Reptilomorpha to scale

Taxa closer to Reptilia
(e.g. Silvanerpeton) than to Lissamphibia (e.g. Rana) are considered Reptilomorpha  by definition (Säve-Söderbergh 1934). Contra tradition and paradigm (see below), in the large reptile tree (LRT, 1440 taxa) most of the following taxa (Fig. 1) are basal (non-reptle) taxa that fulfill this definition. Some outgroup taxa are also shown along with Silvanerpeton, the last common ancestor of all reptiles (= amniotes) in the LRT.

Figure 1. Members of the Reptilomorpha. starting with Caerorhachis and their proximal outgroups illustrated to scale and in their phylogenetic order from top (primitive) to bottom (derived). Not all reptilomrophs have long limbs and large feet, but these traits are generally not found in non-reptilomorphs. Frogs are important convergent exceptions.

The clade Reptilomorpha
includes all members of the clade Reptilia (= Amniota), but we’re going to focus on the stem amniotes today, basically from Caerorhachis to Gephyrostegus.

According to Wikipedia
“As the exact phylogenetic position of Lissamphibia within Tetrapoda remains uncertain, it also remains controversial which fossil tetrapods are more closely related to amniotes than to lissamphibians, and thus, which ones of them were reptiliomorphs in any meaning of the word. These include the diadectomorphs, seymouriamorphs, most or all “lepospondyls”, gephyrostegids, and possibly the embolomeres and chroniosuchians. In addition, several “anthracosaur” genera of uncertain taxonomic placement would also probably qualify as reptiliomorphs, including Solenodonsaurus, Eldeceeon, Silvanerpeton, and Casineria.”

Most of these taxa
nest within the clade Reptilia in the LRT. Taxon exclusion has been the traditional cause of this problem, something the LRT was designed to take care of with high confidence because all candidates are tested.

Origin of amniotes
Wikipedia reports, “Exactly where the border between reptile-like amphibians (non-amniote reptiliomorphs) and amniotes lies will probably never be known, as the reproductive structures involved fossilize poorly, but various small, advanced reptiliomorphs have been suggested as the first true amniotes, including Solenodonsaurus, Casineria and Westlothiana.”

In the LRT, these three taxa nest well within the Reptilia.
Exactly where the last common ancestor of all living reptiles has been known for several years. Phylogenetic analysis makes this easy. Whichever taxon is the last common ancestor all living mammals, birds, lizards and crocodilians marks the border. Here in the LRT, that taxon is Silvanerpeton (Fig. 1) from the Viséan (Early Carboniferous) with an even earlier genesis because several reptile ingroup taxa are coeval in the Viséan.

Figure 1. Subset of the LRT focusing on basal tetrapods and showing those taxa with lobefins (fins) and those with fingers and toes (feet). Inbetween we have no data. By this cladogram and by definition, Microsauria is a clade within Reptilomorpha. 

Not all reptilomrophs have long limbs and large feet,
but these traits are generally not found in non-reptilomorph basal tetrapods. By convergence, frogs (genus: Rana) are important exceptions. Needless to say, longer, stronger limbs are ideal for terrestrial excursions, despite the fact that some reptiles, like snakes and skinks, get along very well without them.

Earlier we talked about the lack of posterior dorsal ribs in the earliest reptiles. This provided additional space for gravid females to grow amniotic eggs prior to laying them. A deep pelvis permitted the expulsion of larger eggs. A deeper pelvis is found in Eusauropleura (Fig. 1) and more derived taxa. Platyrhinops (Fig. 1), in this regard a reptile-mimic, also had a deep pelvis and long legs by convergence.

The earliest fishapods,  
like Panderichthys and Tiktaalik had a wide flat body with dorsal ribs that were much wider than deep — less chance for tipping while walking. This wide, flat morphology is retained up to Utegenia, when the dorsal ribs start curving to enclose a deeper, narrower torso (by convergence with other taxa, like Ichthyostega. At the same time the orbits moved laterally.

Not all reptilomorphs are small,
but the earliest reptilomorphs were no larger than than the juveniles of their ancestor, Greererpeton (Fig. 1), distinct from the giant ‘amphibians’ we looked at earlier.

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References
Carroll RL 1991. The origin of reptiles. Pp. 331–53 in Schultze H-P and Trueb L editors. Origins of the higher groups of tetrapods — controversy and consensus. Ithaca: Cornell University Press.
Gauthier J, Kluge AG and Rowe T 1988. The early evolution of the Amniota. In The Phylogeny and Classification of the Tetrapods: Volume 1: Amphibians, Reptiles, Birds. Edited by MJ Benton. Clarendon Press, Oxford, pp. 103–155.
Ruta M, Coates MI and Quicke DLJ 2003. Early tetrapod relationships revisited. Biological Reviews 78 (2): 251–345.
Säve-Söderbergh G 1934. Some points of view concerning the evolution of the vertebrates and the classification of this group. Arkiv för Zoologi. 26A: 1–20.

The conquest of the land: 9 or 10x and counting…

Traditional paleontology 
has given us a picture of a more or less simple ladder of stem tetrapod evolution that had its key moment when an Ichthyostega-like taxon first crawled out on dry land. Then, according to the widely accepted paradigm, certain lineages returned to the water while others ventured forth onto higher and drier environs.

By contrast,
The large reptile tree (LRT, 1033 taxa) documents a bushier conquest of land, occurring in at least seven Devonian waves until the beachhead was secured by our reptile ancestors.

Dr. Jennifer Clack and her team have shown us that fish/amphibians can have limbs (Acanthostega and Ichthyostega) and not be interested in leaving the water. That comes later and later and, well, seven times all together.

Figure 1. Colosteus relatives according to the LRT scaled to a common skull length. Their sizes actually vary quite a bit, as noted by the different scale bars. Only Pholidogaster and Colosteus are taxa in common with traditional colosteid lists.

The first wave:
simple small fins to simple small limbs
Arising from lobe-fin fish with one nostril migrating to the inside of the mouth, like Osteolepis, the much larger collosteid, Pholidogaster, had small limbs with toes. The smaller, but equally scaly and eel-like Colosteus, reduced those limbs to vestiges, showing they were not that important for getting around underwater in that wriggly clade. Neither shows signs of ever leaving the water and phylogenetically neither led to the crawling land tetrapods. However, like the living peppered moray eel (Gymnothorax pictus, Graham, Purkis and Harris 2009) in search of crabs, these taxa might have made the first landfall without limbs. See terrestrial moray eel video here. 

Figure 2. Greererpeton reduced to a blueprint of body parts. Here there may be one extra phalanx on pedal digit 5 and one missing on pedal digit 2 compared to sister taxa. So an alternate is shown with that repair. The skulls at left are juveniles.

The second wave:
fins to limbs on long flattened bottom feeders
Fully limbed Greererpeton and Trimerorhachis were derived from finny flat taxa like Panderichthys and Tiktaalik. Both Greererpeton and Trimerorhachis were likewise flat- and long-bodied aquatic forms that seem unlikely to have been able to support themselves without the natural buoyancy of water. Their descendants in the LRT likewise look like they were more comfortable lounging underwater like living hellbenders (genus Cryptobranchus. According to Wikipedia: “The hellbender has working lungs, but gill slits are often retained, although only immature specimens have true gills; the hellbender absorbs oxygen from the water through capillaries of its side frills.”  Only rarely do hellbenders leave the water, perhaps to climb on low pond rocks. If the Greererpeton clade was similar, this would have been the second meager and impermanent conquest of the land. And they would not have gone too far from the pond.

Figure 3. Pederpes is a basal taxon in the Whatcheeria + Crassigyrinus clade.

The third wave:
the Pederpes/Eryops clade experimented with overlapping ribs.
Arising from shorter Ossinodus and Acanthostega, a clade that included Pederpes, Ventastega, Baphetes and Eryops arose. This clade looks quite capable of conquering the land for the third time. Their overlapping ribs helped support their short backbone, for the first time lifting their belies off the substrate when doing so, matching Middle Devonian tracks. Some clade members, like Crassigyrinus (with its vestigial limbs) and Saharastega (with its flattened skull) appear to have opted for a return to a watery environment. And who could blame them? In any case, their big lumbering bodies were not well adapted to clambering over dry obstacles, like rocks and plants, that made terrestrial locomotion more difficult. And the biggest best food was still in the water. No doubt limbs helped many of them find new ponds and swamps when they felt the urge to do so, like living crocs. And they probably left the water AFTER some of the smaller and more able taxa listed below.

Figure 4. Proterogyrinus had a substantial neck.

The fourth wave:
a longer neck and a smaller head gave us Proterogyrinus.
Ariising from fully aquatic fish/amphibians with overlapping ribs, like Ichthyostega, basal reptilomorphs, like low-slung, lumbering Proterogyrinus took the first steps toward more of a land-living life. The nostrils shifted forward, but were still tiny, at first. Bur the ribs were slender without any overlap. Perhaps this signaled improvements in lung power. Larger nostrils appeared in more devoted air breathers, like Eoherpeton and Anthracosaurus. All these taxa were still rather large and lumbering and so were probably more at home in the water.

Figure 5. Eucritta in situ and reconstructed. Note the large pes in green.

The fifth wave:
goes small, gets longer legs and gives us Seymouria.
Eucritta is the first of the small amphibians with longer limbs relative to trunk length. This clade also arises from Ichthyostega-like ancestors. One descendant clade begins with a several long-bodied, short-legged salamander-like taxa. Discosauriscus is one of these. It begins life in water, but grows up to prefer dry land. Seymouria is the culmination of this clade. 

Figure 6. Utegenia nests as a sister to Diplovertebron.

The sixth wave:
gives us salamanders and frogs.
Still tied to the water for reproduction and early growth with gills, this clade arises from the seymouriamorph/lepospondyl Utegenia, a short-legged, flat-bodied aquatic taxon. That plesiomorphic taxon gives rise to legless Acherontiscus and kin including modern caecilians. Reptile-mimic microsaurs, like Tuditanus arise from this clade. So do modern salamanders, like Andrias and long-legged, short bodied frogs, like Rana. Their marriage to or divorce from water varies across a wide spectrum in living taxa.

Figure 7. Various stem amniotes (reptiles) that precede Tulerpeton in the LRT. So these taxa likely radiated in the Late Devonian. And taxa like Acanthostega and Ichthyostega are late-survivors of earlier radiations documented by the earlier trackways.

The seventh wave:
gives us the amniotic egg and the reptiles that laid them.
No one should have ever said you have to look like a typical reptile to lay an amnion-covered egg. And if they did, they were not guided by a large gamut phylogenetic analysis. This clade become fully divorced from needing water for reproduction, but basal members still liked the high humidity and wet substrate of the swamp. Arising from basalmost seymouriamorphs like Ariekanerpeton, stem reptiles included Silvanerpeton. These were small agile taxa with relatively long legs that would have had their genesis in the Late Devonian. Their first appearance in the fossil record was much later. The development of the amnion-enclosed embryo may have taken millions of years. The first phylogenetic reptiles appear in the form of amphibian-like Gephyrostegus and Tulerpeton in the Late Devonian, which still had six fingers and scales, but these lacked layers typically found in more fish-like taxa.

So the conquest of the land
by stem and basal tetrapods appears to have occurred seven times, according to the LRT, from distinct clades that were more or less ready to do so and in different ways. And, of course, odd extant fish, like the Peppered moray eel (wave 8) and the mudskipper, (wave 9) and maybe even snakes from stem sea snakes (wave 10) continue this tradition. What will THEY eventually evolve into, given enough time?

References
Clack JA 2006. The emergence of early tetrapods. Palaeogeography Palaeoclimatology Palaeoecology. 232: 167–189.
Clack JA 2009. The fin to limb transition: new data, interpretations, and hypotheses from paleontology and developmental biology. Annual Review of Earth and Planetary Sciences. 37: 163–179.
Coates MI 2014. The Devonian tetrapod Acanthostega gunnari Jarvik: Postcranial anatomy, basal tetrapod interrelationships and patterns of skeletal evolution. Earth and Environmental Science Transactions of the Royal Society of Edinburgh.
Coates MI and Clack JA 1990. Polydactly in the earliest known tetrapod limbs. Nature 347: 66-69.
Graham NAJ, Purkins SJ and Harris A 2009. Diurnal, land-based predation on shore crabs by moray eels in the Chagos Archipelago. Coral Reefs 28(2): 387–397. Online here.
Jarvik E 1952. On the fish-like tail in the ichtyhyostegid stegocephalians. Meddelelser om Grønland 114: 1–90.

wiki/Acanthostega

Anthracosaurus: beware the chimaera!

Figure 1. The complete skull of Anthracosaurus greatly resembles its relative, Neopteroplax.

Anthracosaurus russelli (Huxley 1863, Panchen 1977, Clack 1987; Westphalian, Late Carboniferous, 310 mya, skull length 40cm; Figs. 1, 2) was originally considered a labyrinthodont. The wide, yet pointed, triangular skull and tall orbits recall traits found in labyrinthodonts, like Sclerocephalus, and in the basal tetrapod, Tiktaalik. Here, in the large reptile tree (LRT, 967 taxa),  Anthracosaurus nests with Neopteroplax (Fig. 3) as a derived embolomere, the clade that likely gave rise to Seymouriamorpha, Lepospondyli and Reptilia

At least one orbit
in Anthracosaurus has an inverted teardrop shape. The marginal and palatal fangs are quite large. Although flattened in dorsal view, comparisons suggest the jaw margin was convex, as in Neopteroplax.

Based on its size and nesting,
Anthracosaurus developed a labyrinthodont-like skull by convergence because Proterogyrinus is basal in the Embolomeri. Those giant marginal and palatal fangs indicate a predatory niche.

Figure 2. Left: Anthracosaurus chimaera from Clack 1987. Right: Older tracing in dorsal view of the complete skull and palatal view attributed to Anthracosaurus from an online photo. The narrower skull is made of several different specimens (chimaera) and produces a loss of resolution in the LRT.

Clack 1987
illustrated a lateral and dorsal view of Anthracosaurus (Fig. 2) based on a chimaera of specimens. Unfortunately, plugging that data into the LRT produced loss of resolution over several nodes. Using the older single skull in dorsal view had no such problems.

We looked at the problems chimaera taxa produce
earlier here, and in six blogs that preceded that one.

Figure 3. Neopteroplax has a skull quite similar to the older single skull of Anthracosaurus and they nest together in the LRT.

The clade Anthracosauria has had problems
From Wikipedia: “Gauthier, Kluge and Rowe (1988) defined Anthracosauria as ‘Amniota plus all other tetrapods that are more closely related to amniotes than they are to amphibians” (Amphibia in turn was defined by these authors as a clade including Lissamphibia and those tetrapods that are more closely related to lissamphibians than they are to amniotes).”

In this definition non-amniote Anthracosauria does not include Anthracosaurus, but only Silvanerpeton and Gephyrostegus in the LRT because more basal taxa are also basal to amphibians.

“Similarly, Michel Laurin (1996) uses the term in a cladistic sense to refer to only the most advanced reptile-like amphibians. Thus his definition include the (Diadectomorpha and Solenodonsauridae) and the amniotes.”

In the LRT Diadectomorpha and Solenodonsauridae are amniotes.

“As Ruta, Coates and Quicke (2003) pointed out, this definition is problematic, because, depending on the exact phylogenetic position of Lissamphibia within Tetrapoda, using it might lead to the situation where some taxa traditionally classified as anthracosaurs, including even the genus Anthracosaurus itself, wouldn’t belong to Anthracosauria.

Indeed! And that happened in the LRT.

Laurin (2001) created a different phylogenetic definition of Anthracosauria, defining it as “the largest clade that includes Anthracosaurus russelli but not Ascaphus truei”.

In the LRT Ascaphus, the tailed frog, is derived from the large clade, the embolomeri, that includes Anthracosaurus. However the small clade that includes just Anthracosaurus and Neopteroplax does not include the tailed frog.

“However, Michael Benton (2000, 2004) makes the anthracosaurs a paraphyletic order within the superorder Reptiliomorpha, along with the orders Seymouriamorpha and Diadectomorpha, thus making the Anthracosaurians the “lower” reptile-like amphibians. In his definition, the group encompass the Embolomeri, Chroniosuchia and possibly the family Gephyrostegidae.”

In the LRT the Embolomeri are basal to Eucritta and the Seymouriamorpha, which are basal to the Reptilia (= Amniota) and Lepospondyli (including Amphibia). The Chroniosuchia and Gephyrostegus are both amphibian-like reptiles in the LRT.

The clade Reptilomorpha suffers the same definition problems.
As Wikipedia reported, “As the exact phylogenetic position of Lissamphibia within Tetrapoda remains uncertain, it also remains controversial which fossil tetrapods are more closely related to amniotes than to lissamphibians, and thus, which ones of them were reptiliomorphs in any meaning of the word.”

Wouldn’t it be great if someone could put together
a large gamut phylogenetic analysis that could settle all those controversial issues?

References
Clack JA 1987. Two new speciemens of Anthracosaurus (Amphibia: Anthracosauria) from the Northumberland coal measures. Palaeontology 30(1):15-26.
Huxley TH 1863. Description of Anthracosaurus russelli, a new labyrinthodont from the Lanarkshire coalfield. Quartery Journal of the Geological Society 19:56-58.
Panchen AL 1975. A new genus and species of anthracosaur amphibian from the Lower Carboniferous of Scotland and the status of Pholidogaster pisciformis Huxley. Philosophical Transactions of the Royal Society of London, B. 269: 581-640.
Panchen AL 1977. On Anthracosaurus russelli Huxley (Amphibia: Labyrinthodontia) and the family Anthracosauridae. Philosophical Transactions of the Royal Society B. 279 (968): 447–512.

 

History of reptile Interrelationship hypotheses: Meckert’s PhD thesis

There is a long history
of workers creating hypotheses of reptile interrelationships going back to the mid 18th century (Carl von Linneaus 1758). That history, up until 1995 (Laurin and Resiz 1995 and Meckert 1995), was summarized by Dirk Meckert in his PhD thesis, which otherwise  concentrated on all available specimens of Barasaurus. You can download that thesis here online and read that short but fascinating history for yourself.

Some interesting notes arise from Meckert’s short history:

1. Some studies united pareiasaurs and turtles. Others did not.
2. Other studies united pareiasaurs, diadectids and procolophonids (which happened here just yesterday). Meckert wrote: “The Procolophoniformes contain Procolophonia and Testudinomorpha as sister-groups. Testudines are the sister-group of Pareiasauria within the Testudinomorpha.”
3. Mesosaurs are commonly considered of uncertain affinities. But not here.
4. Many prior studies had the synapsids branch off first. That is incorrect as shown here.
5. No prior studies recognized the original dichotomy of lepidosauromorphs and archosauromorphs.
6. No prior studies recognized Gephyrostegus bohemicus as a sister to the basalmost amniote.
7. Diadectomorpha have been nested in and out of the Amniota. They’re in here.

No studies prior to reptileevolution.com
have included as many as 571 individual species as taxa, not counting the therapsid tree (with 52 additional taxa) and pterosaur tree (with 228 additional taxa) for a total of 851 taxa.

Other studies more recent than 1995
(not included in Meckert’s history) include

1. http://www.palaeos.org/Reptilia and http://palaeos.com/vertebrates/amniota/reptiles.html
2. http://whozoo.org/herps/herpphylogeny.html
3. https://en.wikipedia.org/wiki/Amniote as determined by Benton, M.J. (2004). Vertebrate Paleontology. Blackwell Publishers. xii–452.
4. University of Maryland (John Merck)
5. online pdf, Amniote Origins and Nonavian Reptiles
6. YouTube video by Walter Jahn
7. Tree of Life
8. Hedges 2012
9. Gauthier, Kluge and Rowe 1988 online
10. Hill 2005
11. Mikko’s phylogeny archive
12. ReptileEvolution.com
13. Let me know if I missed any. I’ll add them here.

A while back
we looked at the differences between astronomy and paleontology. As noted earlier, time is never of the essence in paleontology — and that extends to idea acceptance. So many hypotheses of reptile interrelationships are still floating around out there. A definitive and all encompassing demonstration, like the large reptile tree, will probably just float forever with the other several dozen hypotheses out there, hashed, rehashed and rehashed again without end.

This is one of the frustrations of paleontology. And many think it is largely ego driven.

On that note
In astronomy the data, be it observation or spectral analysis, is immediate and widespread. You just have to look up with the right tool in the right direction. Or study the shared data (photos, etc.) Everyone can confirm the observation.

In paleontology the data comes out piecemeal, in low resolution, or imprecise tracings, not from every angle of view. Some key parts are lost and others are hidden beneath other bones or matrix. Sometimes you have to assemble dozens or hundreds of specimens for a proper study. No one is interested in confirming observations or analyses perhaps for years if ever. They’re all too busy with their own projects. Checking the characters and scores of an analysis can take weeks, months or years (as long as it took to build originally), and to do so requires the same amount of globe-hopping to see all the specimens in all the museums. No one is going to do that. They’d rather be making their own discoveries… and adding their taxa to established trees created by hungry PhD candidates, like Dirk Meckert in 1995, done at the nadir or advent of their experience.

The paleo-mantra remains: you must see the specimen!
And even that is no guarantee.

And if you want to break a paradigm or two,
like Ostrom did in the 1960s, you might have to wait for widespread (but never universal) acceptance. Paleontologists like their paradigms. They don’t like to give them up.

References
Benton MJ 2004. Vertebrate Paleontology. Blackwell Publishers. xii–452.
Carroll RL 1988. Vertebrate Paleontology and Evolution, WH Freeman & Co.
Laurin M and Reisz R 1995. A reevaluation of early amniote phylogeny. Zoological Journal of the Linnean Society, 113: 165–223.
Linnaeus C 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.
Meckert D 1995. The procolophonid Barasaurus and the phylogeny of early amniotes. PhD thesis McGill University. Online Barasaurus dissertation

 

 

 

News at the base of the Amniota, part 8: The list of Viséan amniotes has grown.

Earlier (in seven prior blog posts) we looked at the new basalmost amniotes and how they evolved.

With the news
that the Amniota (and amniote eggs) extended back to the Viséan (326-345 mya) let’s take a look at those basalmost amniotes along with the genera that may have been more primitive, but survived more than 30 million years later, into the Westphalian (303-311 mya) and beyond to the Early and Late Permian. That’s a stretch of 80 million years for the most successful taxa. And what made them so successful? Or were they all just as successful, just not found yet in higher strata?

Figure 1. Click to enlarge. Basal amniotes to scale colorized according to the time strata in which their fossils were found. Visean, yellow; Namurian, pink; Westphalian, blue; Permian, tan.

It’s well worth remembering
at this point that in similar fashion, basal primates, like lemurs, also co-exist today with derived primates, like apes and humans. So that happens. At least some of these basalmost amniotes (the Permian forms, like Utegenia, Fig. 1) developed successful traits so well matched to their own niche they survived for tens of millions of years thereafter.

Also remember
that fossilization is a rare event. Even more rare is the discovery of rare fossils. So we’re very lucky to have even single examples of these taxa. They were probably more widespread both across the globe and through time.

Two important points
1) We don’t find amniotes prior to the Viséan. So these Viséan amniotes  (Fig. 1) are likely the earliest representatives of their kind.

2) The Viséan is a short 15 to 35 million years after the very first tetrapods developed limbs from fins some 360 million years ago in the Late Devonian. So evolution was rapid during those first 15 million years. Not so rapid for the next 80 million years, at least for certain taxa.

That’s exciting to think about.

News at the base of the Amniota, part 6: Cladogram of basal lepidosauromorpha

Yesterday we looked at primitive archosauromorpha at the base of the Amniota. Today we’ll look at basal lepidosauromorpha.

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 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.

As mentioned earlier, the Amniota is divided at its base into Lepidosauromorpha (taxa closer to lepidosaurs) and Archosauromorpha. Gephyrotegus bohemicus (Fig. 2) is the last common ancestor and Silvanerpeton is the outgroup anamniote.

Figure 3. Urumqia liudaowanensis (Zhang et al. 1984) ~20 cm snout-vent length, Lower Permian. Formerly considered a sister to Utegenia, an anamniote, it now nests as the basalmost of all lepidosauromorpha.

Urumqia liudaowanensis (Zhang et al. 1984) ~20 cm snout-vent length, Upper Permian, was originally considered a discosaurid seymouriamorph close to Utegenia. Here (Fig. 1) it nests at the base of the lepidosauromorph reptiles despite its late appearance in the fossil record. Note the gastalia are much wider than the dorsal ribs, likely to retain large eggs in gravid females. Distinct from G. bohemicus, Urumqia had shorter limbs, longer posterior dorsal ribs and a robust tail with elongate caudals. The palate included a larger suborbital fenestra, not homologous to later taxa with this trait. The cheek included a small lateral temporal fenestra. The carpals and tarsals are poorly ossified.

Figure 4. Bruktererpeton, a gephyrostegid and a basal lepidosauromorph amniote.

Bruktererpeton fiebigi — (Boy and Bandel, 1973; Fig. 3) is an older (Namurian/ Bashkirian, 320 Ma) sister to Gephyrostegus bohemicus (Ruta, Jefferey and Coates, 2003; Klembara et al., 2014). The pectoral girdle and limbs are more gracile. The scapula is taller. The intercentra are smaller. Other traditional amniote traits, if present, are not preserved.

Figure 5. Thuringothyris. A basal lepidosauromorph.

Thuringothyris  mahlendorffae — (Boy and Martens, 1991; type: MNG 7729; (Müller et al., 2006) referred MNG 10183; Artinskian, Early Permian, 280 Ma) is half the size of Bruktererpeton and documents all traditional amniote traits. Note the derived shape of the humerus and the reduced intercentra.

Traditional amniote traits include:

1. loss/fusion of the intertemporal
2. absence of the otic notch
3. loss/reduction of palatal fangs
4. appearance/expansion of the transverse flange of the pterygoid
5. loss of labyrinthine infolding of the marginal teeth
6. reduction of the intercentra
7. addition of a second sacral vertebra
8. narrowing and elongation of the humeral shaft
9. appearance of the astragalus from fused tarsal elements.

Figure 6. Cephalerpeton. A basal lepidosauromorph.

Cephalerpeton — (Gregory 1948) representing a new sister clade to the Captorhinomorpha, Cephalerpeton had an elongate humuerus with a narrow shaft. The much larger and later Reiszorhinus is a sister.

Figure 7. Two specimens of Concordia, a basal lepidosauromorph.

Concordia — (Müller & Reisz 2005, Stephanian, Late Pennsylvanian, Carboniferous, 4 cm skull length) was considered the oldest known captorhinid, but here (Fig. 1) it nests with Cephalerpeton as a sister to captorhinids.

Figure 8. Romeria texana, a basal capitorhinomorph, lepidosauromorph, amniote.

Romeria texana —(Price1937) Artinskian, Early Permian, ~280 mya, ~25 mm skull length, was the basalmost captorhinid. Here (Fig, 1) the skull is wider and flatter.

Figure 9. Saurorictus, a basal lepidosauromorph in the lineage of Milleretta, compared to sister taxa.

Saurorictus — (Modesto and Smith 2001, SAM PK-8666, skull length ~2.2 cm, estimated total length 15 cm, Late Permian), derived from a sister to Thuringothyris, Concordia and Cephalerpeton, Saurorictus is the taxon basal to all other lepidosauromorpha including diadectomorpha, chelonia and lepidosauria. (sorry, cut off from bottom of cladogram, Fig. 1). It was considered the most complete captorhinid from the Late Permian.

Size comparisons

Figure 10. Basal amniotes to scale. Click to enlarge.

Here, Fig. 10, there is a size reduction in ‘second generation’ basal amniotes/basal lepidosauromorpha. You’ll note that several former anamniotes now nest within the amniota. They were judged anamniotes by the skeletal traits, not by their phylogenetic nesting, which has not been adequately tested until now.

References
Boy JA and Bandel K 1973. Bruktererpeton fiebigi n.gen.n.sp. (Amphibia: Gephyrostegida). Der erste Tetrapode aus dem Rheinisch-Westfälischen Karbon (Namur B; W-Deutschland). Palaeontographica 145: 39–77.
Boy JA and Martens T 1991. Ein neues captorhinomorphes Reptil aus dem thüringischen Rotliegend (Unter-Perm; Ost-Deutschland). Palaeontologische Zeitschrift 65 (3-4): 363–389.
Gregory JT 1948. The structure of Cephalerpeton and affinities of the Microsauria. American Journal of Science 246:550–568
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.
Müller J, Berman DS, Henrici AC, Martens T and Suminda S 2006. The basal reptile Thuringothyris mahlendorffae (Amniota:Eureptilia) from the Lower Permian of Germany. Journal of Paleontology 80:726-739.
Müller J and Reisz RR 2005. An early captorhinid reptile (Amniota: Eureptilia) from the Upper Carboniferous of Hamilton, Kansas. Journal of Vertebrate Paleontology. 25(3): 561-568.
Price LI 1937. Two new cotylosaurs from the Permian of Texas. Proceedings of the New England Zoölogical Club 16:97-102.
Zhang F, Li Y, and Wan X. 1984. A new occurrence of Permian seymouriamorphs in Xinjiang, China. Vertebrate Palasiatica22(4):294-306.

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.

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 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).

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 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.

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 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 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 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 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.

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.

News at the base of the Amniota, part 4: Keratinized epidermal scales

Earlier here, here and here we looked at various aspects of life for basal amniotes in the Viséan to the Westphalian (340-310 mya). Today we’ll look at another trait common to basal amniotes.

Figure 1. Amniote scales from Didelphis (opossum, background) and Iguana. They probably had their origin in the Viséan as basal amniotes spent less and less time in water and needed a form of waterproofing to avoid desiccation.

Phylogenetic Miniaturization and the Genesis of Keratinized Scales
Keratinized scales (Fig. 1) more or less insulate many living amniotes and all living reptiles from evaporative water loss. Anamniotes (frogs, etc.) don’t have a waterproof skin, but may have scattered osteoderms. While many mammals replace scales with fur and birds replace scales with feathers, keratinized scales are retained on the tail of the opossum and the feet of all birds. Scalation also protects against ant and termite bites.

Figure 2. A new reconstruction of Gephyrostegus bohemicus, this most primitive amniote preserves dorsal dermal scales. Ossified ventral scales are more common and sometimes transformed into gastralia rods.

Scale origins
Basal gnathostomes (basal fish) had ossified skin first and an ossified skeleton later. In teleosts and tetrapods the integumentary (skin) skeleton has undergone widespread reduction and/or modification (Vickaryous and Sire 2009). Sarcopterygians (stem tetrapods) had cosmoid scales, characterized by an intrinsic, interconnected canal system with numerous flask-shaped cavities and superficial pores. In Ichthyostega and other basal tetrapods, dentine, enameled, guanine and pore-canal systems were lost, leaving bone (osteoderms) as the remaining dermal element wherever present. Osteoderms are structurally quite variable and are found in a wide variety of tetrapods, including amphibians. Often they have been lost and independently regained. Temnospondyl amphibians, like Greererpeton, have a combination of thin and overlapping scales, granular pellets, and/or robust plates.

Amniote scale origins
While scales or their impressions are rarely preserved, Carroll (1969) reported some indication of dorsal epidermal scales in Gephyrostegus bohemicus (Fig.2). In Cephalerpeton Carroll and Baird (1972) reported that skin impressions had a slightly pebbly texture, but without evidence for discrete scales. Brough and Brough (1967) found ventral scales in tiny Brouffia (their specimen no. 1) and provided a similar description for the developing gastralia of Gephyrostegus. Not sure why the first and most substantial scales first appeared ventrally, rather than dorsally, where the sun shines, except that the ventral surface is in contact with the substrate.

Figure 2. Reptile hatchling about actual size. A larger surface-to-volume ratio increases the danger from desiccation unless ‘waterproofed’ with scales.

The importance of scales
In basalmost amniotes forays onto land likely increased in duration. Dermal protection from desiccation is more important in smaller amniotes due to their larger surface-to-mass ratio (Hedges and Thomas, 2001). This is especially applicable to hatchlings and juveniles, some of which may have been less than 2 cm in snout/vent length because the adults were so small. Basal amniote juveniles would have rivaled certain microsaur juveniles as the smallest tetrapods of their day, and perhaps they competed in similar niches. Scales may have given the advantage to reptiles.

With regard to microsaurs
Carroll and Baird (1968) reported, “Extremely delicate dorsal scales of elongate-oval shape are present between (and occasionally overlapping) the ribs of the better-ossified United States National Museum specimen [of the microsaur Tuditanus]. If allowance is made for their insubstantial nature, the scales of Tuditanus are essentially similar to those of ther Carboniferous microsaurs (Carroll, 1966). There is no evidence of rodlike ventral scales such as occur in other lepospondyls (Baird, 1965), or wheat-shaped gastralia like those of primitive reptiles.”

Going one step further in the middle Jurassic, tiny basal mammals traded scales for insulating hair and thicker fur (assumed by phylogenetic bracketing). On the other hand, the first Jurassic dinosaurs to preserve protofeathers, like slender Sinosauropteryx (Ji and Ji, 1996), were a meter long, so not tiny. Basal volant birds became progressively smaller.

References
Baird D 1965. Paleozoic lepospondyl amphibians.  American Zoologist 5: 287-294.
Brough MC and J Brough 1967. The Genus Gephyrostegus. Philosophical Transactions of the Royal Society London, Series B, Biological Sciences 252:47–165.
Carroll RL 1966. Microsaurs from the Westphalian B of Joggins, Nova Scotia. Proceedings of the Linnean Society of London 177: 63-97.
Carroll RL 1969. Problems of the origin of reptiles. Biological Reviews 44:393–431.
Carroll RL and Baird D 1968. The Carboniferous amphibian Tuditanus (Eosauravus) and the distinction between microsaurs and reptiles. American Museum novitates 2337: 1-50.
Carroll RL. and D Baird 1972. Carboniferous stem-reptiles of the family Romeriidae. Bulletin of the Museum of Comparative Zoology 143:321–363.
Hedges SB and R Thomas 2001. At the lower size limit in amniote vertebrates: A new diminutive lizard from the West Indies. Caribbean Journal of Science 37:168–173.
Ji Q and S Ji 1996. On the discovery of the earliest bird fossil in China (Sinosauropteryx gen. nov.) and the origin of birds. Chinese Geology 10(233): 30–33.
Vickaryous MK and Sire J-Y 2009. The integumentary skeleton of tetrapods: origin evolution, and development. Journal of Anatomy 214:441-464. online here.

News at the base of the Amniota, part 3: The amniotic egg

Earlier we looked at the base of the amniota and the phylogenetic miniaturization that preceded and succeeded basalmost amniotes. Today we’ll take a closer look at the one key trait that defines the Amniota.

Eggs and Embryonic Development
All morphology aside, the single key trait that defines the Amniota is the production of eggs surrounded by extraembryonic membranes and large enough to sustain the development of the developing embryo until it hatches beyond the gilled aquatic stage. Initially such an egg must have been small enough to maintain its shape and integrity out of water during the gradual evolution of those extraembryonic membranes (Carroll, 1969).

Phylogenetic bracketing shows the evolution of the amniotic egg had its genesis in the Viséan (~345 Ma), likely with a sister to Silvanerpeton and Gephyrostegus bohemicus, (the latter known from the Westphalian, 310 mya). Earlier and more derived amniotes are also found in Viséan strata. These include Westlothiana, Casineria and Eldeceeon (Fig. 1). So the origin of the amniotic egg precedes them all.

The anamniote outgroup taxa, Seymouria, Kotlassia and Utegenia (Fig. 1), all known from much later time periods (Permian), had juveniles with external gills (Laurin, 1996; Klembara et al., 2007), and so did not produce amniotic eggs. None of the recovered basal amniotes had juveniles with gills and sensory grooves. Carroll and Baird (1972) considered the small basal amniote Brouffia (Westphalian, Fig. 1) a juvenile. It had no external gills or sensory grooves. Klembara et al. (2014) considered Gephyrostegus watsoni (Fig. 1) a juvenile anamniote, but it, likewise, has no external gills or sensory grooves. Rather it nests between Eldeceeon and Solenodonsaurus in the Archosaurmorpha branch of the Amniota.

Figure 1. Basal amniotes to scale. Click to enlarge.

Basalmost amniotes share three skeletal traits
that indicate larger eggs were likely being produced:

1. reduction to loss of the posterior dorsal ribs permitting expansion of the posterior torso in gravid females;
2. greater depth of the pelvic opening permitting the passage of larger eggs; and
3. unfused pelvic elements providing more pelvic flexibility during egg laying.

Amniotes more derived than G. bohemicus also develop a second sacral vertebra. Since these ‘second generation’ basal amniotes are generally much smaller overall with shorter limbs (Fig. 1), the second sacral rib comes as something of a surprise—unless it was used to help support the greater weight and mass of gravid females.

Certain amniote clades also transform their ossified ventral dermal scales to become elongate gastralia. Perhaps this also helped support the greater weight and width of the egg mass while gravid.

Only female basal amniotes?
Notably, no gender differences have been identified in basal amniote skeletons. Either basal male amniotes also lacked posterior dorsal ribs and had a deeper pelvic opening and/or basal amniotes reproduced by parthenogenesis (reproduction without males), as certain living lizards do (Lutes, et al., 2010). It could go either way.

Figure 2. Click to enlarge. Gephyrostegus watson (Westphalian, 310 mya) in situ and reconstructed. The egg shapes are near the hips as if recently laid. A few insects appear in the matrix. The carpals and tarsals are present, just displaced. So are the tail chevrons. The embryo (E) is hypothetical based on egg shape and size.

Westphalian amniote eggs?
In the basal amniote Gephyrostegus watsoni (Fig. 2, but this taxon needs a new name because it doesn’t nest with the holotype of Gephyrostegus) eight irregular flattened sphere shapes, each 5mm in diameter (five percent of the adult snout/vent length), appear dorsal to the open ‘lumbar’ area. If they were eggs they are the right size to pass through the pelvic opening. Preserved beyond the confines of the mother’s abdomen, the mother could have moved slightly just after depositing her eggs, shortly before burial. No embryonic skeletons should be expected to appear within such eggs. Instead embryos would have developed after egg deposition, as in many living reptiles. No calcified shell should be expected at this early stage of egg evolution. Examples of similar jelly-like soft tissue preservation in the fossil record are known, as in the Triassic lepidosaur, Cosesaurus (Fig. 4), preserved with a medusa (Ellenberger and de Villalta, 1974).

Click to enlarge and see rollover image. Here DGS, digital graphic segregation, enabled the identification of many more bones than firsthand observation, including the displaced carpals and tarsals, along with a few insects and egg-shapes.

Hatchling size
From 1 cm diameter egg sizes (estimated from pelvic openings) curled up Gephyrostegus bohemicus hatchlings would have been ~2.6 cm in length or one-eighth (12 percent) the size of the mother (Fig. 1).

Figure 3. Click to enlarge and see the rollover. Eldeceeon with a strangely expanded belly (defined by gastralia/scales) that could have contained a load of eggs, traced in green here.

A gravid amniote in the Viséan?
The Eldeceeon holotype was preserved with an oddly expanded belly (Fig. 3). Perhaps this was also a gravid female (Fig. 7) with an egg load that pushed out her ossified ventral scales during postmortem decay and/or crushing. I’ve traced some possible eggs shapes found in the matrix.

Smaller ‘second generation’ basal amniotes, like Westlothiana and Casineria (Fig. 3), would have had proportionately smaller eggs.

Figure 4. Extant lizards, A. gravid, B. in the process of laying eggs, C. with egg clutch.

Living examples of gravid females
Extant lizards (Fig. 4) show the extent of belly-stretching in gravid (pregnant) females and the relatively large size of their eggs. A clutch can be about the size of the mother’s eggless torso.

Basal amniote paleobiology
With short, sprawling fore limbs, a weak tail and a large head, Gephyrostegus watsoni (Fig. 2) was likely slow and secretive, like the living Sphenodon, both in leaf litter and in shallow puddles. This would apply even more so to massively burdened gravid females (Fig. 4). Without obvious defenses or weapons, the key to basal amniote success appears to have been an increase in the production of large eggs laid safely out of predator-filled swamps. The East Kirkton (Viséan) and Nyrany Basin (Westphalian) environments were swampy coal forests, so these would have provided the humid air, damp earth, wet leaf litter and abundant puddles needed for basal amniotes to slowly evolve keratinized skin and membrane enclosed eggs. The large orbit of basal amniotes suggests a nocturnal niche. Perhaps they hid and slept during daylight hours, avoiding evaporative sunlight and diurnal predators.

More later.

References
Carroll RL 1969. Problems of the origin of reptiles. Biological Reviews 44:393–431.
Carroll RL and D Baird 1972. Carboniferous stem-reptiles of the family Romeriidae. Bulletin of the Museum of Comparative Zoology 143:321–363.
Ellenberger P and JF de Villalta 1974. Sur la presence d’un ancêtre probable des oiseaux dans le Muschelkalk supérieure de Catalogne (Espagne). Note preliminaire. Acta Geologica Hispanica 9:162–168.
Klembara J, DS Berman, AC Henrici, A Cernansky, R Werneburg and T Martens. 2007. First description of skull of lower Permian Seymouria sanjuanensis (Seymouriamorpha: Seymouriidae) at an early juvenile growth stage. Annals of Carnegie Museum 76:53–72.
Laurin M 1996. A redescription of the cranial anatomy of Seymouria baylorensis, the best known Seymouriamorph (Vertebrata: Seymouriamorpha). PaleoBios 17: 1–16.
Lutes AA, WB Neaves, DP Baumann, W Wiegraebe and P Baumann 2010. Sister chromosome pairing maintains heterozygosity in parthenogenetic lizards. Nature 464:283–286.