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Historical Biology: An International Journal of Paleobiology Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/ghbi20 Pterosaurs, by Mark P. Witton. Princeton University Press, Princeton and Oxford, 2013, 291 pp., ISBN 978-0-691-15061-1 Darren Naisha a Ocean and Earth Science, National Oceanography Centre Southampton, University of Southampton, Southampton SO14 3ZH, UK Published online: 06 Mar 2014.

To cite this article: Darren Naish (2014): Pterosaurs, by Mark P. Witton. Princeton University Press, Princeton and Oxford, 2013, 291 pp., ISBN 978-0-691-15061-1, Historical Biology: An International Journal of Paleobiology, DOI: 10.1080/08912963.2014.882099 To link to this article: http://dx.doi.org/10.1080/08912963.2014.882099

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BOOK REVIEW

Pterosaurs, by Mark P. Witton, Princeton and Oxford, And there are other proto-pterosaurs out there in the Princeton University Press, 2013, 291 pp., ISBN 978-0- literature, just as there are proto-birds and proto-bats. 691-15061-1 Chapter 6 – ‘Flying reptiles’ – summarises current thinking on pterosaur aerodynamics and launch behaviour, Mark Witton’s Pterosaurs is an enormous, lavishly one of Witton’s most interesting contentions being that illustrated encyclopaedia of all things pterosaur. Scholarly pterosaurs have consistently been made too lightweight by but highly readable, fully referenced throughout, and previous authors. This has some impact on wing loading featuring hundreds of excellent photos, diagrams and and hence flight style. As is familiar to pterosaur workers, beautiful, colour life restorations, this volume is a must- Witton’s revised mass estimates and wing loadings make own, whatever your interest in pterosaurs. And there are not pterosaurs more comparable to the birds that they otherwise that many books devoted to pterosaurs to begin with, so resemble in planform (Witton 2008; Witton and Naish another one on the market can only be good. The volume is 2008) – a satisfying picture that has been bolstered by properly titled Pterosaurs: Natural History, Evolution, conclusions from some other workers (but, of course, not all Anatomy. However, this title does not appear anywhere on of them; this is pterosaur science we are talking about). the cover (or spine). More innovative is Chapter 7 (‘Down from the skies’), Pterosaurs is a large book, featuring 26 chapters devoted to terrestrial locomotion. Witton’s pterosaurs can spread over more than 280 pages. Early chapters review bound, swim, climb and most of them walk with narrow the history of our ideas about pterosaurs, the evolutionary gaits (though these generalisations do not apply across the relationships and origin of the group, and what we know of whole of Pterosauria, of course). The pterosaurs of the pterosaur anatomy (bony and soft). prior literature have been imagined as inelegant sprawlers, Many issues covered here remain controversial. The slow, clumsy and lame on the ground, unable even to resist consensus opinion supported by the majority of recent light gusts of wind (recall how the terrestrial locomotion phylogenetic studies is that pterosaurs are crown- depicted for the Walking With Dinosaurs ornithocheirids archosaurs, close to dinosaurs (note that the clade was based on shuffling people, walking on crutches) Ornithodira exists no matter where pterosaurs fall within (Wellnhofer 1988; Unwin 1988, 1997, 2005). Diapsida) (Sereno 1991; Benton 1999; Brusatte et al. In fact, data from trackways, pelvic and limb 2010; Nesbitt 2011). This is the idea that Witton favours, morphology, limb bone thickness and bending strength which is fair enough since there are unpublished studies (Fastnacht 2005), and inferred hindlimb and forelimb showing that the crown-archosaur hypothesis is supported posture (Fujiwara and Hutchinson 2012) indicate very even when oddball taxa considered close to pterosaurs by strongly that pterodactyloid pterosaurs at least were far more Downloaded by [Mr Darren Naish] at 07:15 10 March 2014 some (Peters 2000) are included in the analysis too. proficient on the ground than people have previously given Nevertheless, the crown-archosaur idea does look odd them credit for. Witton weaves all these lines of evidence given that we lack taxa that appear in any way transitional together in a compelling narrative and can be said to be between pterosaurs and other ornithodirans (Faxinalip- leading a revolution on the terrestrial abilities of pterosaurs. terus minima – published in 2010 as a possible sister- I agree with him: it now seems that pterodactyloids were taxon to the rest of Pterosauria – is not a pterosaur, nor proficient and agile on the ground, though non-pterodacty- obviously close to pterosaurs at all (Soares et al. 2013)). loids likely were not. Witton follows the honourable tradition of inventing Readers may also be surprised to see several Witton hypothetical proto-pterosaurs that bridge the gap between illustrations that show pterosaurs swimming and even Scleromochlus-like ornithodirans and known, early pter- diving. Some trackways seemingly show that some osaurs. I am not entirely sold on his use of the innovative pterosaurs foraged in shallow water, their bodies afloat term HyPtA (meaning Hypothetical Pterosaur Ancestor) and their long forelimbs being used as punting tools. Given for these creatures – the acronym is not exactly sexy – but the aquatic habits of many pterosaur species, it is hard to his invention of the creatures themselves is not all that bad: believe that they were incapable of at least some aquatic invoking an anatomy and lifestyle for an ancestral behaviour: despite claims made here and there about organism involves the formulation of a set of hypotheses pterosaurs being too pneumatic to swim or dive, or with a that will be supported or knocked down by future finds. wing form or body shape that would prevent swimming or 2 Book Review

diving, there are no obvious or compelling reasons In fact, details present across the pterosaur skeleton disallowing such behaviour, and Witton proposes that taxa show, Witton argues, that all those different pterosaur like Pteranodon might routinely have floated and dived. groups were doing different things – sometimes subtly Yes, there are frigatebirds (which can swim, if they different, sometimes strikingly different. As an example, let really have to, but generally avoid doing so), but hardly us briefly consider the long-snouted, long-winged ornitho- any pterosaurs are as specialised for a dedicated aerial cheiroid-type pterosaurs (istiodactylids, boreopterids, existence as they are (nyctosaurids are the probable ornithocheirids, nyctosaurids and pteranodontids), vir- exception). Hone and Henderson (2013) have since used tually all of which possess proportionally small, relatively digitally simulated floating models to test the floating weakly muscled hindlimbs. These pterosaurs all look more abilities of pterosaurs. or less alike, and, yes, the general view has been that they Chapter 8 – ‘The private lives of pterosaurs’ – reviews pretty much all did the same thing – they flew over the several areas that will be unfamiliar to the majority of non- surface of the sea and grabbed fish. specialists: what do we know about pterosaur prey items, But when all of their anatomical nuances, proportional pathologies, parasites, sex lives and growth strategies? The differences and so on are examined, a case can be made honest answer is ‘not much’, but we know enough to at that we are, in fact, seeing groups of species that, even least give us an insight into all of these areas. As is the case within this one group, perhaps did such diverse things as throughout the book, diagrams and life reconstructions scavenge from vertebrate carcasses in terrestrial environ- depict the key evidence or show re-enactments of things ments, grab fish from the sea surface, float or swim and that really must have happened. ‘cage’ mouthfuls of small, planktonic prey, float and make The meat and proverbial potatoes of the book are short surface dives, and live a frigatebird-like life that formed by reviews of all pterosaurian groups. Each group really does involve flying over the water surface and (each ‘family’, if you like) gets its own chapter. Once upon grabbing fish. a time, popular books would have given you the In short, Witton’s view of pterosaurs is exciting and impression that pterosaur diversity consisted of Dimor- refreshing in its diversity, complexity and plausibility. phodon, Rhamphorhynchus, Pterodactylus and Pterano- Think of modern animal groups, whether they be gulls, don and not much else. But things have boomed in recent seals or anoles: anatomical differences reflect different decades, such that the chapters here on wukongopterids, feeding and foraging strategies, nuances and niche-specific boreopterids, lonchodectids, chaoyangopterids, thalasso- behaviours are everywhere, and no two species are ever dromids and others will be wholly novel to some readers. really alike. This sort of thing was surely true of extinct On that note, it is well known that views on the animals – I am sure it was, we all are. But this is the first phylogeny, systematics and nomenclature of pterosaurs time that we have seen this many ideas on pterosaur vary quite considerably from one study to the next (though behaviour and lifestyle. note that members of particular research communities do I personally think that more ideas of this sort are tend to converge on similar results). Witton notes this and needed – so long as, that is, that they are internally logical, the arrangement he favours – in general, it is the incorporate data from all available lines of evidence and phylogenetic scheme recovered by Lu¨ et al. (2010) – is of appear grounded in a good understanding of biomecha- course not the one favoured by all pterosaur experts. But nics, ecology and the anatomy, behaviour, diversity and

Downloaded by [Mr Darren Naish] at 07:15 10 March 2014 when you are doing a book you have to come up with an functional morphology of living animals. As should be arrangement of some sort. obvious from the fact that I co-author with him (Witton Anyway, each of these chapters introduces us to the and Naish 2008, 2013), I think that Mark is very good at group in question, reviews and discusses what we know of this sort of thing (certainly better than those palaeontol- its anatomy, discusses the probable flight and grounded, ogists who take just one or two features and concoct an terrestrial behaviour of the group’s members and, finally, unrealistic and unlikely behavioural hypothesis) and I provides an overview of palaeoecology. The last chapter – generally find myself agreeing with his conclusions and the discussion of palaeoecology – is one of the most novel hypotheses. within the book. Tradition and convention would have it However, it is important to note that, at the moment, that pterosaurs flew over the surface of the sea and caught many of the ideas that Witton discusses and explores are, fish, or flew close to watery places and caught dragonflies indeed, just ideas: typically proposed here for the first time and other flying insects. Admittedly, this ‘samey’ view of ever, they now require investigation using rigorous, pterosaurs was challenged by the idea that dsungaripterids quantitative techniques. But you have to start somewhere. might have eaten shelled molluscs (Wellnhofer 1991) and Mark knows this and his hypotheses should serve as an that tapejarids were perhaps frugivorous (Wellnhofer and inspiration for future studies. Kellner 1991). But, otherwise, the overwhelming If there is anything that becomes a take home with impression has always been that pterosaurs caught fish regard to this book, it is that pterosaurs are more diverse in or insects while flying on the wing. anatomy and proportions that has been made obvious Book Review 3

before. Without naming names, the general impression References created by some pterosaur workers is that pterosaurs are Bennett SC. 2013. New information on body size and cranial display basically all alike, only with different skulls. As is made structures of Pterodactylus antiquus, with a revision of the genus. Pala¨ontol Z. 87:269–289. clear, not only through the text, but also thanks to those Benton MJ. 1999. Scleromochlus taylori and the origin of dinosaurs and standardised reconstructions, pterosaurs actually exhibit pterosaurs. Proc R Soc Lond B. 354:1423–1446. notable and significant differences in the length and form Brusatte SL, Benton MJ, Desojo JB, Langer MC. 2010. The higher-level phylogeny of Archosauria (Tetrapoda: Diapsida). J Syst Palaeontol. of the neck, in wing length, in hindlimb size and in, well, 8:3–47. everything, really. Costa FR, Rocha-Barbosa O, Kellner AWA. 2013. A biomechanical The pace of pterosaur research means that, needless to approach on the optimal stance of Anhanguera piscator (Pterodac- tyloidea) and its implications for pterosaur gait on land. Hist Biol, say, many new things have appeared since Pterosaurs doi:http://dx.doi.org/10.1080/08912963.2013.807253 appeared. There are quite a few new, recently named taxa Fastnacht M. 2005. The first dsungaripterid pterosaur from the that would have warranted discussion or mention including Kimmeridgian of Germany and the biomechanics of pterosaur long bones. Acta Palaeontol Pol. 50:273–288. the Tupuxuara-like azhdarchoid Caupedactylus, the small Fujiwara S-I, Hutchinson JR. 2012. Elbow joint adductor moment arm as azhdarchoid Vectidraco, the Patagonian azhdarchid Aero- an indicator of forelimb posture in extinct quadrupedal tetrapods. titan and a raft of new Cambridge Greensand taxa. Witton’s Proc R Soc Lond B. 279:2561–2570. Hone DWE, Henderson DM. 2013. The posture of floating pterosaurs: scepticism about the alleged azhdarchoid identity of ecological implications for inhabiting marine and freshwater Aurorazhdarcho (he has it wrongly spelt Auroazhdarcho habitats. Palaeogeogr Palaeoclimatol Palaeoecol, doi:10.1016/j. palaeo.2013.11.022. [sic]) has since been confirmed by Bennett’s (2013) paper Lu¨ J, Unwin DM, Jin X, Liu Y, Ji Q. 2010. Evidence for modular on Pterodactylus and Ardeadactylus, and then there is the evolution in a long-tailed pterosaur with a pterodactyloid skull. Proc whole awkward situation involving Lonchodraco and R Soc B. 277:383–389. Nesbitt SJ. 2011. The early evolution of archosaurs: relationships and the Lonchodraconidae. New studies on pterodactyloid limb origin of major clades. Bull Am Mus Nat Hist. 352:1–292. posture have appeared (Fujiwara and Hutchinson 2012; Peters D. 2000. A reexamination of four prolacertiforms with Costa et al. 2013), and so on. implications for pterosaur phylogenesis. Riv Ital Paleontol Stratigr. 106:293–336. Overall, the style of the text is friendly and Sereno PC. 1991. Basal archosaurs: phylogenetic relationships and conversational, even witty and vernacular. There are functional implications. Soc Vert Paleontol Mem. 2:1–53. definitely places where I feel that it needed to be a bit Soares MB, Dalla Vecchia FM, Schultz CL, Kellner AWA. 2013. On the supposed pterosaurian nature of Faxinalipterus minima Bonaparte tighter though, and I get the impression that better copy- et al. (2010) from the Upper Triassic of Rio Grande do Sul, Brazil. In: editing was needed. On other negative points, one or two Saya˜o JM, Costa FR, Bantim RAM, Kellner AWA, editors. of the illustrations look a bit rushed (I do not much like the International Symposium on Pterosaurs, Rio Ptero 2013, Short Communications. Universidad Federal do Rio de Janeiro; p. 95–97. theropod on p. 102) and I am not sure that the curly lines Unwin DM. 1988. New remains of the pterosaur Dimorphodon used in a few of the diagrams (Figures 7.8 and 9.2) – they (Pterosauria: Rhamphorhynchoidea) and the terrestrial ability of look like something out of a Tim Burton movie – fit with early pterosaurs. Modern Geol. 13:57–68. Unwin DM. 1997. Pterosaur tracks and the terrestrial ability of pterosaurs. the rest of the book. However, these are exceptions in an Lethaia. 29:373–386. otherwise glorious tour-de-force. Unwin DM. 2005. The pterosaurs from deep time. New York: Pi Press. As if it is not clear enough from what I have already Wellnhofer P. 1988. Terrestrial locomotion in pterosaurs. Hist Biol. 1:3–16. said, Witton’s Pterosaurs is a remarkable visual feast, Wellnhofer P. 1991. The illustrated encyclopedia of pterosaurs. London: packed full of novel art as well as excellent photographs that Salamander Books Ltd. Downloaded by [Mr Darren Naish] at 07:15 10 March 2014 the author clearly worked hard to obtain. There are, in fact, Wellnhofer P, Kellner AWA. 1991. The skull of Tapejara wellnhoferi Kellner (Reptilia, Pterosauria) from the Lower Cretaceous Santana illustrations of some sort on virtually every single page – Formation of the Araripe Basin, northeastern Brazil. Mitteilungen you will never get bored of looking at this book. A major Bayerischen Staatssammlung Pala¨ontol Hist Geol. 31:89–106. Wittonian innovation is the invention of a consistent pose Witton MP. 2008. A new approach to determining pterosaur body mass and its implications for pterosaur flight. Zitteliana. B28:143–158. for pterosaurian reconstructions (both skeletons and life Witton MP, Naish D. 2008. A reappraisal of azhdarchid pterosaur restorations). Reminiscent of Greg Paul’s lateral-view functional morphology and paleoecology. PLoS ONE. 3:e2271. Witton MP, Naish D. 2013. Azhdarchid pterosaurs: water-trawling dinosaurs, they make it easy to compare and contrast the pelican mimics or “terrestrial stalkers”? Acta Palaeontol Pol, doi: many taxa Witton illustrates. And look out for the nods and http://dx.doi.org/10.4202/app.00005.2013 witty homages: without giving the game away, there are references to Batman Begins, the arthropod-filled gorges of Darren Naish Skull Island, and what, exactly, is that marking on the Ocean and Earth Science, National Oceanography Centre lonchodectid’s wing? Southampton, University of Southampton, Southampton If you like or are even vaguely interested in pterosaurs, SO14 3ZH, UK you really need this book. I dearly hope that it inspires Email: [email protected] others to produce similar, spectacularly well-illustrated q 2014, Darren Naish volumes on other tetrapod groups.