Examining Functional Convergence Between Triassic Phytosaurs and Slender-Snouted Crocodylians

A new preprint in PeerJ.

Lemanis R., Jones A.S., Butler R.J., Anderson P.S.L., and E.J. Rayfield. 2019. Comparative biomechanical analysis demonstrates functional convergence between slender-snouted crocodilians and phytosaurs. PeerJ Preprints 7:e27476v1https://doi.org/10.7287/peerj.preprints.27476v1

Abstract - Morphological similarities between the extinct Triassic archosauriform clade Phytosauria and extant crocodilians have formed the basis of long-proposed hypotheses of evolutionary convergence. These hypotheses have informed the reconstructions of phytosaur ecology and biology, including feeding preferences, body mass, soft tissue systems, mating behaviours, and environmental preferences. However, phytosaurs possess numerous cranial apomorphies that distinguish them from modern crocodilians and potentially limit ecomorphological comparisons. Here, we present the first computational mechanical comparison of phytosaur cranial strength to several extant crocodilian taxa using two biomechanical approaches: beam theory and finite element analysis. We demonstrate mechanical convergence between the slender-snouted phytosaur Ebrachosuchus neukami and modern slender-snouted crocodilians. We provide evidence that the phytosaurian premaxillary palate is functionally equivalent to the crocodilian secondary palate. The premaxillary palate is associated with greater resistance to biting induced stress, lower strain energy, higher resistance to bending and torsion, as well as increased performance under tension. In all tests, Ebrachosuchus performed worse than all tested crocodilians, showing higher stress under equivalent loading conditions. These findings have implications for the proposed feeding ecology of slender-snouted phytosaurs and corroborate previous broad assessments of phytosaur ecology based on morphological comparisons to crocodilians; however, we urge caution in overextending those assessments given the current paucity of comparative functional data.

So Long Paleorhinus and Pseudopalatinae

Long needed redescription of the type material of the phytosaur Parasuchus hislopi from India and a revision of the non-mystriosuchin parasuchid phytosaurs. It will take me awhile to abandon the name Pseudopalatinae. 

Kammerer, C. F., Butler, R. J., Bandyopadhyay, S., and M. R. Stocker. 2015. Relationships of the Indian phytosaur Parasuchus hislopi Lydekker, 1885. Papers in Paleontology (early online). 

Abstract: The neotype skull of the Indian phytosaur Parasuchus hislopi Lydekker, 1885 (ISI R42) is re-evaluated and compared with the type material of other basal phytosaurs. Parasuchus hislopi is extremely similar to species previously placed in Paleorhinus (P. bransoni and P. angustifrons), sharing with them such characters as a series of nodes on the lateral surface of the jugal, paired ridges on the squamosal and a frontal depression. Parasuchus hislopi represents a valid species: it can be distinguished from P. bransoni by a relatively low narial eminence and P. angustifrons by the absence of paired nasal depressions. Inclusion of Parasuchus hislopi in a phylogenetic analysis of phytosaurs recovers it in a well-supported clade with P. bransoni and P. angustifrons. Parasuchus is considered the senior synonym of Paleorhinus and Arganarhinus. Parasuchus (here considered to include P. hislopi, P. angustifrons, P. bransoni and P. magnoculus) has a broad circum-Pangaean distribution, with species occurring in the south-western United States, Morocco, central Europe and India. Phytosaur higher-level taxonomy is also revised: Parasuchidae is redefined to include ‘Paleorhinus-grade’ phytosaurs and the later-diverging Mystriosuchinae (the group formerly known as Phytosauridae), and Pseudopalatinae is renamed Mystriosuchini for reason of priority.

Bone Histology of Phytosaur, Aetosaur, and Other Archosauriform Osteoderms

Just in time for Christmas...

Scheyer, T. M., Desojo, J. B., and I. A. Cerda. 2013. Bone histology of phytosaur, aetosaur, and other archosauriform osteoderms (Eureptilia, Archosauromorpha). Anatomical Record (early view) DOI: 10.1002/ar.22849

Abstract - As in other archosauriforms, phytosaurs and aetosaurs are characterized by the presence of well-developed osteoderms. Here we provide a comparative study on the microstructure of phytosaur (five taxa) and aetosaur (thirteen taxa) osteoderms. For outgroup comparison, we sampled osteoderms of the sister taxon to Aetosauria, Revueltosaurus callenderi, and the doswelliid Jaxtasuchus salomoni. Phytosaur, aetosaur, and Jaxtasuchus osteoderms are composed of a diploe structure, whereas the Revueltosaurus osteoderm microanatomy is more compact. The external cortex of phytosaurs, Revueltosaurus and Jaxtasuchus osteoderms is mainly composed of parallel-fibered bone. In aetosaurs, the external cortex mainly consists of lamellar bone, with lines of resorption within the primary bone indicating successive cycles of bone erosion and deposition. The basal cortex in all the specimens is composed of parallel-fibered bone, with the cancellous internal core being more strongly developed in aetosaurs than in phytosaurs. Woven or fibro-lamellar bone was recorded in both phytosaurian and aetosaurian taxa, as well as in Jaxtasuchus. Structural fibers, which at least partly suggest metaplastic origin, were only recorded in the internal core of two phytosaurs and in the
basal cortex of one aetosaur. Osteoderm thickness and cancellous to compact bone ratios appear to be subject to ontogenetic change. Minimum growth mark counts in osteoderms sampled indicate that some aetosaurs and phytosaurs lived for at least two decades. Bone microstructures are more uniform in phytosaur osteoderms and show a higher level of disparity among aetosaur osteoderms, and at least in the latter, histological features are potentially apomorphic for species/genus level.

Redescription of "Paleorhinus" (Phytosauria) Specimens from Germany

Butler, R. J., Rauhut, O. W. M., Stocker, M. R., and R. Bronowicz. 2013. Redescription of the phytosaurs Paleorhinus (‘Francosuchus’) angustifrons and Ebrachosuchus neukami from Germany, with implications for Late Triassic biochronology. Zoological Journal of the Linnean Society Early View. DOI: 10.1111/zoj.12094

 
Abstract - Phytosaurs are a diverse and morphologically distinctive clade of superficially crocodile-like archosauriforms that had a near global distribution during the Late Triassic. Because their remains are among the most abundant vertebrate remains recovered in many Upper Triassic terrestrial formations, phytosaurs are used extensively in long-range biochronological and biostratigraphic correlations. The biochronologically oldest and earliest branching known phytosaurs include an array of nominal species from the early Late Triassic of the United States, Germany, Poland, Morocco, and India that have been synonymized within the genus Paleorhinus, and subsequently used to define a global ‘Paleorhinus biochron’. However, recent phylogenetic work suggested that the North American species previously referred to Paleorhinus are paraphyletic. Here, we reassess the systematics and anatomy of putative specimens of Paleorhinus from southern Germany. Two well-preserved basal phytosaur skulls from the Blasensandstein (Carnian) of Bavaria form the holotypes of Francosuchus angustifrons and Ebrachosuchus neukami, both of which were synonymized with Paleorhinus by previous workers. We demonstrate that Francosuchus angustifrons shares unique synapomorphies with specimens referred to Paleorhinus bransoni from the Late Triassic of Texas, and thus refer the species to Paleorhinus. By contrast, the longirostrine Ebrachosuchus is highly distinctive in morphology, and our new cladistic analysis of Phytosauria demonstrates that it represents a valid taxon that is more closely related to Phytosauridae than to Paleorhinus. We provide the first autapomorphy-based support for a monophyletic but restricted Paleorhinus (supported by a nodal row on the jugal, and low paired ridges on the squamosal) and confirm that previous broader conceptions of Paleorhinus are likely to be paraphyletic.