Monday, October 11, 2010

Pterosaur books to know and love, part 2: The Pterosaurs from Deep Time

Many, many moons ago, the Heroes of Pterosaur.Net were suggested to provide a reading list for people who just can’t get enough of our leathery-winged chums. Longer-term Pterosaur.Net groupies with good memories may recall that our suggested shortlist was, well, very short with only two notable tomes: Peter Wellnhofer’s 1991 Illustrated Encyclopaedia of Pterosaurs and David Unwin’s The Pterosaurs from Deep Time (2005). You were promised reviews of both: we looked at the Encyclopaedia back in January (seriously, where is 2010 going?) and, today, we’re casting our critical eyes over The Pterosaurs from Deep Time. If it sounds like the pterosaur book for you, however, you’re going to have to go Sherlock to find it: it’s been discontinued for a number of years and only loiters now in second-hand stores. However, Mike from Ottawa informed us last time that he found a cheap copy in an online store that only set him back $10, so you may not have to pay through the nose to obtain one. Question is, of course, do you want to own it in the first place?

The best pterosaur book since the last one
Believe it or not, there have only ever been three popular English books on pterosaurs published: Harry Seeley’s Dragons of the Air (1901 – yes, 1901), Wellnhofer’s Encyclopaedia, and the organ under our spotlight today, The Pterosaurs from Deep Time. The former books set a high benchmark for treatment of their subject matter: Seeley’s book is a classic, a slice of pterosaur history that summarises Seeley’s heretical ideas on pterosaur origins, biology and ecology. Seriously diehard pterosaur aficionados will want a copy but, alas, unless you find a lucky copy in a second hand shop, you’ll have to fork out a lot for it (cheapskates like me, though, can download the entire thing, for free, from here). Wellnhofer’s effort, as discussed previously, remains an essential reference tool for anyone with an interest in flying reptiles and has aged with dignity over the last 20 years. In fact, the only thing that Wellnhofer got wrong with his book was his timing: it was published just before the recent pterosaur research bonanza started (fuelled by spiffing new specimens from Brazil and China) and, therefore, doesn’t report the cohesion of ideas that he may have if he wrote it in 2000, say.

So, when Dave Unwin entered this arena with his book in 2005, he wasn’t exactly swamped with competition. With his main contenders being a dusty, ultra-rare historical tome and an excellent but rather old-fashioned encyclopaedia, he could’ve produced a sloppy, poorly-illustrated mess and still stood a chance of writing the best modern pterosaur book. Thankfully for us, The Pterosaurs from Deep Time continues the trend of its predecessors and is an excellent, informative presentation of modern ideas in pterosaur research. Given the prestige of its author, this isn’t a huge surprise: David Unwin is widely recognised as one of the foremost pterosaur experts in the world. Beginning his career with an assessment of the Victorian taxonomic mess that is the Cambridge Greensand pterosaur assemblage, Unwin has since worked on pterosaurs from all over the world and made significant contributions to research on virtually all aspects of their palaeobiology (including [deep breath]: pterosaur phylogeny, several geographical and taxonomic reviews, details of their flight apparatus, pterosaur respiration, reproduction, terrestrial locomotion and, most recently, bringing the world Darwinopterus). Given that his writing style is also extremely breezy and clear, he's surely one of the most suitable pterosaur palaeontologists around to summarise the current status of pterosaur knowledge. Hence, saying Unwin’s book is ‘the best since the last one’ is not only true by default, but also a considerable complement in saying that Deep Time is a worthy follow-on to Wellnhofer’s Encyclopaedia.

The pterosaur story
Although comparisons between Deep Time and the Encyclopaedia are inevitable, they're not entirely fair because the two books have very different formats. The Encyclopaedia is quite schematic in its layout with clearly defined sections and liberal use of detailed, technical diagrams: it feels far more like a textbook than something you would read from cover to cover. Deep Time, by contrast, is far more prosaic, flowing from chapter to chapter in a continuous way that makes it very easy to read in entirety. It begins with an introduction to pterosaurs and the concepts of fossilisation and geological time before moving onto an overview of pterosaur phylogeny (based on Unwin’s 2003 work), their anatomy, locomotion, reproduction and, finally, an attempt to tell the complete tale of pterosaur evolution. The encompassing approach of this latter chapter is very characteristic of Unwin’s work: many of his technical publication feature discussions of the broader implications for whichever topic is under scrutiny, be it eggshells (Unwin and Deeming 2008) or wing membrane distribution and terrestrial locomotion (Unwin 1999). The story presented in the penultimate chapter of Deep Time seeks to tie all the threads from the prior chapters together, explaining how and why pterosaurs took to the skies, the rise and success of pterodactyloids and their eventual extinction. Beyond this lies a wealth of footnotes and references to keep technical pterosaur buffs happy, along with a list of valid pterosaur species. As may be expected, the information was, at the time of writing, entirely up-to-date and we’re given the full benefit of the first 15 pterosaur bonanza years, including pterosaur brain CT scans to the wealth of soft-tissue data revealed by Brazilian and Chinese fossil Lagerstätten. There is, therefore, a wealth of information and interpretation held in Deep Time and, thanks to Unwin’s frequent quips and asides, it’s as enjoyable to read as it is informative.

We’re treated to glossy, full-colour illustrations for much of the book and, while the sources of many images will be known to those familiar with other pterosaur literature, most diagrams and drawings are well-executed and tie in nicely with the text. The excellent photographs, many of which are of previously unpublished details of well-known specimens or entirely new finds, are worthy of mention as are the superb pterosaur paintings by Todd Marshall that are dotted throughout the book (see image, above, of a Marshall Quetzalcoatlus, along with the Sordes on the book cover at the top of this post). I have a soft-spot for Marshall’s work: his animals and environments look refreshingly imperfect, a bit like they’ve been left outside in the wind and rain for some time. Plus, his style is one of the most striking and recognisable among the modern crop of palaeoartists. His punky, scruffy pterosaurs are no exception, and the depth he gives to his environments creates the impression of a broader world beyond the confines of the canvas. Happily, his pterosaur anatomy and postures aren’t too bad either, making his images a handy source of modern pterosaur restorations.

So it’s praise, praise, praise, for Deep Time, then, but it’s not all good. For one thing, while most of the figures are absolutely spiffing, a few are a bit shaky with ugly, blocky lines, flat colours and poorly-defined details. This never becomes so much of a problem that you cannot see what the figure represents and there are only a handful of instances in the entire book, but the difference in quality between some illustrations is marked. The largest issue I have with Deep Time, however, is that sometimes the reader is given the impression that we know more about pterosaurs than we actually do. Take, for instance, Unwin’s arguments that a lack of pterosaur footprints prior to the mid-Jurassic is indicative of poor terrestrial abilities in non-pterodactyloid pterosaurs: is the record of pterosaur footprints really that complete that we can conclude this? I mean, how many trackways do we have of arboreal protosaurs or small, lithe dinosauriforms? Does that mean that they were clumsy, sluggish terrestrial animals, too? Though I can partially see Unwin's point on this one, I'm skeptical of using negative evidence in such a way, particularly with groups like pterosaurs with particularly patchy fossil records. Likewise, the suggestion that lonchodectids were Cretaceous pterosaur generalists seems a little bit of a stretch given that virtually no-one knows what they were really like and their record is very limited indeed.

On a similar note, the digital modelling of pterosaur terrestrial locomotion – the ‘Roborhamphus’ and ‘Robodactylus’ models crafted and worked on with Don Henderson (see Don's animation above, stolen from here) – is presented as accepted fact when, if I’m not mistaken, this work has only been published in a few abstracts (Unwin and Henderson 1999; Henderson and Unwin 1999) and not actually peer reviewed (please correct me if I’m wrong, though). Given that these models are somewhat at odds with other interpretations of pterosaur anatomy (e.g. laterally projecting hindlimbs in the Rhamphorhynchus model [contra. Padian 1983; Bennett 1997] and inflexible wrists in the pterodactyloid variant [contra. Bennett 2001; Wilkinson 2008; plus lots of pterosaur specimens with articulated, flexed wrists orientating depressing the wing metacarpal at least 40 degrees from the radius/ulna]), it may have been nice if a few more caveats about this work had been put in somewhere. Happily, this section of the book is unusual in not citing other, alternative interpretations and, in most instances, alternative hypotheses are mentioned and discussed.

Still, nothing’s perfect
These are relatively minor quibbles when the overall quality of the book is considered, though, and issues like those mentioned above are too rare top be major failings. In sum, then, even if you already own Wellnhofer’s Encyclopaedia, Deep Time is an essential purpose for the more up-to-date information it contains and, if you’re new to pterosaurs, it’s hard to imagine a better introduction. The fact that I've recommended this book to so many students and other pterosaurphiles is further testament to its quality and, frankly, it leaves a hard act to follow for the next guy in line writing a popular pterosaur book. Wait a second...


  • Bennet, S. C. 1997. The arboreal leaping theory of the origin of pterosaur flight. Historical Biology, 12, 265-290.
  • Bennett, S. C. 2001. The osteology and functional morphology of the Late Cretaceous pterosaur Pteranodon. Palaeontographica Abteilung A, 260, 1-153.
  • Henderson, D. and Unwin, D. M. 1999. Mathematical and computational model of a walking pterosaur. Journal of Vertebrate Paleontology, 19, 50A.
  • Padian, K. 1983. A functional analysis of flying and walking in pterosaurs. Palaeobiology, 9, 218-239.
  • Seeley, H. G. 1901. Dragons of the air. Meuthuen and Co., London, United Kingdom, 239 pp.
  • Unwin, D. M. 1999. Pterosaurs: back to the traditional model? Trends in Ecology and Evolution, 14, 263-268.
  • Unwin, D. M. 2003. On the phylogeny and evolutionary history of pterosaurs. In: Buffetaut, E. and Mazin, J. M. (eds.) Evolution and Palaeobiology of Pterosaurs, Geological Society Special Publication, 217, 139-190.
  • Unwin, D. M. 2005. The Pterosaurs from Deep Time. Pi Press, New York, 347 pp.
  • Unwin, D. M. and Deeming, D.C. 2008. Pterosaur eggshell structure and its implications for pterosaur reproductive biology. Zitteliana, B28, 199-207.
  • Unwin, D. M. and Henderson, D. 1999. Testing the terrestrial ability of pterosaurs with computer-based methods. Journal of Vertebrate Paleonotology, 19, 81A.
  • Wellnhofer, P. 1991. The Illustrated Encyclopedia of Pterosaurs. Salamander Books Ltd., London. 192 pp.
  • Wilkinson, M. T. 2008. Three dimensional geometry of a pterosaur wing skeleton, and its implications for aerial and terrestrial locomotion. Zoological Journal of Linnaean Society, 154, 27-69.


  1. I have and like the book, but as a paleo-artist and somebody who really wants to get to know the bones, it absolutely pales in comparison to Wellnhofer's Encyclopedia. I have a similar problem with the blocky, detail-bare line drawings in Deep Time. It's a great "here's where we're at" kind of book, but it doesn't really get into the specifics.

    We need a "Pterosauria, 2nd Edition."

    Or, you know, a 1st edition.

  2. Zachary,

    You're right on every point there. Wellnhofer's book is undeniably more comprehensively illustrated and a more useful reference for palaeoartists. I'm not sure that Unwin's book is less detailed, though: I think it's more detailed about some things, but skimps more over the details of others. Wellnhofer touches on just about everything pterosaurian circa 1991, but doesn't/couldn't go into as much detail on issues like pneumaticity, footprints, phylogeny and the like. Unwin does this, but at the expense of some details like individually describing and illustrating different genera. It's a tough act to juggle: too much information may loose less-technically minded readers, but too little will give the book no clout with the real enthusiasts.

    As for the Pterosauria volume, it is being worked on, but I expect it's still some time away. Bear in mind that the first Dinosauria volume took something like six years to get together: on that timescale, The Pterosauria is still running way ahead of schedule.

  3. "It's a tough act to juggle: too much information may loose less-technically minded readers, but too little will give the book no clout with the real enthusiasts."

    I liked the level of detail and depth of the Unwin book better than the breadth of the Wellnhofer book, but then I had the Wellnhofer book as well. I think it works out better to have different books doing different approaches, though I know not everyone is as fabulously wealthy as I am and thus not all could afford two books on pterosaurs. My own taste is for the depth, but in sufficiently laymanlike terms that I can actually follow it. In various subjects I'm familiar with, good in depth material is harder to find that broad survey info, particularly in this age of the internet.

    BTW, I expect, from all you've written before, that you will provide this but I want to say that stuff along the lines of 'Here's _how_ we know this is especially interesting.

    Love the ptink pterosaur in the ptrevious ptost, BTW.

    Mike from Ottawa

  4. Yes, the two books are I think complimentary. The technical stuff of Wellnhofer (what the bones are like, where things were found, how old they were etc.) has not dated at all and also covers things like the history of research in more detail than does Unwin. But 'Deep Time' really brings the phylogeny, evolution, and ecology up to date. Between the two it's an excellent mix.

    Good luck Mark, no pressure....

  5. I have Seeley 1901, Wellnhofer 1991 and Unwin 2006. Seeley's Dragons of the Air is a first edition. I am a very happy pterosaur fan!