News at the base of the Amniota, part 5: Cladogram of basal archosauromorpha

Earlier here and elsewhere we looked at the origin of the Amniota. Today we’ll take a look at the cladogram (Fig. 1) and some of the taxa no one expected to see on this side of the anamniote/amniote transition series.

Figure 1. Cladogram of basal amniotes, a subset of the large reptile tree. Dots represent phylogenetic size reductions. Bootstrap scores are shown. Archosauromorpha in gray. Lepidosauromorpha in black at the bottom. Figure 1. Cladogram of basal amniotes, a subset of the large reptile tree. Dots represent phylogenetic size reductions. Bootstrap scores are shown. Archosauromorpha in gray. Lepidosauromorpha in black at the bottom.

Figure 1. Cladogram of basal amniotes, a subset of the large reptile tree. Dots represent phylogenetic size reductions. Bootstrap scores are shown. Archosauromorpha in gray. Lepidosauromorpha in black at the bottom.

As before, the Amniota is divided at its base into the new Lepidosauromorpha (taxa closer to lepidosaurs) and the new Archosauromorpha (closer to archosaurs).

Figure 1. A new reconstruction of Gephyrostegus bohemicus. This species lived 30 million years after the origin of the Amniota in the Visean, 340 mya. Note the lack of posterior dorsal ribs. This trait shared by all basalmost amniotes, may provide additional space for massive eggs in gravid females, but is also shared with males, if there were males back then.

Figure 2. A new reconstruction of Gephyrostegus bohemicus. This species lived 30 million years after the origin of the Amniota in the Visean, 340 mya. Note the lack of posterior dorsal ribs. This trait shared by all basalmost amniotes, may provide additional space for massive eggs in gravid females, but is also shared with males, if there were males back then.

Gephyrotegus bohemicus (Fig. 1, Westphalian, 310 mya) is the last common ancestor of all amniotes and Silvanerpeton (Viséan, 340 mya) is the outgroup anamniote (or very possible also an amniote).

Utegenia nests as the common ancestor of frogs, salamanders, caecelians and microsaurs.

Figure 3. Utegenia nests as the common ancestor of frogs, salamanders, caecelians and microsaurs but the only known specimens are from the Earliest Permian.

Note the placement of the seymouriamorph, Utegenia (Fig. 3), at the base of the Lepospondyli, which includes extant amphibians and microsaurs… and just outside the base of the Amniota.

Basal Archosauromorpha

Figure 3. Two specimens attributed to Eldeceeon that nest together.

Figure 4. Two specimens attributed to Eldeceeon that nest together. The lack of posterior dorsal ribs was first noticed in the holotype.

Eldeceeon rolfei  – (Smithson 1994, ~27 cm in total length, Early Carboniferous (Viséan) ~335 mya), is from the same formation that yielded Silvanerpeton and Westlothiana. Eldeceeon is known from two dissimilar specimens that nest together. They have a smaller skull and slightly shorter limbs with smaller girdles while retaining a deep ventral pelvis.

Gephyrostegus-watsoni588

Figure 5. Gephyrostegus watsoni (Westphalian, 310 mya) reconstructed. Embryo is hypothetical. Note the lack of posterior dorsal ribs.

Gephyrostegus watsoni – (Brough and Brough 1967) was originally named Diplovertebron punctatum (Watson 1926, Fig. 5), but reassigned to Gephyostegus bohemicus by Carroll (1970) despite the size difference. Carroll thought G. watsoni was a juvenile. Klembara et al. (2014) agreed. The high arched neural spines, small intercentra, and the extreme lean of the posterior skull mark this small basal amniote/gephyrostegid distinct from all others. Egg shapes were found nearby along with insects. The embryo shown is hypothetical.

Figure 4. Solenodonsaurus reconstructed.

Figure 6. Solenodonsaurus reconstructed. The largest of the basal amniotes, likely aquatic. Note the intertemporal is still present. That doesn’t matter. It still nests with amniotes.

Solenodonsaurus janenschi – (Broili 1924) Early Permian ~13 cm skull length was considered the sister to all other amniotes by all prior workers, but here Solenodonsaurus nests as a basal archosauromorph, basal to chroniosuchids.

Figure 5. Three chorniosuchids to scale.

Figure 7. Three chorniosuchids to scale.

Chroniosuchids – (Tverdokhlebova 1972) Early Permian ~7 cm skull length, were considered aberrant pre-reptiles by all prior workers, but here they nest within the Archosauromorpha. Note the convergent appearance of an antorbital fenestra.

Figure 7. Casineria reconstructed.

Figure 8. Casineria reconstructed.

Casineria kiddi – (Paton, Smithson & Clack 1999) Viséan, Carboniferous, ~335 mya). Tiny Casineria lies at the end of a phylogenetic series of decreasing size beginning with Proterogyrinus.

Figure 8. Westlothiana reconstructed.

Figure 9. Westlothiana reconstructed. The gray area is hypothetical as if gravid.

Westlothiana – (Smithson & Rolfe 1990) lived ~338 mya, earlier than any other known reptile. This reconstruction has longer anterior dorsal ribs and shorter posterior dorsal ribs than originally reconstructed. A longer torso is a different solution to egg containment.

brouffia588overall

Figure 10. Broffia reconstructed. The smallest of all basal amniotes, this could be a juvenile or just a small adult.

Brouffia orientalis – (Carroll and Baird 1972) Westphalian, Late Carboniferous, (CGH IIIB 21 c. 587) and counterpart (MP451), specimen 1 of Brough and Brough (1967) was considered very small Gephyrostegus with two sacrals and an intertemporal. Carroll (1970) considered it not congeneric. Carroll and Baird (1972) considered it a primtiive reptile with a single sacral and no intertemporal. The missing skull of the sister taxon Casineria (Fig. 8) probably looks like this one.

We’ll look at basal Lepidosauromorpha tomorrow.

References
Broili F von 1924. Ein Cotylosaurier aus der oberkarbonischen Gaskohle von Nürschan in Böhmen. Sitzungsberichte der Mathematisch-Naturwissenschaftlichen Abteilung der Bayerischen Akademie der Wissenschaften zu München 1924: 3-11.
Brough MC and Brough J 1967. Studies on early tetrapods. III. The genus Gephyrostegus. Philosophical Transactions of the Royal Society B252: 147-165.
Brough MC and Brough J 1967. The Genus Gephyrostegus. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 252 (776): 147–1
Carroll RL 1970. The Ancestry of Reptiles. Philosophical Transactions of the Royal Society London B 257:267–308. online pdf
Carroll RL 1970. The ancestry of reptiles. Philosophical Transactions of the Royal Society B257: 267-308.
Clack JA and Klembara J 2009. An articulated specimen of Chroniosaurus dongusensis and the morphology and relationships of the chroniosuchids. Special Papers in Palaeontology, 81: 15–42.
Danto M, Witzmann F and Müller J 2012. Redescription and phylogenetic relationships of Solenodonsaurus janenschi Broili, 1924, from the Late Carboniferous of Nyrany, Czech Republic. Fossil Record 15 (2) 2012, 45–59.
Klembara J, Clack J, and Cernansky A 2010. The anatomy of palate of Chroniosaurus dongusensis (Chroniosuchia, Chroniosuchidae) from the Upper Permian of Russia. Palaeontology 53: 1147-1153.
Klembara J, Clack J, Milner AR and Ruta M 2014. Cranial anatomy, ontogeny, and relationships of the Late Carboniferous tetrapod Gephyrostegus bohemicus Jaekel, 1902. Journal of Vertebrate Paleontology 34:774–792.
Laurin M and Reisz 1999. A new study of Solenodonsaurus janenschi, and a reconsideration of amniote origins and stegocephalian evolution. Canadian Journal of Earth Sciences 36:1239-1255.
Paton RL Smithson TR and Clack JA 1999. An amniote-like skeleton from the Early Carboniferous of Scotland. Nature 398: 508-513.
Schoch RR, Voig S and Buchwitz M 2010. A chroniosuchid from the Triassic of Kyrgyzstan and analysis of chroniosuchian relationships. Zoological Journal of the Linnean Society 160: 515–530. doi:10.1111/j.1096-3642.2009.00613.x
Smithson TR 1994. Eldeceeon rolfei, a new reptiliomorph from the Viséan of East Kirkton, West Lothian, Scotland. Transactions of the Royal Society of Edinburgh: Earth Sciences 84 (3-4): 377–382.
Smithson TR & Rolfe WDI 1990. Westlothiana gen. nov. :naming the earliest known reptile. Scottish Journal of Geology no 26, pp 137–138.
Tverdochlebova GI 1972. A new Batrachosaur Genus from the Upper Permian of the South Urals, Paleontol. Zh., 1972: 95–103.

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