More on the Origin of Turtles – Lyson et al. 2010

Lyson et al.  (2010 – available online) put together their hypothesis on the origin of turtles. In their abstract, they wrote, “We reanalysed a recent dataset that allied turtles with the lizard–tuatara clade and found that the inclusion of the stem turtle Proganochelys quenstedti  and the ‘parareptile’ Eunotosaurus africanus  results in a single overriding morphological signal, with turtles outside Diapsida.”

Milleretta (RC14 specimen) and the Lyson et al. 2010 tree on the origin of turtles.

Figure 1. Milleretta (RC14 specimen) and the Lyson et al. 2010 tree on the origin of turtles. Note the broad ribs already developing in Milleretta, a sister to Acleistorhinus and Eunotosaurus. On its face this seems like a slam dunk for Eunotosaurus and turtles. However, according to the large reptile tree the origin of turtles parallleled the origin of Eunotosaurus. Missing from the Lyson et al. 2010 analysis is Romeria primus and Stephanospondylus, which are closer to the lineage of turtles. A sister to Romeria primus is the last common ancestor of Eunotosaurus and turtles.

Unfortunately,
Lyson et al. (2010) did not include Romeria primusOrobates (Fig. 2) and Stephanospondylus, three taxa found to be closer to the origin of turtles than Eunotosaurus, a terminal taxon with only one known sister, Acleistorhinus. Unfortunately we have no post-crania for Romeria primus (other than slender manual digits) or Acleistorhinus. That lack of data makes it less obvious how they are related to other taxa, but still the large reptile tree nested them in that fully resolved tree. Stephanospondylus was also the sister to the pareiasaurs, a derived clade previously and correctly associated with turtles, but only at the bases of both clades.

Click to enlarge. These skulls are arranged phylogenetically according to the results recovered from the large reptile tree.

Figure 2. Click to enlarge. These skulls are arranged phylogenetically according to the results recovered from the large reptile tree. This was first published a few days ago.

Would be nice to find the common ancestor of both pareiasaurs and turtles, something a little less turtle-like than Stephanspondylus. For now, Orobates(in yellow, Fig. 2) is the best candidate, and prior to that, Romeria primus (in pink). Orobates and Stephanospondylus are Early Permian. The two turtles are Late Triassic. That gives 60-70 million years to evolve a carapace and plastron, plenty of time for transitional taxa to be discovered in. 

Eunotosaurus

Figure 3. Eunotosaurus, a milleretid not related to turtles, but converged with them in several ways. Actually Eunotosaurus is closer to Acleistorhinus and the Caseasauria, which makes sense if put these two together, like Clark Kent and Superman.

Lyson et al. 2012 did find turtle genes closer to lizard genes, while others did not.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

References
Broom R 1924. On the classification of the reptiles. Bulletin of the American Museum of Natural History 51:39-45.
Geinitz HB and Deichmüller JV 1882. Die Saurier der unteren Dyas von Sachsen. Paleontographica, N. F. 9:1-46.
Gregory WK 1946. Pareiasaurs versus placodonts as near ancestors to turtles. Bulletin of the American Museum of Natural History 86:275-326
Kissel R 2010. Morphology, Phylogeny, and Evolution of Diadectidae (Cotylosauria: Diadectomorpha). Toronto: University of Toronto Press. pp. 185. online pdf
Li C, Wu X-C, Rieppel O, Wang L-T, Zhao L-J 2008. An ancestral turtle from the Late Triassic of southwestern China. Nature 456: 497-501.
Lyson TR, Bever GS, Bhullar B-AS, Joyce WG and Gauthier JA. 2010. Transitional fossils and the origin of turtles. Biology Letters 2010 6, 830-833 first published online 9 June 2010. doi: 10.1098/rsbl.2010.0371
Lyson TR, Sperling EA, Heimberg AM, GauthierJA, King BL, and Peterson KJ 2011. MicroRNAs support a turtle + lizard clade. Biol Lett 2011 : rsbl.2011.0477v1-rsbl20110477.abstract – online news story
Reisz RR and Head JJ 2008. Turtle origins out to sea. Nature 456, 450–451.
Rieppel O and deBraga M 1996. Turtles as diapsid reptiles. Nature 384:453-454.
Rieppel O and Reisz RR 1999. The Origin and Early Evolution of Turtles. Annual Review of Ecology and Systematics 30: 1-22.
Romer AS 1925. Permian amphibian and reptilian remains described as Stephanospondylus. Journal of Geololgy 33: 447-463.
Stappenbeck R 1905. Uber Stephanospondylus n. g. und Phanerosaurus H. v. Meyer: Zeitschrift der Deutschen Geologischen Gesellschaft, v. 57, p. 380-437.
Williston SW 1917. The phylogeny and classification of Reptilies. Journal of Geology 28: 41-421.

wiki/Stephanospondylus

What is Eunotosaurus?

Eunotosaurus was a wide-bellied reptile with extremely broad ribs and eight dorsal vertebrae…which is how one could also describe a turtle. Several specimens are known. Smaller than a human hand, some were preserved curled up.

With Turtles. Then Not. Then With Again.
Watson (1914) originally considered Eunotosaurus a turtle ancestor.

Romer (1956) separated Eunotosaurus from turtles, with no other idea how Eunotosaurus nested.

Cox (1969) nested Eunotosaurus with Captorhinus and kin.

Cisneros et al. (2008) nested Eunotosaurus with Milleretta. (Bravo!!)

Currently Lyson et al. (2010) puts Eunotosaurus and turtles together again. Unfortunately the support values were chiefly below 0.66 and too few taxa were used compared to the large study found here. On the plus side, Eunotosaurus did nest with Acleistorhinus and Milleretta in that study.

Eunotosaurus and its sister taxa, Acleistorhinus and Milleretta RC14.

Figure 1. Eunotosaurus and its sister taxa, Acleistorhinus and Milleretta RC14.

The Closest Sister Taxa to Eunotosaurus
Here, in the large study, Acleistorhinus was the closest sister taxon to Eunotosaurus (Figure 1). Unfortunately, only the skull is known. The next closest sister taxon was the RC14 specimen of Milleretta, which shares the expanded ribs that characterize Eunotosaurus. More Eunotosaurus details can be found at reptileevolution.com

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

References
Watson DMS 1914. Eunotosaurus africanus (Seeley) and the ancestors of the Chelonia, Proceedings of the Zooogical Society of London 11:1011–1020.
Cisneros JC, Rubidge BS, Mason R and Dube C 2008. Analysis of millerettid parareptile relationships in the light of new material of Broomia perplexa Watson, 1914, from the Permian of South Africa. Journal of Systematic Palaeontology 2008 (6): 453–462. doi:10.1017/S147720190800254X
Cox CB 1969.
 The problematic Permian reptile Eunotosaurus. Bulletin of the British Museum of Natural History 18: 167–196.
Keyser AW and Gow CE 1981. First complete skull of the Permian reptile
Eunotosaurus africanus Seeley. South African Journal of Science 77: 417–420.
Gow CE 1997. A reassessment of Eunotosaurus africanus Seeley (Amniota: Parareptilia). Palaeontologia Africana, 34:33–42.
Gow CE and de Klerk B 1997. First record of Eunotosaurus (Amniota: Parareptilia) from the Eastern Cape. Palaeontologia Africana, 34: 27–31.
Lyson TR, Bever GS, Bhullar B-AS, Joyce WG and Gauthier JA 2010. Transitional fossils and the origin of turtles. Proceedings of the Royal Society B Available online 9 June 2010: 830. online pdf
Modesto SP 2000. Eunotosaurus africanus and the Gondwanan ancestry of anapsid reptiles. Palaeontologia Africana, 36:15–20.

wiki/Eunotosaurus