A deeper extension for the Lepidosauria

By definition
the Lepidosauria includes Rhynchocelphalia (Sphenodon), Squamata (Iguana), their last common ancestor and all descendants. By this definition pterosaurs and kin are lepidosaurs because they nest between rhychocephalians and iguanids in a traditionally unrecognized clade the Tritosauria (Fig. 1).

Figure 1. Subset of the LRT focusing on the Lepidosauria. Now the drepanosaur clade lumps with the rhynchocephalians in the crown group. Extant lepidosaurs are in gray.

Figure 1. Subset of the LRT focusing on the Lepidosauria. Now the drepanosaur clade lumps with the rhynchocephalians in the crown group. Extant lepidosaurs are in gray.

While reviewing
the large reptile tree (LRT, 1087 taxa, subset Fig. 1) following the addition of Avicranium, the base of the Rhynchocephalia  shifted back to include Jesairosaurus, and the drepanosaursSaurosternon and Palaegama, which formerly nested as outgroup Lepidosauriformes now nest basal to the tritosaurs, pro-squamates and squamates within the Lepidosauria, based on the traditional definition.

With this change
the non-lepidosaur Lepidosauriformes are reduced to just the glider clade, Coletta, Paliguana, and Sophineta, taxa with a diapsid skull architecture. These remain stem lepidosaurs. The membership of the clade Lepidosauriformes do not change.

Remember,
despite their diapsid temporal morphology, these are not members of the clade Diapsida, which is restricted to Archosauromorph ‘diapsids’ only. Petrolacosaurus is a basal member of the monophyletic Diapsida. The clade name ‘Lepidosauriformes’ includes all lepidosauromorphs with upper and lateral temporal fenestrae. If you know any traditional paleontologists who still think lepidosaurs are related to archosaurs, please show them the LRT.

Once a definition for a clade is made
then the next step is to see which taxa fall under than definition… and then to see if that definition is a junior synonym for a previously published definition based on clade membership. Remember, traditional traits may not give you monophyly, but phylogenetic analysis always will.

And
yes, I do review all the scores in the LRT and announce updates when they are made.

 

Shringasaurus: new rhynchocephalian lepidosaur with horns

Sengupta, Ezcurra and Bandyopadhyay 2017 bring us
a new, very large, horned rhynchocephalian lepidosaur, Shringasaurus (Fig. 1). Unfortunately, that’s not how the Sengupta team nested it (due to the sin of taxon exclusion, see below). Even so, there is consensus that the new taxon is closely related to the much smaller Azendohsaurus (Fig. 1).

Figure 1. Shringasaurus to scale with Azendohsaurus. Line art modified from Sengupta et al. Color added here. Note the anterior lappet of the maxilla over the premaxilla. The supratemporal  (dark green) remains.

Figure 1. Shringasaurus to scale with Azendohsaurus. Line art modified from Sengupta et al. Color added here. Note the anterior lappet of the maxilla over the premaxilla. The supratemporal  (dark green) remains.

From the abstract:
“The early evolution of archosauromorphs (bird- and crocodile-line archosaurs and stem-archosaurs) represents an important case of adaptive radiation that occurred in the aftermath of the Permo-Triassic mass extinction. Here we enrich the early archosauromorph record with the description of a moderately large (3–4 m in total length), herbivorous new allokotosaurian, Shringasaurus indicus, from the early Middle Triassic of India. The most striking feature of Shringasaurus indicus is the presence of a pair of large supraorbital horns that resemble those of some ceratopsid dinosaurs. The presence of horns in the new species is dimorphic and, as occurs in horned extant bovid mammals, these structures were probably sexually selected and used as weapons in intraspecific combats. The relatively large size and unusual anatomy of Shringasaurus indicus broadens the morphological diversity of Early–Middle Triassic tetrapods and complements the understanding of the evolutionary mechanisms involved in the early archosauromorph diversification.”

Allokotosauria
Shringasaurus was nested in the clade, Allokotosauria, According to Wikipedia, “Nesbitt et al. (2015) defined the group as a  containing Azendohsaurus madagaskarensis and Trilophosaurus buettneri and all taxa more closely related to them than to Tanystropheus longobardicus, Proterosuchus fergusi, Protorosaurus speneri or Rhynchosaurus articeps.” This definition was based on the invalidated hypothesis that rhynchosaurs and allokotosaurs were close to the base of the Archosauriformes as the addition of more taxa will demonstrate. Basically this clade equals Trilophosaurus, Azendohsaurus and now Shringasaurus. In the large reptile tree (LRT, 1049 taxa) this clade nests between Sapheosaurus + Notesuchus and Mesosuchus + Rhynchosauria all nesting within Sphenodontia (=  Rhynchocephalia), so they are all lepidosaurs. All you have to do is add pertinent taxa to make this happen in your own phylogenetic analysis.

Figure 2. Scene from the 1960 film, The Lost World, featuring a giant iguana with horns added presaging the appearance of Shringasaurus.

Figure 2. Scene from the 1960 film, The Lost World, featuring a giant iguana with horns added presaging the appearance of Shringasaurus.

Coincidentally the 1960 film,
The Lost World featured an iguana made up with horns similar to those of Shringasaurus.

References
Sengupta S, Ezcurra MD and Bandyopadhyay S 2017. A new horned and long-necked herbivorous stem-archosaur from the Middle Triassic of India. Nature, Scientific Reports 7: 8366 | DOI:10.1038/s41598-017-08658-8 online here.

No Wiki page yet.

What is the enigmatic Otter Sandstone (Middle Triassic) diapsid?

Updated October 10, 2020
after µCT scans were published revealing hidden data buried in the matrix. Click here to see the new data and new nesting with Dinocephalosaurus.

Coram, Radley and Benton 2017
presented a “small diapsid reptile [BRSUG 29950-12], possibly, pending systematic study, a basal lepidosaur or a protorosaurian.” According to Coram et al. “The Middle Triassic (Anisian) Otter Sandstone was laid down mostly by braided rivers in a desert environment.”

Figure 1. The Middle Triassic Otter Sandstone diapsid BRSUG 29950-12 under DGS nested with basalmost lepidosaurs like Megachirella.

Figure 1. The Middle Triassic Otter Sandstone diapsid BRSUG 29950-12 under DGS nested with basalmost lepidosaurs like Megachirella. Skeleton is exposed in ventral (palatal) view.

The LRT is here to nest and identify published enigmas
The large reptile tree (LRT 1041 taxa) nests BRSUG 29950-12 with the basalmost lepidosaur Megachirella. They are a close match and preserve nearly identical portions of their skeletons (Fig. 2). Megachirella was originally considered a sister to Marmoretta, another basal sphenodontian from the much later Middle/Late Jurassic.

FIgure 2. Megachirella (Renesto and Posenato 2003) is a sister to the BSRUG diapsid.

FIgure 2. Megachirella (Renesto and Posenato 2003), also from Middle Triassic desposits, is a sister to the BSRUG diapsid and provides a good guide for its eventual reconstruction.

At the base of the Lepidosauria
in the LRT nests Megachirella, derived from a sister to Sophineta (Early Triassic) and Saurosternon + Palaegama (Latest Permian) and kin. Sisters to Megachirella within the Lepidosauria include the tritosaurs Tijubina + Huehuecuetzpalli (Early Cretaceous), Macrocnemus (Middle Triassic) and the prosquamate Lacertulus (Late Permian). Also similar and related to Palaegama is Jesairosaurus (Middle Triassic). So the genesis of the Lepidosauria is Late Permian. The initial radiation produced taxa that continued into the Early Cretaceous. The radiation of derived taxa continued with three major clades, only one of which, the Tritosauria, is now completely extinct.

Note
It is important to remember that lepdiosaurs and protorosaurs are not closely related, but arrived at similar bauplans by convergence, according to the LRT. The former is a member of the new Lepidosauromorpha. The latter is a member of the new Archosauromorpha. Last common ancestor: Gephyrostegus and kin.

Nesting at the base of the Lepidosauria
in the Sphenodontia clade makes the BSRUG specimen an important taxon. Let’s see if and when this taxon is nested by academic workers that they include all of the pertinent taxa and confirm or re-discover the Tritosauria. The LRT provides a good list of nearly all of the pertinent taxa that should be included in that future study, many of which are listed above. Based on that list, the BSRUG specimen is a late-survivor of a perhaps Middle Permian radiation of basal lepidosaurs.

References
Coram RA, Radley JD and Benton MJ 2017. The Middle Triassic (Anisian) Otter Sandstone biota (Devon, UK): review, recent discoveries and ways ahead. Proceedings of the Geologists’ Association in press. http://dx.doi.org/10.1016/j.pgeola.2017.06.007

Do gliding lizards (genus: Draco) actually grab their extended ribs?

Figure 1. Extant Draco flying with hands either grabbing the leading edge of the membrane or streamlining their hands on top of it.

Figure 1. Extant Draco flying with hands either grabbing the leading edge of the membrane or streamlining their hands on top of it. Images from Dehling 2016.

Gliding lizards
of the genus Draco (Figs. 1, 2) come in a wide variety of species. Similar but extinct gliding basal lepidosauriformes, like Icarosaurus (Fig. 2), form a clade that arose in the Late Permian and continued to the Early Cretaceous.

Figure 2. Two Draco species fully extending their rib membranes without the use of the hands.

Figure 2. Two Draco species fully extending their rib membranes without the use of the hands.

A recent paper
(Dehling 2016) reported, “the patagium is deliberately grasped and controlled by the forelimbs while airborne.” Evidently this ‘membrane-grab’ behavior has not been noted before. I wondered if the rib skin is indeed grasped, or does the forelimb merely fold back against the leading edge of the patagium in a streamlined fashion? Photographs of climbing Draco specimens (Fig. 2) show that the patagium  can fully extend without the aid of the forelimbs to stretch them further forward.

Figure 3. Icarosaurus. Note the tiny ribs near the shoulders. The bases for the strut-like dermal bones are the ribs themselves flattened and transformed by fusion to act like transverse processes, which sister taxa do not have. Note the length of the hands corresponds to the base of the anterior wing strut.

Figure 3. Icarosaurus. Note the tiny ribs near the shoulders. The bases for the strut-like dermal bones are the ribs themselves flattened and transformed by fusion to act like transverse processes, which sister taxa do not have. Note the length of the hands corresponds to the base of the anterior wing strut, a great place to rest the manus or grab the membrane.

A quick review of prehistoric gliding keuhneosaurs
(Fig. 3) show that the manus unguals are not quite as large and sharp as those of the pes and that the manus in gliding mode extends just beyond the shorter two anterior dermal struts so that the glider -may- have grasped the anterior struts in flight. Or may have rested the manus there. Remember, these are taxa unrelated to the extant Draco, which uses actual ribs to stretch its gliding membrane. The same holds true for the more primitive Coelurosauravus and Mecistotrachelos, which have not been traditionally recognized as basal kuehneosaurs.

* As everyone should know by now…
the so-called transverse processes in kuehneosaurs are the true ribs, only fused to the vertebrae. The ribs remain unfused to the vertebrae in the older and more primitive coelurosauravids. No sister taxa have transverse processes elongate or not.

References
Dehling M 2016. How lizards fly: A novel type of wing in animals.

Earliest(?) stem squamate – SVP abstracts 2016

Klugman and Pritchard 2016
believe they have found the earliest lepidosaur stem squamate (see below). The large reptile tree finds earlier stem squamates (Fig. 1, click here to enlarge).
Earlier we looked at the wider and narrower definitions of the term ‘stem’.
Figure 1. CLICK TO ENLARGE. Stem taxa are closest ancestors to living taxa. Here basal diapsids and marine enaliosaurs are stem archosaurs. Triceratops is a stem bird. Captorhinids are stem turtles. Pterosaurs are stem squamates.

Figure 1. CLICK TO ENLARGE. Stem taxa are closest ancestors to living taxa. Here basal diapsids and marine enaliosaurs are stem archosaurs. Triceratops is a stem bird. Captorhinids are stem turtles. Pterosaurs are stem squamates. The colors here indicate the wider definition of ‘stem’.

From the Klugman and Pritchard abstract (abridged)
“Crown group lepidosaurs are highly diverse: they comprise more than 7,000 globally distributed extant species of lizards and snakes (Squamata), plus the single rhynchocephalian genus Sphenodon. The earliest known lepidosaurs are rhynchocephalians from the Late Triassic of Europe, (1) and this group quickly diversified and achieved a global distribution by the end of the Triassic. In contrast, early squamates have a sparse fossil record; their first representatives are found in the Early-Middle Jurassic of Laurasia (2). Although Rhynchocephalia and Squamata diverged in the Middle Triassic, a 40-50 million years ghost lineage exists for Squamata. Jurassic squamates are already considerably derived, and have already diversified into their extant groups, which testifies to a substantial gap in the known fossil record. Here we report on a new lepidosaur from a Norian microvertebrate site in Petrified Forest National Park, Arizona. This fossiliferous locality is from the Upper Blue Mesa Member of the Chinle Formation, and is dated to 221 mya. The depositional environment is a shallow anoxic lake, where skeletal elements preserved are disarticulated and often fragmentary. The site has yielded a diverse small vertebrate fauna, including the new lepidosaurs and several undescribed rhynchocephalians. Skeletal elements are represented by numerous small, delicate pleurodont maxilla and dentaries. We integrated the material of the new lepidosaurs into phylogenetic analyses of Permo-Triassic Diapsida and Mesozoic Lepidosauromorpha, using maximum parsimony, maximum likelihood, and Bayesian analysis. All analyses support the new taxon as the sister taxon to all other Squamata, (3) substantially reducing the ghost lineage of Squamata. This discovery indicates that the absence of squamate fossils in their early evolutionary history could be caused in part by collection bias towards larger, more robust specimens. This taxon provides a look into the early evolutionary history of squamates. It also adds direct evidence of yet another major lineage of extant terrestrial vertebrates to originate in the Triassic.”
Notes
  1. In the LRT Megachirella (Middle Triassic) is an earlier basal rhynchocephalian. Bavariasaurus (Late Jurassic) and
  2. Lacertulus (Late Permian; Fig. 2) are basal stem squamates. Ascendonanus (Rößler et al. 2012), an Early Permian lepidosaur in the iguanid clade is the earliest lepidosaur I have encountered yet, although this is based on low-rez data. Based on these nestings, the original radiation of lepidosaurs must have occurred in the Permian and then enjoyed very long period of stasis.  Lacertulus is the oldest known lepidosaur and older than any Late Triassic Petrified Forest taxa. It does not have pleurodont (fused to the jaw) teeth.
  3. The LRT is an analysis that includes a long list of pro or proto-squamates and tritosaurs that are sisters to the Squamata. Palaegama (Late Permian), Tridentinosaurus (Early Permian) and Saurosternon (Latest Permian) are sisters to the Lepidosauria and they are basal to the highly derived Late Permian taxon, Coelurosauravus. So the original radiation of lepidosaurs and their lepidosauriform sisters must have been in the Early Permian. If one deletes Sphenodon, then another stem squamate would be Macroleter (Middle Permian). Earlier than this and you get into stem turtles.
Figure 1. Lacertulus, a basal squamate from the Late Permian

Figure 2. Lacertulus, a basal squamate from the Late Permian

References
Klugman B and Pritchard AC 2016. Earliest stem-squamate (Lepidosauria) from the Late Triassic of Arizona. Abstract from the 2016 meeting of the Society of Vertebrate Paleontology.
Rößler R, Zierold T, Feng Z, Kretzschmar R, Merbitz M, Annacker V and Schneider JW 2012. A snapshot of an early Permian ecosystem preserved by explosive volcanism: New results from the Chemnitz Petrified Forest, Germany. PALAIOS, 2012, v. 27, p. 814–834

What is Triopticus? (It’s not what they think it is…)

Updated July 13, 2017 with new bone identifications for Triopticus. These further cement the sisterhood to Tanytrachelos. 

It’s been a long time
since an interesting ‘reptile’ showed up in the literature. Especially an ‘enigma’ like this one.

A recent paper by Stocker et al. 2016
reports on a domed and expanded Late Triassic cranium that they identify as an archosaur, but it’s unlike that of any other archosaur. Triopticus primus was named for its three eyes, with a big one on top (Fig. 1). The authors compared the domed appearance of the cranium in Triopticus with a Cretaceous dome-headed ornithischian dinosaur, Stegoceras.  They also discussed convergence in general and provided a CT scan brain endocast of Triopticus.

Unfortunately 
the authors employed a prior cladogram (Fig. 2) by Nesbit et al. 2015, expanded from Pritchard et al. 2015. that was shown to not recover sister taxa that looked alike and did not provide a gradual accumulation of derived traits at several nodes. In their cladogram Triopticus nested without resolution among basal archosauriforms like Proterosuchus, which looks nothing like it. By contrast, the LRT was able nest and fully resolve Triopticus elsewhere.

Figure 1. In round 1 I added characters shown here to the LRT in two passes. One recovered a sisterhood with mesosaurs. The other nested with Tanytrachelos, among the tanystropheid tritosaur lepidosaurs. Both shown here for comparison.

Figure 1. In round 1 I added characters shown here to the LRT in two passes. One recovered a sisterhood with mesosaurs. The other nested with Tanytrachelos, among the tanystropheid tritosaur lepidosaurs. Both shown here for comparison. Triopticus would be 2x the size of the giant Tanyrachelos from New Mexico.

From the Stocker et al. abstract:  “Exemplifying this extreme morphological convergence, we present here a new dome-headed taxon from the assemblage, which further illustrates the extraordinary range of morphological disparity present early in the Late Triassic.” That ‘extraordinary range’ should be — and will be — chopped down substantially with the right sister taxa.

A few problems with the archosauriform hypothesis include:

  1. No other archosauriforms, until you get to pachycelphalosaurs in the Cretaceous, expand the cranium deleting the upper temporal fenestra.
  2. The entire rostrum and mandible is absent, so no naris, antorbital fenestra or teeth are known, even in part.
  3. They dubiously identified an antorbital fenestra and fossa at the edge of the fossil.
  4. …and they were not aware that Tanytrachelos and kin, including pterosaurs within the – Lepidosauria -, also have an antorbital fenestra, but without a fossa.
  5. A large pineal opening is present, but never present at such a size in archosauriforms.
  6. The extreme angle of the rostrum coupled with the large orbit are traits not found in basal archosauriforms that typically have a long boxy rostrum.
Figure 2. Stocker et al. 2106 cladogram nesting Triopticus uncertainly within a set of unresolved basal archosauriforms. The LRT completely resolves that node.

Figure 2. Stocker et al. 2106 cladogram nesting Triopticus uncertainly within a set of unresolved basal archosauriforms and far from the Tanystropheidae. The LRT completely resolves all nodes. Note how this cladogram mixes Lepidosauromorpha with Archosauromorpha and separates the protorosaurus, Protorosaurus and Prolacerta.

This is a perfect problem
for the large reptile tree (LRT) which now provides then 820, now 1036 opportunities for Triopticus to nest in. With that large number of taxa, unfortunately I had to split the matrix in two, even for a simple Heuristic Search. By contrast, the Stocker et al. matrix included 30 taxa and 247 characters.

Stocker et al. report,
“We chose this dataset because the following combination of character states in Triopticus are also present in some archosauromorph taxa:

  1. presence of a single occipital condyle;
  2. ossified laterosphenoid;
  3. presence of a metotic strut of the otoccipital;
  4. presence of upper and lower temporal fenestrae;
  5. presence of an antorbital fenestra and fossa formed by the lacrimal.”

They provided no reconstructions of included taxa.

First,
I tested Triopticus against basal tetrapods and the new Lepidosaurmorpha and found that Triopticus nested with the aquatic, long-necked tritosaur Tanytrachelos (Fig. 1), large specimens of which were recently found in New Mexico (Fig. 3). Like Triopticus the rostrum descends at a high angle from a tall cranium in Tanytrachelos, which also shares a large orbit and a large pineal foramen (at present known only from sister taxa). Like related fenestrasaurs and langobardisaurs, Tanytrachelos also had a small antorbital fenestra without a fossa, but that would have been beyond the rim of the broken skull in Triopticus (Fig. 5).

Figure 2. A large incomplete Tanytrachelos from New Mexico compared to the smaller more complete East Coast specimen. Triopticus would be twice as large as the New Mexico specimen.

Figure 3. A large incomplete Tanytrachelos from New Mexico compared to the smaller more complete East Coast specimen. Triopticus would be twice as large as the New Mexico specimen.

Second,
I tested Triopticus with the rest of the matrix, the new Archosauromorpha, and found that Triopticus nested with the mesosaurs (Fig. 4), an aquatic enaliosaur clade close to thalattosaurs and ichthyosaurs, all derived from basal pachypleurosaurs. It did not nest with archosauriforms. While basal mesosaurs have typical diapsid temporal regions, Mesosaurus, like Triopticus, closes up the upper temporal fenestra, then the lateral temporal fenestra with bone expansion.  Mesosaurs also retain a relatively large pineal foramen and have large eyes, but they don’t have a sharply descending preorbital region.

Mesosaurus

Figure 4. Mesosaurus, like Triopticus, has a large pineal foramen and expands the skull bones to obliterate former temporal fenestrae.

Digital Graphic Segregation
was applied to the cranial lump that is Triopticus (Fig. 5) and the skull suture patterns, perhaps overlooked by those with firsthand access due to the expansion of the cranial bones, revealed a Tanytrachelos-like morphology (Fig. 2). I illustrate this interpretation here with the hope that this hypothesis can be either confirmed or falsified. This is a tough assignment.

Figure 5. Triopticus reconstructed along the bauplan of Tanytrachelos.  The upper temporal fenestra is the top half of a divided lateral temporal fenestra. At 72 dpi this is 90 percent of actual size.

Figure 5. Triopticus reconstructed along the bauplan of Tanytrachelos. The upper temporal fenestra is the top half of a divided lateral temporal fenestra. At 72 dpi this is 90 percent of actual size.

Tanystropheids
have been reported from the Hayden Quarry of northern New Mexico (Chinle Formation) far from the west central Texas location of Otis Chalk. Stocker et al. included Tanytrachelos in their study, even though they have not provided a reconstruction of it, so it is difficult to imagine how they interpreted it. Tanystropheids, in general, have widely varying skull shapes. Triopticus appears to have expanded the morphospace just a little, not a lot.

The loss of ventral material in the Triopticus fossil
appears to have occurred at the roof the narial/oral opening.

So what other long-necked animal
expands the cranium like Triopticus? Giraffa, the giraffe. Maybe it will turn out to be a better analogy than short-necked Stegoceras?

References
Nesbitt SJ, Flynn JJ, Pritchard AC, Parrish JM, Ranivoharimanana L and Wyss AR 2015. Postcranial osteology of Azendohsaurus madagaskarensis (?Middle to Upper Triassic, Isalo Group, Madagascar) and its systematic position among stem archosaur reptiles. Bulletin of the American Museum of Natural History 899, 1-125.
Pritchard AC, Turner AH, Nesbitt SJ, Irmis RB and Smith ND 2015. Late Triassic tanystropheid (Reptilia, Archosauromorpha) remains from northern New Mexico (Petrified Forest Member, Chinle Formation): insights into distribution, morphology, and paleoecology of Tanystropheidae. Journal of Vertebrate Paleontology, 10.1080/02724634.02722014.02911186.
Stocker MR, NesbittSJ, Criswell KE, Parker WG, Witmer LM, Rowe TB, Ridgely R  Brown MA 2016. A Dome-Headed Stem Archosaur Exemplifies Convergence among Dinosaurs and Their Distant Relatives. Current Biology (advance online publication)DOI: http://dx.doi.org/10.1016/j.cub.2016.07.066   pdf

So, is it Leptosaurus? Kallimodon? Or neither?

Rauhut and Lopez-Arbarello (2016)
compare several complete specimens of Late Jurassic rhynchocephalians from Germany (Fig. 1) with a newer specimen from Schamhaupten, JME-Scha 100 (lower left of Fig. 1).  that was earlier described by Renesto and Viohl 1997 as Leptosaurus pulchellus, JME-Scha 40. (Why the number change?) The holotype, Leptosaurus neptunium (Fig. 1b, was described earlier by Fitzinger 1837, but referred to Kallimodon by Cocude-Michel 1963. So that sets the stage for the study.

We looked at this taxon earlier here when Renesto and Viohl published on it.

Figure 1. Taxa considered by Rauhut and López-Arbarello include Homoeosaurus, Leptosaurus, Kallimodon and rhynchocephaian specimen from Schamhaupten, JME-Scha 100.

Figure 1. Taxa considered by Rauhut and López-Arbarello include Homoeosaurus, Leptosaurus, Kallimodon and rhynchocephaian specimen from Schamhaupten, JME-Scha 100.

From the Rauhut and Lopez-Arbarello abstract:
“Unfortunately, the taxonomy of the rhynchocephalians from these units has not been satisfactorily established so far, which hampers studies of their evolutionary importance.” 

Actually
the large reptile tree (LRT) has taken the first steps toward that goal (Fig. 2). Though not aware of the Leptosaurus holotype, I used instead the JME-Scha specimen in its place based on the work and nomenclature of Renesto and Viohl 1997. Not sure, but the holotype of Leptosaurus looks quite a bit like the Homoeosaurus specimens I used for data. 

Figure 2. Subset of the large reptile tree, the Rhynchocephalia. This clade also includes Rhynchosauria, Azendohsaurus and Trilophosaurus.

Figure 2. Subset of the large reptile tree, the Rhynchocephalia. This clade also includes Rhynchosauria, Azendohsaurus and Trilophosaurus.

From the Rauhut and Lopez-Arbarello abstract:
“Differences to the type of Kallimodon pulchellus include the morphology of the maxillary teeth, the phalangeal formula of the manus, and the shape of the posterior process of the second sacral rib. An important difference with the type of Leptosaurus neptunius is the higher number of premaxillary teeth in the specimen from Schamhaupten (four versus two), despite a significantly larger body size, whereas there is rather a tendency to reduce the number of premaxillary teeth through fusion during ontogeny in rhynchocephalians.” 

Rauhut and Lopez-Arbarello conclude
that the JME Scha specimen cannot be referred to either Kallimodon or Leptosaurus, but they do not rename the JME Scha specimen.

Unfortunately
the authors do not realize that the tiny JME Scha specimen is the branching off point in the LRT for the toothed rhynchocephalian, Azendohsaurus and its sister Trilophosaurus, which has no premaxillary teeth because these taxa are not included on the Rauhut and Lopez-Arbarello family tree (Fig. 3). They also did not realize that Priosphenodon is the branching off point for the origin of rhynchosaurs.

Figure 3. Rhynchocephalian cladogram from Rauhut and López-Arbarello lacks many pertinent taxa and includes one, Homoeosaurus, that belongs elsewhere.

Figure 3. Rhynchocephalian cladogram from Rauhut and López-Arbarello lacks many pertinent taxa and includes one, Homoeosaurus, that belongs elsewhere. The LRT nests Kallimodon with Sphenodon and Saphenosaurus with Noteosuchus.

Other problems with the Rauhut and Lopez-Arbarello family tree
include the unwarranted inclusion of Homoeosaurus. In the LRT Homoeosaurus nests within the protosquamates the only matrix to test it on a large gamut of taxa. The authors also include several taxa that were not included in the LRT, like Eilenodon, represented only by a posterior jaw fragment. On the other hand, missing taxa from the Rauhut and Lopez-Arbarello rhynchocephalian tree (Fig 3) include:

  1. Megachirella
  2. Marmoretta
  3. Ankylosphenodon
  4. Heleosuchus
  5. Sphenotitan
  6. Noteosuchus
  7. Trilophosaurus
  8. Azendohsaurus
  9. Eohyosaurus
  10. Mesosuchus
  11. Rhynchosaurus
  12. Bentonyx
  13. Hyperodapedon

You might not like this, if you like traditional studies
but these taxa all nest within the clade Rhynchocephalia in the LRT. And Gephyrosaurus is no longer the most primitive of the lot.

Perhaps 
Rauhut and Lopez-Arbarello will someday expand their taxon list. That’s why the LRT is here… to make taxon selection simple, complete and verifiable, not just traditional. The hard work has already been done for you. All you have to do is focus on your clade of interest!

References
Cocude-Michel M 1963. Les Rhynchocéphales et les Sauriens des Calcaires lithographiques (Jurassique supérieur) d’Europe occidentale.– Nouvelles Archives du Muséum d’Histoire Naturelle de Lyon 7: 1–187.
Fitzinger LJF 1837. Vorläufiger Bericht über eine höchst interessante Entdeckung -Dr. Natterers in Brasil. Oken’s Isis.
von Meyer H 1850. Mittheilungen an Professor Bronn gerichtet: Neües Jahrbuch fur Mineralogie, Geologie und Palaontologie, Bd 18, p. 195-204.
Rauhut OWM and Lopez-Arbarello A 2016. Zur Taxonomie der Brückenechse aus dem oberen Jura von Schamhaupten (On the taxonomy of the rhynchocephalian from the Late Jurassic of Schamhaupten). Archaeopteryx 33: 1-11; Eichstätt 2016.
Renesto S and Viohl G 1997. A sphenodontid (Reptilia, Diapsida) from the late Kimmeridgian of Schamhaupten (Southern Franconian Alb, Bavaria, Germany). Archaeopteryx 15:27-46

 

Tridentinosaurus antiquus: a glider ancestor, not a protorosaur

I had never heard of this one before. 
Evidently this Early Permian reptile is famous for being fossilized between volcanic layers and for preserving more skin than bone. Using DGS I was able to tease out some of the bone (Fig. 1) and nest Tridentinosaurus not with the protorosaurs, as Leonardi (1959) proposed, but with basal lepidosauriforms. Tridentinosaurus nests in the large reptile tree as an Early Permian descendant of the late-surviving Palaegama and an ancestor to the Late Permian ‘rib’ glider, Coelurosauravus and the Late Triassic ‘rib’ glider, Icarosaurus along with other glider clade members.

Figure 1. Tridentinosaurus at 26.5 cm long is an Early Permian ancestor to Late Permian Coelurosauravus and Late Triassic Icarosaurus.

Figure 1. Tridentinosaurus at 26.5 cm long is an Early Permian ancestor to Late Permian Coelurosauravus and Late Triassic Icarosaurus. Here two images taken in different light conditions were superimposed, then traced. An arboreal lifestyle is suspected here, based on the long limbs and toes.

Tridentinosaurus antiquus (Early Permian, Dal Piaz 1932, Leonardi 1959, 26.5cm long; Museum of Paleontology of the University of Padua 26567). Ronchi et al. described the specimen as “a beautiful but biochronologically useless specimen of which only the out−line of the soft tissues is well preserved.” The volcanic sediments in Sardinia occur in Cisuralian / Sakmarian deposits 291 million years old.

Although known for more than 50 years, 
and with quite a story to tell, this genus was not famous enough to merit its own Wikipedia page when I wrote this. Based on phylogenetic bracketing, the tail may have been twice as long originally.

Most prior workers do not nest 
Coelurosauravus and kin with Kuehneosaurus and kin (including Xianglong from the Cretaceous. Here they do nest together and Tridentinosaurus provides clues to the clade’s arboreal origin. Apparently this is a novel hypothesis, a by-product of having so many (694) taxa in the large reptile tree (subset Fig. 2).

Figure 2. Subset of the large reptile tree showing the nesting of Tridentinosaurus at the base of the gliders, close to the drepanosaurs.

Figure 2. Subset of the large reptile tree showing the nesting of Tridentinosaurus at the base of the gliders, close to the drepanosaurs.

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.
Ronchi, A., Sacchi, E., Romano, M., and Nicosia, U. 2011. A huge caseid pelycosaur from north−western Sardinia and its bearing on European Permian stratigraphy and palaeobiogeography. Acta Palaeontologica Polonica 56 (4): 723–738.

Saurodektes: Filling in the missing parts

Saurodektes rogersorum
is a small owenettid lepidosauromorph (BP/1/6025, Early Induan, Early Triassic) originally named “Saurodectes” by Modesto et al. (2003), but that name was preoccupied by a fossil louse. The holotype of Saurodektes is known from a partial skull and anteriormost postcrania (Fig. 1).

Figure 1. Saurodektes (originally Saurodectes) by Modesto et al. 2003 (black/white). Missing parts in color based on phylogenetic bracketing.

Figure 1. Saurodektes (originally Saurodectes) by Modesto et al. 2003 (black/white). Missing parts in color based on phylogenetic bracketing.

The missing parts of the skull
can be restored using phylogenetic bracketing after phylogenetic analysis. In the large reptile tree Saurodektes nests at the base of the Owenetta clade, between the Nyctiphruretus and Barasaurus clades.

Procolophonomorpha?
No. All of these taxa nest far from Procolophon and kin, which nest with Diadectes and kin.

Lepidosauriformes?
Almost. These taxa were ancestral to Paliguana and the Lepidosauriformes, which gave rise to all living lizards and a host of extinct relatives, including pterosaurs.

Nascent upper temporal fenestra?
No. While the tiny space between the parietal and large supratemporal appears to be creating an upper temporal fenestra in Saurodektes, in lepidosauriformes the supratemporal is reduced and migrates to the back as it it is replaced by the squamosal, which comes to rim the upper temporal fenestra. Best to consider those cranial holes as damaged goods here in Saurodektes.

References
Modesto SP, Damiani RJ, Neveling J,Yates AM 2003. A new Triassic owenettid parareptile and the Mother of Mass Extinctions. Journal of Vertebrate Paleontology 23 (3): 715.

Jesairosaurus and the drepanosaurs leave the Tritosauria :-(

My earlier reconstruction
of the basal lepidosauriform, Jesairosaurus (Fig. 1; contra Jalil 1997, not a protorosaur/prolacertiform) included several errors based on attempting to create a chimaera of several specimens of various sizes based on scale bars. In this case, scale bars should not have been used. Rather fore and hind parts had to be scaled to common elements, like dorsal vertebrae, as shown below (Fig. 2). I think this version more accurately reflects the in vivo specimen, despite its chimeric origins. All of the partial skeletons assigned to this genus were discovered at the same Early to Middle Triassic sandstone site and two were touching one another. A larger skull, ZAR 7, shows the variation in size from the skull to shoulders remains of the ZAR 6 specimen.

Figure 1. New reconstruction of the basal lepidosauriform, Jesairosaurus (Jalil 1993).

Figure 1. New reconstruction of the basal lepidosauriform, Jesairosaurus (Jalil 1997). The wide and flat ribs are interesting traits for a likely arboreal reptile.

Mother of all drepanosaurs
The Drepanosauria is an odd clade of slow-moving arboreal reptiles that includes Hypuronector, Vallesaurus, Megalancosaurus and Drepanosaurus (Figs. 2, 3). Jesairosaurus was not a drepanosaur, but nested basal to this clade before the present revisions. It remains basal to the Drepanosauria now with revisions.

The revised reconstruction of Jesairosaurus 
shifts this clade away from Huehuecuetzpalli, Macrocnemus and the rest of the Tritosauria. Now Jesairosaurus and the drepanosaurs nests between Saurosternon, Palaegama and the so-called “rib” gliders, beginning with Coelurosauravus.

A short history of Jesairosaurus
Shortly after their discovery Lehman 1971 referred the several hematite encrusted specimens to the Procolophonida. Further preparation showed that they were referable to the Diapsida, according to Jalil (1990) and the, more specifically, to the Prolacertiformes (Jalil 1997) as a sister to Malerisaurus with Prolacerta as a common ancestral sister. Jalil did not include the closest sisters of Jesairosaurus as revealed by the present analysis.

With a much larger list of taxa,
the large reptile tree nests Malerisaurus between the Antarctica specimen assigned to Prolacerta (AMNH 9520) and the holotype of Prolacerta. Jesairosaurus, as mentioned above, nests with the basal lepidosauriformes. Any traits shared with protorosaurs are by convergence. Deletion of Jesairosaurus does not affect the nesting of the Drepanosauria as basal lepidosauriformes.

Figure 3. Drepanosaurs and their ancestor sisters, Jesairosaurus and Palaegama to scale.

Figure 3. Drepanosaurs and their ancestor sisters, Jesairosaurus and Palaegama to scale.

Arboreal
This new nesting shifts drepanosaurs closer to kuehneosaurs (Figs. 3, 4), another notably arboreal clade.

Figure 3. The new nesting for Jesairosaurus and the drepanosaurs as sisters to the Kuehneosaurs, several nodes away from Huehuecuetzpalli and the tritosaurs.

Figure 3. The new nesting for Jesairosaurus and the drepanosaurs as sisters to the Kuehneosaurs, several nodes away from Huehuecuetzpalli and the tritosaurs.

Certainly
there will someday be more taxa to fill in the current large morphological gaps in and around Jesairosaurus, but here’s what we have at present (Fig. 3) with regard to the origin of the so-called “rib” gliders (actually dermal rods, not ribs, as shown by Coelurosauravus) and the origin of the drepanosaurs.

Figure 4. Jesarosaurus to scale with sisters Palaegama and Coelurosauravus.

Figure 4. Jesairosaurus to scale with sisters Palaegama and Coelurosauravus.

The shifting of a clade
like Jesairosaurus + Drepanosauria occurred due to inaccurate reconstructions used for data. Science builds on earlier errors and inaccuracies. I let the computer figure out where taxa nest in a cladogram of 606 possible nesting sites, minimizing the negative effects of bias and tradition.

It’s sad
to see the drepanosaurs leaving the Tritosauria as it contains several oddly Dr. Seuss-ian variations on the tritosaur theme.

Also note the nesting
of the basal Rhynchocephalians, Megachirella and Pleurosaurus, between the palaegamids and the tritosaurs (Fig. 4). In the course of this study, both also received updates to their skull reconstructions. The former was difficult to interpret without knowing where it nested. What appeared to be an odd sort of a squamosal in Megachirella now appears to be a pair of displaced pleurosaur-like premaxillae. For Pleurosaurus I should not have trusted a prior line drawing by another worker. Here I used DGS to create what appears to be a more accurate skull without so many apparent autapomorphies.

References
Jalil N 1990. Sur deux cranes de petits Sauria (Amniota, Diapsida) du Trias moyen d’ Algerie. Comptes Rendus de I’ Academic des Sciences, Paris 311 :73 1- 736.
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.
Lehman JP 1971. Nouveaux vertebres du Trias de la Serie de Zarzai’tine. Annales de Paleontologic (Vertebres) 57 :71-93.