Short-necked azhdarchid pterosaurs - say what?

LPV (FGGUB) R.2395, our unnamed short-necked azhdarchid from Maastrichtian deposits of the Hațeg basin. Prints of this chap are available.

Odds are that most regular readers of this blog are familiar with azhdarchid pterosaurs, the toothless, often gigantic flying reptiles which increasingly dominated pterosaur evolution in the Cretaceous. With a better fossil record than most pterosaur lineages, they are among the best understood and most researched of all flying reptiles. In recent years our understanding of their anatomy, distribution and palaeobiology has advanced considerably.

Since azhdarchids were recognised as a group in the 1980s it has been realised that their most striking and characteristic features pertain to their neck anatomy. In both relative and absolute terms they have the longest necks of any pterosaurs, stretching their mid-series cervical vertebrae to long tubes with reduced features. Cervicals IV - VI are especially long and tubular, with the mid-portions of the neural spines reduced to such a degree that they are effectively split into anterior and posterior sections. The only significantly developed features of these cervicals are bulbous zygopophyses and condyles, which nestle together so snugly that neck articulation seems limited. Regular readers will know that this has considerable bearing on the likely habits of these animals, probably precluding strenuous lifestyles such as skim-feeding or pelican-like scooping. The extremes of their neck bones - cervicals I - III, and the somewhat 'dorsalised' VIII - IX - are less modified, although some azhdarchid weirdness infects these too in terms of length, neural spine shape, or both.

With azhdarchid cervicals being so diagnostic, they can be identified when found in isolation and even when only poorly preserved. Indeed, many azhdarchid occurrences are represented by isolated cervicals - they compete with jaw tips for the most common type of azhdarchid fossil. In recent years, the complete neck osteology of the Santonian, central Asian azhdarchid Azhdarcho lancicollis has been documented in some detail (Averianov 2010), permitting some insight into which specific part of the neck sequence isolated neck bones represent. It can be a little tough to tell a cervical IV and V apart - they seem very similar, except that V is invariably the longer of the two (indeed, the longest of the entire neck) - but we can now at least tentatively identify bones from the rest of the sequence. This is a major breakthrough, meaning that isolated cervicals can tell us a lot more than just where an azhdarchid was preserved: their potential for taxonomic and biomechanical studies has been increased considerably.

Eyes to Romania

A lot of new azhdarchid material, including said isolated cervicals, has recently been emerging from various Maastrichtian deposits of Romania. These sediments represent the rivers and lakes which once ran through Hațeg island, an ancient setting famous for its dwarf dinosaurs and the enormous azhdarchid Hatzegopteryx thambema. The Hațeg pterosaurs, and their neighbours from Transylvania, have been the focus of a number of recent pterosaur papers and, this week, a team of researchers (including Mátyás Vremir, myself, Darren Naish, Gareth Dyke, Stephen Brusatte, Mark Norell, and Radu Totoianu) published another in American Museum Novitates (Vremir et al. 2015). It reports the discovery of just one bone* - the near-complete azhdarchid cervical LPV (FGGUB) R.2395, from Hațeg Basin red beds - but it's enough to cast new light on these otherwise familiar azhdarchid fossils, as well as the general evolution of azhdarchids themselves.

*So yes, the picture at the top is 1% fossil data, 99% palaeoart polyfiller.

Line drawing of LPV (FGGUB) R.2395, isolated azhdarchid cervical IV or V. It don't look like much, but it's got it where it counts. From Vremir et al. 2015.

R.2395 (above) is not a large vertebra, being an estimated 100 mm long and 44 mm across the prezygapophyses when complete (it's missing the posterior portion, including most of the condyle and postzygagpophyses, as well as the left prezygapophysis). Our paper provides a long discussion about the likely portion of the neck represented by R.2395, concluding that it is probably a cervical IV or V. In azhdarchids, these vertebrae are among the longest of all, sometimes being eight times longer than wide. R.2395 was clearly a lot stouter than this however, barely being twice longer than broad. This proportion is unique among azhdarchid CIV and Vs, and there was some discussion among our team as to whether or not this was sufficient to erect a new taxon. In the end we decided the material was too scant to support a name of its own, but it is almost certainly a new species.

The width of R.2395 suggests the neck owner was not a tiny animal. It's difficult to get a size estimate from a single neck bone, especially with cervical length varying so much taxonomically across Pterosauria, as well as ontogenetically. However, vertebral width is a little more stable with respect to overall body size, and that of R.2395 indicates an animal on the small size for an azhdarchid - an arm-wavy wingspan estimate of 3 m seems about right. When we plugged the length of R.2395 into a database of near-complete azhdarchid necks, we found the estimated CIII - CVIII length was a paltry 352–419 mm (the range depends on whether it represents a CIV or CV): 23-41% shorter than the estimated neck length of the similarly-sized Transylvanian azhdarchid Eurazhdarcho, langendorfensis and notably shorter than neck of the smaller (2.5 m wingspan) Chinese azhdarchid Zhejiangopterus linhaiensis  (measured neck length 502 mm). It seems R.2395 did indeed have a short, robust neck for an azhdarchid of its size.

The possibility that R.2395 represents a short-necked juvenile of a long-necked species was something we looked into. After all, pterosaur necks seem to increase in length disproportionately to body size, and a 3 m wingspan would leave a lot of growing room for several azhdarchid species. However, the bone texture of R.2395 is characteristically 'polished' and avascular where well preserved, which has been suggested for some pterosaurs as an indicator of skeletal maturity. This observation is bolstered by the sharply ossified anatomy of R.2395: young pterosaur skeleltons tend to have rounded, poorly defined features, but our vertebra shows a surprising amount of sharply-defined detail in its superficially simple form. We don't know how old R.2395 was when it died, but it did seem to have a very well-ossified skeleton: it was likely at, or very near to, full size, and further neck growth seems unlikely. 

What does it mean to find short necks in a clade thought to be defined by long necks? We have no idea how this plugs into azhdarchid evolution, although the retention of all azhdarchid cervical features apart from the enhanced length is of interest there. We might also conclude that suggestions azhdarchids were all anatomically similar (e.g. Witton and Naish 2008) are questionable. There are doubtless some functional and palaeoecological implications of this discovery as well - we speculate that neck mechanics and strength likely differed between long- and short-necked azhdarchids - but further remains are clearly needed to say anything substantial about specific functionality. And... in truth, I'm biting my tongue here: there's tons to say about this, but I don't want to scoop other papers which are in prep and review. Both myself and Darren Naish have been hinting at some of this short-necked azhdarchid stuff for a while now, and heavily implying that we have things to say about Haztegopteryx as well as this much smaller animal. Some readers may, therefore, be wondering why there's no discussion of giant pterosaurs here. We were both hoping that this would be out by now as well, but it turns out that this parallel-running project has been published first - that's just how these things work out sometimes! Further details on these finds is coming, with other publications in states of progress which will add to this picture. Watch this space, in other words.

To end on a less cagey note, it is worth briefly mentioning why the seemingly maximum size of R.2395 is rather interesting. It is often said that small pterosaurs are absent from the upper Cretaceous, a fact sometimes controversially attributed to competition with birds. If R.2395 really is an adult, it joins Eurazhdarcho and the 2.5 m wingspan Montanazhdarcho in suggesting that smaller pterosaurs were not absent, and perhaps even not uncommon, in the latest Cretaceous. Granted, these species are not as small as some pre-Cretaceous pterosaurs, but they are certainly size-consistent with taxa found in Lower Cretaceous Lagerstätten of China and Brazil. I do wonder if the lack of sites of exceptional preservation in the Late Cretaceous has resulted in under-sampling of smaller pterosaur species in Late Cretaceous rocks, and that drawing conclusions about the Cretaceous decline of diminutive pterosaurs, and associated competition with birds, is premature. An elephant in the room here is the scarcity of small, Late Cretaceous juvenile pterosaurs: we know they had to exist, and yet they are exceptionally rare fossils. There is clearly a preservation bias against these small individuals - can we rule out that the same bias was not acting against small adults, too? That might be nonsense, but I do wonder if  we sometimes take the - knowingly poor - fossil record of pterosaurs a bit too literally.

A quick plug for a good cause

Finally, some readers will know that I like to bang drums about Supporting Original Palaeoart, and that it's often independent artists who're providing some of the more interesting and creative palaeoart projects out there. One of these comes from David and Jennie Orr (David is perhaps best known around these parts for founding Love in the Time of Chasmosaurs, as well as a his unique, stylish artwork), who have launched an Indiegogo campaign to fund their book Mammoth is Mopey. It's a wonderfully illustrated book for younger readers showing a different prehistoric animal for each letter of the alphabet. The animals in question aren't the same tried-and-tested taxa we see in every kids book however: they're the likes of Brontomerus, Jeholopterus, Gorgonops and so on, and each is wonderfully illustrated in David's Bézier-curve-loving style. It looks great, and I've already pledged enough for two copies: one for my nephew, and another for me (hey, you're never too old to have an alphabet refresher, right?). An example image from the book is below - I can't wait to see the rest.

Artistic Ankylosaurus is artistic. From Mammoth is Mopey, which you can support here. Illustration by David Orr.

References

• Averianov, A. O. (2010). The osteology of Azhdarcho lancicollis Nessov, 1984 (Pterosauria, Azhdarchidae) from the late Cretaceous of Uzbekistan. Proceedings of the Zoological Institute RAS, 314(3), 264-317.
• Witton, M. P., & Naish, D. (2008). A reappraisal of azhdarchid pterosaur functional morphology and paleoecology. PLoS One, 3(5), e2271.
• 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 (Haţeg Basin, Transylvania, Romania). American Museum Novitates 3827, 1-16.