Turtles and pterosaurs tested again, nine years later with 1300 more taxa

In 2011,
back when the large reptile tree (LRT, 1786+ taxa) was much smaller (only about 400 taxa) I attempted a rather odd test. I wondered if turtles and pterosaurs (both Lepidosauromorpha in the LRT) would nest together while playing taxon deletion games. Here’s the 2011 link:

https://pterosaurheresies.wordpress.com/2011/07/25/pterosaurs-and-turtles-say-it-aint-so/

Click here to see those 2011 turtle-pterosaur results, still posted online.

Other workers interested in pterosaurs
(most recently Ezcurra et al. 2020) also like to play taxon deletion games as they attempt to cherry-pick preferred sisters close to dinosaurs while omitting tested and validate sisters far from dinosaurs.

The backstory
Peters 2000 added four tritosaur tanystropheid, fenestrasaur pterosaur precursors, Langobardisaurus, Cosesaurus, Sharovipteryx, and Longisquama (Fig. 1) to four previously published analyses and in every case these four nested closer to pterosaurs than any archosaur, archosauriform or archosauromorph. Unfortunately those taxa were omitted from more recent analysis, like those of Kellner 2003, Unwin 2003, Hone and Benton 2007, 2008, Bennett 2012 and Ezcurra et al. 2020.

A few years later, but still 14 years ago,
Peters 2007 added the lepidosaur, Huehuecuertzpalli and it attracted the four fenestrasaurs + pterosaurs. The LRT nested turtles within the Lepidosauromorpha here in 2011, updated here in 2014.

Now that many more taxa are present in the LRT,
let’s rerun that test and its various deletion subunits.

Today, in 2020, repeating the experiment with more taxa
deleting all lepidosauromorphs, other than turtles (and their ancestors back to Stephanospondylus) and pterosaurs, and keeping all archosauromorphs and enaliosaurs. Outgroups retained = Gephyrostegus and Silvanerpeton.

Results: Pterosaurs nest with turtles and basal sea reptiles rather than archosaurs and archosauromorphs.

Adding back all basal diapsids and protosaurs

Results: Basal diapsids as the first large clade, followed by protorosaurs with tritosaurs based on the convergence found there.

Adding back all tritosaurs
(= Macrocnemus as the last common ancestor) nests turtles and pareiasaurs as the first large clade, tritosaurs (including pterosaurs) as the next large clade, followed by archosauromorophs (including Lagerpeton and Scleromochlus).

Figure 1. Squamates, tritosaurs and fenestrasaurs in the phylogenetic lineage preceding the origin of the Pterosauria.

Deleting all non-fenestrasaur tritosaurs
(= Cosesaurus as the last common ancestor)

Results: Fenestrasaurs nest with basal diapsids. Orovenator is the proximal outgroup.

Deleting all non-pterosaur fenestrasaurs
(Bergamodactylus as the last common ancestor).

Results: Pterosaurs nest between turtles and choristoderes.

Figure 2. Basal diasids and proto-diapsids. Largely ignored these putative synapsids actually split from other synapsids while retaining the temporal fenestra trait that serves as the basis for the addition of upper temporal fenestra in diapsids. Included here are Protorothyris, Archaeovenator, Mycterosaurus, Heleosaurus, Mesenosaurus, Broomia, Milleropsis, Eudibamus, Petrolacosaurus, Spinoaequalis, and Tangasaurus.

Deleting all basal diapsids
(only turtles, pterosaurs and archosauromorphs are ingroup taxa).

Results: Pterosaurs nest between turtles and choristoderes, far from Scleromochlus, dinosaurs and Lagerpeton.

Deleting all turtle ancestors 
(= deleting Stephanospondylus through pareiasaurs)

Results: Pterosaurs nest between turtles and choristoderes, far from Scleromochlus, dinosaurs and Lagerpeton.

Figure 3. Faxinalipterus matched to Scleromochlus. The former is more primitive, like Gracilisuchus, in having shorter hind limbs and more robust fore limbs. The maxilla with fenestra and fossa, plus the teeth, are a good match.

Re-inserting terrestrial younginiforms and protorosaurs
to this last taxon list.

Results: Two large clades follow the turtle clade. Pterosaurs nest between three basal Youngina specimens and the clade Protorosauria, apart from the terrestrial younginiformes (other Youngina specimens + Pararchosauriformes (= Proterosuchus as the last common ancestor and choristoderes) and Euarchosauriformes (= Euparkeria as the last common ancestor and Lagerpeton and Scleromochlus).

Figure 4. The closest kin of Tropidosuchus are the much larger Chanaresuchus (matching Nesbitt 2011) and the smaller Lagerpeton.

Bottom line:
With or without tritosaurs and fenestrasaurs, pterosaurs prefer to nest with terrestrial younginiforms, choristoderes or turtles rather than lagerpetids, dinosaurs or Scleromochlus. Taxon exclusion remains the problem in traditional cladograms (like the recent Ezcurra et al. 2020).

Please send this post to anyone who still believes or protects
the outmoded clades ‘Ornithodira’ or ‘Avemetatarsalia’. Too many professors and their students are clinging to invalidated myths based on taxon exclusion — which is not what real scientists do. Real scientists test all competing candidates without cherry-picking or omitting taxa to suit their personal whims and traditions, in fear of their professors or colleagues.

If you would like to play taxon deletion games with the LRT,
click here, then click on the yellow CLICK HERE for LRT MacClade.nex file box with your request.

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References
Bennett SC 1996. The phylogenetic position of the Pterosauria within the Archosauromorpha. Zoological Journal of the Linnean Society 118:261-308.
Bennett SC 2012. The phylogenetic position of the Pterosauria within the Archosauromorpha re-examined. Historical Biology. iFirst article, 2012, 1–19.
Benton MJ 1983. The Triassic reptile Hyperodapedon from Elgin, functional morphology and relationships. Philosophical Transactions of the Royal Society of London, Series B, 302, 605-717.
Brusatte SL , Benton MJ , Desojo JB and Langer MC 2010. The higher-level phylogeny of Archosauria (Tetrapoda: Diapsida), Journal of Systematic Palaeontology, 8:1, 3-47.
Evans SE 1988. The early history and relationships of the Diapsida. In: M. J. BENTON (Ed.), The Phylogeny and Classificationof the Tetrapoda. 1. Amphibians, Reptiles, Birds. Systematics Symposium Association Special Volume; Oxford (Clarendon Press), 221–260.
Ezcurra MD et al. (17 co-authors) 2020. Enigmatic dinosaur precursors bridge the gap to the origin of Pterosauria. Nature (2020). https://doi.org/10.1038/s41586-020-3011-4
Hone DWE and Benton MJ 2007. An evaluation of the phylogenetic relationships of the pterosaurs to the archosauromorph reptiles. Journal of Systematic Palaeontology 5:465–469.
Hone DWE and Benton MJ 2008. Contrasting supertree and total evidence methods: the origin of the pterosaurs. Zitteliana B28:35–60.
Jalil N-E 1997. A new prolacertiform diapsid from the Triassic of North Africa and the interrelationships of the Prolacertiformes. Journal of Vertebrate Paleontology 17(3), 506-525.
Kellner AWA 2003. Pterosaur phylogeny and comments on the evolutionary history of the group. Geological Society Special Publications 217: 105-137.
Nesbitt SJ 2011. The early evolution of archosaurs: relationships and the origin of major clades. Bulletin of the American Museum of Natural History 352: 292 pp.
Peters D 2000b. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.
Peters D 2000. A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis. Rivista Italiana di Paleontologia e Stratigrafia 106 (3): 293–336.
Peters D 2007. The origin and radiation of the Pterosauria. In D. Hone ed. Flugsaurier. The Wellnhofer pterosaur meeting, 2007, Munich, Germany. p. 27.
Sereno PC 1991. Basal archosaurs: phylogenetic relationships and functional implications. Journal of Vertebrate Paleontology 11 (Supplement) Memoire 2: 1–53.
Unwin DM 2003. On the phylogeny and evolutionary history of pterosaurs. Pp. 139-190. in Buffetaut, E. & Mazin, J.-M., (eds.) (2003). Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217, London, 1-347.

https://pterosaurheresies.wordpress.com/2012/09/27/bennett-2012-still-barking-up-the-wrong-pterosaur-tree/

https://pterosaurheresies.wordpress.com/2020/12/10/new-pterosaur-precursor-study-excludes-all-pterosaur-precursors/

https://pterosaurheresies.wordpress.com/2012/04/13/a-supertree-of-pterosaur-origins-hone-and-benton-2007-2009/

Megaevolutionary dynamics in reptiles: Simoes et al. 2020

Simoes et al 2020 discuss
“rates of phenotypic evolution and disparity across broad scales of time to understand the evolutionary dynamics behind the origin of major clades, or how they relate to rates of molecular evolution.”

“Here, we provide a total evidence approach to this problem using the largest available data set on diapsid reptiles.”

Unfortunately not large enough to understand that traditional ‘diapsid’ reptiles are diphyletic, splitting in the Viséan and convergently developing two

“We find a strong decoupling between phenotypic and molecular rates of evolution,”

Yet another case of gene-trait mismatch in analysis.

“and that the origin of snakes is marked by exceptionally high evolutionary rates.”

Taxon exclusion is the reason for this exclusion.

Figure 1. Cladogram from Simoes et al. 2020. Gray tones added to show Lepidosauromorpha in the LRT.

“Here, we explore megaevolutionary dynamics on phenotypic and molecular evolution during two fundamental periods of reptile evolution: i) the origin and early diversification of the major lineages of diapsid reptiles (lizards, snakes, tuataras, turtles, archosaurs, marine reptiles, among others) during the Permian and Triassic periods,”

In the LRT the new archosauromorphs split from new lepidosauromorphs in the Viséan (Early Carboniferous).

“as the origin and evolution of lepidosaurs (lizards, snakes and tuataras) from the Jurassic to the present.”

In the LRT lepidosaurs had their origin in the Permian and the Simoes team ignores the Triassic radiation of lepidosaurs leading to tanystropheids and pterosaurs.

So without a proper and valid phylogenetic context,
why continue? How can they possibly discuss ‘rates of change’ if they do not include basal taxa from earlier period?

“Our results indicating exceptionally high phenotypic evolutionary rates at the origin of snakes further suggest that snakes not only possess a distinctive morphology within reptiles,  but also that the first steps towards the acquisition of the snake body plan was extremely fast.”

In the LRT many taxa are included in the origin of snakes from basal geckos. These are missing from Simoes list of snake ancestor.

Figure 1.  Subset of the LRT focusing on lepidosaurs and snakes are among the squamates.

In the LRT all sister taxa resemble one another
and document a gradual accumulation of derived traits.

If you have any particular evolutionary questions,
they were probably answered earlier in previous posts. Use the keyword box at upper right to seek your answer.

 

Early Triassic Elessaurus: another overlooked Cosesaurus sister!

Cosesaurs (basal fenestrasaurs)
(Figs. 1–3) are popping up everywhere lately!

Figure 1. MIddle Triassic Cosesaurus. Double its size and it would be close to Early Triassic Elessaurus. See figure 6.

Earlier we looked at a tiny Early Cretaceous cosesaur skull in amber
originally mistaken for a bird/dinosaur: Oculudentavis.

Figure 2. Elessaurus hind limb elements gathered together to scale. Some original scale bars were off by 2x.

Today De-Oliveira et al. 2020 bring us
a new Early Triassic cosesaur, Elessaurus gondwanoccidens (UFSM 11471, Figs. 2, 3) known from a hind limb, pelvis, partial sacrum and proximal caudal vertebrae. Cosesaurus was a derived tanystropheid, close to Langobardisaurus, but was not included in the De-Oliveira taxon list.

Figure 3. Elessarus pes compared to Cosesaurus to scale and x2. Note differences between original tracing and DGS tracing. Digit 5 was not lost. It is tucked beneath the metatarsals and was not scored in the LRT.

In the De-Oliveira et al. published cladogram
(Fig. 4) Elessaurus nests basal to the Tanystropheidae (= Macrocnemus, Amotosaurus, Tanystropheus, Tanytrachelos and Langobardisaurus) a clade they derive from Protorosaurus and Trilophosaurus. in one cladogram (Fig. 4), but not in the other (Fig. 5). The authors reported, “In addition, the new specimen presents some features only found in more specialized representatives within Tanystropheidae, such as the presence of a well-developed calcaneal tuber with a rough lateral margin.” 

Figure 4. Published cladogram by De-Oliveira et al. 2020. Note different nesting than their SuppData cladogram in figure 5.

By contrast, in the De-Olveira et al. SuppData cladogram
Elessaurus nests without resolution within the Rhynchosauria (Fig. 5). Distinct from the first analysis (Fig. 4), this (Fig. 5) included the drepanosaur ancestor, Jesairosaurus, and the derived macrocnemid, Dinocephalosaurus. The authors reported, “In this second analysis, Elessaurus adopts different positions among the MPTs, it is recovered, e.g. within Archosauriformes, as a sister-taxa of Allokotosauria+Archosauriformes and an early rhynchosaur.” This is a red flag indicating major problems in scoring and taxon exclusion.

Figure 5. Cladogram from DeOliveira et al. 2020 with colors added to show distribution and mixing of Lepidosauromorpha and Archosauromorpha clades in the LRT. Many of these purported sister taxa do not look alike! Strangely, here Elessaurus nests with rhynchosaurs, not tanystropheids (Fig. 2). The taxon in pink had to be looked up and revised here.

By contrast, 
in the large reptile tree (LRT, 1661+ taxa), Elessaurus nests as a derived tanystropheid, alongside Cosesaurus (Fig. 1). a smaller Middle Triassic taxon omitted from the original study. The authors mistakenly considered tanystropheids to be archosauromorphs, again due to taxon exclusion. Despite the many traits that converged with archosauromorph protorosaurs, tanystropheids are tritosaur lepidosaurs, derived from Huehuecuetzpalli (Fig. 7) and Tijubina. In the LRT, adding taxa does not create chaos. New taxa neatly take their place within the current tree topology. By omitting Cosesaurus, the authors omitted the most similar taxon to Elessaurus and all the added information included herein.

The cladogram by De-Oliveira et al. shuffles lepidosauromorphs
with archosauromorphs without an understanding of their Viséan split. As a result, several purported ‘sisters’ in figure 5 do not look alike, but apparently nested together by default. Based on these ‘odd bedfellows’ I suspect the authors borrowed another worker’s cladogram without checking scores or examining results.

From the abstract:
“The origin and early radiation of Tanystropheidae, however, remains elusive.”

This is false. We know the origin of Tanystropheidae back to Cambrian chordates. Taxon exclusion by the authors prevents them from recovering both distant and proximal sister taxa.

“Here, a new Early Triassic archosauromorph is described and phylogenetically recovered as the sister-taxon of Tanystropheidae.”

By contrast, in the LRT Elessaurus is a derived fenestrasaur close to Cosesaurus, a taxon excluded by the authors. We know cosesaur tracks (= Rotodactylus, Fig. 6) go back to the Early Triassic and some were much larger than Cosesaurus.

Figure 6. Click to enlarge. Scaling a quadrupedal Cosesaurus to the larger Rotodactylus tracks from Haubold 1983. Quadrant represents center of balance in the closeup foot. Graphic representation of a butt joint is nearby.

More from the abstract:
“The new specimen, considered a new genus and species, comprises a complete posterior limb articulated with pelvic elements. It was recovered from the Sanga do Cabral Formation (Sanga do Cabral Supersequence, Lower Triassic of the Parana Basin, Southern Brazil), which has already yielded a typical Early Triassic vertebrate assemblage of temnospondyls, procolophonoids, and scarce archosauromorph remains. This new taxon provides insights on the early diversification of tanystropheids and represents further evidence for a premature wide geographical distribution of this clade. The morphology of the new specimen is consistent with a terrestrial lifestyle, suggesting that this condition was plesiomorphic for Tanystropheidae.”

Likewise, Cosesaurus and related fenestrasaurs in the LRT are terrestrial taxa, distinct from other tanystropheids, all arising from tritosaur lepidosaurs like Tijubina and Huehuecuetzpalli.

Figure 7. The father of all pterosaurs and tanystropheids, Huehuecuetzpalli, a late survivor in the Early Cretaceous from a Late Permian radiation.

Larger quadrupedal cosesaurs, 
like Elessaurus, had two sacrals (Fig. 1). Smaller bipedal cosesaurs (Fig. 1) had four. Both had anterior processes on the ilium, not longer than the acetabulum width, distinct from non-fenestrasaur tanystropheids.

PS
Figure 2 in De-Oliveira has a scale bar problem in their figure 2 (explained here in Fig. 2).

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References
De-Oliveira TM, Pinheiro FL, Da-Rosa AAS, Dias-Da-Silva S and Kerber L 2020.
A new archosauromorph from South America provides insights on the early diversification of tanystropheids. PLoS ONE 15(4): e0230890

Oculudentavis in more incredible detail! (thanks to Li et al. 2020)

Li et al. 2020 bring us
higher resolution scans of the putative tiny toothed ‘bird’ (according to Xing et al. 2020) Oculudentavis (Fig. 1). Following a trend started here a week ago, Li et al. support a generalized lepidosaur interpretation, but then tragically overlook/deny details readily observed in their own data (Fig.1).

FIgure 1. CT scan model from Li et al. 2020, who denied the presence of a quadratojugal and an antorbital fenestra, both of which are present. Colors applied here. The previously overlooked jugal-lacrimnal suture becomes apparent at this scale and presentation.

Li et al. deny the presence of a clearly visible antorbital fenestra.
They report, “One of the most bizarre characters is the absence of an antorbital fenestra. Xing et al. argued the antorbital fenestra fused with the orbit, but they reported the lacrimal is present at the anterior margin of the orbit. This contradicts the definition of the lacrimal in birds, where the lacrimal is the bone between the orbit and antorbital, fenestra. In addition, a separate antorbital fenestra is a stable character among archosaurs including non-avian dinosaurs and birds, and all the known Cretaceous birds do have a separate antorbital fenestra.”

Contra Li et al.
a standard, ordinary antorbital fenestra is present (Fig. 1 dark arrow) and the lacrimal is between the orbit and antorbital fenestra. This is also the description of the antorbital fenestra and fenestrasaurs, like Cosesaurus (Fig. 2), Sharovipteryx and pterosaurs (Peters 2000).

Li et al. report,
“The ventral margin of the orbit is formed by the jugal.”

Contra Li et al.
the lacrimal is ventral to half the orbit (Fig. 1). The jugal is the other half. The suture becomes visible at the new magnification.

Li et al. report,
“Another unambiguous squamate synapomorphy in Oculudentavis is the loss of the lower temporal bar.” 

Contra Li et al.
the lower temporal bar is created by the quadratojugal, as in Cosesaurus, Sharovipteryx and pterosaurs. In Oculudentavis the fragile and extremely tiny quadratojugal is broken into several pieces. DGS (coloring the bones) enables the identification of those pieces (Fig. 1).

Figure 2. Cosesaurus with colors applied. Compare to figure 1.

Li et al. conclude,
“Our new morphological discoveries suggest that lepidosaurs should be included in the phylogenetic analysis of Oculudentavis.” 

Contra Li et al.
these are all false ‘discoveries’.

Also note that Li et al. cannot discern
which sort of lepidosaurs should be tested in the next phylogenetic analysis of Oculudentavis. That’s because lepidosaur tritosaur fenestrasaurs, like Cosesaurus (Fig. 2), are not on their radar. That’s because pterosaur referees have worked to suppress the publication of new data on Cosesaurus and kin. And that’s what scientists get for not ‘playing it straight.’

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References
Li Z, Wang W, Hu H, Wang M, Y H and Lu J 2020. Is Oculudentavis a bird or even archosaur? bioRxiv (preprint) doi: https://doi.org/10.1101/2020.03.16.993949
Xing L, O’Connor JK,; Schmitz L, Chiappe LM, McKellar RC, Yi Q and Li G 2020. Hummingbird-sized dinosaur from the Cretaceous period of Myanmar. Nature. 579 (7798): 245–249.

wiki/Oculudentavis

 

Was Vellbergia really a juvenile basal lepidosaur? Let’s check…

Earlier we looked at tiny Vellbergia
(Sobral, Simoes and Schoch 2020; Middle Triassic) represented by a disarticulated tiny skull (Fig. 1). The large reptile tree (LRT) nested this hatchling with the much larger adult Prolacerta (Fig. 1). The MPT was 20263 steps for 1654 taxa.

The LRT nesting ran counter to the SuppData cladogram
of Sobral, Simoes and Schoch 2020, who nested Vellbergia among basal lepidosaurs, the closest of which are shown here (Fig. 1). Earlier I did not show the competing lepidosaur candidates. That was an oversight rectified today.

Figure 1. Vellbegia compared to the lepidosaurs it would nest with if Prolacerta and all Archosauromorpha were deleted. Gray areas on Vellbergia indicate restored bone that is lost in the fossil.

To test the lepidosaur hypothesis of relationships,
I deleted all Archosauromorph taxa, including Prolacerta, from the LRT to see where among the Lepidosauromorpha Vellbergia would nest. With no loss of resolution, Vellbergia nested between Palaegama and Tjubina + Huehuecuetzpalli at the base of the Tritosauria plus Fraxinisaura + Lacertulus (Fig. 1) at the base of the Protosquamata. The resulting MPT was 20276 steps, only 13 more than the Prolacerta hypothesis of interrelationships.

That is a remarkably small number considering the great phylogenetic distance between these taxa in the LRT.

Rampant convergence
is readily visible among the competing taxa (Fig. 1). No wonder Prolacerta was named “before Lacerta“, the extant squamate. According to Wikipedia, “Due to its small size and lizard-like appearance, Parrington (1935) subsequently placed Prolacerta between basal younginids and modern lizards. In the 1970s (Gow 1975) the close link between Prolacerta and crown archosaurs was first hypothesized.” That was prior to cladistic software and suffered from massive taxon exclusion.

Allometry vs. Isometry
One of the lepidosaurs shown above, Huehuecuetzpalli (Fig. 1), is known from both an adult and juvenile. The older and younger specimens were originally (Reynoso 1998) considered identical in proportion. Such isometry is an ontogenetic trait shared with other tritosaur lepidosaur clade members, including pterosaurs. On the other hand, if Vellbergia was a hatchling of Prolacerta, some measure of typical archosauromorph allometry should be readily apparent… and it is… including incomplete ossification of the nasals, frontals and parietals along with a relatively larger orbit and shorter rostrum, giving Vellbergia a traditional ‘cute’ appearance appropriate for its clade.

Size
Sobral, Simoes and Schoch considered Vellbergia a juvenile, but it is similar in size to the adult lepidosaurs shown here (Fig. 1). On the other hand, Vellbergia is appropriately smaller than Prolacerta, in line with its hatchling status.

Time
Remember also that Vellbergia is from the Middle Triassic. Prolacerta is from the Early Triassic. They were not found together and some differences are to be expected just from the millions of years separating them.

For comparison: another juvenile Prolacerta,
this time from Early Triassic Antarctica (Spiekman 2018; AMNH 9520), is much larger than Vellbergia from Middle Triassic Germany (Fig. 2), but just as cute. Note the relatively larger orbit and shorter rostrum compared to the adult Prolacerta (Fig. 1), traits likewise found in Vellbergia.

Figure 2. Small Prolacerta specimen AMNH 9520 from Spiekman 2018 compared to scale with Vellbergia. Sclerotic rings (SCL) identified by Spiekman 2018 are re-identified as pterygoids here.

Generally
crushed, disarticulated and incomplete juvenile specimens of allometric taxa are difficult to compare with adults. Even so, what is left of hatchling Vellbergia tends to resemble the larger juvenile and adult specimens of Prolacerta more than hatchling Vellbergia resembles the similarly-sized adult lepidosaurs it nests with in the absence of Prolacerta from the taxon list.

Phylogenetic analysis is an inexact science.
Nevertheless no other known method breaks down and rebuilds thousands of taxa more precisely. Only taxon exclusion appears to trip up workers at present.

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References
Gow CE 1975. The morphology and relationships of Youngina capensis Broom and Prolacerta broomi Parrington. Palaeontologia Africana, 18:89-131.
Parrington FR 1935. On Prolacerta broomi gen. et sp. nov. and the origin of lizards. Annals and Magazine of Natural History 16, 197–205.
Reynoso V-H 1998. Huehuecuetzpalli mixtecus gen. et sp. nov: a basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodríguez, Central México. Philosophical Transactions of the Royal Society, London B 353:477-500.
Sobral G, Simoes TR and Schoch RR 2020. A tiny new Middle Triassic stem-lepidosauromorph from Germany: implications fro the early evolution of lepidosauromorphs and the Vellberg fauna. Nature.com Scientific Reports 10, Article number: 2273.
Spiekman SNF 2018. A new specimen of Prolacerta broomi from the lower Fremouw Formation (Early Triassic) of Antarctica, its biogeographical implications and a taxonomic revision. Nature.com/scientificreports (2018)8:17996

wiki/Prolacerta

Carbonodraco enters the LRT alongside Kudnu

Mann et al. 2019 bring us new taxon
Carbonodraco lundi (Fig. 1; CM 23055; Middle Pennsylvanian, Moscovian), formerly considered a type of Cephalerpeton ventriarmatum (Moodie 1912), a basal lepidosauromorph reptile close to captorhinids. The authors considered their find the oldest (Middle Pennsylvanian, Moscovian) parareptile, an invalid and paraphyletic clade in the large reptile tree (LRT, 1613 taxa).

Mann et al. start with a redescription of Cephalerpeton,
a taxon needing no such redescription. This illustrates the narrow focus of these workers and paleontologists in general. They continue to cherry-pick taxa and use freehand illustrations of reconstructions to support their views. The bones in their reconstruction do not match their tracing of the in situ bones and several bones were left unidentified.

In order to decide what a new taxon is or isn’t
there is only one way to do it. You need to add your taxon to a wide gamut phylogenetic analysis, like the LRT. Then let the software tell you what you have. The LRT minimizes bias and taxon exclusion by including such a wide gamut and large number of taxa. Mann et al. did not mention a long list of pertinent taxa in their study.

Figure 1. Carbonodraco in situ (lower right), as originally reconstructed freehand (upper right) and using DGS methods to reconstruct the skull by coloring the bone drawing. Based on the drawings autapomorphic upper temporal fenestrae appear to be present, convergent with other clades in which this occurred.

Mann et al.
considered Carbonodraco a member of the Acleistorhinidae. That’s why they gave it a hypothetical lateral temporal fenestra, misidentified the prefrontal, did not identify the squamosal, quadratojugal, squamosal, supratemporal and postparietal in their freehand drawing. The DGS method minimizes such bias-generated errors by coloring the bones and moving them, ‘as is’, to the reconstruction (Fig. 1).

By contrast
the LRT nested Carbonodraco with Kudnu (Bartholomai 1979; Figs. 2, 3). These two nest at the base of the Pareiasauria + turtles, taxa not included in Mann et al. Both Cephalerpeton and Acleistorhinus are taxa included in the LRT and both nest far from Carbonodraco.

Figure 2. Carbonodraco enters the LRT alongside another recent addition, Kudnu, at the base of the pareiasaurs + turtles.Figure 1. Carbonodraco enters the LRT alongside another recent addition, Kudnu, at the base of the pareiasaurs + turtles.

Mann et al. are working from an old and invalidated hypothesis
when they report, “Amniotes can be divided into two major lineages, synapsids (mammals and their extinct relatives) and reptiles (crocodiles, birds, and lepidosaurs, and their eAmniotes can be divided into two major lineages, synapsids (mammals and their extinct relatives) and reptiles (crocodiles, birds, and lepidosaurs, and their extinct relatives).”

No, the LRT indicates the first dichotomy splits Archosauromorpha (including Synapsida) from Lepidosauromorpha.

Figure 3. Kudnu colorized using DGS and slight restored postcranially, shown 10x natural size at a 72 dpi standard screen resolution. Here’s a taxon basal to Stephanospondylus, pareiasaurs and turtles. Prior workers excluded Stephanospondylus from their studies.

“The origin and early diversification of these groups are believed to have occurred during the early Carboniferous because the oldest amniotes, the reptile Hylonomus lyelli and the putative synapsid Protoclepsydrops haplous, are known from the classic locality of Joggins, Nova Scotia, Canada (313–316 Ma).”

No, the LRT indicates Silvanerpeton (Viséan, 335 mya) is the last common ancestor of all amniotes (= Reptilia).

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References
Mann A, McDaniel EJ, McColville ER and Maddin HC 2019. Carbonodraco lundi gen et sp. nov., the oldest parareptile, from Linton, Ohio, and new insights into the early radiation of reptiles. Royal Society Open Science, 27 November 2019. https://royalsocietypublishing.org/doi/10.1098/rsos.191191#d3e1049

Excellent YouTube Video on Mosasaurs

This one features paleoartist, Brian Engh
working with the Mike Triebold’s fossil company (Triebold Paleontology, Inc).

Sadly
Triebold’s Pteranodons are leaping on their forelimbs and their Jeholopterus has giant scleral rings (eyes) in the antorbitral fenestra. Both were based on my originals, which did not have these invalid modifications. Those are market forces at work, veering away from evidence.

References
DontMessWithDinosaurs.com
TrieboldPaleontology.com

The most basal lepidosauriforms and lepidosaurs to scale

Lepidosauriform fossils are extremely rare in the Mesozoic and Paleozoic.
In the Earliest Permian we find Tridentinosaurus (Fig. 1; Dal Piaz 1931,1932; Leonardi 1959), a taxon ancestral to the pseudo-rib-gliders of the Late Permian (Coelurosauravus) through the Early Cretaceous (Xianlong) and close to the origin of all other lepidosauriforms, including living snakes, lizards and the tuatara (genus: Sphenodon).

Figure 1. Basal lepidosauriformes to scale from Tridentinosaurus (Earliest Permian) to Huehuecuetzpalli (Early Cretaceous). Subtle differences lump and split these taxa into their various clades.

 

Sometime during the Early Permian
the Lepidosauria split between the Sphenodontia + Drepanosauria and the Tritosauria + Protosquamata in the large reptile tree (LRT, 1381 taxa).

Short-legged
Jesairosaurus, in the Early Triassic, nests basal to the clade of slow-moving, arboreal drepanosaurs. On another branch, Megachirella (Middle Triassic) and Gephyrosaurus (Early Jurassic) are basal members of the Sphenodontia.

Long-legged
and probably arboreal Saurosternon and Palaegama, (both Late Permian) are the earliest known Lepidosauria, but they are basal to the Tritosauria + Protosquamata clades.

Figure 5. Subset of the LRT focusing on the Tritosauria. Note the separation of one specimen attributed to Macrocnemus.

Late-surviving, long-legged basal Tritosauria
include tiny Tijubina and Huehuecuetzpalli (both Early Cretaceous). This clade gave rise to giant Tanystropheus, exotic Longisquama and volant Pteranodon.

Tiny and long-legged Late Permian
Lacertulus is the basal taxon in the previously unrecognized clade Protosquamata, the parent clade to the extant Squamata. This taxon documents the antiquity of this clade.

Going back to the Early Permian
we have a long-torso, short-legged specimen, MNC TA-1045, that nests in the LRT just outside the extant Squamata (Iguana). MNC TA-1045 was found alongside the genus Ascendonanus (MNC-TA0924), a basal archosauromorph diapsid with a shorter torso you can see here. The MNC TA-1045 specimen pushes the genesis of the lepidosaurs back to the Early Permian, nearly coeval with the basalmost lepidosauriform shown in figure 1, Tridentinosaurus.

The Lepidosauromorph-Archosauromorph dichotomy
was already present in the Viséan (Early Carboniferous, 330 mya), so the new Lepidosauromorpha had 30 million years to diverge into captorhinomorphs, diadectomorphs, millerettids and lepidosauriforms by the time Tridentinosaurus first appears in the Earliest Permian (300 mya).

Late surviving,
but basalmost lepidosauromorphs include Sophineta , Paliguana and Coletta (all Early Triassic). These taxa have an upper temporal fenestra not seen in outgroup taxa.

Proximal outgroups for the Lepidosauriforms
include the late-surviving owenettids: Barasaurus (Late Permian) and kin, Owenetta (Late Permian) and kin, and the late-surviving macroleterids (Middle Permian) and nycteroleterids (Middle Permian) before them.

At least that’s what the data says so far.
With every new taxon the tree grows stronger and more precise, so the odds of changing the tree topology with additional taxa continue to drop. Looking forward to seeing more Paleozoic arboreal lepidosauromorph discoveries as they arrive.

References
Dal Piaz Gb. 1932 (1931). Scoperta degli avanzi di un rettile (lacertide) nei tufi compresi entro i porfidi quarziferi permiani del Trentino. Atti Soc. Ital. Progr. Scienze, XX Riunione, v. 2, pp. 280-281. [The discovery of the remains of a reptile (lacertide) in tuffs including within the Permian quartz porphyry of Trentino.]
Leonardi P 1959. Tridentinosaurus antiquus Gb. Dal Piaz, rettile protorosauro permiano del Trentino orientale. Memorie di Scienze Geologiche 21: 3–15.

www.reptileevolution.com/reptile-tree.htm

 

 

 

 

Miscellaneous notes #1: Limnoscelis and Saurorictus

A few loose ends
from the large reptile tree (LRT) that I failed to mention earlier:

#1
Saurorictus (Fig. 1; Late Permian; Modesto and Smith 2001; SAM PK-8666), nesting at the base of the captorhinids and their basal lepidosauromorph sisters, is the proximal outgroup taxon in the LRT for Limnoscelis. Except for size, the resemblance is striking and, so far, unreported in traditional paleontology.

Sadly,
Wikipedia still thinks of Limnoscelis as a “reptile-like diadectomorph (a type of reptile-like amphibian).” Romer 1946 and Willistion 1911 pegged it as a reptile.

Figure 1. Limnoscelis and its outgroup sister, Saurorictus.

While we’re on the subject of the Limnoscelidae
the 2011 nesting of Tetraceratops (Fig. 2) with Tseajaia and Limnoscelis in the LRT has not been challenged. Relying on taxon exclusion, Tetraceratops was nested as a basal therapsid most recently by Amson and Laurin 2011 and earlier by other studies that excluded other clades. The thin squamosal broken into smaller pieces that drifted into the orbit were overlooked and ignored by earlier studies. The convergence is interesting, though. Spindler 2014 did question the nesting of Tetraceratops as a basal therapsid, but only on the basis of a trait list, not by expanding their taxon list to Tseajaia and its sisters. 

Figure 2. Click to Enlarge. The skull of Tetraceratops (middle column) compared to candidate sisters, Limnoscelis and Tseajaia (left) and Haptodus and Biarmosuchus (right). In the boxed area are other interpretations of Tetraceratops by Matthew 1908 and Amson and Laurin 2011.

 

Limnoscelis 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: Pp. 185-210 in: Carboniferous-Permian Transition in Canon del Cobre, Northern New Mexico, edited by Lucas, Schneider and Spielmann, New Mexico Museum of Natural History & Science, Bulletin 49.
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

Tetraceratops References
Amson E and Laurin M 2011. On the affinities of Tetraceratops insignis, an Early Permian synapsid. Acta Palaeontologica Polonica 56(2):301-312. online pdf 
Conrad J and Sidor CA 2001. Re−evaluation of Tetraceratops insignis (Synapsida: Sphenacodontia). Journal of Vertebrate Paleontology 21: 42A.
Matthew WD 1908. A four-horned pelycosaurian from the Permian of Texas.
Bulletin of the American Museum of Natural History 24:183-185.
Laurin M and Reisz RR. 1996. The osteology and relationships of Tetraceratops insignis, the oldest known therapsid. Journal of Vertebrate Paleontology 16:95-102. doi:10.1080/02724634.1996.10011287.\
Sidor CA and Hopson JA 1998. “Ghost lineages and “mammalness”: Assessing the temporal pattern of character acquisition in the Synapsida”. Paleobiology 24: 254–273.
Spindler F 2014. Reviewing the question of the oldest therapsid. Paläontologie, Stratigraphie, Fazies (22) Freiberger Forschungshefte C 548: 1–7.

wiki/Tetraceratops

 

Enigmatic Teraterpeton understood, at last, with better data

Finally
some photographic skull material has appeared online for Teraterpeton (Fig. 1). Not sure when these first appeared. Could have been years ago. I have not been searching until a day or two ago (see below).

First added
to the large reptile tree (LRT, 1371 taxa) on the basis of drawings by Sues 2003, the long rostrum and antorbital fenestra + the infilling of the lateral temporal fenestra of Teraterpeton  are traits that don’t go together anywhere else on anyone’s cladogram. Sues considered Teraterpeton an archosauromorph nesting with short-snouted Trilophosaurus (Figs. 1, 3).

In the LRT
Trilophosaurus is a rhynchocephalian lepidosaur nesting between derived sphenodontids, like short-snouted Sapheosaurus, and primitive rhynchosaurs, like short-snouted Mesosuchus. All related taxa have a diapsid-like temple architecture, even though the new clade Diapsida (Petrolacosaurus and kin) is restricted to members of the Archosauromorpha in the LRT. Lepidosaurs with a diapsid architecture have their own clade name: “Lepidosauriformes.” Details here and here.

The latest thinking identifies the large hole in the rostrum of Teraterpeton
that extends nearly to the orbit as a naris alone, not a combination of naris + antorbital fenestra. Here (Fig. 1) a broken strut-like bone lying atop the slender maxilla appears to have separated a naris from an antorbital fenestra in vivo. Even so, the present scoring for Teraterpeton with an antorbital fenestra without a fossa does not nest it with other taxa having an antorbital fenestra with or without a fossa.

Figure 1. Skulls of Teraterpeton and Trilophosaurus compare well aft of the orbit, not so much below the orbit or in the rostrum.

This would not be the first time
an antorbital fenestra appeared in a lepidosaur. Pterosaurs and their ancestors, the fenestrasaurs, also have this trait by convergence with several other tetrapod taxa.

Comparing Trilophosaurus to Teraterpeton
(Fig. 1) is difficult until you get to the postorbital region of the skull. Then it’s a good match. Trilophosaurus has a reduced rostrum, a small naris, a robust maxilla and no hint of an antorbital fenestra. But like Teraterpeton alone, the lateral temporal fenestra found in all related taxa, is infilled with a large flange of the quadrate. Even so, the crappy character list for the LRT is able to nest these two taxa together.

Trilophosaurus and Teraterpeton nest with
the distinctively different Shringasaurus and Azendohsaurus in the LRT. The large variety in their morphologies hints at a huge variation yet to be found here. This is yet one more case where a list of traits may fail you, but a suite of several hundred traits will eliminate all other possibilities by a statistical process known as maximum parsimony. While the parsimony is minimal in this clade, it is still more than any other candidate taxa can offer from a list that has grown to over 1300.

A recent abstract on Teraterpeton
by Pritchard and Sues 2016 bears a review.

“Teraterpeton hrynewichorum, from the Upper Triassic (Carnian) Wolfville Formation of Nova Scotia, is one of the more unusual early archosauromorphs, with an elongate edentulous snout, transversely broadened and cusped teeth, and a closed lateral temporal fenestra. Initial phylogenetic analyses recovered this species as the sister taxon to Trilophosaurus spp. New material of Teraterpeton includes the first-known complete pelvic girdle and hind limbs and the proximal portion of the tail. These bones differ radically from those in Trilophosaurus, and present a striking mosaic of anatomical features for an early saurian.”

I agree completely, which makes solving this mystery so intriguing, and one perfectly suited to the wide gamut of the LRT.

The ilium has an elongate, dorsoventrally tall anterior process similar to that of hyperodapedontine rhynchosaurs. 

In all cladograms trilophosaurs are close to rhynchosaurs.

The pelvis has a well-developed thyroid fenestra, a feature shared by Tanystropheidae, Kuehneosauridae, and Lepidosauria. 

These taxa all nest within the Lepidosauriformes in the LRT. Mystery solved.

Figure 2. Azendohsaurus skull reconstructed with two premaxillary teeth, not four.

“The calcaneum is ventrally concave, as in Azendohsaurus”. 

Teteraterpeton and Trilophosaurus nest as sisters to Azendohsaurus (Fig. 2)  in the LRT.

The fifth metatarsal is proximodistally short, comparable to the condition in Tanystropheidae.” 

This condition is also found in lepidosaur tritosaur fenestrasaurs, including pterosaurs. Tanystropheidae nest as tritosaurs in the LRT. Mystery solved.

“Much as in the manus, the pedal unguals of Teraterpeton are transversely flattened and dorsoventrally deep.” 

The unguals of Trilophosaurus are also exceptionally deep and transversely flat.

“Phylogenetic analysis of 57 taxa of Permo-Triassic diapsids and 315 characters supports the placement of Teraterpeton as the sister-taxon of Trilophosaurus in a clade that also includes Azendohsauridae and, rather unexpectedly, Kuehneosauridae.”

Add taxa and the unexpected kuehneosaurs will drift to a more basal node.

“The mosaic condition in Teraterpeton underscores the importance of thorough taxon sampling for understanding the dynamics of character change in Triassic reptiles and the use of apomorphies in identifying fragmentary fossils.”

The term ‘mosaic’ is misleading. In the LRT there are no closer sisters to Teraterpeton than Trilophosaurus, and then, rather obviously, the similarities are immediately obvious only in the cheek region, distinct from all other taxa in the LRT.

Figure 3. Trilophosaurus has filled in the lateral temporal fenestra, reduced the orbit and increased the upper temporal fenestra, among other differences with Azendohsaurus.

Forcing Teraterpeton
back to long-snouted clades with an antorbital fenestra, like the Diandongosuchus clade, adds a minimum of 11 extra steps to the LRT.

Key to understanding
the lepidosaur nature of these taxa involves first understanding that the first dichotomy in the clade Reptilia separates the new Archosauromorpha from the new Lepidosauromorpha. Until someone else does this and it becomes consensus, we will continue to experience the confusion exhibited by Pritchard and Sues 2016 (above). This has been documented online for the last seven years.

References
Pritchard AC, Sues H-D 2016. Mosaic evolution of the early saurian post cranium revealed by the postcranial skeleton of Teraterpeton hrynewichorum (Archosauromorpha, Late Triassic). Abstract from the 2016 meeting of the Society of Vertebrate Paleontology.
Sues H-D 2003. An unusual new archosauromorph reptile from the Upper Triassic Wolfville Formation of Nova Scotia. Canadian. Journal of Earth Science 40(4): 635-649.

Thanks to
reader NP for bringing this taxon back to my attention.