You heard it here in 2011: diadectids are amniotes

Co-author, David S. Berman,
has been saying diadectids are amniotes since the 1990s, but not with a comprehensive taxon list, and, apparently nobody listened. The consensus apparently prefers their diadectids with tadpoles.

Here’s what Wikipedia reports
“Diadectes (meaning crosswise-biter) is an extinct genus of large, very reptile-like amphibians that lived during the early Permian period (ArtinskianKungurian stages of the Cisuralian epoch, between 290 and 272 million years ago[1]). Diadectes was one of the very first herbivorous tetrapods, and also one of the first fully terrestrial animals to attain large size.”

Skeleton of Diadectes. Perhaps unnoticed are the broad dorsal ribs of this taxon, basal to Stephanospondylus, Procolophon and pareiasaurs.

Figure 1 Skeleton of Diadectes. Perhaps unnoticed are the broad dorsal ribs of this taxon, basal to Stephanospondylus, Procolophon and pareiasaurs.

Klembara et al. 2019 report
on the inner ear morphology of diadectids and seymouriamorphs. From the abstract:
“Two pivotal clades of early tetrapods, the diadectomorphs and the seymouriamorphs, have played an unsurpassed role in debates about the ancestry of amniotes for over a century, but their skeletal morphology has provided conflicting evidence for their affinities. Both maximum parsimony and Bayesian inference analyses retrieve seymouriamorphs as derived non‐crown amniotes and diadectomorphs as sister group to synapsids.”

Figure 2. Cladogram from Klembara et al. 2019. Green shows reptile taxa in the LRT.

Figure 2. Cladogram from Klembara et al. 2019. Green shows reptile taxa in the LRT.

Dr. David Marjanovic wrote in the DML:
“Amniota is a crown-group; there’s technically no such thing as a “stem-amniote”, because if it’s on the stem, it’s not an amniote.”

Unfortunately, Klembara et al. don’t have enough taxa
to understand that Amniota is a junior synonym for Reptilia. So Repitilomorpha works well for pre-reptiles. More importantly, for the subject at hand, Diadectes (Fig. 1) and kin have been deeply nested within the large reptile tree (LRT, 1583 taxa) since 2011. This is an online resource you can use to double check your taxon list, just to make sure it is up to date. The Klembara et al. taxon list (Fig. 2) is so inadequate it nests several reptiles apart from one another and omits dozens of others pertinent to this issue.

Figure 2. Subset of the LRT focusing on basal lepidosauromorphs and Diadectes.

Figure 3. Subset of the LRT focusing on basal lepidosauromorphs and Diadectes.

Bottom line, when you add enough taxa
diadectomorphs are not close to synapsids, but arise from millerettids.

At least the Klembara team
moved diadectomorphs inside the Amniota. That’s a minor victory. Add the above taxa to your cladogram (Fig. 2) and see where Diadectes nests. That’s what the LRT is here for… to help workers avoid taxon exclusion.


References
Klembara J, Hain M, Ruta M, Berman DS,  SEPierce and Henrici AC 2019. Inner ear morphology of diadectomorphs and seymouriamorphs (Tetrapoda) uncovered by highâresolution xâray microcomputed tomography, and the origin of the amniote crown group. Palaeontology (advance online publication) Future publication date: August 5, 2020
doi: https://doi.org/10.1111/pala.12448
https://onlinelibrary.wiley.com/doi/full/10.1111/pala.12448

wiki/Diadectes

Walking Orobates video on YouTube and in Nature

This is a wonderful experiment/demonstration/simulation
and a wonderful fossil, matching a digitized 300 million-year-old Orobates skeleton to Orobates tracks.

This time the problem comes from basal tetrapod experts
One writes in the Dinosaur Mailing List, “As so much else in amniote phylogeny, it remains unclear whether the diadectomorphs are just outside Amniota or just inside. The question has never been tested in an analysis with enough taxa and enough characters; some matrices may have had close to enough of one, but definitely not of both, and there haven’t been many in total in the first place.”

Another comes from SmithsonianMag.com
“At first glance, the 300 million-year-old Orobates pabsti might look like a chunky lizard. In actuality, this animal from the Permian period is what experts know as a stem amniote—a vertebrate that’s part of the evolutionary lineage between amphibians, which reproduce in the water, and the last common ancestor of mammals and reptiles, which lay eggs on land.”

A third comes from the title of the Nature paper (Nyakatura et al. 2019)
“Reverse-engineering the locomotion of a stem amniote.” Clearly the authors have never heard of an amphibian-like reptile.

This is an example of wrong thinking.
The large reptile tree (LRT, 1378 taxa) nests Orobates deep inside the Reptilia (the clade Amniota is a junior synonym, see below). There are those who say the LRT needs more characters to… what? The LRT already lumps and splits virtually all of its taxa in complete resolution documenting a gradual accumulation of traits at every node. That is the yardstick by which all cladograms should be judged. How will adding characters suddenly shift taxa on the tree topology? Taxa nest where they do because they most closely resemble their sisters. And that clade most closely resembles their more distant sisters in a series that ultimately includes all tested taxa. All other possible nesting sites for taxa are provided in a wide gamut analysis.

Apparently basal tetrapod experts are hoping the LRT is somehow wrong. If so, which taxa are wrongly nested and where should they nest instead? They simply need to add the pertinent taxa listed in the LRT to their own analyses with their own character lists to find out for themselves and report the results. This is how science works. Anyone can repeat the experiment, but the experiment in this case, requires pertinent taxa, not more characters.

Case in point:
Remember how Hone and Benton 2007, 2009 deleted pertinent taxa in their purported quest to test two competing hypotheses on pterosaur origins? When they found out the fenestrasaurs were attracting pterosaurs, they deleted the fenestrasaurs and never did find out where pterosaurs originated.

Comment to Nature
“This is a wonderful experiment/demonstration/simulation. The problem comes from the systematics of Orobates. Testing a wide gamut of tetrapod taxa nests Orobates between Limnoscelis and Tseajaia + Tetraceratops. This clade nests between Saurorictus + Captorhinidae and Milleretta, all traditional amniotes. Phylogenetic analysis nests the amphibian-like reptiles Gephyrostegus (Late Carboniferous) and Silvanerpeton (Early Carboniferous) as the last common ancestors of all included amniote taxa. Three nodes of amphibian-like reptiles nest between these two and the Saurorictus + Captorhinidae clade. So Orobates nests well within the Amniota, now a junior synonym for the clade Reptilia. The wide gamut cladogram is found online here: http://reptileevolution.com/reptile-tree.htm”

References
Nyakatura JA, et al. (11 co-authors) 2019. Reverse-engineering the locomotion of a stem amniote. Nature.com PDF online.

Articles from the popular press:

https://www.smithsonianmag.com/science-nature/scientists-used-robot-study-how-prehistoric-lizards-evolved-walk-land-180971283/

Continuing problems in vertebrate paleontology – part 1

A quick glance through various paleontology topics
on various Wikipedia pages reveals a rather long list of antiquated views that remain as false paradigms. These were falsified by testing in the large reptile tree. We’ll just bring up a few of these at a time while waiting for more interesting paleo-news to break.

  1. The first amniotes, referred to as “basal amniotes”, resembled small lizards and evolved from the amphibian reptiliomorphs about 312 million years ago. Move that back to 340 mya for phylogenetically widespread fossil evidence of basal amniotes and to 360 mya for their hypothetical origins.
  2. The first dichotomy within the Amniotes produced the clades Synapsida and Sauropsida. In the LRT several amniote clades precede the advent of the Synapsida. The last common ancestors of all amniotes are Silvanerpeton and Gephyrostegus bohemicus. The first dichotomy produced the new Lepidosauromorpha and the new Archosauromorpha. Synapsids nest deeply within the new Archosauromorpha.
  3. Diadectomorpha is a clade of large reptile-like amphibians. The LRT nests all diadectomorphs deep within the new Lepidosauromorpha. 
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 1. A new reconstruction of Gephyrostegus bohemicus, a late-surviving basalmost amniote. This species lived 30 million years after the other amniotes 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 known in all specimens.

And while we’re at it…
This comes courtesy of Ben Creisler at the DML TheSociety of Vertebrate Paleontology 2016 Meeting Program and Abstract Book is available here:

PLEASE NOTE! Content is embargoed until the actual presentation.
“Unless specified otherwise, coverage of abstracts presented orally at the Annual Meeting is strictly prohibited until the start time of the presentation, and coverage of poster presentations is prohibited until the relevant poster session opens for viewing.”
There is one abstract in there
that confirms something I discovered four years ago, which brings some satisfaction that it was discovered again.

Former reptile: Gymnarthrus. Former reptile, former amphibian: Diadectes. Both from Case 1910.

Case 1910
described several skulls from what he presumed were Permian deposits in Archer County, Texas. Yes, they are Early Permian and home to many a Dimetrodon.

Among the several skulls
was Gymnarthrus willoughbyi (Fig. 1), known from a tiny 1.6cm skull. Case reported: “It was thought at first that both the basisphenoid and the parasphenoid process constituted the the parasphenoid bone and that the animal was an amphibian, but this is impossible… the animal approaches the intermediate form between the amphibians and reptiles.” Today we know Gymnarthrus to be one of the lizard mimics, the lepospondyl microsaurs. Case also wrote, “The nearest approach to this form is the small amphibian skull described by Broili as Cardiocephalous sternbergii, but this is described as having the skull complete, no parietal foramen, teeth regularly diminishing in size anteriorly but with cutting edges and lyra present.” I don’t know what lyra are in this context.

Figure 1. Gymarthrus willougbyi, drawn by Case 1910 on the left and von Huene 1913 on the right.

Figure 1. Gymarthrus willougbyi, drawn by Case 1910 on the left and von Huene 1913 on the right. These are apparently freehand sketches and, judging by the perspective implied by the large orbit on the right, sketched from two distances.

Carroll and Gaskill 1978
allied Gymnarthrus with Cardiocephalus, another microsaur.

Figure 2. Diadectes phaseolinus in situ, as originally illustrated and as reillustrated above according to phylogenetic bracketing.

Figure 2. Diadectes phaseolinus in situ, as originally illustrated and as reillustrated above according to phylogenetic bracketing. Case reported the tail was as long as the presacral portion of the column, but did not illustrate it that way for this specimen. No intercentra were present.

Case also identified Diadectes as a reptile
(order Cotylosauria), but later authors (and currently Wikipedia, taken from a PhD thesis by R Kissel 2010) considered it a reptile-like amphibian. The large reptile tree nests Diadectes as derived from Milleretta and all the “Diadectomorpha” listed in Wikipedia are reptiles. Limnoscelis, Orobates and Tseajaia do not nest with Diadectes in the large reptile tree, but bolosaurids and procolophonids do. So we’ve got some housecleaning to do at that node.

The interesting thing about this Diadectes specimen,
according to Case 1910, is the set of expanded dorsal ribs beneath the scapulae. He writes, “The ribs of the third, fourth and fifth vertebrae show a well defined articular end with a distinct neck. The bodies of these ribs are expanded into thin triangular plates, with the front edge straight and the posterior edge drawn out into a point which overlaps the succeeding rib; this forms a strong protection for the anterior thoracic region. The sixth, seventh and eighth [ribs] are overlain by thin, narrow, plates which continue backward the protection of the thoracic region to a point opposite the posterior end of the scapula.” Some, but not all Diadectes specimens have such expanded ribs.

Case presumed
that gastralia (his ‘abdominal ribs’  were present. They are not. Case notes “the animal was distinctly narrow chested, with the bones of the the girdle strongly interlocked. Diadectes had practically no neck.”

Based on the mounted skeleton, Case reiterated
“the suggestions previously made by the author that these animals are the nearest discovered forms to the ancestors of turtles.” That old hypothesis has not been confirmed by the large reptile tree, as noted earlier.

References
Carroll RL and Gaskill P 1978. The Order Microsauria. Memoirs of the American Philosophical Society 126:1-211 [J. Mueller/T. Liebrecht/T. Liebrecht]
Case EC 1910.
 New or little known reptiles and amphibians from the Permian (?) of Texas. Bulletin of the American Museum of Natural History 28 (17):163-181.
Huene FRF von and Gregory WK 1913. The skull elements of the Permian Tetrapoda in the American Museum of Natural History, New York. Bulletin of the AMNH ; v. 32, article 18.: 315-386.

Another, more complete Colobomycter adds data to this enigma

Revised June 10, 2016 with a new reconstruction and nesting with Eothyris. 

A new paper
by MacDougall et al. 2016 introduces Colobomycter vaughni (BRMP 2008.3.1, Fig. 1) a new toothy specimen that adds much needed data to the former enigma taxon, Colobomycter pholeter. They report on the synapomorphies, “enlarged premaxillary tooth and paired enlarged maxillary teeth, unique dentition that grants it an appearance quite distinct from other parareptiles at Richards Spur. This new material differs from that of C. pholeter in that it possesses at least three more teeth on its maxilla, the enlarged premaxillary and maxillary teeth are more gracile than those in C. pholeter, and the lacrimal is restricted externally to the orbital margin and does not exhibit an extra lateral exposure.” 

Figure 1. The new Colobomycter compared to the original pasted on a ghost of the new material. We're learning more about this genus!

Figure 1. The new Colobomycter compared to the original pasted on a ghost of the new material. We’re learning more about this genus!

Unfortunately, 
MacDougall et al. considered Colobomycter a member of the Lanthanosuchoidea. According to MacDougall et al. taxa in that clade include Feeserpeton, Lanthanosuchus, Acleistorhinus and Delorhynchus.

In the large reptile tree 
(Fig. 2) Lanthanosuchus
 nests with Bashkyroleter, Macroleter and Emeroleter.

On the other hand (and this is revised from the original posting)
Colobomycter pholeter
(Vaughn 1958, Modesto and Reisz 2008, UWBM 95405), Lower Permian ~278 mya, was originally considered a caseid pelycosaur, like Eothyris. (But note that Eothyris is not considered a pelycosaur in the large reptile tree (subset Fig. 2). Later, Modesto and Reisz (2008) considered Colobomycter a “parareptile” close to Acleistorhinus. After further consideration, it turns out that Colobomycter is indeed quite similar to Eothyris, as Vaughn 1958 indicated with much less data and fewer optional candidate taxa to consider. Hats off to Vaughn!

Figure 2. Revised cladogram of Colobomycter nesting this genus with Eothyris in an unnamed clade that includes caseasauria.

Figure 2. Revised cladogram of Colobomycter nesting this genus with Eothyris in an unnamed clade that includes caseasauria.

Sharp-eyed observers will note
that earlier I nested the rostrum of Colobomycter with procolophonids based on a smaller portion of rostrum. Clearly. I’m not as sharp as Vaughn was.

At this point
Colobomycter likely had a lateral temporal fenestra.

Herbivore or carnivore?
There are herbivores, carnivores and omnivores related to Colobomycter. It looks like the anterior dentary teeth could scrape off or collect whatever the premaxillary tusks had stabbed into. Eothyris had similar large maxillary teeth.

Figure 3. Eothyris skull in three views. This taxon is the closest known relative to Colobomycter.

Figure 3. Eothyris skull in three views. This taxon is the closest known relative to Colobomycter.

This is only one of tens of thousands of errors I have made
I’m only embarrassed by the ones that have yet to surface. Science and scientists don’t always have all the answers, but if the formula (or in this case cladogram) recovers a sticking point, as it did earlier, it will reward you to go back in and figure out where the errors were made. In this case several little errors among several taxa added up, but are corrected here, resulting once again in a completely resolved tree, hopefully more closely echoing Nature.

References
MacDougall MJ, Modesto SP and Reisz RR 2016. A new reptile from the Richards Spur Locality, Oklahoma, USA, and patterns of Early Permian parareptile diversification, Journal of Vertebrate Paleontology (advance online publication). www.tandfonline.com/doi/

Orobates in 3D

Wikipedia considers Orobates an anamniote (not a reptile = amniote) close to Diadectidae, which are also considered anamniotes. So does a recent paper by Nyakatura et al. 2015 who un-deformed the MNG 10181, the holotype, using CT scans and software (Fig. 1).

Figure 1. A 3D Orobates produced from CT scans by Nyakatura et al. 2015.

Figure 1. A 3D Orobates produced from CT scans by Nyakatura et al. 2015.

From the Nyakaturs et al. 2015 introduction
“Orobates pabsti (Holotype MNG 10181) is one of the most basal diadectids, from the lower Permian of Germany (Bromacker quarry, Tambach Formation, Tambach-Dietharz, Thuringia). Because of its phylogenetic position, this nearly complete, articulated, comparatively well-preserved specimen can be considered a key fossil. Additionally, fossil trackways from the ichnospecies Ichnotherium sphaerodactylum of the same locality were unequivocally produced by Orobates pabsti rendering this combination of body fossil and fossil trackways the oldest known track-trackmaker association and offering direct evidence of its locomotor behavior.”

It appears that Nyakatura et al. did everything right 
to produce a good virtual model of Orobates.

Figure 1. Limnoscelis and two suitable ancestral taxa, Orobates and Milleretta, all shown to scale (below) and to fit (above).

Figure 2. Limnoscelis and two suitable ancestral taxa, Orobates and Milleretta, all shown to scale (below) and to fit (above).

Their only fault
was trusting an old cladogram that gave them a mistaken nesting outside the Reptilia. The large reptile tree nests Orobates inside the Reptilia, inside the Lepidosauromorpha, with the clade Diadectidae derived from the undisputed reptile, Millleretta (Fig. 2). No stem (=pre) amniotes more closely match the morphology.

Once again
an old, traditional sour matrix seems to be at fault.

References
Berman, DS et al. 2004. A new diadectid (Diadectomorpha), Orobates pabsti, from the Early Permian of Central Germany. Bulletin of Carnegie Museum of Natural History 35 :1-36. doi: 10.2992/0145-9058(2004)35[1:ANDDOP]2.0.CO;2
Nyakatura JA, Allen VR, Lauströer J, Andikfar A, Danczak M, Ullrich H-J, et al. 2015. A Three-Dimensional Skeletal Reconstruction of the Stem Amniote Orobates pabsti (Diadectidae): Analyses of Body Mass, Centre of Mass Position, and Joint Mobility. PLoS ONE 10(9): e0137284. doi:10.1371/journal.pone.0137284

wiki/Orobates

Adding taxa to the Diadectes clade

Adding a few
and distinct Diadectes specimens (no two appear to be conspecific) opens the door to new insights into that corner of the cladogram. Some of the data are from 3D skull images with sutures delineated. Others are from firsthand observation. Some data are from drawings. Berman et al. 1992 made an interesting observation that prior authors illustrated the skull roof of Diadectes in a variety of ways (Fig. 1). The caption does not indicate that all were drawn from the same specimen. I suspect they were not.

Figure 1. How Berman et al. copied the illustrations of prior authors who each figured the skull roof of Diadectes. Perhaps these were several distinct specimens, not just one.

Figure 1. How Berman et al. copied the illustrations of prior authors who each figured the skull roof of Diadectes. Perhaps these were several distinct specimens, not just one. Not sure, at this point, which illustrations represent which specimens.

The Berman et al. phylogenetic analysis
included seven taxa, including two suprageneric taxa, Pelycosauria and Captorhinomorpha. They included only nine characters. The anamniote, Seymouria, was the outgroup. The first clade included Pelycosauria + (Limnoscelis +(Tseajaia and Diadectes). The second clade included Captorhinomorpha + Petrolacosaurus. The large reptile tree includes hundreds more taxa and characters. The pertinent subset is shown here (Fig. 2). It is also clear from the Berman et al. taxon set that they thought they were dealing with a small set of basal reptiles and pre-reptiles. In 2015 it is clear that they did not include the pertinent taxa they should have as some of these taxa are not related to any of the others except distantly.

Figure 2. How the large reptile tree lumps and splits the several Diadectes specimens now included here. Note that bolosaurids, including Phonodus, now nest within other Diadectes specimens.

Figure 2. How the large reptile tree lumps and splits the several Diadectes specimens now included here. Note that bolosaurids, including Phonodus, now nest within other Diadectes specimens.

Now, with current data
it is becoming increasingly clear that both bolosaurids and procolophonids nest within  a fully reptilian Diadectes clade. It is also clear that the genus Diadectes needs to be further split, as Kissel (2010) started to do by renaming Silvadectes and Oradectes from former Diadectes species.

Skeleton of Diadectes. Perhaps unnoticed are the broad dorsal ribs of this taxon, basal to Stephanospondylus, Procolophon and pareiasaurs.

Figure 3. Skeleton of Diadectes (UC 706, UC 1078). Perhaps unnoticed are the broad dorsal ribs of this taxon, basal to Stephanospondylus, Procolophon and pareiasaurs.

Also note
the placement of Stephanospondylus as a proximal sister taxon to the diadectids nesting at the base of the pareiasaurs (including turtles). Turtles are sisters to pareiasaurs and they ARE pareiasaurs because they are derived from pareiasaurs, just as birds are derived from theropod dinosaurs.

Figure 4. Click to enlarge. Stephanospondylus based on parts found in Stappenbeck 1905. Several elements are re-identified here. Note the large costal plates on the ribs, as in Odontochelys. The pubis apparently connected to a ventral plastron, not preserved. The interclavicle was likely incorporated into the plastron.

Figure 4. Click to enlarge. Stephanospondylus based on parts found in Stappenbeck 1905. Several elements are re-identified here. Note the large costal plates on the ribs, as in Odontochelys. The pubis apparently connected to a ventral plastron, not preserved. The interclavicle was likely incorporated into the plastron.

Like everyone who studies prehistoric reptiles
there is a day when you don’t know anything about a taxon and later there is a day when you are making contributions to Science. Those days keep on coming.

References
Berman DS, Sumida SS and Lombard E 1992. Reinterpretation of the Temporal and Occipital Regions in Diadectes and the Relationships of Diadectomorphs. Journal of Vertebrate Paleontology 66(3):481-499.
Kissel R 2010. Morphology, Phylogeny, and Evolution of Diadectidae (Cotylosauria: Diadectomorpha). Thesis (Graduate Department of Ecology & Evolutionary Biology University of Toronto).