Secondly, Campanian and Maastrichtian pterosaurs are, without exception, pretty big animals. Many species from this time are renowned for their gigantic size: it's these stages which give us the famous 10 m wingspan, 250 kg colossi like Quetzalcoatlus, Arambourgiania and Hatzegopteryx, as well as a number of other giant azhdarchids which are too poorly known for generic titles. Coinciding with the evolution of the giants is a loss of small pterosaur taxa - those animals less than 2.5 m across the wings which are present, more-or-less, throughout the rest of pterosaur history. This shift in body size is sometimes interpreted as pterosaurs demonstrating 'Cope's Rule', the somewhat controversial proposal that species evolve towards large body size over time (Hone and Benton 2007; Benson et al. 2014). It's argued by some that competition from birds may be the driver behind this trend, as early avians fought small flying reptiles for ecological space and ultimately forced pterosaurs into larger sizes (e.g. Benson et al. 2014). Note that this concept is not without its detractors, including myself - I won't go into my reasons now but I plan to outline them here eventually.
Whether you agree with the bird-pterosaur competitive displacement hypothesis or not, we can't disagree that the end of the Cretaceous is almost entirely devoid of small pterosaur remains. Only a handful of specimens record small pterosaurs in the Campanian and Maastrichtian, and they're all tricky to work with. Aside from being highly fragmentary, some are controversially identified (such as Piksi barbarulna, an alleged small pterosaur from the Two Medicine Formation - see Agnolin and Varricho 2012 for the pro-pterosaur case) and others represent probable juvenile individuals (Godfrey and Currie 2005). Whatever it signifies, the lack of diminutive pterosaur specimens from the close of the Mesozoic is a real phenomenon of our fossil record, and any new specimen of a small, latest Cretaceous flying reptile has to be something to get excited about.
Enter: a new small, latest Cretaceous pterosaur specimen to get excited about
|Title slide of my SVPCA 2016 talk, discussing the findings of Martin-Silverstone et al. 2016, out today. If you don't get the reference, you clearly get out too much, have too many friends and aren't watching enough crap TV.|
|RBCM.EH.2009.019.0001, a fragmentary azhdarchoid pterosaur from the Campanian Northumberland Formation, British Columbia. It's, er, not the prettiest pterosaur specimen you'll ever see. Combination of figures from Martin-Silverstone et al. 2016.|
We've identified the Hornby specimen as an azhdarchoid, and noted several features indicative of, but not conclusive to, an azhdarchid ID. We suspect the specimen is an azhdarchid because of its provenance and its basic anatomical characteristics, but the specimen does not contain the right bits to confirm an azhdarchid identity. Nonetheless, narrowing the specimen down to Azhdarchoidea allows us to estimate its body proportions and confirm that the specimen was indeed a small animal when it died. We estimated its wingspan using two methods factoring both the humerus and vertebrae, and each pointed to a wingspan between 1.4 and 1.6 m. That puts our pterosaur at a comparable size to a good sized-seagull and, while these are respectably-sized modern birds, this is small for a latest Cretaceous pterosaur. Rather than poking giraffes in the face, our little chap would only just be beyond predation risk from an average housecat (below). The only contemporary pterosaur competing with the Hornby azhdarchoid for size is Piksi, a poorly known possible pterosaur from the western US. Our new study lists a number of reasons why the pterosaurian characterisation of Piksi is problematic however: in short, its morphology is all wrong for a flying reptile and we suspect a non-pterosaurian ID is more likely. The Hornby specimen is thus a contender for the smallest latest Cretaceous pterosaur currently known.
|A 1.5 m wingspan azhdarchoid next to one (SI) MrTiddlesmetre. From Martin-Silverstone et al. (2016).|
|What's inside the RBCM.EH.2009.019.0001 humerus? A mix of things, but among them are features indicative of late-stage juvenility/subadulthood. Please see the paper for details of this figure. From Martin-Silverstone et al. 2016.|
A small pterosaur amongst the pigeons
There's obviously a limit to what a single fragmentary specimen can tell you about the evolution of a group, but what the Hornby specimen means for pterosaur evolution is interesting and - if we've interpreted it correctly - potentially significant. Most obviously, it suggests that small pterosaurs may have been present in the Campanian stage of the Late Cretaceous after all, at least in one part of the world. Regular readers will be aware that there's growing evidence for Late Cretaceous pterosaur faunas being less uniform than previously realised (e.g. Vremir et al. 2013, 2015), and our new specimen plugs into this picture nicely: it increasingly seems that the end Cretaceous wasn't just a stage for large-to-giant long-necked azhdarchids. What's more, while the specimen only provides one data point against the idea that birds ousted small pterosaurs, the presence of at least two types of bird in the Northumberland Formation seems to indicate small pterosaurs and birds coexisted in at least this palaeoenvironment. We might see this as a continuation of the coexistence pterosaurs and birds demonstrate in Jurassic and Early Cretaceous localities: maybe pterosaurs and birds got along OK after all.
|...except when pterosaurs stole their eggs. Our PR art for the new paper, where a group of Hornby azhdarchoids perform guerrilla raids on shore-living Campanian bird nests. Take THAT, birds.|
With all this said, the most important message of the paper has to be this: we need more data on small pterosaurs in the latest Cretaceous. The specimens we have are scrappy, hard to work with and offer limited scope for analysis. Thus, any small Late Cretaceous pterosaur material is significant, and whether they're lying unnoticed in museum collections or pulled straight out of the field, they are noteworthy specimens which need to be put on record. Curators and researchers, please keep your eyes peeled!
And that, in a nutshell, is our new paper: be sure to check it out if you want more details. You can also read Liz's take on the study over at The Conversation and other experts have been chiming in at news sites covering the story. With a bit of luck, this is not the only news you'll be hearing about Late Cretaceous pterosaurs from these quarters this year - more on these projects as they move along. All that's left to do is to thank Liz and Victoria for inviting me to collaborate with them on the new specimen - I learned a huge amount trying to get my head around this challenging material and its histology, and had a blast working with them.
This blogpost, paper and artwork are sponsored by PatreonRegular readers will know that this blog and its art are sponsored by a suite of awesome Patrons, but this post is proof that this support goes further than mere internet tomfoolery and contributes to papers and outreach, too. Supporting my blog from $1 a month not only helps keep this blog ticking over, but helps me contribute thoughts, words and illustrations to scientific research. In return you get access to bonus blog content: additional commentary, in-progress sneak-previews of paintings, high-resolution artwork, and even free prints. For this post, we'll be talking about the PR art I've done for this research: how was the azhdarchoid reconstructed from that
- Agnolin, F. L., & Varricchio, D. (2012). Systematic reinterpretation of Piksi barbarulna Varricchio, 2002 from the Two Medicine Formation (Upper Cretaceous) of Western USA (Montana) as a pterosaur rather than a bird. Geodiversitas, 34(4), 883-894.
- Bennett, S. C. (1993). The ontogeny of Pteranodon and other pterosaurs. Paleobiology, 19(01), 92-106.
- Benson, R. B., Frigot, R. A., Goswami, A., Andres, B., & Butler, R. J. (2014). Competition and constraint drove Cope's rule in the evolution of giant flying reptiles. Nature communications, 5, 3567.
- Butler, R. J., Benson, R. B., & Barrett, P. M. (2013). Pterosaur diversity: untangling the influence of sampling biases, Lagerstätten, and genuine biodiversity signals. Palaeogeography, Palaeoclimatology, Palaeoecology, 372, 78-87.
- Godfrey, S. J., & Currie, P. J. (2005). Pterosaurs. Dinosaur Provincial Park: A Spectacular Ancient Ecosystem Revealed, 292-311.
- Hone, D. W. E., & Benton, M. J. (2007). Cope's Rule in the Pterosauria, and differing perceptions of Cope's Rule at different taxonomic levels. Journal of Evolutionary Biology, 20(3), 1164-1170.
- Kellner, A. W. (2015). Comments on Triassic pterosaurs with discussion about ontogeny and description of new taxa. Anais da Academia Brasileira de Ciências, 87(2), 669-689.
- Martin-Silverstone, E., Witton, M. P., Arbour, V. M, & Currie, P. J. (2016). A small azhdarchoid pterosaur from the latest Cretaceous, the age of flying giants. Royal Society Open Access, 3, 160333.
- McGowen, M. R., Padian, K., De Sosa, M. A., & Harmon, R. J. (2002). Description of Montanazhdarcho minor, an azhdarchid pterosaur from the Two Medicine Formation (Campanian) of Montana. PaleoBios, 22(1), 1-9.
- Prondvai, E., Stein, K., Ősi, A., & Sander, M. P. (2012). Life history of Rhamphorhynchus inferred from bone histology and the diversity of pterosaurian growth strategies. PLoS One, 7(2), e31392.
- Vremir, M., Kellner, A. W., Naish, D., & Dyke, G. J. (2013). A new azhdarchid pterosaur from the Late Cretaceous of the Transylvanian Basin, Romania: implications for azhdarchid diversity and distribution. PLoS One, 8(1), e54268.
- Vremir, M., Witton, M., Naish, D., Dyke, G., Brusatte, S. L., Norell, M., & Totoianu, R. (2015). A Medium-Sized Robust-Necked Azhdarchid Pterosaur (Pterodactyloidea: Azhdarchidae) from the Maastrichtian of Pui (Haţ eg Basin, Transylvania, Romania). American Museum Novitates, (3827), 1-16.