Aetosaurs: New Phylogenetic Analysis, New Taxon; and New Technique to Analyze Incongruent Character Datasets

My new paper in PeerJ features a new phylogenetic analysis of the Aetosauria (Archosauria: Pseudosuchia). I've added many new characters and feature all known valid aetosaur taxa. In the Supplemental Materials, each character is described and figured to clarify them for future use.


This paper also introduces a new aetosaur, Scutarx deltatylus, from the Chinle Formation of Arizona. To date I've collected four partial Scutarx skeletons, the first in 2002, from Petrified Forest National Park. The holotype specimen includes a partial skull, the first good aetosaur skull material collected from Arizona since the 1930s. In previous papers I have assigned all of these specimens to Calyptosuchus (Stagonolepis) wellesi; however, when I undertook a revision of that taxon I noted that my Petrified Forest specimens (all from the Sonsela Member of the Chinle Formation) all possessed raised triangular bosses on the posteromedial corner of the paramdian osteoderms.  These are not present in the type specimen from Texas, or from the large amount of material from the Placerias Quarry of Arizona. Interestingly the boss-bearing and non-boss-bearing specimens were separated stratigraphically. Presently the specimens from the Blue Mesa Member of Arizona and the Tecovas Formation of Texas are those that lack the boss (Calyptosuchus wellesi), and those from the Sonsela Member, and middle Cooper Canyon Formation are those that possess the boss (Scutarx deltatylus). The skull and ilium of Scutarx are also autapomorphic as discussed in the paper.

Finally I adapted the method to compute Partioned Bremer Support to look at the possibility of character conflict between anatomical partitions in aetosaurs. Essentially does the armor have a different phylogenetic signal than the rest of the skeleton. Bremer Support is used in all phylogenetic analyses to show branch support; however, what is generally not realized is that the individual support values of characters from each anatomical partition combine to total this number. But if these character sets actually support a different tree than the one recovered in the total character analysis, they will reduce the Bremer Support value (negative Bremer Support). Some character sets are ambivalent and actually provide no support to the branches.  This analytic tool allows you to look at your tree support node by node and see what character datasets support the topology.  This technique has been used since the late 1990s to compare molecular vs. morphological character sets, but this is the first time it has been employed to look at anatomical partitions in a purely morphological study.  

That said, I am a little disappointed with my final discussion section in the paper.  While I am excited about the potential for Partitioned Bremer Support, I still am learning all that it demonstrates, thus my discussion is not as strong as I would have liked. I hope that others will find it of interest and utility and that we can see if if it works for other taxonomic groups. 

Parker, W.G. 2016. Revised phylogenetic analysis of the Aetosauria (Archosauria: Pseudosuchia); assessing the effects of incongruent morphological character sets. PeerJ 4:e1583

Abstract- Aetosauria is an early-diverging clade of pseudosuchians (crocodile-line archosaurs) that had a global distribution and high species diversity as a key component of various Late Triassic terrestrial faunas. It is one of only two Late Triassic clades of large herbivorous archosaurs, and thus served a critical ecological role. Nonetheless, aetosaur phylogenetic relationships are still poorly understood, owing to an overreliance on osteoderm characters, which are often poorly constructed and suspected to be highly homoplastic. A new phylogenetic analysis of the Aetosauria, comprising 27 taxa and 83 characters, includes more than 40 new characters that focus on better sampling the cranial and endoskeletal regions, and represents the most comprenhensive phylogeny of the clade to date. Parsimony analysis recovered three most parsimonious trees; the strict consensus of these trees finds an Aetosauria that is divided into two main clades: Desmatosuchia, which includes the Desmatosuchinae and the Stagonolepidinae, and Aetosaurinae, which includes the Typothoracinae. As defined Desmatosuchinae now contains Neoaetosauroides engaeus and several taxa that were previously referred to the genus Stagonolepis, and a new clade, Desmatosuchini, is erected for taxa more closely related to Desmatosuchus. Overall support for some clades is still weak, and Partitioned Bremer Support (PBS) is applied for the first time to a strictly morphological dataset demonstrating that this weak support is in part because of conflict in the phylogenetic signals of cranial versus postcranial characters. PBS helps identify homoplasy among characters from various body regions, presumably the result of convergent evolution within discrete anatomical modules. It is likely that at least some of this character conflict results from different body regions evolving at different rates, which may have been under different selective pressures.

Cranial Anatomy of the Aetosaur Paratypothorax andressorum

Schoch, R. R., and J. B. Desojo. 2016. Cranial anatomy of the aetosaur Paratypothorax andressorum Long & Ballew, 1985, from the Upper Triassic of Germany and its bearing on aetosaur phylogeny. Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen 279(1):73-95. DOI:

Abstract -
The large aetosaur Paratypothorax andressorum has so far been known only by its osteoderms. Here we describe for the first time the skull of a complete, articulated specimen of this taxon that was found in the type horizon at Murrhardt, southwestern Germany. Paratypothorax
andressorum has the following cranial autapomorphies: (1) upper jaw margin with deep notch between premaxilla and maxilla, (2) maxilla-lacrimal suture with finger-like projection, (3) upper
temporal fenestra triangular, and (4) first paramedian cervical osteoderms narrow and oval, much smaller than second row. Apart from these features, the skull of P. andressorum closely resembles that of the small aetosaur Aetosaurus ferratus known from the same horizons, despite major differences in the morphology of osteoderms. Both taxa share (1) the pointed, beak-shaped premaxilla which expands only gently anterior to the nasal, (2) maxilla and lacrimal excluding jugal
from margin of antorbital fenestra, (3) exclusion of squamosal from margin of infratemporal fenestra, and (4) posterior part of jugal not downturned. Phylogenetic analysis reveals poorly resolved relationships within Aetosauria, but exclusion of a problematic taxon Coahomasuchus results in a much better resolution, with Paratypothorax to nest with Rioarribasuchus, Tecovasuchus, Typothorax, and Redondasuchus within a monophyletic Typothoracinae. Interestingly, Aetosaurus and Stenomyti form successive sister taxa of this clade rather than fall within an aetosaurine grade of basal aetosaurs, as suggested by previous authors. The resemblance of Paratypothorax and Aetosaurus in many cranial features, their close relationship as suggested by the present analysis, and the immature state of all available Aetosaurus specimens suggest two new alternative hypotheses: (1) Aetosaurus is the juvenile of a close relative of Paratypothorax or (2) it is itself the juvenile of Paratypothorax.

Archosauromorph from the Middle Permian of South America

Martinelli, A. G., Francischini, H., Dentzien-Dias, P. C., Soares, M. B., and C. L. Schultz. 2016.
The oldest archosauromorph from South America: postcranial remains from the Guadalupian (mid-Permian) Rio do Rasto Formation (Paraná Basin), southern Brazil. Historical Biology.

Abstract -
In this contribution, we report a distal portion of a left humerus that likely belongs to an indeterminate basal archosauromorph from the Guadalupian (mid-Permian) Rio do Rastro Formation (Paraná Basin) of southern Brazil. A precise taxonomy of the fragmented and isolated humerus UFRGS-PV-0546-P is not warranted at generic nor familiar level but, likely, this specimen belongs to an Archosauromorpha due to the lack of both the entepicondylar and the ectepicondylar foramina. The narrow distal end of the humerus, the rounded radial and ulnar condyles, and the moderately developed supinator process with a shallow ectepicondylar groove (not notched) are features reminiscent of tanystropheids rather than that of other archosauromorphs. This material likely represents the first and oldest Permian archosauromorph from South America and indicates the presence of this lineage before the P/T boundary.

The Early Evolution of Rhynchosaurs

Congratulations to Max Langer for being honored with his own rhynchosaur genus.

Ezcurra, M. D., Montefeltro, F., and R. J. Butler. 2016. The Early Evolution of Rhynchosaurs. Frontiers in Ecology and Evolution 3:142. DOI: 10.3389/fevo.2015.00142.

Abstract -
The rhynchosaurian archosauromorphs are an important and diverse group of fossil tetrapods that first appeared during the Early Triassic and probably became extinct during the early Late Triassic (early Norian). Here, the early evolution of rhynchosaurs during the Early and early Middle Triassic (Induan-Anisian: 252.2-242 Mya) is reviewed based on new anatomical observations and their implications for the taxonomy, phylogenetic relationships and macroevolutionary history of the group. A quantitative phylogenetic analysis recovered a paraphyletic genus Rhynchosaurus, with “Rhynchosaurusbrodiei more closely related to hyperodapedontines than to Rhynchosaurus articeps. Therefore, a new genus is erected, resulting in the new combination Langeronyx brodiei. A body size analysis found two independent increases in size in the evolutionary history of rhynchosaurs, one among stenaulorhynchines and the other in the hyperodapedontine lineage. Maximum likelihood fitting of phenotypic evolution models to body size data found ambiguous results, with body size evolution potentially interpreted as fitting either a non-directional Brownian motion model or a stasis model. A Dispersal-Extinction Cladogenesis analysis reconstructed the areas that are now South Africa and Europe as the ancestral areas of Rhynchosauria and Rhynchosauridae, respectively. The reconstruction of dispersal events between geographic areas that are broadly separated paleolatitudinally implies that barriers to the dispersal of rhynchosaurs from either side of the paleo-Equator during the Middle Triassic were either absent or permeable.

Archosauriform Remains from Late Triassic of San Luis Province, Argentina

Gianechini, F. A., Codorniú, L., Arcucci, A. B., Elías, G. C., and D. Rivarola. 2015. Archosauriform remains from the Late Triassic of San Luis province, Argentina, Quebrada del Barro Formation, Marayes–El Carrizal Basin. Journal of South American Earth Sciences. DOI:10.1016/j.jsames.2015.12.012

Here we present archosauriform remains from ‘Abra de los Colorados’, a fossiliferous locality at Sierra de Guayaguas, NW San Luis Province. Two fossiliferous levels were identified in outcrops of the Quebrada del Barro Formation (Norian), which represent the southernmost outcrops of the Marayes–El Carrizal Basin. These levels are composed by massive muddy lithofacies, interpreted as floodplain deposits. The specimens consist of one incomplete maxilla (MIC-V718), one caudal vertebra (MIC-V719), one metatarsal (MIC-V720) and one indeterminate appendicular bone (MIC-V721). The materials can be assigned to Archosauriformes but the fragmentary nature and lack of unambiguous synapomorphies preclude a more precise taxomic assignment. The maxilla is remarkably large and robust and represents the posterior process. It preserved one partially erupted tooth with ziphodont morphology. This bone shows some anatomical traits and size match with ‘rauisuchians’ and theropods. MIC-V719 corresponds to a proximal caudal vertebra. It has a high centrum, a ventral longitudinal furrow, expanded articular processes for the chevrons, a posteriorly displaced diapophysis located below the level of the prezygapophyses, and short prezygapophyses. This vertebra would be from an indeterminate archosauriform. MIC-V720 presents a cylindrical diaphysis, with a well-developed distal trochlea, which present resemblances with metatarsals of theropods, pseudosuchians, and silesaurids, although the size matches better with theropods. MIC-V721 has a slender diaphysis and a convex triangular articular surface, and corresponds to an indeterminate archosauriform. Despite being fragmentary, these materials indicate the presence of a diverse archosauriforms association from Late Triassic beds of San Luis. Thus, they add to the faunal assemblage recently reported from this basin at San Juan Province, which is much rich and diverse than the coeval paleofauna well known from Los Colorados Formation in the Ischigualasto–Villa Union Basin.