Palaeopleurosaurus and Pleurosaurus are not sisters. >GASP!<

Earlier we looked at the phylogenetic connection between Marmoretta, Megachirella, Ankylosphenodon and the aquatic rhynchocephalians, the pleurosaurs (Figs. 1, 2). Continuing work with the large reptile tree (still not updated) illuminates new relationships among Palaeopleurosaurus and Pleurosaurus (Figs. 1, 2), which turns out to be pretty obvious on hindsight.

Figure 1. Pleurosaurus and Palaeopleurosaurus to scale with sisters.

Figure 1. Pleurosaurus and Palaeopleurosaurus to scale with sisters along with arrows indicating evolutionary pathways. 

The distinct skulls
of conventional pleurosaurs give them away (Fig. 2). Palaepleurosaurus looks like many basal rhynchocephalians. Pleurosaurus looks more streamlined. By convention that meant it was more derived, better adapted to an aquatic niche.

Figure 2. Pleurosaurus and Palaeopleurosaurus skulls compared to those of sister taxa.

Figure 2. Pleurosaurus and Palaeopleurosaurus skulls compared to those of sister taxa. They are more closely related to the graphically associated taxa than to each other. 

Actually just the opposite is true,
according to phylogenetic analysis. Pleurosaurus has a low, long rostrum because its closest sisters, Palaegama, Megachirella and Marmoretta share this skull shape. Elsewhere on the tree, Palaeopleurosaurus nests with rhynchocephalians with a similar skull morphology, including Ankylosphenodon, succeeding Gephyrosaurus and preceding Planocephalosaurus.

In case you didn’t notice…
Long-legged Palaegama is a VERY basal lepidosauriform. One branch of descendants, the lepidosaurs, retained a terrestrial niche. These included basal tritosaurs, like Tijubina, and rhynchocephalians. Another branch produced the Permian to Cretaceous so-called ‘rib’ gliders, like Icarosaurus. A third branch produced aquatic forms in the form of Pleurosaurus.

Figure 3. Palaegama, close to the origin of all Lepidosauriformes.

Figure 3. Palaegama, close to the origin of all Lepidosauriformes. This generalized taxon was close to the origin of arboreal climbers and gliders, terrestrial lizard-like forms and aquatic pleurosaurs.

Earlier I mistakenly nested the Marmoretta, Megachirella, Palaeopleurosaurus, Pleurosaurus clade between Gephyrosaurus and Planocephalosaurus. Less reliance on published drawings and more reliance on specimen photographs was helpful in working out the problems that arose with the data from the drawings. This new nesting sheds light on the origin of the Lepidosauria and requires the redefinition of the Pleurosauridae because the two former pleurosaur genera are more closely related to other taxa than to each other.

References
Carroll RL 1985. A pleurosaur from the Lower Jurassic and the taxonomic position of the Sphenodontida. Palaeontographica Abteilung A, 189:1-28.
Dupret V 2004. The pleurosaurs: anatomy and phylogeny. Revue de Paléobiologie, Geneve 9:61-80.
Meyer H 1831. IV Neue Fossile Reptilien, aud der Ordnung der Saurier.
Reynoso VH 1996. Early Cretaceous Lepidosaurs (Reptilia: Diapsida) from Central Mexico and the Phylogeny of Lepidosauromorphs. 369 pp. Unpublished Ph.D. Thesis, McGill University, Montreal, Canada.

wiki/Pleurosaurus

Megachirellla and Marmoretta are basal to Pleurosaurs

Earlier we looked at pleurosaurs (Fig. 1, elongate, aquatic rhynchocephalians). Pleurosaurus goldfussi (Meyer 1831) was discovered first. Palaeopleurosaurus is a more primitive taxon with a distinct premaxillary tooth. Note the retraction of the nares, common to many aquatic reptiles.

The present blogpost updates their origins with phylogenetic analysis, adding these two taxa to the large reptile tree.

Dupret (2004) nested pleurosaurs (Fig. 1) with Sapheosaurus. Adding pleurosaurs to the large reptile tree (not updated yet) nested them with Marmoretta and Megachirella (Figs. 2-5), helping to remove the ‘enigma’ status from the latter. Dupret (2004) did not include these two taxa in analysis.

The pleurosaurs

Figure 1. The pleurosaurs, Pachypleurosaurus and Pleurosaurus, known rhynchocephalians, now nesting with Marmoretta and Megachirella.

Pleurosaurs are yet one more clade of “return to the water” reptiles, and probably the last one anyone thinks of. They’re just not often reported on. Wiki reports, Pleurosaurus fossils were discovered in the Solnhofen limestone formation of BavariaGermany and CanjuersFrance.” The limbs were reduced. The torso and tail were elongated. Pleurosaurs probably swam in an eel-like or snake-like undulating pattern.

But where did they come from?

Figure 2. Marmoretta, a basal rhynchocephalian in the lineage of pleurosaurs

Figure 2. Marmoretta, a basal rhynchocephalian in the lineage of pleurosaurs

Marmoretta oxoniensis (Evans 1991) Middle/Late Jurassic, ~2.5 cm skull length, orginally considered a sister of kuehneosaursdrepanosaurs and lepidosaurs. Here Marmoretta was derived from a sister to GephyrosaurusMarmoretta was a sister to Planocephalosaurus and Megachirella. 

Distinct from Gephyrosaurus, the skull of Marmoretta was flatter overall with a larger orbit. The parietals were longer. The naris was larger and more dorsal. The prefrontal was narrower. The lacrimal was still visible. The jugal was reduced.

A flat-headed rhynchocephalian, Marmoretta nests near the base of that clade, prior to the fusion of teeth together and to the jaws in many derived taxa, including pleurosaurs.

Figure 1. Megachirella, a flat-headed rhynchocephalian close to Marmoretta and basal to pleurosaurs.

Figure 3. Megachirella, a flat-headed rhynchocephalian close to Marmoretta and basal to pleurosaurs.

Megachirella wachtleri (Renesto and Posenato 2003, Renesto and Bernardi 2013) KUH-1501, 2 cm skull length, Middle Triassic, was a tiny lepidosauromorph with a moderately elongated neck and flattened skull. The teeth were short and stout. Megachirella was originally nested with Marmoretta and the large study confirms it, but it is also basal to the aquatic pleurosaurs.

Figure 4. Megachirella in situ with bones colorized. Some bones are represented by impressions of the lost bone.

Figure 4. Megachirella in situ with bones colorized using DGS techniques. Some bones are represented by impressions of the lost bone. The yellow premaxilla tooth is represented by a questionable impression/crack. The nasal may not be a bone, according to S. Renesto. Scale bar = 1 cm.

 

Shifting the pleurosaurs to Gephyrosaurus adds 13 steps. To Planocephalosaurus adds 23 steps. More steps are added with a shift to other rhynchocephalians.

Figure 5. Skull elements of Megachirellla traced in color (Fig. 4) then transferred to line art in three views.

Figure 5. Skull elements of Megachirellla traced in color (Fig. 4) then transferred to line art in three views. Reconstructions are important in such roadkill taxa.

Megachirella is a Middle Triassic rhynchocephalian. That leaves plenty of time for a sister to evolve into a Late Jurassic pleurosaur. The retracted naris common to pleurosaurs is clear on both Marmoretta and Megachirella. All three had an open lateral temporal fenestra.

If you find any mistakes here, please let me know. Such specimens are at or a little beyond the edge of my experience.

References
Carroll RL 1985. A pleurosaur from the Lower Jurassic and the taxonomic position of the Sphenodontids.
Dupret V 2004. The pleurosaurs: anatomy and phylogeny. Revue de Paléobiologie, Geneve 9:61-80.
Evans SE 1991. A new lizard−like reptile (Diapsida: Lepidosauromorpha) from the Middle Jurassic of Oxfordshire. Zoological Journal of the Linnean Society 103:391-412.
Fraser NC and Sues H-D 1997. In the Shadows of the Dinosaurs: early Mesozoic tetrapods. Cambridge University Press, 445 pp. Online book.
Heckert AB 2004. Late Triassic microvertebrates from the lower Chinle Group (Otischalkian-Adamanian: Carnian), southwestern U.S.A. New Mexico Museum of Natural History and Science Bulletin 27:1-170.
Meyer H 1831. IV Neue Fossile Reptilien, aud der Ordnung der Saurier.
Renesto S and Posenato R 2003. A new lepidosauromorph reptile from the Middle Triassic of the Dolomites (northern Italy). Rivista Italiana di Paleontologia e Stratigrafia 109(3) 463-474.
Renesto S and Bernardi M 2013. Redescriptions and phylogenetic relationships of Megachirella wachtleri Renesto et Posenato, 2003 (Reptilia, Diapsida). Paläontologische Zeitschrift, DOI 10.1007/s12542-013-0194-0

The Odd Swimming Sphenodontids

Updated November 11, 2014 with the Dnesting of pleurosaurs with Megachirella and Marmoretta. And updated December 3, 2014 with the division of the pleurosaurs into two convergent clades. 

Hard to believe that our favorite New Zealand “Living Fossil,” Tuatara (Sphenodon), had some aquatic sisters, but here they are.

Figure 1. Pleurosaurus and Palaeopleurosaurus to scale with sisters.

Figure 1. Pleurosaurus and Palaeopleurosaurus to scale with sisters.

Pleurosaurus goldfussi (Meyer 1831), Late Jurassic. 60 cm in length. The Triassic terrestrial sphenodontians produced a Late Jurassic marine lineage known as the pleurosaurs after the first of these to be discovered, Pleurosaurus and also one of the most derived. Palaeopleurosaurus appears to be a stretched out version of its terrestrial antecedent, Planocephalosaurus, and was a transitional form to later, longer, more streamlined pleurosaurs.

Added Dec 03, 2104: > That was the traditional nesting. New analyses indicate that Pleurosaurus nested between Palaegama, Megachirella and Marmoretta at the base of the Lepidosauria, which radiated in the Middle to Late Permian. Palaeopleurosaurus had a convergent return to an aquatic niche as it nested between Gephyrosaurus and Planocephalosaurus. The similar Ankylosphenodon was a sister taxon.

Figure 2. Pleurosaurus and Palaeopleurosaurus skulls compared to those of sister taxa.

Figure 2. Pleurosaurus and Palaeopleurosaurus skulls compared to those of sister taxa.

Pleurosaurs were Late Jurassic aquatic sphenodontids, characterized by a long, streamlined and elongated body (with a short neck), small limbs and (as in most aquatic reptiles) nares that were displaced from the snout tip to closer to the orbits. The premaxilla of Palaeopleurosaurus was ventrally elongated to form a sharp spike that would have snared prey. They swam with snake-like undulations of the entire body. Their neural spines grew to become large rectangles, as in snakes. Pleurosaurs produced no Cretaceous descendants.

There is not much that is controversial about these lepidosaurs. They are not often studied and they are rarely on anyone’s Top 10 list, so I thought I’d toss them out for a little publicity.

Added Dec. 03, 2014: > Well, perhaps I spoke too soon as the two traditional pleurosaurs are not that closely related to one another. That’s a little bit of news!

References
Carroll RL 1985. A pleurosaur from the Lower Jurassic and the taxonomic position of the Sphenodontids.
Fraser NC and Sues H-D 1997. In the Shadows of the Dinosaurs: early Mesozoic tetrapods. Cambridge University Press, 445 pp. Online book.
Heckert AB 2004. Late Triassic microvertebrates from the lower Chinle Group (Otischalkian-Adamanian: Carnian), southwestern U.S.A. New Mexico Museum of Natural History and Science Bulletin 27:1-170.
Meyer H 1831. IV Neue Fossile Reptilien, aud der Ordnung der Saurier.

wiki/Planocephalosaurus
wiki/Pleurosaurus