Wednesday, January 27, 2010

Pterosaur books to know and love, part 1: The Illustrated Encyclopedia of Pterosaurs

Your local bookshop is probably full of more popular books on dinosaurs than you would ever want to own. There’s the ‘spotters guide’ type, more encyclopaedic efforts, swish-looking coffee table tomes and even handy pocket-sized variants that ensure wherever you’re are, whatever you're doing, you can always find out if it was Ankylosaurus or Euoplocephalus that had the coolest dermal scutes. Of course, if you want to expand your knowledge of fossil forms beyond terrible lizards, you rapidly find that there’s a very short list of books to own. What popular books are there on marine reptiles, stem-mammals, basal archosaurs or the hundreds of other fossil tetrapods you may want to know about? Heck, what about fossil fish or even invertebrates, the forms that really make up the bulk of the fossil record? It’s no secret that there’re virtually no popular texts about these critters at all: despite the fact that they’re just as interesting as dinosaurs, publishers ignore them and push out the same rehashed texts on Tyrannosaurus and his chums again and again.

Dinosaur domination of bookshelves has meant that even extraordinary, spectacular fossil animals with a heavy palaeocult following – such as pterosaurs – are rarely given page space to shine. One of the first E-mails received was a suggestion from reader Bob Meek that we compile a list of recommended pterosaur books and, while it’s an excellent idea, I’m afraid the list of books we could recommend is very short. This isn't because we're snobbish and lazy (well, not entirely), but more reflective of the scant number of accessible pterosaur books. In fact, I can only think of two books that are really essential items for casual pterosaurophile bookshelves: the classic 1991 Illustrated Encyclopedia of Pterosaurs by Peter Wellnhofer and David Unwin’s more recent The Pterosaurs from Deep Time (2005). Both are out of print and will require both sleuthing of online second-hand booksellers and probably a fair amount of cash to purchase, but, as I'll try to explain across my next two posts, there are plenty of reasons why budding pterosaur nerds should bother.

The Godfather of pterosaur books, by the Godfather of pterosaurs

We begin, then, with Wellnhofer’s 1991 encyclopaedia (cover shown above, stolen from here). You may be wondering why I’m recommending that you track down a volume that was written almost two decades ago, saw it’s last print run at the turn of the millennium and, in many respects, reports virtually no discoveries from the Golden Age of pterosaur research we’re now enjoying. Circa 1991, there was practically no consensus on pterosaur terrestrial locomotion, most of Azhdarchoidea was unheard of, the Araripe and Jehol Groups were only just beginning to yield pterosaur remains and modern methods of phylogenetic analysis were still being born. It's a valid question, then, to ask why you should bother buying a book that misses out so much new and exciting information. The answer is a wholehearted 'yes', and here's why.

Firstly, it's hard to think of an author who would be more suited to write a pterosaur encyclopaedia than Peter Wellnhofer. For those who don't know, Wellnhofer is the chap who virtually single-handedly resurrected pterosaur research with his landmark monographs on Solnhofen and Santana Formation pterosaurs (Wellnhofer 1970, 1975, 1985, 1991b), pterosaur handbook (the pterosaur researcher’s equivalent of a car owners manual; Wellnhofer 1978) and much else besides. In essence, Wellnhofer created the landscape on which modern pterosaur researchers would begin to work and, having watched that landscape begin to blossom throughout the 80s, it seems only fitting that Wellnhofer should be the man to summarise work on pterosaurs up to that point. What’s more, Wellnhofer’s thoroughness, comprehensive knowledge and accessible writing style – the same attributes that make his technical work so useful decades after their publication – are all evident throughout the book. The prestige of the author alone, then, tells you that The Illustrated Encyclopedia of Pterosaurs won't be garbage.

Wellnhofer’s encyclopedia was originally published as the sister volume to David Norman’s 1988 Illustrated Encyclopedia of Dinosaurs and the two would eventually be bound together in 2000. While both books devote their first chapters as introductions to their respective groups (discussing their ancestry, anatomy, role in popular culture and a history of their research), the pterosaur encyclopedia does not take a phylogenetic approach to reviewing genera like the dinosaur volume, instead progressing through geological time to introduce pterosaur taxa in their relevant stratigraphic positions. Wellnhofer does make comments about possible pterosaur relationships in his text (they’re quite archaic: Dimorphodon is suggested to be the direct evolutionary ancestor of Anuroganthus, for instance) but they have no relation to the structure the book. Most pterosaur genera known in the early 90s are mentioned and, by discussing and illustrating the majority of these with specimen photographs and life restorations (such as the adjacent John Sibbick illustration of Solnhofen pterosaurs found on p. 86-87, and lifted wholesale from here), this section makes up the meat of the book. The end chapters overview thoughts on pterosaur lifestyles, locomotion, extinction, reconstruction and key museums to visit to see pterosaur remains. As an overview of pterosaurs, then, it’s a pretty comprehensive piece of work and, to my knowledge, there’s not been a volume of similar scope published since.

Imagery is almost always given precedence over text with plenty of large, colour pterosaur paintings and sketches by John Sibbick, pterosaur restorations and technical drawings by the author, buckets of photographs and schematic diagrams of pterosaur anatomy adorning every page. The reliance on imagery gives the impression that the book is more suited to pterosaur enthusiasts or even children than researchers, but complementing the imagery is text loaded with citations of technical pterosaur literature. Granted, the citations are only listed once on context specific pages and finding a particular reference can be awkward but, seeing as some parts of the book stand out as the best available synopses of given topics (the chapter summarising the history of pterosaur research is worth pointing to here), these citations make the Illustrated Encyclopedia of Pterosaurs an incredibly useful research aid. Equally, some of the specimen images are the best – and occasionally the only - published photographs available of certain material (like the non-Quetzalcoatlus Javelina Formation azhdarchid, for instance [p. 144]*) and, to my knowledge, no resource published since has offered so many useful pterosaur images in a single volume.

*This rarely discussed specimen, TMM 42489-2, is actually really intriguing: represented only by a fragmentary rostrum and mandible (but still 80 cm long despite its incompleteness), Wellnhofer’s book mistakenly refers it to Quetzalcoatlus but, as demonstrated by Kellner and Langston (1996), it clearly belongs to another taxon. It’s since been called a ‘tupuxuarid’ (Kellner 2004, Martill and Naish 2006) but more recently referred to Azhdarchidae (Lű et al. 2008; Witton 2009). Whatever it is, it’s clearly distinct from
Quetzalcoatlus and shows that another large, and apparently snub-nosed, pterosaur was present in the Javelina Formation. I reckon it should form the subject of a blogpost sometime, in fact.

The bad bits

Of course, it was be foolish to say that the book hasn’t dated: it has. The last two decades has seen so much pterosaur research published that many ideas familiar to modern readers are noticeably absent from The Illustrated Encyclopedia of Pterosaurs. There’s no sexy images of UV-illuminated pterosaurs, quad-launches, flamboyantly crested forms like Tupuxuara or Tupandactylus, no istiodactylids outside of southern England, no cladograms of pterosaur relationships, precious little on pterosaur trackways and much else besides. Equally, Sibbick’s restorations, though beautifully painted, are rife with anatomical inaccuracies such as pterosaurs demonstrating twiglet-like limbs, hanging, bat-like from tree branches by their feet, a considerably overly muscled tail on the Rhamphorhynchus (see image, above), a duck-billed Istiodactylus, an Anhanguera that is disarticulating its neck vertebrae to grab a fish (see Sibbick Santana Formation pterosaur spread from pages 128 - 129 of Wellnhofer [1991a], above. Swiped from here)... you get the idea. Many of these inaccuracies are classic problems with pterosaur palaeoart but, crucially, most could’ve been avoided if more attention had been paid to pterosaur anatomy. You couldn’t, for instance, fit the pterosaur arm skeleton within the restored forelimbs of Sibbick's pterosaurs, meaning it violates one of the most fundamental requirements of any palaeoart: the restored animal's flesh should fit around it's skeleton, right? The biggest gear grind, though, comes from the text: virtually every pterosaur in the book is suggested to eat fish regardless of skull moprhology, dentition, wing anatomy and everything else. Like, seriously: practically all of them. And the jaw morphology of Dsungaripterus is even favourably compared to that of modern skim-feeders. Sheesh.

But that’s OK

Still, these are minor niggles compared to what is offered in the rest of the book. Simply put, the comprehensiveness and straightforwardness of its approach mean it is the best summary of the first 200 years of pterosaur research out there, full stop. No serious pterosaur enthusiast or researcher should be without a copy and, although out of print for 10 years, you can still find some cheap(ish) copies online. Top buyer’s tip: if you’re not fussed about buying an original 1991 print, the combined Norman/Wellnhofer volume is often cheaper and, aside from the page numbers, presents exactly the same content as the standalone 1991 version. You can then use the money you save buying that to purchase Dave Unwin’s 2005 book, The Pterosaurs from Deep Time, which is what I’ll try to persuade you to read next time.


  • Kellner, A. W. A. 2004. New information on the Tapejaridae (Pterosauria, Pterodactyloidea) and discussion of the relationships of this clade. Ameghiniana, 41, 521-534.
  • Kellner, A. W. A. and Langston, W. Jr. 1996. Cranial remains of Quetzalcoatlus (Pterosauria, Azhdarchidae) from Late Cretaceous sediments of Big Bend National Park. Journal of Vertebrate Paleontology, 16, 222-231.
  • Lü, J., Unwin, D. M., Xu, L., and Zhang, X. 2008. A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution. Naturwissenschaften, 95, 891-897.
  • Martill, D. M. and Naish, D. 2006. Cranial crest development in the azhdarchoid pterosaur Tupuxuara, with a review of the genus and tapejarid monophyly. Palaeontology, 49, 925-941.
  • Norman, D. 1988. The Illustrated Encyclopedia of Dinosaurs. Salamander Books, London. 208 pp.
  • Unwin, D. M. 2005. The Pterosaurs from Deep Time. Pi Press, New York, 347 pp.
  • Wellnhofer, P. 1970. Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Bayerische Akademie der Wissenschaften, Mathematisch- Wissenschaftlichen Klasse, Abhandlugen, 141, 1-133.
  • Wellnhofer, P. 1975. Die Rhamphorhynchoidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Palaeontographica A, 148, 1-33, 132-186, 149, 1-30.
  • Wellnhofer, P. 1978. Handbuch der Paläoherpetologie. Teil 19: Pterosauria. Gustav Fischer Verlag, Stuttgart. 82 pp.
  • Wellnhofer, P. 1985. Neue pterosaurier aus der Santana-Formation (Apt) der Chapada do Araripe, Brasilien. Palaeontographica. Abteilung A, 187, 105-182.
  • Wellnhofer, P. 1991a. The Illustrated Encyclopedia of Pterosaurs. Salamander Books Ltd., London. 192 pp.
  • Wellnhofer, P. 1991b. Weitere pterosaurierfunde aus der Santana-Formation (Apt) der Chapada do Araripe, Brasilien (Translated title: Additional pterosaur remains from the Santana Formation (Aptian) of the Chapada do Araripe, Brazil). Palaeontographica Abt. A, 215. 43-101.
  • Witton, M. P. 2008. The Palaeoecology and Diversity of Pterosaurs. Unpublished Ph.D. thesis, University of Portsmouth. 307 pp.


  1. As a quite add-on, I know of translated versions that are available in German and Spanish and I belive French too. There are also at least two different editions in English with different colours and slightly different content.

    Not that this should affect anyone going out to buy a copy, but there are several options, especially for non-English speakers.

  2. Hi Ian,

    That's a forgiveable error: the antler-like crests of Nyctosaurus were first published in 2003, over a decade after the book was written and post-dating both publication dates. That said, Salamander books didn't have the book updated for their 2000 reprint, either, so it seems unlikely that they'd have changed it anyway.

  3. I was able to peruse a copy in the Naturkunde Musuem in Zurich, and was impressed by the informational layout of many of the graphics which help the layman like myself to associate a species with time and place. After reading this post, I found a second-hand copy for under 30 Euros. Thanks for the review!

  4. I've got both the Wellnhofer and Unwin books. It's fascinating to see the changes over the time between the two books. Unwin's book is simply one of the best, if not the best, popular books on any extinct animals ever. Helps, of course that it's on the most fascinating extinct vertebrates ever.

    Anyone who's interested enough in pterosaurs to have read this far and lacks either of the books should stop reading this comment and hunt up their own copies. Unwin's book can be had, used, for about $10 US in various places, which is a ludicrous price considering the only sane reason to part with a copy is death. You can get the Wellnhofer book, together with the Norman's dinosaurs (also a book I like a lot) for a ridiculous $3 US. What is wrong with people!

    OK, rant aside, get these books. Now.

    Mark: it is good to read a review even when I have the books, to hear what stands up, what is wrong and what has been superseded. Thanks.

    Mike from Ottawa

  5. Huh, a cover of the crappy 2nd edition is used above, oh the shame. True pterosaurophiles all own the valuable and superior first edition, surely? Like an idiot, I bought this at full price before the remaindered 2nd edition appeared.