A slight adjustment to an OMNH Cotylorhynchus reconstruction

The AMNH specimen
of Cotylorhynchus is a spectacular sight to see (Fig. 1). It’s huge! And complete! It’s bigger than a Galápagos tortoise with a skull just as small.

Figure 1. Cotylorhynchus AMNH specimen. Note the angle of the ribs.

Figure 1. Cotylorhynchus AMNH specimen. Note the angle of the ribs. Is that a whip-lash tail? Compare to Datheosaurus (Fig. 4). Just look at those massive elbows!

Romer and Price 1940
pictured Cotylorhynchus with vertical dorsal ribs (Fig. 2 lateral view).

Cotylorhynchus romeri

Figure 2. Cotylorhynchus romeri

The Sam Noble Museum Oklahoma’s Museum of Natural History
in Norman, Oklahoma, USA, has a mount of Cotylorhynchus (Fig. 3) that follows the Romer and Price illustration with vertical ribs. Here, in this 2-frame GIF animation, I have angled them back to match the in situ specimen and most other quadrupedal tetrapods.

Figure 3. Cotylorhynchus mount in the Sam Noble Museum of Natural History with vertical ribs modified here to have diagonal ribs more typical of tetrapods and reflective of the in situ fossil.

Figure 3. Cotylorhynchus mount in the Sam Noble Museum of Natural History with vertical ribs modified here to have diagonal ribs more typical of tetrapods and reflective of the in situ fossil.

Cotylorhynhcus romeri (Stovall 1937) Kungurian, Middle Permian, ~265 mya, ~6 m in length, was the largest sister to Casea and Ennatosaurus. It was the largest land animal of its time.

Figure 2. Milleretta, caseasaurs and kin. The LRT nests these taxa together apart from the Synapsida, with which they share a lateral temporal fenestra.

Figure 4. Milleretta, caseasaurs and kin. The LRT nests these taxa together apart from the Synapsida, with which they share a lateral temporal fenestra. Note the angle of the ribs in the Milleretta reconstruction, similar to the suggestion for Cotylorhynchus. Casea and Ennatosaurus continue to have invalid vertical ribs in the above figure due to my laziness.

All prior and other current reports
nest Cotylorhynchus with the synapsid pelycosaurs, but here in the large reptile tree (LRT. 1315 taxa) the caseid clade nests more parsimoniously with Milleretta, Feeserpeton and Australothyris and other plant-eaters, many of which share a lateral temporal fenestra in the new Lepidosauromorpha, opposite to the coeval pelycosaurs nesting in the new Archosauromorpha.

We looked at this traditional mistake
based on taxon exclusion here back in 2011. Even so, synapsid workers continue to follow this outdated tradition without testing validated alternatives proposed here.

References
Romer AS and Price LI 1940. Review of the Pelycosauria. Geological Society of America Special Papers 28: 1-538.
Stovall JW 1937. Cotylorhynchus romeri, a new genus and species of pelycosaurian reptile from Oklahoma. Arnerican Journal of Science (5) 34: 308-313.
Stovall JW, Price LI and Romer AS 1966. The Postcranial Skeleton of the Giant Permian Pelycosaur Cotylorhynchus romeri. Bulletin of the Museum of Comparative Zoology 135 (1): 1-30. online pdf

wiki/Cotylorhynchus

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SVP abstracts – Ennatosaurus

Romano et al. 2017
brings us a new reconstruction of Ennatosaurus (Fig. 1, btw – this is not it.)
Figure 1. An old reconstruction of Ennatosaurus, still not a synapsid, closer to millerettids and Eunotosaurus.

Figure 1. An old reconstruction of Ennatosaurus, still not a synapsid, closer to millerettids and Eunotosaurus.

As you read the abstract,
bear in mind the only thing wrong here is the author’s insistence that Ennatosaurus is a pelycosaur and a synapsid. It is neither, as the addition of taxa to a cladistic analysis would have informed the Romano team. Ennatosaurus was derived from the similarly built milllerettids. This was demonstrated several years ago in the large reptile tree by the simple addition of taxa to the inclusion set.

From the abstract:
The Russian caseid Ennatosaurus tecton (Synapsida Caseasauria) is an important member of the group, being among the few “pelycosaurs” occurring in the Middle Permian, thus making caseids among the longest-surviving groups of non-therapsid synapsids. Although the cranial skeleton has been recently restudied in detail, the descriptions currently available for the postcranium are essentially limited to the original short account on the holotype provided by the original description from the 1950s. This contribution represents a new analysis of the postcranium of this taxon, using several different approaches. The postcranium of Ennatosaurus is informative with respect to both the taxonomy and phylogeny, with autapomorphic characters present particularly in the vertebral column. In addition, we conducted eight principal component analyses to investigate the position of the various appendicular elements of Ennatosaurus within the caseid morphospace. Members of all major groups of “pelycosaurs” were included in the morphometric analysis (along with selected outgroup taxa), allowing us to make some broader preliminary inferences regarding postcranial morphospace occupation of these basal synapsids for each individually-considered element. From the results of the principal component analyses, a major decoupling among the morphological patterns of stylopodial and zeugopodial elements is detected. Whereas femora and humeri exhibit a shared common pattern (with a wider overlap in their respective morphospace), the ulnae, radii, tibiae and fibulae show well-separated regions of morphospaces in the different clades. This result indicates the importance of such long bones also for taxonomic differentiation (in addition to their use for classical functional and biomechanical studies). Finally, a 3D photogrammetric model of the mounted specimen at the Paleontological Institute of Moscow has been used to obtain the first in vivo reconstruction of Ennatosaurus tecton, providing for the first time a potentially realistic picture of the Russian caseid in life.

For all this great work
resistance to taxon inclusion doomed any conclusions drawn. Sadly this basic problem is similar to workers who resist adding fenestrasaurs to pterosaurs studies, thalattosaurs to Vancleavea studies, tenrecs and desmostylians to whale studies, etc. etc…

References
Romano M, Brocklehurst N and Fröbisch J 2017. Redescription of the postcranial skeleton of Ennatosaurus tecton (Synapsida, Caseasauria, Caseidae) and its first in vivo restoration. Abstrcts from the 2917 meeting of the Society of Vertebrate Paleontology in Calgary.

Just ran the numbers: Eocasea is not a sister to Casea

Reisz and Fröbisch 2014
considered Eocasea martini (Late Pennsylvanian, Fig. 1) the basalmost caseid, despite its long slender appearance and small size.

Figure 1. Eocasea in situ with anterior skull imagined.

Figure 1. Eocasea in situ with anterior skull imagined based on phylogenetic bracketing. This long, low taxa does not nest with large, big-bellied caseasaurs but with more similar sister, including Delorhynchus. These are not wide dorsal ribs, but belong to a specimen with a standard, narrow torso.

And at the time (2 years ago), I wrote in ReptileEvolution.com,
“[Eocasea] had a long narrow torso and short legs. Note the resemblance to millerettids like Australothyris and Oedaleops.” 

Reisz and Fröbisch did not include those two
in their phylogenetic analysis because they were so sure they had a caseasaur. They also did not include Eunotosaurus, Acleistorhinus, Microleter, Delorhynchus, or Feeserpeton for the same reason.

But they should have done so, because
that’s where Eocasea nests in the large reptile tree (Fig. 2) close to, but not with caseasaurs.

Figure 2. Eocasea nests between Feeserpeton + Australothyris and Delorhynchus in this subset of the large reptile tree, not close to Casea.

Figure 2. Eocasea nests between Feeserpeton + Australothyris and Delorhynchus in this subset of the large reptile tree, not close to Casea.

The problems with the Reisz and Fröbish data set
is that it was too small. The authors presumed Eocasea would be a caseasaur and so included only the following taxa: Reptilia, Diadectes, Limnoscelis, Mycterosaurus, Varanops, Oromycter, Casea, Cotylorhynchus, Angelosaurus, Ennatosaurus and Eocasea. As you can imagine, providing scores to a clade named, “Reptilia” is inappropriate and, frankly, dangerous, whether cherry-picking traits or scoring all zeroes. Diadectes, Limnoscelis, Mycterosaurus, Varanops are not related to each other or to caseasaurs. Reisz and Fröbisch were making guesses without having a large gamut study from which to draw subsets.

Reisz and Fröbisch report,
“Eocasea changes significantly our understanding of the evolutionary history of both caseids and caseasaurs.” Not with the current nesting. The discovery of Eocasea extends the fossil record of Caseasauria and Caseidae significantly, well into the Pennsylvanian, in line with the fossil record of other early synapsid clades, indicating that the initial stages of synapsid diversification were well under way by this time.” Eocasea is not a caseid and caseids are not synapsids, so it doesn’t extend anything related to Caseidae.

“More significantly, Eocasea also allows us to re-evaluate the origin and evolution of herbivory within this clade, and terrestrial vertebrates in general.” It’s not an herby and it’s not a clade member. “Thus, we can identify Paleozoic herbivores because their rib cages are typically significantly wider and more capacious than those of their closest insectivorous or carnivorous relatives. Not in Eocasea. “Nevertheless, it is likely that the ability to process this kind of plant matter precedes the skeletal correlates that can be found in the fossil record.” This is an unsupported supposition in light of the new nesting. “It is therefore possible that we are underestimating the extent of herbivory that existed in the Paleozoic, but this does not invalidate our results because the clades of herbivores that we examine here are widely separated by successive clades of non herbivorous vertebrates.” There is no ‘therefore’ when the setup if invalid. 

To their credit, 
Reisz and Fröbisch did not nest Edaphosaurus or Protorothyris as outgroup taxa to the Caseasauria (Eothyris at its base). Perhaps that is so because they did not include these taxa! And I wonder why? But they did nest the unrelated Varanops and Mycterosaurus as caseasaur outgroups. Both nest about twenty nodes away on the other major branch of the Reptilia, the new Archosauromorpha.The Reisz and Fröbisch tree is bogus because their outgroup taxon list was based not on testing, but on tradition.

If you’re looking for
osteological evidence for herbivory in the ancestry of the Caseasauria, you won’t find big bellies and flat teeth, but you will find several herbivores arising from Milleretta (late Permian late survivor of a Carboniferous radiation) in the clade Lepidosauromorpha. These include diadectids and their allies bolosaurids (post-crania unknown) and procolophonids, pareiasaurs and their allies turtles, along with caseasaurs.

Working on their ‘wish list’
Reisz and Fröbisch continue with their hypothesis: “Whereas other caseids also show dental specializations, with leaf-like large serrations being present in the marginal dentition, Eocasea, Oromycter, and the undescribed Bromacker Quarry caseid lack these serrations. Interestingly, both Oromycter, and the Bromacker caseid show skeletal evidence for herbivory, raising the possibility that oral processing in the form of puncturing vegetation may have evolved within Caseidae after the acquisition of herbivory.”  Only the latter two are indeed caseasaurids. Eocasea definitely is not one. You can’t derive homolog conclusions from unrelated taxa.

Reisz and Fröbisch continue
“Late Pennsylvanian and Early Permian diadectids also show convincing evidence of dental and skeletal adaptations for herbivory. These enigmatic
[not any more] Paleozoic forms are part of Diadectomorpha, a sister group to crown Amniota  [not any more]. A preliminary phylogeny of diadectids indicates that Ambedus, a small diadectid from the Early Permian, tentatively identified as omnivorous because of its labiolingually expanded cheek teeth (but no evidence of dental wear) is the sister taxon to all other diadectids. Ambedus may not be a diadectid as noted here. However, the oldest known diadectid from the late Pennsylvanian of Oklahoma is already clearly an herbivore and older than the edaphosaur Edaphosaurus novomexicanus. As is the case with the caseid and edaphosaur synapsids, the sister taxon of Diadectidae, the Early Permian Tseajaia from New Mexico, was faunivorous.” That oldest known diadectid is not identified.

Reisz and Fröbisch add to their unsupported hypothesis with,
“Although the holotype of Eocasea certainly represents a juvenile individual [actually, and you can check this, it is the same size as sister taxa, but smaller than basal and other caseasaurs], it is diminutive, with an estimated snout-vent length of 125 mm. In contrast, the smallest known herbivorous caseid with a comparable ontogenetic age, based on level of ossification of the vertebrae and pedal elements, is a basal, undescribed form from Germany and has an estimated snout-vent length of 400 mm.” Not sure which specimen this is…

If Reisz and Fröbisch had just
increased the size of their taxon list, they would/could have correctly nested Eocasea, and avoided making the many subsequent mistakes based on that bad nesting, including the unfortunate and inappropriate naming of the taxon and the bogus headline that got tacked to the article and all the PR that attended it.

We don’t have a name yet
for the enanticaseasaurs or paracaseasaurs (Fig. 2), but we need one!

References
Reisz R and Fröbisch J 2014. The oldest caseid synapsid from the Late Pennsylvanian of Kansas, and the evolution of herbivory in terrestrial vertebrates. PLoS ONE 9(4): e94518. doi:10.1371/journal.pone.0094518

 

Vaughnictis (Brocklehurst et al. 2016): a new last common ancestor of birds and bats

Recently Brocklehurst et al. 2016
renamed ‘Mycterosaurus’ smithae (Lewis and Vaughn 1965; early Permian, MCZ 2985). The new name is Vaughnictis smithae. The specimen was originally considered a varanopid close to the holotype of Mycterosaurus (Fig. 1). Now the Brocklehurst team nest the MCZ 2985 specimen between Eothyris and Oedaleops (Fig. 1) at the base of the Caseasauria, which they consider a clade at the base of the Synapsida.

Unfortunately, 
the large reptile tree nests Vaughnictis at the bases of two major clades: between Protorothyris and the Synapsida (represented here (Fig. 1) by Elliotsmithia) + the Prodiapsida (represented here by Mycterosaurus). The Caseasauria, as noted five years ago here, does not nest with the Synapsida when the taxon list is expanded, and the Prodiapsida (former varanopids) split from the Synapsida at their base when the taxon list is expanded.

Figure 2. Vaughnictis nests between Protorothyris and the Synapsida (Elliotsmithia) + Prodiapsida (Mycterosaurus) in the large reptile tree - not the Caseasauria (Eothyris + Oedaleops).

Figure 1. Vaughnictis nests between Protorothyris and the Synapsida (Elliotsmithia) + Prodiapsida (Mycterosaurus) in the large reptile tree – not the Caseasauria (Eothyris + Oedaleops). Despite the many similarities, the narrow skull with parallel sides, upturned mandible tip and longer rostrum are a few traits that split Vaughnictis from caseasaurs and lump it with prosynapsids.

Lewis and Vaughn got it right.
Vaughnictis is a sister to Mycterosaurus in the LRT. It is not a caseasaur.

Brocklehurst et al. got it right
in that Vaughnictis is distinct enough from Mycterosaurus to warrant its own genus.

Despite their phylogenetic distance
only 22 additional steps are needed when Vaughnictis is force shifted over to Eothyris. This is largely due to convergence. Both clades developed lateral temporal fenestrae in similar patterns, had large eyes and a short rostrum at this stage.

Figure 2. Vaughnictis skull in situ with color tracings. See figure 3 for reconstruction.

Figure 2. Vaughnictis skull in situ with color tracings. See figure 3 for reconstruction. The jaws shifted posteriorly during taphonomy The parietal and its opening are difficult to read.

The phylogenetic importance of Vaughnictis
was overlooked by Brocklehurst et al. It is the most primitive known specimen in the lineage of synapsids and diapsids to have a lateral temporal fenestra. That’s why it is the last common ancestor of bats and birds, an honor formerly earned by Protorothyris, but now superseded by a taxon with lateral temporal fenestrae.

Figure 1. Color tracings of bones moved to their in vivo positions and traced.

Figure 3. Color tracings of bones moved to their in vivo positions and traced. Note the anterior shifting of the jaws to their in vivo positions based on posterior dentary and jugal positions common to most if not all candidate sister taxa. 

I wish that Brocklehurst et al. had 

  1. created a multi-view reconstruction
  2. showed candidate sisters side by side compared to Vaughnictis
  3. not excluded pertinent taxa (diapsids for the former varanopids and millerettids for the caseasaurs)
  4. used colors to identify bones, rather than lines, which helps when bones overlap. They did color the teeth (Fig. 4), but not all the teeth.
Figure 6. Teeth scanned by Brocklehurst fit to dorsal view of skull. Premaxillary and maxillary teeth were not published. Note the scale bar for the teeth appears to be off by a factor of 2.

Figure 4. Teeth scanned by Brocklehurst fit to dorsal view of skull. Premaxillary and maxillary teeth were not published. Note the scale bar for the teeth appears to be off by a factor of 2. Also note the premaxillary teeth appear to be jammed back from their in vivo position. 

The interesting thing about their cladogram
is that the Brocklehurst team nested Captorhinus, Limnoscelis and Tseajaia as outgroup taxa — which is correct for Caseasauria, but not for Synapsida. Protorothyris is also listed as the proximal outgroup to the Caseasauria (incorrect) + Synapsida (correct). It is clear they rely on tradition, rather than testing for their inclusion set.

In Vaughnictis, as opposed to Eothyris, note the 

  1. relatively narrow skull
  2. the rising mandible tip
  3. the lack of maxilla/orbit contact
  4. the shorter temporal length
  5. the lower rostrum

These traits ally Vaughnictis with Elliotsmithia to the exclusion of basal caseasaurs.

Brocklehurst et al. note:

  1. Vaughnictis lacks these mycterosaurine and varanopid traits (but it is not a member of either of these clades in the LRT) :
    a. slender femur
    b. linguo-labially compressed and strongly recurved teeth – I disagree, the teeth are indeed recurved
    c. lateral boss on the postorbital – I don’t see this on candidate taxa
  2. Vaughnictis has these caseasaur traits:
    a. coronoid teeth – plesiomorphic for synapsids, but they have been lost in derived caseids, ophiacodontids, varanopids and sphenacodontians. Most workers do not include these in their tracings of pertinent taxa, so are rarely noted.
    b. large supratemporal – actually they are long, as in synapsids, not large (and wide) as in caseasaurs
    c. large pineal foramen – unable to confirm, but Elliotsmithia and Mycterosaurus also have a large pineal foramen.

Teeth
Broklehurst et al. published synchrotron scans of the palatal, dentary and coronoid teeth (perhaps the scale bar should be 1 cm, not 5mm, Fig. 4), but did not publish scans of the maxillary teeth. All of the palatal teeth form shagreen fields, not single rows. That’s different than all candidate sister taxa, whether caseasaurid or protorothyrid. What they label as “vomerine teeth” may be premaxillary teeth based on the posterior displacement of the jaws. The dentary teeth are recurved and robust. Coronoid and parasphenoid teeth are present.

Figure 6. Subset of the large reptile tree showing the nesting of Vaughnictis at the base of the Synapsida and Prodiapsida.

Figure 5. Subset of the large reptile tree showing the nesting of Vaughnictis at the base of the Synapsida and Prodiapsida. Also note that the Synapsida is NOT the first clade to branch off from the base of the Amniota. Far from it.

This Brocklehurst team was led by
the venerable Robert Reisz, who has made dozens of great discoveries, but has resisted testing candidates suggested by the large reptile tree. And that sort of paleoxenophobia is unfortunate. Outsiders can make valuable contributions.

Finally, kudos and credit to the Brocklehurst team,
for finding the one best specimen closest to the advent of the Synapsida + Prodiapsida. It should be in every textbook from here on out.

References
Brocklehurst N, Reisz RR, Fernandez V and Fröbisch 2016. A Re-Description of ‘Mycterosaurus’ smithae, an Early Permian Eothyridid, and Its Impact on the Phylogeny of Pelycosaurian-Grade Synapsids. PLoS ONE 11(6):e0156810. doi:10.1371/journal.pone.0156810
Lewis GE, Vaughn PP 1965. Early Permian vertebrates from the Cutler Formation of the Placerville Area, Colorado. Geological Survey Professional Paper 500C: 1–50.
Reisz RR, Dilkes DW and Berman DS 1998. Anatomy and relationships of Elliotsmithia longiceps Broom, a small synapsid (Eupelycosauria: Varanopseidae) from the late Permian of South Africa. Journal of Vertebrate Paleontology 18(3):602-611.

Another, more complete Colobomycter adds data to this enigma

Revised June 10, 2016 with a new reconstruction and nesting with Eothyris. 

A new paper
by MacDougall et al. 2016 introduces Colobomycter vaughni (BRMP 2008.3.1, Fig. 1) a new toothy specimen that adds much needed data to the former enigma taxon, Colobomycter pholeter. They report on the synapomorphies, “enlarged premaxillary tooth and paired enlarged maxillary teeth, unique dentition that grants it an appearance quite distinct from other parareptiles at Richards Spur. This new material differs from that of C. pholeter in that it possesses at least three more teeth on its maxilla, the enlarged premaxillary and maxillary teeth are more gracile than those in C. pholeter, and the lacrimal is restricted externally to the orbital margin and does not exhibit an extra lateral exposure.” 

Figure 1. The new Colobomycter compared to the original pasted on a ghost of the new material. We're learning more about this genus!

Figure 1. The new Colobomycter compared to the original pasted on a ghost of the new material. We’re learning more about this genus!

Unfortunately, 
MacDougall et al. considered Colobomycter a member of the Lanthanosuchoidea. According to MacDougall et al. taxa in that clade include Feeserpeton, Lanthanosuchus, Acleistorhinus and Delorhynchus.

In the large reptile tree 
(Fig. 2) Lanthanosuchus
 nests with Bashkyroleter, Macroleter and Emeroleter.

On the other hand (and this is revised from the original posting)
Colobomycter pholeter
(Vaughn 1958, Modesto and Reisz 2008, UWBM 95405), Lower Permian ~278 mya, was originally considered a caseid pelycosaur, like Eothyris. (But note that Eothyris is not considered a pelycosaur in the large reptile tree (subset Fig. 2). Later, Modesto and Reisz (2008) considered Colobomycter a “parareptile” close to Acleistorhinus. After further consideration, it turns out that Colobomycter is indeed quite similar to Eothyris, as Vaughn 1958 indicated with much less data and fewer optional candidate taxa to consider. Hats off to Vaughn!

Figure 2. Revised cladogram of Colobomycter nesting this genus with Eothyris in an unnamed clade that includes caseasauria.

Figure 2. Revised cladogram of Colobomycter nesting this genus with Eothyris in an unnamed clade that includes caseasauria.

Sharp-eyed observers will note
that earlier I nested the rostrum of Colobomycter with procolophonids based on a smaller portion of rostrum. Clearly. I’m not as sharp as Vaughn was.

At this point
Colobomycter likely had a lateral temporal fenestra.

Herbivore or carnivore?
There are herbivores, carnivores and omnivores related to Colobomycter. It looks like the anterior dentary teeth could scrape off or collect whatever the premaxillary tusks had stabbed into. Eothyris had similar large maxillary teeth.

Figure 3. Eothyris skull in three views. This taxon is the closest known relative to Colobomycter.

Figure 3. Eothyris skull in three views. This taxon is the closest known relative to Colobomycter.

This is only one of tens of thousands of errors I have made
I’m only embarrassed by the ones that have yet to surface. Science and scientists don’t always have all the answers, but if the formula (or in this case cladogram) recovers a sticking point, as it did earlier, it will reward you to go back in and figure out where the errors were made. In this case several little errors among several taxa added up, but are corrected here, resulting once again in a completely resolved tree, hopefully more closely echoing Nature.

References
MacDougall MJ, Modesto SP and Reisz RR 2016. A new reptile from the Richards Spur Locality, Oklahoma, USA, and patterns of Early Permian parareptile diversification, Journal of Vertebrate Paleontology (advance online publication). www.tandfonline.com/doi/

Microleter mckinzieorum Tsuji et al., 2010

Microleter (Fig. 1) was described a few years ago (Tsuji et al. 2010) as an Early Permian parareptile (an invalid multiphyletic assembly of early reptiles). Tsuji et al. nested Microleter between millerettids and Acleistorhinus + Lanthanosuchus (another unnatural assembly).

Figure 1. Microleter in situ and reconstructed with a larger lateral temporal fenestra than originally reconstructed. The skull is 3 cm long. That's a pair of fused vomers and a left pterygoid (dorsal view) at lower right. Freehand original reconstruction by Tsuji et al. 2010 at upper left.

Figure 1. Microleter in situ and reconstructed with a larger lateral temporal fenestra than originally reconstructed. The skull is 3 cm long. That’s a pair of fused vomers and a left pterygoid (dorsal view) at lower right. Freehand original reconstruction by Tsuji et al. 2010 at upper left. Note the expansion of the quadratojugal/squamosal in the freehand drawing compared to the in situ tracing. Note the reduction of the postorbital in the freehand drawing. Note the absence of the splenial in the freehand drawing.

Character analysis
Tsuji et al. used the matrix of Modesto et al. (2009) which was based on Mülller and Tsuki (2007) consisting of 30 taxa and 137 characters. Both numbers are too small. The analysis recovered six trees in which Microleter nested in an unresolved polygamy with Australothyris and Acleistorhinus  + Lanthanosuchus at the base of the ‘ankyramorphan parareptiles’ (another unnatural assembly).

The large reptile tree (575 taxa, completely resolved) found Microleter nested between Delorhynchus and Eunotosaurus + Acleistorhinus. The clade Australothyris + Feeserpeton is the proximal outgroup. The caseasaurs and millerettids are more distant.

Figure 2. The nesting of Microleter with Delorhynchus, Acleistorhinus and Eunotosaurus.

Figure 2. The nesting of Microleter with Delorhynchus, Acleistorhinus and Eunotosaurus.

With insight Tsuji et al report, “As it is becoming increasingly clear, temporal fenestration is actually a common phenomenon among parareptiles, quite a departure for a group once termed Anapsida.”

Oddly,
Tsuji et al. include mesosaurs in their parareptilia and do not give them temporal fenestra. Oddly Tsuji et al nest Procolophon with Owenetta. Oddly they nest Eudibamus with Belebey. Oddly Tsuji et al nest Acleistorhinus with Lanthanosuchus, but not Eunotosaurus.They think the anapsid condition re-evolved in pareiasaurs. That’s not true. The ‘parareptile’ pseudoclade is a mess. It’s time for a thorough cleaning with more taxa.

Notably
the pterygoids produced a circular opening between them, as in Eunotosaurus, but not so exaggerated. Acleistorhinus does not have this trait. Here (Fig. 1), based on self-evident transfer techniques, the lateral temporal fenestra is reconstructed larger than Tsuji et al. drew it freehand. The lacrimal may not have contacted the naris according to the reconstruction where the maxilla contacts the nasal.

References
Linda A. Tsuji; Johannes Muller; Robert R. Reisz (2010). Microleter mckinzieorum gen. et sp. nov. from the Lower Permian of Oklahoma: the basalmost parareptile from Laurasia”Journal of Systematic Palaeontology 8 (2): 245–255.

The origin of Limnoscelis

Limnoscelis, according to Wikipedia, “is a genus of large (1.5 m in total length), very reptile-like diadectomorph (a type of reptile-like amphibian) from the Early Permian of North America. Contrary to other diadectomorphans, Limnoscelis appear to have been a carnivore. Though the post cranial skeleton is very similar to the early large bodied reptiles like pelycosaurs and pareiasaurs, the digits lacked claws, and the bones of the ankle bones were fused like in other reptile-like amphibians. This would not allow them to use their feet actively in traction, but rather as holdfasts, indicating Limnoscelis primarily hunted slow moving prey.”

Figure 1. Limnoscelis and two suitable ancestral taxa, Orobates and Milleretta, all shown to scale (below) and to fit (above).

Figure 1. Limnoscelis and two suitable ancestral taxa, Orobates and Milleretta, all shown to scale (below) and to fit (above).

The large reptile tree nested Limnoscelis well within the Lepidosauromorpha branch of the Reptilia/Amniota along with the smaller Orobates and not far from tiny Milleretta (Fig. 1). The latter two are the most suitable ancestral morphologies yet found on the large reptile tree.

Limnoscelis and Orobates do not nest with Diadectes and other diadectomorphs, but also, not too far away from that clade. The Limnoscelis clade still nests with Tseajiaia and Tetraceratops.

Are those carnivorous teeth in Limnoscelis?
Most sister taxa in surrounding clades are likely herbivores. Some related taxa had canines, but not Limnoscelis.

When are we going find consensus
on the nesting of Limnoscelis? We need a competing large gamut phylogenetic analysis to confirm or refute the topology recovered by the large reptile tree. Either that, or let the results of the large reptile tree get published.

Added Janurary 10, 2019
Saurorictus (Fig. 2; Late Permian; Modesto and Smith 2001; SAM PK-8666), nesting at the base of the captorhinids and their sisters, is the proximal outgroup taxon in the LRT now. Except for size, the resemblance is striking.

Figure 1. Limnoscelis and its outgroup sister, Saurorictus.

Figure 2. Limnoscelis and its outgroup sister, Saurorictus.

References
Berman DS, Reisz RR and Scott D 2010. Redescription of the skull of Limnoscelis paludis Williston (Diadectomorpha: Limnoscelidae) from the Pennsylvanian of Canon del Cobre, northern New Mexico: In: Carboniferous-Permian Transition in Canon del Cobre, Northern New Mexico, edited by Lucas, S. G., Schneider, J. W., and Spielmann, New Mexico Museum of Natural History & Science, Bulletin 49, p. 185-210.
Modesto SP and Smith RMH 2001. A new Late Permian captorhinid reptile: a first record from the South African Karoo. Journal of Vertebrate Paleontology 21(3): 405–409.
Romer AS 1946. The primitive reptile Limnoscelis restudied American Journal of Science, Vol. 244:149-188
Williston SW 1911. A new family of reptiles from the Permian of New Mexico: American Journal of Science, Series 4, 31:378-398.

wiki/Saurorictus
wiki/Limnoscelis