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The origin of marine crocs re-revised

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Figure 1. Several Jurassic sea crocs, apparently derived from Late Triassic Dyoplax.

Dr. Andrea Cau 2019
recently revised the affinities of the extinct marine crocs (Figs. 1,2). Here (Fig. 3), with more outgroup taxa, the affinities of the outgroups are more refined by adding taxa omitted by Cau. The in-group marine croc clade of Cau 2019 continue as is untested.

Figure 2. Reduced from Cau 2019 showing Fruitachampsa as the proximal outgroup for marine and river crocs. The outgroup Postosuchus is not related to Crocodylomorpha in the LRT.

As we learned earlier
choosing outgroup taxa is not as scientific as letting a wide gamut phylogenetic analysis, like the large reptile tree (LRT, 1549 taxa, subset Fig. 3), choose outgroup taxa for you. 

Figure 3. Subset of the LRT focusing on Crocodylomorpha. Here, with more outgroup taxa, Fruitachampsa nests far from marine and river crocs. Why is the Fruitachampsa score <50? It was scored without post-crania. Saltopus has no crania. So when post-cranial traits are added to Fruitachampsa (soon) expect that score to increase. 

Strangely,
the proximal outgroup taxon for marine crocs recovered by Dr. Cau (Fig. 2) was tiny Fruitachampsa (Figs. 3, 4), a small, gracile Late Jurassic biped sprinter that nests with the small, gracile, Late Triassic biped sprinter Scleromochlus (Figs. 4, 5) in LRT. Fruitachampsa would seem to have few traits in common with river and marine crocs.

Figure 4. Fruitachampsa reconstructed. Note the many homologies with Scleromochlus and the few with marine crocs.

Cau did not include Scleromochlus
in his cladogram (Fig. 1), nor did he include Dyoplax.

Figure 5. Fruitachampsa skull to scale with Scleromochlus skull. No antorbital fenestra is present according to Clark 2011, but phylogenetic braceting with Scleromochlus indicates otherwise. Apparently the jawline has phylogenetically eroded. The vomers are missing. The chonae are conjoined medially, contra Clark 2011.

While we’re on the subject of Fruitachampsa,
it appears to have an antorbital fenestra conjoined with the jawline rather than a very deeply emarginated jawline when put into a phylogenetic context. For similar reasons, the central fenestra in the palate is likely the conjoined choanae of primitive crocodylomorphs like Scleromochlus, rather than the posteriorly shifted choanae of derived eucrocs.

Figure 6. Faxinalipterus matched to Scleromochlus. The former is more primitive, like Gracilisuchus, in having shorter hind limbs and more robust fore limbs. The maxilla with fenestra and fossa, plus the teeth, are a good match.

Despite appearances
Dyoplax (Fig. 7) is not considered a crocodylomorph, let alone an outgroup to marine crocs, as we learned earlier here.

Figure 7. Dyoplax arenaceus Fraas 1867 is a mold fossil recently considered to be a sphenosuchian crocodylomorph. Here it nests as a basal metriorhynchid (sea crocodile) in the Late Triassic.

In the LRT
(subset Fig. 3) Dyopolax is the outgroup taxon to marine crocs while Dibrothosuchus (Fig. 8) is basal to this clade + river crocs.  

Figure 8. Dibothrosuchus nests basal to all later quadrupedal crocs, including marine crocs, in the LRT. The hind limbs are not known. Phylogenetic bracketing suggests shorter legs are more likely.

Once again,
a wide gamut phylogenetic analysis is key to recovering interrelationships.


References
Cau A 2019. A revision of the diagnosis and affinities of the metriorhynchoids (Crocodylomorpha, Thalattosuchia) from the Rosso Ammonitico Veronese Formation (Jurassic of Italy) using specimen-level analyses. PeerJ, DOI 10.7717/peerj.7364
Clark JM 2011. A new shartegosuchid crocodyliform from the Upper Jurassic Morrison Formation of western Colorado. Zoological Journal of the Linnean Society, 2011, 163, S152–S172. doi: 10.1111/j.1096-3642.2011.00719.x


Source: https://pterosaurheresies.wordpress.com/2019/07/28/the-origin-of-marine-crocs-re-revised/


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