Among the long-necked ctenochasmatids
close to Gegepterus, is Gladocephaloideus (Lü et al. 2012), a taxon originally considered a cycnorhamphid. A new, smaller specimen with an equally impressive set of long cervicals was recently assigned to the same genus (Lü et al. 2016, Fig. 1) as a juvenile.
Figure 1. Gladocephaloideus (the holotype) compared to the new specimen referred to Gladocephaloideus and its two sister taxa in the large pterosaur tree. Long necks in ctenochasmatids made several appearances by convergence. Of particular interest, note the size of the pelvis in the JPM specimen, no larger than that of the much smaller MB.R. specimen. Lü et al considered the pelvis incomplete and it may be. Sister taxa are illustrated here from figure 2.
the two specimens do not nest together in the large pterosaur tree (Fig. 2). Close, but separated by several nodes.
Before leaving Figure 1,
note the size of the relatively tiny pelvis in the JPM specimen, no larger than that of the much smaller MB.R. specimen with a relatively large pelvis of similar shape. This reconstruction was built from scraps that give the appearance of complete bones. Lü et al. 2016 considered the pelvis incomplete and did not attempt a reconstruction. No sacrum is associated with the pelvic elements to confirm or refute the present size reconstruction. New conspecific specimens will help.
Figure 2. Ctenochasmatids arise from these dorygnathids.
Lü et al. amended the diagnosis of Gladocephaloideus
to accommodate the new smaller specimen. That’s not a good idea before determining that they are indeed conspecific. In order to obviate that prospect, in phylogenetic analysis Lü et al. created a chimaera of the two specimens eliminating any possibility of testing one against the other and against all other pterosaur taxa. In my experience it is extremely rare to find conspecific pterosaurs. That is why I try to nest only specimens, not chimaeras.
Figure 3. (Left) The pes of Gladocephaloideus compared to (right) the pes of Ctenochasma elegans (a smaller, more primitive Ctenochasma with fewer teeth). Compare to the pes in figure 1.
of the holotype and referred specimen are similar but not the same in proportion or appearance. They score differently. In pterosaurs, feet are like fingerprints, enabling one to lump conspecific taxa and split convergent look-alikes.
Figure 4. The JPM specimen in situ along with a reconstruction of its skull compared to the holotype of Gladocephaloideus. The anterior mandible was glued on in the wrong direction, as noted by Lü et al. 2016.
Lü et al. did not test
the MBR stem ctenochasmatid. Lü et al. nest ctenochasmatids with cycnorhamphids, among other odd yet traditional nestings. This may be due to the low number of included taxa (67) vs. 215 in the large pterosaur tree.
The taxonomy and systematics get a little confusing….
the original Gladocephaloideus was assigned by Lü et al. 2012 to the Gallodactylidae (cycnorhamphids), but in consideration of the smaller specimen Lü et al. 2016 shifted it to the Ctenochasmatidae where it nests with Pterofiltrus, which nests as a cycnorhamphid in the large pterosaur tree. Moreover, Lü et al. include as related taxa, Elanodactylus (a derived germanodactylid, basal to pteranodontids), Beipiaopterus (a basal azhdarchid), Feilongus and Moganopterus (both cycnorhamphids). At the next level of closest kin Lü et al. include Pterodactylus longicollum (a pterodactylid), Gnathosaurus (a ctenochasmatid) and Cearadactylus (an ornithocheirid nesting far from other ornithocheirids).
This buckshot phylogeny only get worse, but I’ll stop here.
I applaud Lü et al.
for re-identifying the holotype Gladocephaloideus as a ctenochasmatid, as first reported here several years ago. But the rest of their phylogenetic analysis has to add taxa to get up to speed with current research. Their pterosaur tree recovered over 3000 MPTs compared to the fully resolved single tree at ReptileEvolution.com.
Juvenile? Big question that needs new insight to answer:
Lü et al. 2016 report, “The unfused contact between the extensor tendon process and the proximal end of wing phalange 1, as well as the poorly ossified epiphyses of the wing phalanges, indicates that JPM-2014-004 is an early juvenile individual.” It is hard to consider the JPM specimen a juvenile because in phylogenetic analysis it is larger than both proximal adult taxa. The more primitive MB.R. specimen, despite its size is an adult having undergone phylogenetic miniaturization from larger Dorygnathus and Angustinaripterus ancestors, an evolutionary process that often gives rise to new morphologies, in this case, the clade Ctenochasmatidae with all of its synapomorphies. Phylogenetically miniaturized pterosaurs retain, through neotony, juvenile bone microstructure. Lü et al. 2016 report woven bone structure and no LAGs or zones. This may occur in rapidly growing tiny (sparrow-to-hummingbird-sized) adults less than one year in age. Tiny pterosaurs, like tiny birds may have matured in far less than a year, just like small birds. Lü et al. 2016 note that LAGs are uncommon in pterodactyloids, but may be seen in larger specimens that presumably lived longer. Size matters in a phylogenetic context. This has to be considered before making statements about ontogenetic age estimates.
Tibial bone wall thickness
With a radius of 16 units (holding a cm scale up to my monitor) the tibial bone thickness was 5 units, leaving 11 units of hollow cavity. That is same cortex ratio as in the 2x larger (9x heavier) holotype specimen.
But wait! What’s this?? A long rostrum on a juvenile???
That can’t be so, IF you follow the work of Bennett, Witton, Wellnhofer and others. That would be internally inconsistent! However, if you follow ReptileEvolution.com and the Pterosaur Heresies you’ll note that many tiny adults AND embryos had long rostra and small eyes. No problem under the isometric growth hypothesis, which I hope will someday gain a little acceptance because it is demonstrably factual.
Lü J-C, Ji Q, Wei X-F and Liu Y-Q 2012. A new ctenochasmatoid pterosaur from the Early Cretaceous Yixian Formation of western Liaoning, China. Cretaceous Research in press. doi:10.1016/j.cretres.2011.09.010.
Lü J, Kundrát M, Shen C 2016. New Material of the Pterosaur Gladocephaloideus Lü et al., 2012 from the Early Cretaceous of Liaoning Province, China, with Comments on Its Systematic Position. PLoS ONE 11(6): e0154888. doi:10.1371/ journal.pone.0154888