Above: Male (foreground) and female Geosternbergia sternbergi, mounted casts at the Royal Ontario Museum. Photo by Kenn Chaplin, licensed.
This paper is interesting in that it attempts to reverse some of the trends of the past few decades concerning pterosaur diversity, in some cases (in others it's merely a re-calibration of the old genericometer). Kellner erects two new species and ressurects a genus, Geosternbergia.
I first encountered Pteranodon sternbergi in Dave Peters' awesome picture book A Gallery of Dinosaurs and other Ancient Reptiles (which also probably instilled my OCD towards drawing scale diagrams). It took my young mind a while to register that this huge, tall/broad crested pterosaur belonged to the same genus as the familiar, backward-pointing-crested Pteranodon lonciceps that flew beside it. I have to admit that I though P. sternbergi just looked... cooler. In this kind of side-by-side comparison, they really don't looks like they belong to the same genus. But as always, genera are subjective things, and nobody doubted that P. longiceps and P. sternbergi were each others closest relatives (they even overlapped in time and geologic range).
P. sternbergi was first described by Harksen in 1966 as a species of Pteranodon, based on a skull which differed from other species by its tall, vertical crest. In 1972, Miller placed it rather arbitrarily in its own subgenus, as Pteranodon (Sternbergia) sternbergi. The name Sternbergia turned out to be pre-occupied (as far as I know, subgenenus names compete with genera for priority). Miller amended the name to Geosternbergia in 1978.
However, this designation fell out of favor by the 1990s, when Chris Bennett published a couple of hefty reviews of Pteranodon from the Niobrara and related formations in Kansas. Bennett stramlined Pteranodon taxonomy, taking the several species that had been considered valid up to that time and showing that much of the variation was likely due to age and/or sexual dimorphism. For example, Bennett re-affirmed the idea that some small-crested Pteranodon specimens represent females (and some juveniles) of the same species as the longer-crested adult males. With this variation in mind, he suggested that all Pteranodon specimens could fit into two species: P. lonciceps and P. sternbergi. Because differences between species are limited almost entirely to the skull and crest (though Kellner 2010 suggests that some consistent differences may be found in the skeleton with further study), Bennett had to rely mainly on stratigraphic position to decide which species a specimen belonged to. While the two did overlap in time, it was only for a very brief period, so even though skulls are very rare compared to skeletons, a specimen from the lower Niobrara could be confidently referred P. sternbergi, while one from higher in the formation probably came from P. longiceps. (Image at left: illustration of various Pteranodon skulls by Matt Martyniuk, licensed).
One problem with this method, which Kellner points out in his new paper, is that many specimens (especially those recovered back during the days of Cope and Marsh) lack information about their provenence detailed enough to allow such an assignment. Bennett himself simply referred these to Pteranodon sp., but if Kellner's new work holds up, they'd need to be assigned even more broadly, only to indeterminate Pteranodontidae.
That's because Kellner has re-assessed the variation within the traditional grouping known as Pteranodon, and found some specimens that seem to represent new species among them. For me, the most interesting is Dawndraco kanzai, or "Kanza Dawn dragon" named for Dawn, apparently an Iroquois sky goddess, not the English word. The type and so far only specimen is UALVP 24238, a really interesting nearly complete skull and skeleton usually attributed to P. sternbergi (as in Bennett, 1994). Aside from being one of the most complete (former) Pteranodon specimens, it has always struck me as very, very odd. One of the primary reasons for making this a new species it its upper jaw. In most Pteranodon skulls (though none are as complete as you may assume based on its ubiquity in paleoart), the jaws can be seen to curve upward toward the tip and taper off into a needle-sharp projection at the tip. In the Dawndraco holotype, the preserved portion of the jaws are extremely long relative to the rest of the skull, but show no signs of tapering. In fact, the top and bottom margins of the upper jaw form essentially completely parallel lines up until the break. Letting your imagination fill in the rest, this must represent either a phenomenally long-billed animal, or one with a very unusual fat, somewhat flattened tip. The bone within this tall, long bill looks like a loose, honeycomb mesh of very thin struts. Taken together, this does seem like it probably comes from something fairly different than Pteranodon proper. (Image at right: Skull of Dawndraco, from Kellner 2010. Scale bar = 500mm).
The next new species is Geosternbergia maysei. Kellner considers Geosternbergia a distinct genus based on its unique skull characteristics, but again, there is currently no analysis to suggest that it is any more or less closely related to Pteranodon than anything else, so it remains a subjective decision (though it would be interesting to see someone perform an analysis using all of Kellner's species and Nyctosaurus). Anyway, G. maysei is named for a partial skull (KUVP 27821) from the South Dakota Sharon Springs formation. It was a large individual that Bennett previously referred to P. longiceps. However, Kellner notes that the crest is inclined further upward than it should be for that species, and that the premaxilla is arranged differently in forming part of the crest. It also appears to have a larger nasoantorbital fenestra, and a lower and larger temporal fenestra, than in G. sternbergi.
How well either of these identifications remains to be seen. I'm more inclined to be convinced by Dawndraco than G. maysei, simply because the very strange bill of the former seems harder to explain by age or gender variation. Kellner has also tended to be the 'leader' of one 'camp' when it comes to pterosaur taxonomy, usually opposed to Dave Unwin -- see their drastically different recent taxonomies of the ornithocheirids, for example. It will be interesting to see not only if these new species are accepted, but by whom.
While on the subject of new pterosaurs, two new species have also just been reported from the mid-Jurassic Tiojishan formation, some with soft tissue: Kunpengopterus sinensis and Darwinopterus linglongtaensis seem to add support for a monophyletic group of Tiaojishan pterosaurs somewhat intermediate between pterodactyloids and "rhamphorhynchoids", the Wukongopteridae. If I have time to read the paper more closely I'll try to follow up with a full post on these guys.
Kellner, A.W.A. (2010. "Comments on the Pteranodontidae (Pterosauria, Pterodactyloidea)
with the description of two new species." Anais da Academia Brasileira de Ciências, 82(4): 1063-1084. doi: 10.1590/S0001-37652010000400025.