Showing posts with label Thalassodromeus. Show all posts
Showing posts with label Thalassodromeus. Show all posts

Wednesday, 13 August 2014

Lies, damned lies, and 'Thalassodromeus sebesensis'

Yesterday, a huge team of authors called out the science behind 'Thalassodromeus sebesensis', an alleged new pterosaur species 40 million years and thousands of miles out of time and space (Grellet-Tinner and Codrea 2014). As with many outlandish palaeontological claims, the evidence behind 'T. sebesensis' really falls apart rapidly under scrutiny, principally because the alleged pterosaur remains actually represent an unremarkable piece of turtle plastron (Dyke et al. 2014).

A, the plastron of the fossil Romanian turtle Kallokibotion magnificum, compared with B, the alleged holotype 'cranial crest' of 'Thalassodromeus sebesensis'. For further details, see yesterday's post.
Since then, the response to our comment has been published (Codrea and Grellet-Tinner 2014). I'll admit to being surprised that Codrea and Grellet-Tinner maintain the specimen as a pterosaur, and consider the arguments raised against our points as weak, hypocritical and problematic, but whatever: the two arguments are now out, and the palaeontological community can judge for themselves. CT scanning is apparently planned for the specimen (Codrea and Grellet-Tinner 2014), which should put 'T. sebesensis' to bed once and for all.

This post isn't really about that, though: it's about correcting a mistruth in Codrea and Grellet-Tinner's response. Their comment shows little decorum or professionalism, attempting to undermine our response with ad hominem potshots at some authors of Dyke et al. (2014), including criticism of their editorial skills and the taxonomic confusion surrounding specimens described by the authors. Moreover, they criticise us for not examining the specimen, UBB ODA-28, before publishing our response. They state that:
"...UBB ODA-28 is housed in an official and recognized Romanian institution, thus available for examinations to anyone interested. This includes Dyke’s July 2nd 2014 written request to examine UBB ODA-28, which was immediately granted, although, Dyke went on writing its hasty comment without examining UBB ODA-28."
Codrea and Grellet-Tinner, 2014, p. 3-4 (my emphasis)

Well, this isn't really true. Some of it is: Gareth Dyke did write to ask for permission to look at the specimen this year - specifically between July and September - but 'immediate' access was not granted. Rather, eventual access was promised following on-going studies, including CT scanning of the specimen, the dates of which was not disclosed. This is not, as Codrea and Grellet-Tinner describe, 'immediately' granting access, but nebulously promising access at an undetermined future date. 

This may not seem like a big deal, but our integrity is being questioned for having not seen the specimen, so we - the authors of Dyke et al. (2014) - think the record should be set straight. There's no doubt that examining specimens is the way forward in any research. But it was clear from Gareth's correspondence that accessing UBB ODA-28 was going to be difficult for the immediate future, and all the while the science behind 'T. sebesensis' remained extremely problematic and in need of swift rebuttal. Why? In short: none of us concerned with pterosaurs or European palaeontology want to deal with this outrageous, nonsensical claim in future publications. Hence, we fell back on using the published accounts of UBB ODA-28 to construct an argument against the pterosaur identification. Given that our authorship team has collectively amassed thousands of hours examining actual thalassodromid pterosaurs, as well as turtle plastrons, and how obvious the turtle affinities of the specimen are, this method seemed more than sufficient for the task at hand. Despite allegations from Codrea and Grellet-Tinner, these were not the actions of a team hastily assembling a rebuttal, but a collective of experienced individuals succinctly calling out obvious flaws in bad science.

So there we go: that's our side of that mistruth. Hopefully, that's the last we'll hear of 'T. sebesensis' around these parts, for there are much more interesting and exciting things to cover: palaeoart guides, Triassic fuzzy saltating xerocoles, dinosaur fat humps... all coming soon.

References

  • Codrea, V. A., & Grellet-Tinner, G. (2014). Reply to Comment by Dyke et al. on "Thalassodromeus sebesensis, an out of place and out of time Gondwanan tapejarid pterosaur" by Grellet-Tinner and Codrea (July 2014)"  Gondwana Research. IN PRESS
  • Dyke, G. J., Vremir, M., Brusatte, S., Bever, G., Buffetaut, E., Chapman, S., Csiki-Sava, Z, Kellner, A. W. A., Martin, E, Naish, D, Norell, M, Ősi, A, Pinheiro, F. L., Prondvai, E, Rabi, M, Rodrigues, T., Steel, L., Tong, H, Vila Nova B. C. & Witton, M. (2014). Thalassodromeus sebesensis-a new name for an old turtle. Comment on" Thalassodromeus sebesensis, an out of place and out of time Gondwanan tapejarid pterosaur", Grellet-Tinner and Codrea. Gondwana Research. IN PRESS.
  • Grellet-Tinner, G., & Codrea, V. A. (2014). Thalassodromeus sebesensis, an out of place and out of time Gondwanan tapejarid pterosaur. Gondwana Research. IN PRESS

Tuesday, 12 August 2014

'Thalassodromeus sebesensis': pterosaur out of time and space? Nope, just a misidentified chunk of turtle.

Thalassodromeus sethi after some worms. Note: not a turtle. From Witton (2013)
Today sees the publication of an article challenging an exciting claim made in recent pterosaurology (Grellet-Tinner and Codrea 2014). If you missed it, the article concerned identifies a thalassodromid pterosaur in uppermost Cretaceous rocks of Romania and the erects a new species, Thalassodromeus sebesensis Grellet-Tinner and Codrea, 2014. At the centre of this is ODA-28, an (alleged) fragmentary cranial crest only fully exposed on one surface. None of this may not sound like a big deal, except that other thalassodromids - including the alleged sister species, Thalassodromeus sethi - are only known from the Lower Cretaceous Araripe Group of Brazil. T. sebesensis thus is about 40 million years out of time and thousands of miles out of place, and also occurring when azhdarchid pterosaurs basically represent the entire diversity of Pterosauria (Grellet-Tinner and Codrea 2014). Suddenly, the routine act of naming of a new animal is rewriting our understanding of pterosaur evolution.

There's more. Despite having only a scraps of bone to work with, Grellet-Tinner and Codrea (2014) suggested the T. sebesensis crest anchored muscles to form a ‘sizeable fleshy crest’, acted as a rudder in flight, that it somehow highlighted co-evolution between Romanian pterosaurs and angiosperms, and ecological segregation between azhdarchids and thalassodromids. All of these ideas are pretty radical in one way or another, especially considering the fossil material they are based on.

Blah blah blah… extraordinary claims, extraordinary evidence etc. When T. sebesensis was published it raised the collective eyebrows of pterosaur workers for all the wrong reasons. ODA-28 has no obvious ties to Thalassodromidae (or Thalassodrominae, if that’s how you roll - see Witton 2009), Pterosauria, or even to a cranial crest. Today, I and 19(!) other authors have said this in print (Dyke et al. 2014), noting that ODA-28 lacks any pterosaurian synapomorphies or even features typical of the group. As anyone who has handled pterosaur fossils can attest, pterosaur remains are distinctive at gross and microscopic level, and ODA-28 lacks any features expected in pterosaur bone (e.g. extremely thin bone walls separated by trabeculae). Any resemblance to the Thalassodromeus sethi holotype is entirely superficial, and shared characters between the two specimens - notably the ‘fossae’ at the base of the ‘T. sebesensis’ crest - are really incomparable on detailed examination. A clear lack of symmetry in ODA-28 shows it is not a medial skeletal element either, and thus not the cranial crest of anything. In short, cancel the text-book revisions: the temporal and palaeobiogeographical anomaly of ‘Thalassodromeus sebesensis’ is just a fairly major misidentification of a scrappy fossil (Dyke et al. 2014).

The 'flying turtle': the holotype of 'T. sebesensis' compared with the plastron of the turtle Kallokibotion. A, NHMUK R4930, the lectotype plastron of Kallokibotion magnificum with the portion corresponding to ODA-28 outlined in black (photo supplied by S. Chapman, Natural History Museum, London); B) ODA-28 (modified from Grellet-Tinner and Codrea, 2014). Abbreviations: hypo, hypoplastron; hxc, hypoplastron-xiphiplastron suture; ihc, intra-hypoplastral suture; ib, inguinal buttress; ps, pubic scar; meso, mesoplastron; mhc, meso-hypoplastral contact; pll, posterolateral lip; xiphi, xiphiplastron. Scale bar for A equals 50 mm. From Dyke et al. (2014).

Is ODA-28 anything exciting at all? Well, not especially. The specimen is clearly a piece of turtle plastron, exactly matching the internal structure of the hypoplastron and xiphiplastron of the Maastrichtian, Romanian genus Kallokibotion (above, Dyke et al. 2014). The anatomy of Kallokibotion has been documented fairly thoroughly and known for about 100 years (e.g. Gaffney and Maylan 1992), allowing us to be confident in this identification. Ergo, 'T. sebesensis’ offers nothing other than new a piece of fossil turtle and a name for the Kallokibotion synonymy list.

In all, a bit of an anticlimax. How did our short paper end up with 20 authors? The response was started by experts in the terrestrial faunas of upper Cretaceous Romania, who asked me and the Natural History Museum’s Lorna Steel if we could contribute a few paragraphs targeting the flawed pterosaur identity of the specimen. While we were working, it emerged that pterosaur experts from Brazil were also planning a response. The editors of Gondwanan Research, who published Grellet-Tinner and Codrea (2014), understandably only wanted one response, so the two teams joined forces. By the time experts in turtles, Romanian fossils and pterosaurs were all on board, we ended up with a truly international background: the USA, UK, Brazil, Romania and France are all represented.

A final note: this is not the first time thalassodromids have been pulled to the top of the Cretaceous. Kellner (2004) and Martill and Naish (2006) argued that a partial skull and mandible from the Maastrichtian Javelina Formation of Texas represented a thalassodromid based on perceived similarities with the thalassodromid Tupuxuara. While others have argued against this idea (the mandible and premaxillary morphology are more similar to those of azhdarchids - Lü et al. 2008; Witton 2013) - these claims have not been met with a sledgehammer response because the suggestions are not unreasonable. Sure, I don’t think the Javelina material in question is thalassodromid, but I can see why others might. ‘T. sebesensis’ has been swiftly rebutted by a crowd of experts because the underlying science is so clearly bogus that all concerned with pterosaur and Romanian palaeontology wanted it’s impact nipped in the bud. A response from Grellet-Tinner and Codrea will be published soon, so we'll see what they make of our rebuttal. To end on a high: there are exciting pterosaur remains coming out of Romania, and some of them are in the review/publication system already. Hopefully, we'll have some news on these out soon.

References

  • Dyke, G. J., Vremir, M., Brusatte, S., Bever, G., Buffetaut, E., Chapman, S., Csiki-Sava, Z, Kellner, A. W. A., Martin, E, Naish, D, Norell, M, Ősi, A, Pinheiro, F. L., Prondvai, E, Rabi, M, Rodrigues, T., Steel, L., Tong, H, Vila Nova B. C. & Witton, M. (2014). Thalassodromeus sebesensis-a new name for an old turtle. Comment on" Thalassodromeus sebesensis, an out of place and out of time Gondwanan tapejarid pterosaur", Grellet-Tinner and Codrea. Gondwana Research. IN PRESS.
  • Gaffney, E. S., & Meylan, P. A. (1992). The Transylvanian turtle, Kallokibotion, a primitive cryptodire of Cretaceous Age. American Museum novitates; no. 3040.
  • Grellet-Tinner, G., & Codrea, V. A. (2014). Thalassodromeus sebesensis, an out of place and out of time Gondwanan tapejarid pterosaur. Gondwana Research.
  • Kellner, A. W. A. (2004). New information on the Tapejaridae (Pterosauria, Pterodactyloidea) and discussion of the relationships of this clade. Ameghiniana, 41, 521-534.
  • Lü, J., Unwin, D. M., Xu, L., & Zhang, X. (2008). A new azhdarchoid pterosaur from the Lower Cretaceous of China and its implications for pterosaur phylogeny and evolution. Naturwissenschaften, 95(9), 891-897.
  • Martill, D. M., & Naish, D. (2006). Cranial crest development in the azhdarchoid pterosaur Tupuxuara, with a review of the genus and tapejarid monophyly. Palaeontology, 49(4), 925-941.
  • Witton, M. P. (2009). A new species of Tupuxuara (Thalassodromidae, Azhdarchoidea) from the Lower Cretaceous Santana Formation of Brazil, with a note on the nomenclature of Thalassodromidae. Cretaceous Research, 30(5), 1293-1300.
  • Witton, M. P. (2013). Pterosaurs: natural history, evolution, anatomy. Princeton University Press.

Tuesday, 2 July 2013

Praise for Pterosaurs: Natural History, Evolution, Anatomy

Before there were pterosaur fossils, there were rotting pterosaur corpses. This here is the rotting skull of Dsungaripterus weii, one of the largest dsungaripteroid pterosaurs known. His eye socket seems particularly interesting for some reason. Full, uncropped version of this painting, from Witton (2013).
Time's been a bit short for the last week, which means I'm falling behind my ideal blogging frequency. What better way to catch up, then, than to have some others write a post for me? Pterosaurs: Natural History, Evolution, Anatomy has been released in the wilds for several weeks now, which means that reviews are starting to trickle in. I'm happy to say that all legitimate reviews of the book have come up very well indeed, and there's praise all around. But you don't have to take my word for it: summary statements and links to reviews are provided below, along with some additional artwork from the book, just for fun. In case you missed it, an overview of Pterosaurs content is provided here and via the shiny new Pterosaurs widget on the top right of the page. The nice folks at Princeton have also recently released samples of 12 double-page spreads of the book, so be sure to check those out if you've not seen a copy yet. Kindle and dead-tree versions of the book can be ordered from Amazon. On to the review summaries!
Witton’s combination of style and substance makes Pterosaurs a true treasure and an absolute must for anyone curious about the extinct flyers... 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.
Brian Switek, Laelaps (National Geographic Phenomena)
This really is the ultimate guide to pterosaurs, providing us with a richer view of pterosaur diversity and behaviour than allowed in the two previous great volumes on the group (Wellnhofer 1991, Unwin 2005) and containing a substantial amount of review and analysis of pterosaur ecology and functional morphology.
Darren Naish, Tetrapod Zoology (Scientific American blog network)
Whatever the intent of the author, the book does succeed at a number of levels. While probably a tricky read for those very unfamiliar with fossils, it should be easily accessible for anyone with a passing interest in palaeo as well as providing a solid review of the whole of the Pterosauria that’ll be genuinely useful for researchers for many years. I’m sure I’ll be typing “Witton, (2013) stated….” quite a lot in the future and that, if anything, should be a good measure of how I rate this as a scientific text. Now go buy a copy and read it, it really is very good. 
Dave Hone, Pterosaur.Net
This book is both academically interesting and truly fun to read. That is a difficult balance to reach, but Witton does an excellent job of it by using a lighthearted, informal writing style in combination with a well-referenced, serious scientific review. An invaluable reference.
Michael Habib, endorsement at Princeton University Press 
(Mike's summary response as referee to the book text)   

Thalassodromeus manufacturing pterosaur feeding traces, which are known from a number of pterosaur tracksites. Was Thalassodromeus always this placid? Maybe not, according to imagery at this post. From Witton (2013). 


 
This book does a good job of summarizing several of the diverse arguments that fly — pardon me — around ...Witton walks a fine line between presenting a technical review and providing an introductory text for students unfamiliar with the group in question or students unfamiliar with scientific discourse at all. ... Recommended? Yes, with kudos.
Jaime Headden, The Bite Stuff 
Mark... presents the uncertainties of science but never shies away from making his opinion clear. [He] respects the complexities [of scientific writing] without allowing them to clump up the text. ... Mark isn’t the craftiest of illustrators, but I wouldn’t change a single of his drawings for any pile of slickness. Mark understands illustration, and he illustrates. ...  I can wholeheartedly recommend the book already."
David Mass, DRIP 
PTEROSAURS would make an excellent addition to any reference collection and especially that of an advanced (adult or young adult) lay-reader.
Greg Leitich Smith, GSL BLog 
I can tell you that it is not only a fascinating bit of text, its illustrations will leave you gaping in awestruck amazement.
John E. Riutta, The Well-read Naturalist   
I thoroughly enjoyed this book. Witton's style is rather informal, but his coverage of the subject is academically rigorous, and his excellent illustrations convey both his knowledge of and passion for his subject. It tells a great story of the history of extraordinary animals, and should appeal to anyone interested in science, let alone palaeontology. It is worth its cover prices for the illustrations alone, many of which are the work of the author and presented with a quirky sense of humour.
Richard Forest,  Amazon.co.uk book review

Just how do you make a pterodactyloid pterosaur? Follow these instructions, bake for an undisclosed number of millions
of  years, and viola! Preondactylus, Darwinopterus and Pterodactylus skeletals from Witton (2013).

Thanks very much to those who've taken time to review Pterosaurs or sing its praises online in blog comments and social media. An additional big thanks to those who've personally written to me to express their satisfaction with the book. Hammy as it is to say, it's really great to hear that so many people like it.

Next time (hopefully very soon): mummified pterosaurs (promise number 3 for that one) and the most tenacious of all proposed pterosaur habits.

Reference
    • Witton, M. P. 2013. Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.

    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.

    References
    • 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.