Trimerorhachis: a late survivor of the fin/finger transition?

Figure 1. Trimerorhachis was considered a dvinosaurian temnospondyl. Here both Trimerorhachis and Dvinosaurus nest low on the basal tetrapod tree, close to the fin/finger transition.

Figure 1. Flattened Trimerorhachis was considered a dvinosaurian temnospondyl. Here both Trimerorhachis and Dvinosaurus nest low on the basal tetrapod tree, close to the fin/finger transition, not within the Temnospondyli. Both are late survivors of a Devonian radiation.

Wikipedia reports:
Trimerorhachis (Early Permian, (Cope 1878, Case 1935, Schoch 2013; up to 1m in length) is an extinct genus of dvinosaurian temnospondyl within the family Trimerorhachidae. The trunk is long and the limbs are relatively short. Many bones are poorly ossified, indicating that Trimerorhachis was poorly suited for movement on land. The presence of a branchial apparatus indicates that Trimerorhachis had external gills in life. The body of Trimerorhachis is also completely covered by small and very thin osteoderms, which overlap and can be up to 20 layers thick. The scales were more similar to fish scales than they were to reptile scales, according to Colbert 1955. However, Olson 1979 disputed that interpretation. Specimens are often preserved as masses of bones that are mixed together and densely packed in slabs of rock”

Figure 2. Trimerorhachis forelimb and hind limb in situ and reconstructed.

Figure 2. Trimerorhachis forelimb and hind limb in situ and reconstructed. Pawley 1979 did not report metacarpals or a pubis. It is possible and perhaps likely that only 4 metacarpals were present along with two phalanges, but its worth exploring all possibilities. 

As a late (Early Permian) survivor of a Late Devonian radiation
Trimerorhachis evolved by convergence certain traits found in other more derived tetrapods, like a longer femur and open palate (narrow, bowed pterygoids). Testing all possibilities while minimizing assumptions is the most valuable benefit of a large gamut phylogenetic analysis conducted by unbiased software. Workers used to eyeball specimens in the pre-computer days.

Figure 2. Trimerorhachis pelvis. The pubis is not ossified.

Figure 3. Trimerorhachis pelvis. The pubis is not ossified here, according to Pawley 1979, but see Fig. 1.

Like other workers,
Pawley 1979 considered Trimerorhachis close to Dvinosaurus (Fig. 7) and both thought to be derived from the basal temnospondyl Balanerpeton and Dendrerpeton. The large reptile tree (LRT) nests both taxa at the base of the Lepodpondyli, not closely related to Trimerorhachis and distinct from Temnospondyli. Pawley supports the hypothesis that aquatic ‘temnospondyls,’ like Trimerorhachis, had terrestrial ancestors. By contrast, the LRT nests Trimerorhachis with weak-limbed taxa more primitive than any temnospondyl.

Additionally
the LRT nests Batrachosaurus and Gerrothorax in the Dvinosaurus / Trimerorhachis clade. This clade features horizontally opposed dorsal ribs and an equally flattened skull. Another flattened taxon, Ossinodus, is closely related. I have not seen limb material for any of these taxa. Acanthostega is the closest taxon that preserves limbs.

Figure 3. Trimerorhachis hind limb and pes from Pawley 1979.

Figure 4. Trimerorhachis hind limb and pes from Pawley 1979 and reconstructed here.

Pawley 1979 noted,
“The vast majority of the [Trimerorhachis] specimens consists of ornamental cranial and pectoral girdle bones, intercentra, and larger elements of the appendicular skeleton. Neural arches, pleurocentra, ribs and distal limb elements are rare.” No sacrals were found by Pawley. No dorsal ribs had uncinate processes (like those in Ichthyostega and Eryops). The chevrons were long and tapered distally (creating a fin?). The interclavicle was diamond-shaped with a longer anterior portion.

Figure 4. Trimerorhachis humerus changes during ontogeny

Figure 5. Trimerorhachis humerus changes during ontogeny

The humerus
(Fig. 5) was  L-shaped and the degree of torsion varied between specimens from 45º to 90º. The distal end always exhibited a low degree of ossification.

Figure 6. Trimerorhachis cladogram. Gray area is the Temnospondyli clade.

Figure 6. Trimerorhachis cladogram. Gray area is the Temnospondyli clade.

Pawley considered
Trimerorhachis a secondarily adapted aquatic temnospondyl. All workers have noted the wide open palate vacuities that characterize most, but not all members of the Temnospondyli. By contrast, the LRT nests Trimerorhachis with taxa that had not yet left the water completely and shared a flat morphology with Tiktaalik and Panderichthys.

This is the second time
elongate limbs and digits have appeared by convergence in basal tetrapods. Earlier Pholidogaster and kin provided the first exceptions to the rule. Note that all known specimens of Trimerorhachis are Early Permian, some tens of millions of years later than the Late Devonian radiation of that clade. The Ichthyostega line is the one that ultimately produced crown Tetrapoda via a sister to Eucritta.

FIgure 8. Dvinosaurus nests with Trimerorhachis and also has ceratobranchial (gill) bones.

FIgure 7. Dvinosaurus nests with Trimerorhachis and also has ceratobranchial (gill) bones. The loss of the intertemoral is shown here in light green merging to the postorbital in orange. 

If these nestings are not correct
and Trimerorhachis ultimately nests higher on the basal tetrapod tree, then we’re witnessing massive convergence of another sort, convergence that allies Trimerorhachis with tetrapods at the fin/finger transition. I’d like to see limbs for Gerrothorax or any other plagiosaur, if available.

Figure 9. Ossinodus is a close relative of Trimerorhachis in the LRT.

Figure 8. Ossinodus is a close relative of Trimerorhachis in the LRT. 

By the way, I find this fascinating…
week after week, far and away the most popular page(s) on this blog continue to be on the origin of bats.

References
Berman DS and Reisz RR 1980. A new species of Trimerorhachis (Amphibia, Temnospondyli) from the Lower Permian Abo Formation of New Mexico, with discussion of Permian faunal distributions in that state. Annals of the Carnegie Museum. 49: 455–485.
Case EC 1935. Description of a collection of associated skeletons of Trimerorhachis. University of Michigan Contributions from the Museum of Paleontology. 4 (13): 227–274.
Colbert EH 1955. Scales in the Permian amphibian Trimerorhachis. American Museum Novitates. 1740: 1–17.
Olson EC 1979. Aspects of the biology of Trimerorhachis (Amphibia: Temnospondyli). Journal of Paleontology. 53 (1): 1–17.
Pawley K 2007. The postcranial skeleton of Trimerorhachis insignis Cope, 1878 (Temnospondyli: Trimerorhachidae): a plesiomorphic temnospondyl from the Lower Permian of North America. Journal of Paleontology. 81 (5):
Williston SW 1915. Trimerorhachis, a Permian temnospondyl amphibian. The Journal of Geology. 23 (3): 246–255.
Williston SW 1916. The skeleton of Trimerorhachis. The Journal of Geology. 24 (3): 291–297.

wiki/Trimerorhachis

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2 thoughts on “Trimerorhachis: a late survivor of the fin/finger transition?

  1. So you’ll never get to the comments? You’re dooming yourself to repating your mistakes forever and to never learning anything. You said you wanted to learn.

    The body of Trimerorhachis is also completely covered by small and very thin osteoderms, which overlap and can be up to 20 layers thick. The scales were more similar to fish scales than they were to reptile scales, according to Colbert 1955. However, Olson 1979 disputed that interpretation.

    Olson was, of course, wrong: these are ordinary dermal scales as plesiomorphically found in tetrapods (Witzmann 2010 “2011”).

    Testing all possibilities while minimizing assumptions is the most valuable benefit of a large gamut phylogenetic analysis conducted by unbiased software.

    This only works if the character list is any good. Yours isn’t, as I’ve explained in several comments that are apparently too long for you to read.

    Pawley 1979

    Both Olson 1979 and Pawley 2007. And everyone else, actually.

    Pawley supports the hypothesis that aquatic ‘temnospondyls,’ like Trimerorhachis, had terrestrial ancestors. By contrast, the LRT nests Trimerorhachis with weak-limbed taxa more primitive than any temnospondyl.

    That’s a sure sign it has succumbed to redundant characters that are all instances of incomplete ossification, an adaptation to aquatic lifestyles.

    I agree that Trimerorhachis didn’t have any terrestrial ancestors. However, I do think it had ancestors that occasionally walked on land when fully adult, from one body of water to the next. That fits the trees I find, too.

    horizontally opposed dorsal ribs

    What is that?

    Another flattened taxon, Ossinodus, is closely related. I have not seen limb material for any of these taxa.

    Whoa. All three of the papers that have described parts of Ossinodus describe limb material. One of them, Bishop (2014), is entirely on the forelimb and shoulder girdle, mostly on an exceptionally well preserved (but very incompletely ossified) humerus. It’s very different from that of Trimerorhachis or of any temnospondyl.

    Concerning Gerrothorax, which is a very common animal, Hellrung’s (2003) thesis about it is lying around on the Internet. It’s in German, but check out the pictures.

  2. Apologies again for just getting to this in mid April. Unable to put my hands on anything you mentioned but the PLOS paper on the Ossinodus radius. re: my character list – it does lump and separate all the taxa. That’s all I want it to do. I’ve added three in the last few months. More will show up when they become necessary or curiosities. I crave limb material for Gerrothorax. If you have it, please send it.

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