A slightly overdue look at a new major technical volume on the amazing winged reptiles of the Mesozoic…
Caption: the cover of Hone et al. (2018) features this fine painting of the Early Jurassic pterosaur Dimorphodon macronyx. It’s a ‘new look’ for this animal relative to previously published efforts.
Pterosaur news is, these days, almost as hard to keep up with as is dinosaur news, a fact not aided by persistence of the tradition that any card-carrying pterosaur expert must seemingly regard themselves as an expert on all the pterosaurs, not just on the members of one or two specific lineages or clades. Come on, I can’t be the only one struggling to keep up with all those new papers on ctenochasmatoids or archaeopterodactyloids or euctenochasmatians or whatever the hell they are. Ah yes… exacerbating the pterosaur situation these days is the fact that we have competing nomenclatural schemes for the group…
Anyway, it’s time to review Hone et al.’s New Perspectives on Pterosaur Palaeobiology, a multi-authored 2018 Geological Society of London volume which mostly presents results of the 2015 Flugsaurier meeting held at Portsmouth, UK (my former academic base). My review here is substantially overdue but, as I’m sure I’ve said before, better late than never. And I’m in the slightly awkward position of reviewing a work where all three of the editors – Dave Hone, Mark Witton and Dave Martill – are sometime associates and co-authors of mine. Guys, please go lightly on the beating when next we meet.
Caption: a reminder of events of the Flugsaurier 2015 meeting, held at the University of Portsmouth. I wrote about the meeting here at TetZoo ver 3 (though all images have been stripped). The montage here features a slide from Witton’s talk on Dimorphodon (the results of which have since been published at PeerJ), the robust holotype skull of Parapsicephalus, and Dave Martill - one of the conference organisers - talking about Ludodactylus from Brazil.
New Perspectives on Pterosaur Palaeobiology features a brief introduction and 17 individual contributions, these starting off with works on inferred ecology and feeding behaviour, muscular anatomy, flight behaviour, growth and ontogeny before switching to descriptions of new taxa, reinterpretations of old ones, and the reporting of new specimens. There’s a lot here to obsess about if you’re really into pterosaurs but… my main take-home on the book is that – while it’s fine, and a perfectly respectable contribution to the pterosaur literature – it doesn’t really live up to its title. Those 17 individual papers don’t really provide ‘new perspectives on pterosaur palaeobiology’: they are, instead, very much ordinary small contributions of the sort that would go into any number of assorted provincial publications, and there certainly isn’t anything here which might lead you to think about any particular aspect of pterosaurs or their biology in a new way. This doesn’t mean that the book is bad, though; there’s lots here to like.
Caption: Mark Witton reconstruction of Rhamphorhynchus swimming and fish eating, aspects of behaviour informed directly by fossils. This image appears in Witton’s 2018 paper in the volume but can also be seen in a blog article here. Image: Mark Witton.
Mark Witton’s paper on pterosaur diet, as gleaned from preserved stomach contents, feeding traces and so on is a fairly solid review. It’s inevitable that much of the content reproduces that included in Witton’s book on pterosaurs (Witton 2013); whatever, it’s good to get this stuff into the technical literature. Don Henderson’s paper on pterosaur bite strength is good in that the results mostly agree with other lines of evidence on pterosaur biology, but an inevitable criticism (which Don does take time to counter) is that some of the 3D models he uses for his calculations are extrapolations made from flattened specimens and the illustrations of pterosaur expert Peter Wellnhofer.
Caption: excellent fossils like this one from the Solnhofen Limestone show that pterosaurs were sometimes (or often?) killed by other animals. This shows how the ray-finned fish Aspidorhynchus sometimes snagged or captured the gull-sized Rhamphorhynchus. Image: Frey & Tischlinger (2012).
Rachel Frigot’s examination of the 3D pelvis of Vectidraco – a raven-sized, Tapejara-like azhdarchoid from the Isle of Wight Greensand which I and colleagues named in 2013 (Naish et al. 2013) – appears sound and I’m pleased that this specimen is providing insight into detailed aspects of pterosaur biology (see also Martin-Silverstone et al. 2017). Colin Palmer’s analysis of wing membrane properties is also welcome; he finds that pterosaur membranes must have been quite different from those of bats in their distal regions at least, and that those parts would have needed reinforcement, a view consistent with suggestions that aktinofibrils were keratinous.
The peculiar striping present on the rostral bones of some Pteranodon specimens is described by Chris Bennett and Paul Penkalski. It seemingly represents ‘waves’ of bone resorption and deposition linked to a reshaping of the animal’s snout as it grew; Bennett and Penkalski further argue that this striping is characteristic of immature males, and immature males alone. What this seems to mean is that males underwent a profound increase in the depth and length of the front part of the upper jaw and snout as they matured, this presumably meaning that the enlargement of this region was sexually dimorphic, and potentially driven by sexual selection pressure. This idea might have been implied on previous occasions, but I’m not sure that it’s been specifically stated. To my mind, it strengthens the idea that pteranodontid specimens with massively enlarged premaxillary rostra – the posterchild here is UALVP 24238, the holotype of the ostensible taxon Dawndraco kanzai Kellner, 2010 – should be interpreted as well-developed specimens of known taxa (Martin-Silverstone et al. 2017), not as new taxa.
Caption: Dawndraco kanzai - most likely a junior synonym of Pteranodon sternbergi (and an ‘extreme male’ example of that species) - as reconstructed by Dean Schnabel (twitter account here). Used with permission.
Another very interesting paper here (to me) is Junchang Lü et al.’s on the new Chinese anurognathid Vesperopterylus lamadongensis from the Jiufotang Formation, based on an essentially complete and mostly articulated skeleton. Its key novelty is in its foot, since the ungual of its first toe is ‘reversed’ relative to that of the other toes, opposing them in the direction of its curvature. If this is genuine (it looks like it is, based on their photos), it’s hard not to interpret it as a climbing or grasping feature, and this is exactly what the authors do. Their accompanying life reconstruction (produced by Zhao Chuang of PNSO* fame) is based on the idea that anurognathids looked like pseudo-bat nightmare monsters, not like the cryptically patterned flying muppets or pterosaurian frogmouths of some other artists.
* A toy company which produces a substantial number of model dinosaurs, pterosaurs and other prehistoric animals.
Caption: Vesperopterylus lamadongensis in life, as reconstructed by Zhao Chuang.
Steve Vidovic and Dave Martill fire another volley in their campaign to give Pterodactylus, the original and archetypal pterodactyl, a good and thorough seeing to. Their contention is that Pterodactylus of tradition is over-lumped and they use morphometrics and cladistics to tease out the various taxa, all of which have been recognised as valid on previous occasion but then lumped back into Pterodactylus or somehow confused with it. They argue here that Seeley’s Diopecephalus (Pterodactylus kochi of many previous authors) warrants distinction, and that ‘Germanodactylus’ rhamphastinus needs a new name, so hello Altmuehlopterus (no love for you, Daitingopterus Maisch et al., 2004, since they argue that it’s a nomen nudum and should be ignored).
Caption: the holotype of Altmuehlopterus, BSP AS I 745 a, at the Jura Museum, Eichstätt. Image: Ghedoghedo, CC BY-SA 3.0 (original here).
Some of Vidovic and Martill’s arguments are a bit difficult to understand and I found myself leaving their chapter unconvinced that they’re right on all of their contentions. Buried in the middle of this paper is an argument that anurognathids are paedomorphic scaphognathines. This is one of those things that’s perhaps possible (I say this with the extreme ontogeny of certain ray-finned fishes in mind), but I can’t say that I find it likely based on anatomical data. There’s some substantial amount of redundant wording in this contribution plus technically incorrect use of directional terms (you shouldn’t use ‘distal’ and ‘proximal’ on pelvic bones). And sorry, again, for being that guy, but ’monophyletic clade’ is also a big fat no (it’s a tautology: a clade – by definition – can only be monophyletic).
With those papers out of the way, we have additional contributions on a pterosaur embryo from Argentina (Codorniú et al.), pterosaur bits and pieces from the Morrison Formation (McLain and Bakker), the Asian dsungaripterid Noripterus (Hone et al.), probable elements of the Jordanian azhdarchid Arambourgiania (Martill and Moser), Oxford Clay pterosaurs of the UK (O’Sullivan), a Jurassic rhamphorhynchid humerus from Thailand (Unwin and Martill), Turonian wing bones from Utah (Bennett), possible chaoyangopterid vertebrae from the Brazilian Cretaceous (Leal et al.), an Italian wing metacarpal from the Late Cretaceous (Dalla Vecchia), and lonchodectid remains from the English Wealden (Rigal et al.).
Caption: Noripterus complicidens, as reconstructed by Hone et al. (2018a) on the basis of the material described by C. C. Young. Diagram by Rebecca Gelertner.
Like all GSL volumes, this work is beautifully produced, with excellent print quality, robust binding and great reproduction of its figures and photos. But I regret to say that the editing is sloppy; there are typos all over the place, including even in abstracts (case study: a shocking four typos on p. 42 alone, tsk tsk). There’s also a weird bit of inconsistency: the volume’s introduction says (Hone et al. 2018b, p. 3) that Rigal et al.’s paper includes the naming of a new lonchodectid genus (potentially a big deal given the obscure nature of this group)… but it totally doesn’t. I’m guessing that the submitted version included said new name but that its recognition didn’t survive the grim reaper that is peer review.
These grand, ex-symposium GSL works (which the GSL has been publishing for 56 years now) are all the same shape and format, so they look great on a shelf if you own a few of them. As always, my primary concern is that their price makes them unaffordable to a great many of the researchers (amateur and professional) who would most benefit from owning them. The good news is that they’re sometimes offered, for a limited time, at a substantially reduced price.
Caption: the GSL volumes I have here in the TetZoo library. I was a co-editor for the ‘Dinosaurs and Other Extinct Saurians’ one. Image: Darren Naish.
Our understanding of pterosaurs has changed enough in recent decades that there really is a (short!) list of must-have volumes which chart the exploding knowledge of the group, among them Peter Wellnhofer’s 1991 Illustrated Encyclopedia of Pterosaurs, Eric Buffetaut and Jean-Michel Mazin’s 2003 conference volume Evolution and Palaeobiology of Pterosaurs, and Mark Witton’s 2013 Pterosaurs: Natural History, Evolution, Anatomy. Is New Perspectives on Pterosaur Palaeobiology a must-have volume of this sort? Well, no, not really. But if you’re a pterosaur researcher – or are simply really, really interested in this remarkable and charismatic Mesozoic group – there’s certainly enough here to make it worth owning, and you’ll likely check various of the contributions at least occasionally. I’ll repeat again – in an effort to avoid being surprised in a dark alley by any combination of the volume’s editors and club-shaped objects – that there’s a lot here which I did like, and will be citing.
Hone, D. W. E., Witton, M. P. & Martill, D. M. (eds) 2018. New Perspectives on Pterosaur Palaeobiology. Geological Society, London, Special Publications 455, pp. 238. Hardback. ISBN 978-1-78620-317-5. £67.50 from Amazon, £90 from publishers. Here from the publishers. Here from amazon.co.uk.
For previous TetZoo articles on pterosaurs, see…
Daisy’s Isle of Wight Dragon and why China has what Europe does not, March 2013
In Rio for the 2013 International Symposium on Pterosaurs, May 2013
Quetzalcoatlus: the evil, pin-headed, toothy nightmare monster that wants to eat your soul, August 2013
Were azhdarchid pterosaurs really terrestrial stalkers? The evidence says yes, yes they (probably) were, December 2013
Some Azhdarchid Pterosaurs Were Robust-Necked Top-Tier Predators, February 2017
Postcranial Palaeoneurology and the Lifestyles of Pterosaurs, August 2018
Refs - -
Hone, D. W. E., Jiang, S. & Xu, X. 2018a. A taxonomic revision of Noripterus complicidens and Asian members of the Dsungaripteridae. In Hone, D. W. E., Witton, M. P. & Martill, D. M. (eds) New Perspectives on Pterosaur Palaeobiology. Geological Society, London, Special Publcations, pp. 149-157.
Hone, D. W. E., Witton, M.P. & Martill, D. M. 2018b. New perspectives on pterosaur palaeobiology. In Hone, D. W. E., Witton, M. P. & Martill, D. M. (eds) New Perspectives on Pterosaur Palaeobiology. Geological Society, London, Special Publcations, pp. 1-6.
