and pointed, toothy jaws. Imagined like this, it really looks like some sort of horrific demon. Richard Orr’s painting – shown below – always makes me think of one of Hieronymus Bosch’s scenes of hell; truly it’s a horrific, terrifying scene (and I don’t mean that as a slight to Orr’s good artwork). Where on earth did this view of Quetzalcoatlus come from?

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Orr’s painting comes from a Dougal Dixon book (Prehistoric Reptiles) that was originally published in 1984 (I only have a 1993 edition). It’s obvious from many of the scenes included in the book that Orr had been heavily inspired by Giovanni Caselli’s reconstructions from L. B. Halstead’s The Evolution and Ecology of the Dinosaurs (first published in 1975). Sure enough, if we look at this book, we see another version of the same creature [shown here]. Caselli didn’t show the animals in as much detail as Orr did in 1984 or Michel did in 1979, but we can see a comparatively short skull and a blunt, knob-like crest at the back of the head. The animals are also shown as being red, just like Orr’s creature: as I’ve said before (in connection with phorusrhacids and astrapotheres: it’s pet peeve # 112), artists frequently follow the colour schemes of their predecessors, meaning that a certain ‘uniform’ becomes associated with a certain creature for no good reason. Anyway… Caselli’s version came first – but where did it come from?

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Alas, this is where I draw a blank. I used to correspond with Giovanni Caselli: we’ve fallen out of touch, but I emailed him today and will let you know what he says. I’ve assumed that the short-headed, toothy nightmare version of Quetzalcoatlus was inspired by an early rendition published in a newspaper article or something, but the problem with this idea is that the early versions of Quetzalcoatlus that accompany papers, magazine articles and such correctly show the animal with a long, toothless rostrum. Wann Langston was illustrating Quetzalcoatlus like this as early as 1977 (recall that Quetzalcoatlus was discovered in 1972 and described – tersely – in 1975) [adjacent image is taken from a magazine article, published in 1977 by Langston: I don't have the complete reference, but the article was titled 'The great pterosaur']. Incidentally, some of you might know that Quetzalcoatlus was so tersely described that there are some doubts about its validity…

Anyway… until I hear back from Giovanni, I’m stumped. I’ve mentioned this freakish version of Quetzalcoatlus before (here on Pterosaur.net), but have never seen it mentioned by anyone else, ever, nor have I met anyone who knows anything about it. If you know more, please do share!

For previous Tet Zoo articles on azhdarchids and other Cretaceous pterosaurs see…

One more thing: Tet Zoo has covered hypothetical flightless pterosaurs a few times (so has the Pterosaur.net blog: go here and here). A new paper in Journal of Vertebrate Paleontology (by Don Henderson) finds Quetzalcoatlus to exceed 500 kg in mass, leading to the serious suggestion that it must have been secondarily flightless. I think there’s been a mistake here (Witton (2008) estimated Quetzalcoatlus to be 225 kg, Paul (2002) suggested 250 kg, and Sato et al. (2009) provided an estimate of 279 kg: even these have been regarded as surprisingly heavy by some workers), but I’ll withold judgement until I’ve read the paper.

Refs – -

Dixon, D. 1993. Prehistoric Reptiles. Evans Brothers Limited, London.

Gilbert, J. 1979. Dinosaurs Discovered. Hamlyn, London.

Halstead, L. B. 1975. The Evolution and Ecology of the Dinosaurs. Peter Lowe, London.

Paul, G. S. 2002. Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Johns Hopkins University Press, Baltimore.

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. doi:10.1371/journal.pone.0005400

Witton, M. P. 2008. A new approach to determining pterosaur body mass and its implications for pterosaur flight. Zitteliana B28, 143-158.

Witton MP, & Naish D (2008). A reappraisal of azhdarchid pterosaur functional morphology and paleoecology. PloS one, 3 (5) PMID: 18509539 [available here]

Comments

#1 Ian
May 25, 2010

Funny you should mention the way some artists follow a color scheme of a prehistoric creature for no reason. I’ve noticed that with Maiasaura, which tends to be given a light brown/orange/beige color.

That being said, I agree with you about how these depictions look like demons. Orr’s painting in particular, if you replace the Mesozoic landscape with fire, brimstone, etc., looks like a scene from prehistoric Hell, with demon pterosaurs feasting on a dinosaurian sinner.
#2 RStretton
May 25, 2010

Ah that picture brings back happy memories. I used to get annoyed with these unrealistic pics when I was a kid but now nostalgia has kicked in and I can accept just how marvelous some of them are. As I recall this book had many other masterpieces in too.
#3 Gray Stanback
May 25, 2010

I’ve noticed it too. Ever since Walking With Dinosaurs, Liopleurodon is ALWAYS black-and-white-spotted
#4 Aaron Wells
May 25, 2010

I remember pink quetzelcoatlus from the kids’ books I read in the 80s. One book even said it might have been pink due to its diet, the same as flamingos.
#5 Abyssal
May 25, 2010

I don’t think I’ve ever seen an image of *Camptosaurus* that wasn’t white with red stripes.
#6 Rory
May 25, 2010

I remember pink quetzelcoatlus from the kids’ books I read in the 80s. One book even said it might have been pink due to its diet, the same as flamingos.

Posted by: Aaron Wells | May 25, 2010 4:47 PM

Would that not have been pteradustro? Quetzalcoatlus was widely believed to have been a carrion eater.

Even as a kid myself I was brought up on outdated books on mesozoic animals-like the dinosaurs that really looked like mean,angry lizards leaning on their hindquarters. Walking with Dinosaurs was my first look at “proper” dinosaurs.

Incidentally is Quetzalcoatlus still the largest pterosaur? or am I severely outdated? I know it is an azdharcid but I don’t know whether it features in Mark Witton’s famous artwork.
#7 Felipe Pinheiro
May 25, 2010

I’ve got the complete reference of “The Great Pterosaur”.. I can send you by e-mail if you like.. It’s a low-quality scan but it works! Send me an e-mail and I’ll send you back the reference right away.
#8 Trish
May 25, 2010

See, when I read Dinosaurs Discovered for an upcoming blog post (it’ll be up Thursday, see link) that scary-arse Quetzel didn’t even register because there’s so much trippy stuff in the illustrations. I’ll be sharing some of my favorites. Of note is the fact that every single Iguanodon in the book is emerald green and has a creepy long giraffe-tongue for some reason.
#9 kris
May 25, 2010

Absolutely Hieronymus Bosch (both Caselli and Orr). I think it’s as much due to the colour as to something in the way of perspective — the background somehow looks flat, and the creatures themselves don’t really have that much depth either. Absolutely creepy and fascinating in effect. The Garden of Mesozoic Delights??
#10 Jerzy
May 25, 2010

It just shows how much good did internet. For decades, paleo artists simply copied one of the few books avialable. Now they have instant access to almost all human knowledge – plus amateurs became involved.

I still remember the time, when most dinosaur illustrations could be tracked to a copy of Knight paintings or Burian paintings. Any new position or interpretation was unusual.
#11 Jerzy
May 25, 2010

“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. doi:10.1371/journal.pone.0005400″

Thank you for bringing up this paper! Lots of fun! A real life example of that apocryphal scientist who was so out of touch with reality, that calculated that bumblebee could not fly!
#12 Andrew
May 25, 2010

Wow, that bulge at the back of the skull, combined with the ear position, makes them look oddly sapient. I remember Halstead’s book well but didn’t realise Caselli’s art was so influential.

Interestingly, and this parallels the conservative images of dinosaurs in cryptozoological lore, fantasy fiction and RPG’s also have some bizarre ideas on Pterosaur anatomy. Check out Fighting Fantasy gamebook “Battleblade Warrior” as an example:

http://fightingfantasy.wikia.com/wiki/File:Figfan31.jpg

A bat-winged toothed Pteranodon with opposing foot claws, and a rhamphorhynchid finned tail, all from 1988! Woohoo!
#13 Steve P
May 25, 2010

I had a jigsaw puzzle with a Quetzalcoatlus (labelled as the “Texas Pterosaur”) which looked very much like these reconstructions but was bright purple and didn’t (as far as I can remember) have any teeth…

These paintings have a really eerie feel to them, despite the inaccuracies!
#14 Dartian
May 26, 2010

I get nostalgic whenever I see Richard Orr’s illustrations. Although I must admit that I’ve never seen his pterosaur artwork before; I’m more familiar with his illustrations of extant animals and extinct mammals. (A Closer Look at Prehistoric Mammals, written by L. B. Halstead and illustrated by Orr and others, was a particular childhood favourite of mine.)

Abyssal:

I don’t think I’ve ever seen an image of *Camptosaurus* that wasn’t white with red stripes.

Surely you’ve seen this?
#15 Tim Morris
May 26, 2010

I’ve started illustrating demons for fun of late, this one definately makes the cut!
#16 Tamara Henson
May 26, 2010

Boy does this bring back memories! The scavenging toothy azdarchids can even be found in kids books published as recently as the late 1990′s. My favorite was the one in the first edition of Gary Gygax’s (the original) Monster Manual for Dungeons & Dragons. The “Giant Pterosaur” illustrated was clearly inspired by the pictures given here.

Speaking of “alays colored the same dinosaurs” do you know it is a tradition to depict stegosaurus as orange in comic books? I read that once in a book on Comic book art in the 1980s and never forgot it. I have also noticed that ever since Anne McCaffrey published the book “Dinosaur Planet” that Pteranodon has a tendancy to be golden brown.

P.S: Darren: someone has hacked into my hotmail address so if you get anything from tamarahenson from that site please disregard and delite it as whoever stole my address is sending spam to those in my mailbox.
#17 Darren Naish
May 26, 2010

Thanks to all for comments. The process whereby prehistoric animals possess a particular colour scheme that is repeatedly depicted by different artists _could_ be regarded as a sort of plagiarism, but I think it’s also a matter of following something that (in cases) looks good. Stegosaurus with bright red plates, for example. In other cases, it’s just lazy, however, and it’s particularly common among artists who just aren’t very good: quite a few recently created crappy CG dinosaurs have been given colour schemes identical to the ones used in Walking With Dinosaurs, for example.

I also agree with everyone who speaks of nostalgia with reference to those kid’s dinosaur books of the 70s and 80s. I think that many dinosaur pics from the 1950s and 60s are truly ugly and offensive to the eye (e.g., Geis’s The How and Why Wonder Book of Dinosaurs), but much of the art from the 70s and 80s was fantastic and wonderful: I mean this in the fantasy or comic book sense, not in the naturalistic/scientifically accurate one. I might do an article on this subject some time.

WRT the Sato et al. paper (comment 11), a response is in the works, I understand. All I will say is that all flying animals aren’t procellariiforms. I’ve now had a quick read of Don Henderson’s paper on mass estimates for pterosaurs. Don makes the point that – at an estimated 544 kg – Quetz. is far removed from the next biggest pterosaur he looked at (Tupuxuara at 22.8 kg), and he uses this point to argue for the possibility of flightlessness for the big bastard. It’s interesting that – while there are two or three azhdarchids that seem to have been similar to Quetz. in size, like Arambourgiania and Hatzegopteryx – most azhdarchids are in a different size class, with wingspans of 3.5-4 m (e.g., Bakonydraco, Azhdarcho, Phosphatodraco and Quetz. sp.). Where are the mid-sized azhdarchids that bridge the gap?

Tamara (comment 16): thanks for clarification. Last night I received an email from ‘you’ saying that you’d been mugged at gun-point while in London, had lost everything, and needed more than $3000 to be wired to you for help. Tone and I became convinced that it was a scam (several things didn’t add up), but if I was more stupid and less poor I could well have fallen for this.
#18 Tamara Henson
May 26, 2010

I’m glad you didn’t fall for it. I have been working overtime and have not been online for almost a week. Then a friend got that bogus e-mail and I found out I was locked out of my account.

Regarding pterosaur art I remember some drawings (especally of Dimorphodon and Pteranodon) during my childhood that were just hideous. I like the more recent puffin and pelican inspired art though I do think that way to many artist take Walking With Dinosaurs to seriously. By the way have you seen the Pteranodon art in the book “In the Presence of Dinosaurs” by John Colagrande, Larry Felder, and Jack Horner? Just lovely, though Pteranodon is my favorate pterosaur and I hope to see one in a movie some day that actually looks like the real thing and not like a Ludodactyus x dragon hybrid.
#19 David Marjanović
May 26, 2010

Aaah, Halstead’s book… memories…

Of note is the fact that every single Iguanodon in the book [...] has a creepy long giraffe-tongue for some reason.

The reason is Louis Dollo, who found a hole in the tip of the lower beak of one specimen, somehow didn’t think it was due to damage to the fossil, and thought a giraffe-like tongue must have passed through that hole. The hole was soon shown to be spurious, but few people knew that it was the only evidence for the tongue!

A real life example of that apocryphal scientist who was so out of touch with reality, that calculated that bumblebee could not fly!

That bumblebees cannot fly like planes, you mean. And indeed they don’t. They fly like hummingbirds instead.

Hummingbirds have remarkably short upper arms and forearms, BTW. The hands are normal, and the wing feathers are again as long as the rest of the wing.
#20 Darren Naish
May 26, 2010

Hummingbird wings: the carpometacarpus accounts for about 60% of the wing skeleton’s length and the humerus makes up about 20% (a carpometacarpus less than 40% of wing skeleton length is more usual). The wing skeleton in hummingbirds (and swifts) contributes less than 40% to total wing length (55-60% is more typical; in albatrosses it’s an exceptionally high 70%), so, yes, their feathers are long and their wing skeletons short.
#21 Anonymous
May 26, 2010

Another Quetzalcoatlus reconstruction that always stumped me was the model that appeared in the Carnegie Collection. Having grown up with the “proper” restoration of the azdarchid, i.e. a long stiff neck, bony crest, and whatnot (though the stork-like pterosaur theory hadn’t come out at the time), I was stumped as to why their Quetzalcoatlus model has such a ridiculously short neck, as short as their Pteranodon.
#22 Stu of the Peak
May 26, 2010

I still have my original copy of Halstead’s book which I received Christmas 1977, and I love the illustrations in it. I did a talk at school on dinosaurs and Quetzalcoatlus got a mention there as up until that point many people were still thinking Doug McClure films when it came to pterosaurs. Another favourite was (and is) Barry Cox’s Prehistoric Animals.
#23 Michele
May 26, 2010

Don’t forget about this version from the unintentionally hilarious film, “Q” http://www.imdb.com/media/rm3931021056/tt0084556
#24 wazza
May 26, 2010

When you’re talking about popular depictions of Quetzalcoatlus, you really can’t go past the Skybax riders in the Dinotopia books, though I’ve no idea how many mistakes were made in their depiction (apart from the obvious one of all dinosaurs being sapient)
#25 Hai~Ren
May 26, 2010

Let’s not forget that Quetzalcoatlus also appeared in Dino-Riders. I’m sure John Sibbick’s reconstruction has also influenced many other artists.
#26 Tamara Henson
May 26, 2010

Q doesn’t count, the Q in the name was meant to refer to the Aztec god Quetzalcoatl not the pterosaur, and in any case the monster turned out to be a female dragon.

Boy do I remember Dinoriders! I actually have their Quetzalcoatlus model that they gave out when you joined their fan club. It was/is a plastic toy with an opening in the belly that a wire attached to a stand fit in so that the animal looked like it was soaring. Interestingly it is a light brown with a darker back.
#27 Darren Naish
May 26, 2010

Back to ‘mid-sized’ azhdarchids… I’ve just looked at the PLoS ONE paper on the new taxon Alanqa saharica. It’s suggested to have a wingspan of 6 m or more, so here’s a taxon that helps ‘bridge the gap’ (see comment 17).
#28 Jaime A. Headden
May 27, 2010

Darren,

Mike Habib remarks that the volumetric model Don used is apparently too large, and would account for the double-mass relative to the other three estimates that have come up (Sato, you and Witton, and Paul).

Don’s conclusion of flightlessness would likely be dismissible were the volumetric model shrunk to something half-again as small, especially since it’s a projection from scaling up a Q. sp. and Habib, Cunningham, and I think you and Witton both agree the model has variable metrics (a different metric per region, etc.) scaling up.
#29 Carlos
May 27, 2010

They look like something horrid that came out of The New Dinosaurs!
#30 Jason
May 27, 2010

Out of curiosity, I checked my collection for any early representations of Quetzalcoatlus. While I didn’t find any additional short-headed or toothed versions other than Halstead (my 1975 edition is titled The World of Dinosaurs, by the way), I did find two other interesting early mentions (links to images included):

1. All New Dinosaurs and their friends (1975) by Robert A. Long and Samuel P. Wells, illustrated by Gregory Irons; http://www.flickr.com/photos/cryptonaut/4644209937/in/set-72157624147189456/

2. Flying Reptiles In the age of Dinosaurs (1976) written and illustrated by John Kaufman; http://www.flickr.com/photos/cryptonaut/4644208183/in/set-72157624147189456/ (The Kaufman book includes several more illustrations of Quetzalcoatlus. See my Flickr set for the rest.)

Looking at these this morning, I noticed for the first time that Kaufman’s picture of Quetzalcoatlus scavenging a dead sauropod seems to be lifted directly from Irons’ earlier illustration! The shape of the feeding Q‘s head is virtually identical, and the sauropod (Alamosaurus, in the original) is essentially a straight copy, right down to the head being obscured by foliage!
#31 Benjamin
May 27, 2010

Good to see some people beyond myself have some love for the illustrations of the 70s and 80s. It seemed as though better artists were being asked to illustrate the books, and, although they had little in the way of accuracy, they excelled at creating plenty of dynamic or moody scenes for a kid to stare at during a rainy afternoon.

Sometimes I wonder how much influence Zallinger had in other artists’ color choices. The red Ramphorhynchus, the blue Camptosaurus, Stegosaurus, Plateosaurus. (I think Peter kept a lot of Rudolph’s color schemes when illustrating his prehistoric animal books from the 80s.)
#32 Mark Witton
May 27, 2010

There are a number of gap-bridging azhdarchids: Zhejiangopterus spans up to 5 m; some Azhdarcho individuals were 6 m; Padian and Smith (1992) reported an azhdarchid somewhere ‘between the large and small morphs of Quetzalcoatlus‘ (5 – 10 m, then); Q. sp. itself is 5 m or so; Godfrey and Currie reported a 5 m span jobbie from Dinosaur Provincial Park and some French material is estimated around the 9 m mark. There may be more.

Regarding Sato et al., Mike Habib and I have something in review at the moment: watch this space.

Don’s recent work on Quetzalcoatlus, as suggested by Jamie, is based on a very outdated (and, even for that time, poorly reconstructed) Quetz. that has a torso twice as long as it should do. If you scale it appropriately, the adjusted mass hits the same rough area as those of Greg Paul, Mike Habib, Jim Cunnigham and me.
#33 Jerzy
May 27, 2010

BTW – what was the MINIMUM flight speed of Quetzalcoatlus? Maybe it could glide very slow just over ground, almost at walking speed?
#34 Jerzy
May 27, 2010

BTW – no pteroid bone, I see. Paleontologists could think again – this bone must have been pretty useful to evolve.

BTW – crazy question. Could pteroid bone of some pterosaurs support a notch or a fully separated little wing membrane, acting like alula in birds?
#35 Warren B.
May 27, 2010

I’ve seen my share of kids’ books – including that Quetzalcoatlus illustration, somewhere… – but most of my memories are from the Orbis ‘Dinosaurs!’ partworks in the early nineties. A lot of the illustrations in that ended up on the London NHM site.
Even for a 10yo it was easy to see who was good and who was taking the proverbial, and some illustration briefs obviously included colour schemes for the sake of uniformity. Red and beige camptosaurs, green and orange allosaurs, white Quetzalcoatlus with red and yellow heads, etc. I still see some old palaeoart for the first time, but recognise it from bad-to-middling copies in that mag.

On the topic of gaming, tabletop miniatures are for the most part still stubbornly outdated and plain sloppy. A picture is worth a thousand words: these examples were sculpted by a ‘palaeoartist’, and released only weeks ago.
http://tiny.cc/xknrc
AFAIK only David Krentz is currently producing good archosaur models at those kinds of scales; probably because he’s approaching it as a palaeoartist (a good one) rather than as a miniature sculptor.
#36 scidog
May 28, 2010

maybe we should just go back to showing them all in gray and put the emphasis on the details.
#37 David Marjanović
May 28, 2010

Could pteroid bone of some pterosaurs support a notch or a fully separated little wing membrane, acting like alula in birds?

Whenever soft tissue is preserved in pterosaurs, the pteroid supports the propatagium. So, no.
#38 Tamara Henson
May 28, 2010

“On the topic of gaming, tabletop miniatures are for the most part still stubbornly outdated and plain sloppy.”
Posted by: Warren B.

No doubt about it, but I’m a collector and the dinorider mini is surprisingly acurate for a 1980′s toy, even though it is built more like Pteranodon. At least it doesn’t have teeth which is more than you can say for Jurassic Park’s “Pteranodons”. Oh well, I can just pretend that they are Ludodactylus .

As for David Krentz, I just saw one of his models in Paleo Times magazine. Very very nice.

“BTW – what was the MINIMUM flight speed of Quetzalcoatlus? Maybe it could glide very slow just over ground, almost at walking speed?”

Posted by: Jerzy

That’s a good question has there been any work done regarding the speeds of the various pterosaurs on the ground or in the air?
#39 kris
May 31, 2010

maybe we should just go back to showing them all in gray and put the emphasis on the details.
Posted by: scidog | May 28, 2010 12:20 AM

if that means darren’s gorgeous drawings, i’m all in favour!
#40 Steve P
June 11, 2010

Is it just me, or has the artist mistaken the naris for the orbit when looking at (something like) a Pterodactylus skull? The position of the ear in the top painting would seem to suggest this; could this explain why the skull has been reconstructed in this way? Doesn’t explain the intensely long neck though…
#41 ranggaw0636
July 18, 2010

Don’t forget about this version from the unintentionally hilarious film, “Q” http://www.imdb.com/media/rm3931021056/tt0084556

I’m remember watching this when i’m still a small kid
#42 Isis the Scientist
July 18, 2010

We just learned about quetzocoatlus on Dinosaur Train this morning. I can’t wait to show Little Isis your blog post when he gets up tomorrow.