To cite this article: J. Michael Parrish (1993): Phylogeny of the Crocodylotarsi, with reference to archosaurian and crurotarsan monophyly, Journal of Vertebrate Paleontology, 13:3, 287-308To link to this article: http://dx.

During the Triassic, archosauromorphs became one of the first groups of diapsid reptiles to diversify in terms of body size and morphological disparity in both terrestrial and marine ecosystems across Pangaea. This seemingly rapid divergence, and the numerous unique body plans stemming from it, concomitantly has confounded reconstructions of archosauromorph relationships. Teasing apart homology from homoplasy of anatomical characters in this broad suite of body types remains an enormous challenge with the current sample of taxa. Here, we present the postcranial anatomy of Azendohsaurus madagaskarensis, an early archosauromorph from ?Middle to Upper Triassic strata of Madagascar. Azendohsaurus madagaskarensis is known from nearly the entire skeleton in an ontogenetically variable sample. The holotype locality consists of a monotypic bone bed; preservation ranges from complete but disarticulated bones to articulated sections of the skeleton. Azendohsaurus madagaskarensis embodies an aberrant constellation of archosauromorph features, including an elongated neck, a short, stocky tail, robust limbs, and unexpectedly short digits terminating in large recurved unguals on the manus and pes. Together with the cranium, the postcrania reveal A. madagaskarensis to be another representative of a growing coterie of highly apomorphic and bizarre Triassic archosauromorphs. At the same time, recovery and description of the full anatomy of A. madagaskarensis helps to identify a monophyletic grouping of specialized taxa that includes the North American Late Triassic-aged archosauromorphs Trilophosaurus, Spinosuchus, and Teraterpeton, Indian Pamelaria, and Moroccan Azendohsaurus laaroussii. Moreover, information derived from the skeleton of A. madagaskarensis solidifies the systematic position of these taxa among other archosauromorphs. Using the most comprehensively sampled phylogenetic analysis of early archosauromorphs, we found the clade encompassing the aforementioned taxa as the nearest outgroup of Prolacerta broomi + Archosauriformes. The newly recognized clade containing Azendohsaurus, Trilophosaurus, Spinosuchus, Pamelaria, and Teraterpeton demonstrates high morphological disparity even within a closely related group of archosauromorphs, underscores the polyphyly of protorosaurs (5 prolacertiforms), and suggests that most major divergences within this group occurred in the Triassic. Furthermore, our results indicate that craniodental character states ascribed to a herbivorous diet were much more pervasive across Triassic Archosauromorpha than previously conjectured.

Articulated digital reconstructions of two diplodocid sauropods revealed cervical poses and feeding envelopes. The necks of Diplodocus and Apatosaurus were nearly straight but gently declined such that the heads, which were themselves angled downward relative to the neck, were close to ground level in their neutral, undeflected posture. Both necks were less flexible than conventionally depicted, and Diplodocus was less capable of lateral and dorsal curvature than Apatosaurus. The results suggest that these sauropods were adapted to ground feeding or low browsing, contrary to the view that diplodocid sauropods were high browsers.

Here, we describe a new species of Azendohsaurus from the Middle-Late Triassic of Madagascar, extending the geographical range of a taxon known otherwise only by a single species from Morocco. Although Azendohsaurus has consistently been regarded as an early dinosaur (based on various advanced dental and gnathic features resembling those characterizing certain dinosaur subgroups), the relatively complete skeletal material, now available from Madagascar, argues strongly against its dinosaurian affinities. Rather, the retention of numerous primitive cranial and postcranial features indicates a surprisingly early divergence of Azendohsaurus within Archosauromorpha and an unusual mosaic of characters in this taxon. Features considered diagnostic of Sauropodomorpha thus are inferred to occur homoplastically in at least one clade of nondinosaurian archosauromorphs, indicating a complex evolution and distribution of features traditionally thought to be derived within archosaurs. Azendohsaurus has teeth resembling those of both early sauropodomorph and ornithischian dinosaurs, yet also possesses numerous inarguable basal archosauromorph cranial and postcranial attributes. This highlights the risk of uncritically referring isolated, Middle-Late Triassic (or even later), 'leafshaped' teeth with denticles to the Dinosauria. Similarly, the occurrence of such teeth in an early diverging archosauromorph indicates that specializations for herbivory originated more frequently within this clade than conventionally assumed. For example, Azendohsaurus and numerous basal sauropodomorph dinosaur taxa share an array of convergently acquired features associated with herbivory, including tooth denticles, expanded tooth crowns, a downturned dentary and the articular located at the ventral margin of the mandible. Some of these features (denticles, expanded crowns and the ventrally deflected articular) are even more widespread among archosauromorphs, including aetosaurs, silesaurs and ornithischian dinosaurs. A downturned dentary also occurs in Trilophosaurus, a taxon further marked by unique specializations for herbivory, including transversely lophate, tricuspid teeth. An array of features associated with herbivory also occurs in rhynchosaurs and certain crocodilians (e.g. Simosuchus). This distribution suggests that craniodental features associated with herbivory were much more pervasive across the archosauromorph clade than previously recognized, possibly evolving at least six to eight times independently.

The morphology of the tarsi, hindlimbs, and pelves of the earliest crocodilians and their nearest relatives, Hallopus and the “sphenosuchians,” indicates that these animals had adaptations for erect posture. The widespread distribution of apparently homologous adaptations for erect gait among the archosaurs with crocodile-normal tarsi suggests that those structures are plesiomorphic for this group, which comprises the Aetosauria, “rauisuchians,” “sphenosuchians,” Hallopus, and the Crocodylia. Adaptations for erect posture are seen most clearly in the structure of the proximal tarsus (astragalus and calcaneum).An important implication of this argument is that the most primitive crocodylomorphs, comprising the “protosuchian” crocodiles, the “sphenosuchians,” and Hallopus, had an erect stance and gait. The sprawling stance and associated gait used by modern crocodilians during swimming and upon entering the water can be viewed as secondary adaptations to an aquatic existence. The environments of deposition and faunal associations of “sphenosuchians” and “protosuchian” crocodiles are consistent with primarily terrestrial habits. Living crocodilians have two types of step cycles, sprawling and erect; the sprawling pattern is overprinted onto the inferred ancestral “high-walk,” and onto the gallop sometimes used by juvenile crocodilians.