Where is the rest of Lanthanolania?

It was back in 2011
when the post-crania of Lanthanolania (Fig. 1) was reported in an abstract by Modesto and Reisz. Prior to that, in 2003, only the skull was described by the same authors. Over the last six years the post-crania of Lanthanolania has not been published.

From the 2011 SVPCA abstract:
“The evolutionary history of Diapsida during the Palaeozoic Era is remarkably poor. Following the reclassification of the Early Permian Apsisaurus witteri as a synapsid last year, only a handful of taxa span the large temporal gap between the oldest known diapsid Petrolacosaurus kansensis and the Late Permian neodiapsid Youngina capensis. These include two Middle Permian neodiapsids, the recently described Orovenator mayorum from Oklahoma, USA, and Lanthanolania ivakhnenkoi from the Mezen region, northern Russia. A recently collected, nearly complete skeleton of Lanthanolania permits a thorough reexamination of the phylogenetic relationships of these two taxa.

“Phylogenetic analysis of 188 characters and 30 diapsid taxa positions these two small forms as stem saurians and the oldest known neodiapsids (recently redefined by the authors as the sister taxon of Araeoscelidia). Interestingly, our results suggest that the lower temporal bar was lost by the ancestral neodiapsid relatively soon after the evolution of the diapsid temporal morphology, and conversely, that the temporal configuration of the Late Permian Youngina capensis is a secondary condition. In addition, the skeletal anatomy of Lanthanolania provides evidence of limb proportions that suggest that this small reptile is the oldest known bipedal diapsid.”

Figure 1. Kuehneosaurid skulls from Palaegama to Coelurosauravus and Mecistotrachelos, and to Lanthanolania, Pamelina, Kuehneosaurus, Icarosaurus and Xianglong. Some of these taxa were not previously recognized as kuehneosaurids or their ancestors.

Figure 1. Kuehneosaurid skulls from Palaegama to Coelurosauravus and Mecistotrachelos, and to Lanthanolania, Pamelina, Kuehneosaurus, Icarosaurus and Xianglong. Some of these taxa were not previously recognized as kuehneosaurids or their ancestors.

Earlier (2011) the large reptile tree (LRT) nested Lanthanolania with the so-called rib gliders between Coelurosauravus and Icarosaurus. Back then we looked at those issues here.

Modesto and Reisz (2003) had a hard time
nesting Lanthanolania and considered it ‘enigmatic’. The closest they came was to nest Lanthanolania at the base of the lepidosauriformes (Rhynchocephalia + Squamata) and in other tests, with Coelurosauravus, which they split apart from the lepidosauriformes by adding intervening unrelated ‘by default’ taxa.

Unfortunately
with their small taxon list, Modesto and Reisz (2003) did not recover the basal split among reptiles that had occurred between the new Lepidosauromorpha and Archosauromorpha at Gephyrostegus + kin at the earliest Carboniferous. Thus the formerly monophyletic clade Diapsida is diphyletic in the LRT. Modesto and Reisz  mixed taxa from the two major clades and that muddied their results. Parts of their results were essentially correct, just unintelligible due to the addition of unrelated intervening archosauromorph basal diapsids.

Traditional paleontology
has likewise never nested coelurosauravids with kuehneosaurids, like Icarosaurus, perhaps based in part on the rib/dermal rod issue.

Problems and guesses:

  1. “Phylogenetic analysis of 188 characters and 30 diapsid taxa positions these two small forms as stem saurians and the oldest known neodiapsids (recently redefined by the authors as the sister taxon of Araeoscelidia).” — Sauria (= last common ancestor of archosaurs and lepidosaurs), is a junior synonym for Reptilia in the LRT. Neodiapsida (= includes all diapsids apart from araeoscelidians (= Petrolacosaurus and Araeoscelida)) or all taxa more closely related to Youngina than to Petrolacosaurus. Thus, in their thinking, Sauria is a clade within Neodiapsida. Modesto and Reisz do not yet recognize that Diapsida is no longer a monophyletic clade. In the LRT Orovenator and Lanthanolania are not related. The former is a basal diapsid archosauromorph. The latter is a basal lepidosauriform lepidosauromorph.
  2. “Interestingly, our results suggest that the lower temporal bar was lost by the ancestral neodiapsid relatively soon after the evolution of the diapsid temporal morphology,” — According to the LRT, the lower temporal bar was not lost nor was it present in the lepidosauromorph ‘rib’ gliders, including Lanthanolania. By contrast, Orovenator is one of the most basal archosauromorphs with an upper temporal fenestra.  Petrolacosaurus is older.
  3. “and conversely, that the temporal configuration of the Late Permian Youngina capensis is a secondary condition.” — In the LRT, it is not a secondary configuration, but is derived from basal diapsid taxa like Orovenator.
  4. “In addition, the skeletal anatomy of Lanthanolania provides evidence of limb proportions that suggest that this small reptile is the oldest known bipedal diapsid.” — I can only guess why they promoted this hypothesis: short torso and long hind limbs? Icarosaurus has such proportions. So does Kuehneosaurus. So does their last common ancestor, Palaegama (Fig. 2) which lacks wire-like dermal ossifications.
Figure 3. Palaegama, close to the origin of all Lepidosauriformes.

Figure 2. Palaegama, close to the origin of all Lepidosauriformes.

The question today is
where is the paper that describes the above-mentioned post-crania of Lanthanolania? Is the post-crania definitely referable?

If the referred specimen came from similar sediments
the matrix was described in 2003 as ‘extremely hard to work with’. Perhaps it is still being worked on. Or it has been shelved.

Phylogenetic bracketing
indicates that the new specimen might or should have wing-like wire/rod dermal elements, like those found in both Coelurosauravus and Icarosaurus, but traditionally considered ribs in Icarosaurus. They are not ribs, as we learned earlier here. The real ribs are short and fused to the vertebrae, appearing to be long transverse processes, but no related taxa have long transverse processes and not all of the ribs are fused to the vertebrae, betraying their identity. Since a mass of dermal rods was not mentioned in the abstract, one  wonders if the new specimen was actually closer to Palaegama than to Lanthanolania?

Late news from Sean Modesto about Lanthanolania:
“The project is currently in the hands of Dr. Reisz. No “ETA” as yet!”

Problems like this one
are a good reason to include the taxa the LRT suggests one include in smaller, more focused studies.

References:
Modesto SP and Reisz RR 2003. An enigmatic new diapsid reptile from the Upper Permian of Eastern Europe. Journal of Vertebrate Paleontology 22 (4): 851-855.
Reisz RR and Modesto SP 2011. The neodiapsid Lanthanolania ivakhnenkoi from the Middle Permian of Russia, and the initial diversification of diapsid reptiles.SVPCA abstract published online.

 

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