Friday 7 June 2013

Pterosaurs: Natural History, Evolution, Anatomy: out at last

Ornithocheirus and Anhanguera welcome you to Pterosaurs: Natural History, Evolution, Anatomy. Double spread from Witton (2013).
So, a rather unexpected and heavy package arrived in my office this week holding copies of Pterosaurs: Natural History, Evolution, Anatomy. The Tweet on the Street is that preorders are already being dispatched. Given that I thought we wouldn't be handling actual copies of this thing until late June, these were pleasant surprises indeed. Slight slop with delivery dates around the world aside, I think it's about time to declare this thing as 'published', which I'm very excited about to say the least. I'm not alone in being happy with this development, however. The first review of Pterosaurs hit the web on Tuesday, courtesy of Brian Switek at Laelaps. I'm happy to report that Pterosaurs emerged rather well from it's first wash:
"Witton’s new tribute to pterosaurs gives these fantastic fossil creatures a much-needed makeover... If you’re truly invested in learning about pterosaurs, Witton’s book is a wealth of information that will be of great use to both specialists and curious general readers."
Nice words indeed, and hopefully a sign that the 2.5 years(!) spent on this project were not wasted. I've been deliberately cagey about many of the details of Pterosaurs. A breakdown of the book chapters was revealed a couple of years back, but many of my favourite bits of the book have been held back so as not to pre-empt it's publication. Now that the book is available, I guess it's time to tell people what to expect and, perhaps more importantly, why you should fork out £19.46 for a copy when you could track down, or may already own, Dave Unwin's (2005) The Pterosaurs From Deep Time or Peter Wellnhofer's (1991) Encyclopaedia of Pterosaurs

What is a Pterosaurs?
Pterosaurs is meant to provide an interesting read for researchers and diehard enthusiasts, while still being approachable for those who are yet to really acquaint themselves with flying reptiles. If you're familiar with the Unwin and Wellnhofer books, you know the tone I've aimed for. (Those interested in reading a sample of the text will want to download the first chapter from Princeton University Press, and check out an early draft [essentially unchanged in the published text] of Chapter 17.) Pterosaurs is, of course, more up to date than either of these books. Only seven years passing between this book and the last, but the differences are quite pronounced. Despite both Unwin's and Wellnhofer's books dating very well, whole groups of pterosaurs have been discovered since their publications (e.g. 'boreopterids', chaoyangopterids, wukongopterids, and many more in the case of Wellnhofer's tome) and ideas of pterosaur lifestyles and habits have changed considerably. It's of small significance in this field of three modern pterosaur books but, by default, Pterosaurs is the most up to date synthesis on these animals currently available.

Thalassodromeus sethi, a pterosaur with a most unfortunate name, showing a baby Brazilian spinosaur that the food chain works both ways. One of my favourite paintings from Witton (2013).
Pterosaurs is meant to combine the best aspects of preceding pterosaur books into one package, putting Unwin's terrific introduction to the group together with Wellnhofer's coverage of all pterosaur species and important fossils. This results in nine chapters covering the broad-strokes of pterosaur research: the history of their discovery, evolutionary origins, osteology, soft-tissues, locomotion (flight and terrestrial locomotion are discussed separately), palaeoecology and extinction. The other 16 chapters focus on specific pterosaur groups, each featuring a history of discovery, distribution maps, overviews of anatomy (including soft-tissues, where known) and discussions of palaeoecology. These latter chapters broadly follow the phylogenetic scheme of Lü et al. (2010) but, because that will not please everyone, alternative taxonomic proposals are mentioned and discussed where relevant (though hopefully not at expense of readability!). Attempts to present different sides to contentious issues are continual throughout the book. As readers will discover, there is still a lot to learn about these animals and it would be foolish to present only a single view as 'right' when pterosaur science continues to evolve and change. The drive to give everyone fair hearing resulted in a reference list of over 500 works and, hopefully, this will make the book a useful starting point for students new to pterosaurs and wanting to hit the primary literature. (Incidentally, Lü Junchang needs to take a bow as probably the most prolific modern pterosaur worker, his portion of the citation list dwarfing virtually everyone else's despite only beginning in the mid-nineties. Way to go, JC!)

Shiny new things
Pterosaurs is certainly not just a straight review of pterosaur literature, however. Some aspects of the book present wholly new information and ideas, or provide alternatives to existing hypotheses. This particularly applies to the 'palaeoecology' sections of the later chapters, as pterosaur lifestyles are frequently poorly researched. In the worst cases, no lifestyle hypotheses have ever been proposed or are half-sentence afterthoughts thrown onto the end of descriptive papers, so are of little scientific merit. In such instances, I've inserted my own ideas about what these animals may have done based on their gross anatomy and form (including, as depicted above, the proposal Thalassodromeus was a predator of moderately-sized terrestrial prey, following numerous lines of evidence that it's proposed skim-feeding habits are likely incorrect [see Humphries et al. 2007] and its unusually robust, peculiar skull).

Old vs. New. What did pterosaur ancestors look like? We don't know, but the traditional view of them as generic flying reptiles (left, inspired by Wellnhofer 1991) has to go all the same. Right, a newly imagined pterosaur ancestor inspired by recent work into pterosaur origins, representing the third stage ('HyPtA C') of five proposed stages of pterosaur evolution. Both images from Witton (2013). 

One of the highlights of these 'new proposals', for me at least, is a complete retooling of the 'protopterosaur' idea first proposed by Rupert Wild (1978 and others), and then popularised by Wellnhofer (1991) and Unwin (2005). Because of the ambiguity about pterosaur ancestry, Wild and his followers proposed a fairly-generic, lizard-like animal as a hypothetical pterosaur ancestor (above left), which doesn't really fit with modern notions of pterosaur evolution. Although there is still some mileage left in the controversy over pterosaur origins, the idea that pterosaurs were close relatives of dinosaurs is the current hypothesis to beat and, with that in mind, I reworked the likely form of their hypothetical ancestor. Indeed, I tried to imagine a whole series of ancestral species, the 'HyPtAs' (Hypothetical Pterosaur Ancestors), and descibe how they may have developed from a small, sprightly terrestrial reptile to the first actively flying vertebrate. The animal shown at right, above, is a 'stage C' HyPtA, 3 of 5 in this sequence.

More than just words
Part of the reason this book took 2.5 years to put together was because of the amount of new illustrations it warranted. The vast majority of diagrams and graphics - ranging from labelled anatomies (below), skeletal reconstructions, myologies for all major body parts and others - are new, but the book is also well stocked with photographs taken by myself and some very generous colleagues. Of course, the book also features a high number of life restorations of many pterosaur species, sometimes set in backgrounds (as per the painting of Thalassodromeus, above) or in more informative lateral views. Most of these were produced specifically for the book, so most should be new to readers. It's hoped that the abundance of skeletal diagrams and muscle reconstructions should be helpful to artists, and, indeed, some bits of text and imagery are almost provided with artists in mind (Fig. 7.6 may be particularly helpful). The paintings of this book could easily have dissolved into a series of images of pterosaurs flying, but efforts were made to render pterosaurs in never-seen-before guises. There's a bunch of Pteranodon diving several metres into water, Lacusovagus performing a mating dance, an azhdarchid struggling against the 'nuclear winter' of the K/T exinction, Dsungaripterus fighting with one another, and a couple of species just sitting the hell down (quite unintentionally, there's a spirit of All Yesterdays running through a lot of the paintings). The intention was to capture some of the possible diversity in landscape, habits and scale represented by these animals and represent them not just as scientific concepts, but as genuine individuals of real, once-existent species

What fossils reveal about pterosaur wings. Note the differences in wing construction in non-pterodactyloids (left) and pterodactyloids. One of the most informative and detailed diagrams in Witton (2013).
And if I say much more, you won't need to buy it
So that's what to expect from Pterosaurs then, folks, available now in hardback and Kindle editions. I'm very pleased to hear from social media and elsewhere that numerous friends have copies already on order, and hope that you enjoy reading it once it arrives. This message particularly extends to those who've supported the project since I announced it back in August 2010. It was certainly a lot of fun, and very educational putting it together, and I look forward to hearing what everyone thinks once they see it.

  • Humphries, S., Bonser, R. H., Witton, M. P. and Martill, D. M. 2007. Did pterosaurs feed by skimming? Physical modelling and anatomical evaluation of an unusual feeding method. PLoS biology, 5, e204.
  • Lü, J., Unwin, D. M., Jin, X., Liu, Y. and Ji, Q. 2010. Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proceedings of the Royal Society B: Biological Sciences, 277, 383-389.
  • Unwin, D. M. 2005. The Pterosaurs from Deep Time. Pi Press, New York, 347 pp.
  • Wellnhofer, P. 1991. The Illustrated Encyclopaedia of Pterosaurs. Salamander Books Ltd., London. 192 pp.
  • Wild, R. 1978. Die Flugsaurier (Reptilia, Pterosauria) aus der Oberen Trias von Cene bei Bergamo, Italien. Bolletino della Societa Paleontologica Italiana, 17, 176-256.
  • Witton, M. P. 2013. Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.


  1. I have pre-ordered it,I can't wait !
    I'm wondering how similar are Pterosaurs and Dinosauria integuments ?


  2. Hey Oliver,

    "how similar are Pterosaurs and Dinosauria integuments?"

    That's a key question for those interested in archosaur evolution. There are researchers who have said dinosaur and pterosaur fluff is very similar or even the exact same thing (i.e. pterosaurs have early versions of feathers) and those who say the exact opposite (pterosaurs developed fluffy 'pycnofibres' uniquely of other reptiles). As has been said time and again here and elsewhere, if they are the same, and pterosaurs and dinosaurs are sister groups, the implication is for an ancestrally fluffy Ornithodira and scaly dinosaurs would have lost their fluff, instead of select species developing it.

    1. That reminds me of something similar: Based on the available evidence, any idea whether the air-sac system was more likely 1) inherited from the common dino-ptero ancestor & lost secondarily in ornithischians, or 2) developed by dinos & pteros independently? Either way, I'm assuming that dinos & pteros share a close common ancestor.

    2. This is another million dollar question. There's some tentative evidence that some non-dinosaurian dinosauriforms may have had post-cranial air sacs like those of birds, which might support the idea that they developed once in a dinosaur + pterosaur clade, with ornithischians losing them secondarily. There's still a lot of discussion on this issue, though.

  3. Mike from Ottawa8 June 2013 at 11:28

    It would be really amusing if it turned out that not only were birds not the first flying vertebrates but that they didn't even 'invent' 'feathers'.

    Got my copy of Pterosaurs on pre-oder. You must be pleased as Punch to finally have copies of your own book in your hands after all that work. I offer you my heartiest contrafibularities!

    1. "It would be really amusing if it turned out that not only were birds not the first flying vertebrates but that they didn't even 'invent' 'feathers'."

      I guess we already know that birds didn't develop feathers themselves, as they're clearly apparenent in theropods which are only distantly related to Aves. But yes, there's a chance that feathers may not even be a 'dinosaur thing'.

      " You must be pleased as Punch to finally have copies of your own book in your hands after all that work. I offer you my heartiest contrafibularities!"

      Thanks muchly. It's still a little surreal. I was getting quite used to it as a ethereal concept rather than an actual object, but now it's on my bookshelf. Am looking forward to seeing what you all think of it. Preorders are already being dispatched, so there the wait shouldn't be too long.

  4. Congrats on your pterosaur book being published. Any plans for a dino book in the near future? In any case, it's already proven very helpful to me (I was looking for something on Google Books & it came up).

    "Dave Unwin's (2005) The Pterosaurs From Deep Time or Peter Wellnhofer's (1991) Encyclopaedia of Pterosaurs."

    I have a couple of questions about that.

    1stly, are you comparing your new book only to those in the above quote b/c they're the next-best pterosaur books to yours or b/c they're the most similar to yours (I.e. Post-Dino Renaissance books for "the enthusiast": )? I ask b/c of the absence of Seeley's "Dragons of the Air" & Buffetaut/Mazin's "Evolution and Palaeobiology of Pterosaurs" (although I figured they might have been too old or too technical).

    2ndly, how is your new book different from "Pterosaurs: Flying Contemporaries of the Dinosaurs"? Is the latter just a more technical version of the former or is there more to it?

    "Right, a newly imagined pterosaur ancestor inspired by recent work into pterosaur origins, representing the third stage ('HyPtA C') of five proposed stages of pterosaur evolution."

    Reminds me of Paul's Proavis ( )? Out of curiosity, did have PP in mind when coming up w/your protoptero? Also, are Scleromochlus & Lagosuchus discussed in the ancestry chapter?

    1. "(although I figured they might have been too old or too technical)"

      Exactly. Both Dragons of the Air and the Buffetaut and Mazin volume are only of interest to specialists because of their age and technical nature, respectively. Seeley's book would be understandable to general readers, but it's over 100 years old and doesn't reflect modern thoughts on pterosaurs.

      "2ndly, how is your new book different from "Pterosaurs: Flying Contemporaries of the Dinosaurs"? Is the latter just a more technical version of the former or is there more to it?"

      Pterosaurs is a lot more detailed and comprehensive than P:CoftD. The Veldmeijer et al. book is a fairly general look at the group as a whole, but Pterosaurs gets right under their skin. Literally, in chapter 5.

      "did have PP in mind when coming up w/your protoptero?"

      No. The HyPtA concepts were developed in direct response to Wild's lizard-like 'protopterosaur' because I didn't think it was a useful model for modern pterosaur researchers.

      "Also, are Scleromochlus & Lagosuchus discussed in the ancestry chapter?"

      Scleromochlus gets a fair hearing (and figuring) as a possible pterosaur ancestor, but Lagosuchus doesn't get much of a look in because of its likely affinities towards the dinosaur end of the tree rather than the pterosaur one.

    2. When you say 'figuring', are there new photographs of the molds?

    3. Sadly, no. Just a skeletal reconstruction. Mike Benton's paper on Scleromochlus remains, as far as I know, the place to go for information on it.

  5. I'm now even more hyped for your book (I see it's on Kindle now, by the way). Not only is your artwork gorgeous, but you certainly have a way with words-- I quite liked the first chapter's introductory discussion of real vs. "hokum" pterosaurs.

    1. I took a lot of the more flippant remarks out towards the end of the writing process: there simply wasn't space to keep them all in. Thankfully, a few of them remain. There's a great line from American Dad in Chapter 24.

  6. Looks amazing Mark, I'll definitely have to look into getting it! Pterosaurs have always intrigued me, as they were certainly more interesting and bizarre than the leathery and skinny demonic-looking creatures which have been prominent in popular culture. They must have been a very unique group of animals which would be spectacular to behold, and I adore the thought of giant, semi-terrestrial and fuzzy azdarchids. I also really enjoyed the fact that you made it clear in the first chapter that these weren't Hollywood monsters, but rather animals. I really love that many people are pushing awareness of this, as I've always hated the popular conception of prehistoric animals as monsters in constant battle.

  7. My copy arrived today. Just... wow. It's even better than I expected, and I did expect a lot. There's all the information one could wish for, and the book looks absolutely beautiful from cover to cover.

    I also love the picture captions.

  8. Late to the party here, but I just finished reading this book and enjoyed it greatly. It's just the kind of book I've wanted about Pterosaurs for a long time. Thanks!