Sterling J. Nesbitt

The Early Evolution of Archosaurs: Relationships and the Origin of Major Clades

Abstract Archosaurs have a nearly 250 million year record that originated shortly after the Permian-Triassic extinction event and is continued today by two extant clades, the crocodylians and the avians. The two extant lineages exemplify two bauplan extremes among a diverse and complex evolutionary history, but little is known about the common ancestor of these lineages. Renewed interest in early archosaurs has led to nearly a doubling of the known taxa in the last 20 years. This study presents a thorough phylogenetic analysis of 80 species-level taxa ranging from the latest Permian to the early part of the Jurassic using a dataset of 412 characters. Each terminal taxon is explicitly described and all specimens used in the analysis are clearly stated. Additionally, each character is discussed in detail and nearly all of the character states are illustrated in either a drawing or highlighted on a specimen photograph. A combination of novel characters and comprehensive character sampling has bridged previously published analyses that focus on particular archosauriform subclades. A well-resolved, robustly supported consensus tree (MPTs  =  360) found a monophyletic Archosauria consisting of two major branches, the crocodylian-line and avian-line lineages. The monophyly of clades such as Ornithosuchidae, Phytosauria, Aetosauria, Crocodylomorpha, and Dinosauria is supported in this analysis. However, phytosaurs are recovered as the closest sister taxon to Archosauria, rather than basal crocodylian-line archosaurs, for the first time. Among taxa classically termed as “rauisuchians,” a monophyletic poposauroid clade was found as the sister taxon to a group of paraphyletic “rauisuchians” and monophyletic crocodylomorphs. Hence, crocodylomorphs are well nested within a clade of “rauisuchians,” and are not more closely related to aetosaurs than to taxa such as Postosuchus. Basal crocodylomorphs such as Hesperosuchus and similar forms (“Sphenosuchia”) were found as a paraphyletic grade leading to the clade Crocodyliformes. Among avian-line archosaurs, Dinosauria is well supported. A monophyletic clade containing Silesaurus and similar forms is well supported as the sister taxon to Dinosauria. Pterosaurs are robustly supported at the base of the avian line. A time-calibrated phylogeny of Archosauriformes indicates that the origin and initial diversification of Archosauria occurred during the Early Triassic following the Permian-Triassic extinction. Furthermore, all major basal archosaur lineages except Crocodylomorpha were established by the end of the Anisian. Early archosaur evolution is characterized by high rates of homoplasy, long ghost lineages, and high rates of character evolution. These data imply that much of the early history of Archosauria has not been recovered from the fossil record. Not only were archosaurs diverse by the Middle Triassic, but they had nearly a cosmopolitan biogeographic distribution by the end of the Anisian.

A Jurassic ceratosaur from China helps clarify avian digital homologies

Theropods have traditionally been assumed to have lost manual digits from the lateral side inward, which differs from the bilateral reduction pattern seen in other tetrapod groups. This unusual reduction pattern is clearly present in basal theropods, and has also been inferred in non-avian tetanurans based on identification of their three digits as the medial ones of the hand (I-II-III). This contradicts the many developmental studies indicating II-III-IV identities for the three manual digits of the only extant tetanurans, the birds. Here we report a new basal ceratosaur from the Oxfordian stage of the Jurassic period of China (156–161 million years ago), representing the first known Asian ceratosaur and the only known beaked, herbivorous Jurassic theropod. Most significantly, this taxon possesses a strongly reduced manual digit I, documenting a complex pattern of digital reduction within the Theropoda. Comparisons among theropod hands show that the three manual digits of basal tetanurans are similar in many metacarpal features to digits II-III-IV, but in phalangeal features to digits I-II-III, of more basal theropods. Given II-III-IV identities in avians, the simplest interpretation is that these identities were shared by all tetanurans. The transition to tetanurans involved complex changes in the hand including a shift in digit identities, with ceratosaurs displaying an intermediate condition.

Ecologically distinct dinosaurian sister group shows early diversification of Ornithodira.

The early evolutionary history of Ornithodira (avian-line archosaurs) has hitherto been documented by incomplete (Lagerpeton) or unusually specialized forms (pterosaurs and Silesaurus). Recently, a variety of Silesaurus-like taxa have been reported from the Triassic period of both Gondwana and Laurasia, but their relationships to each other and to dinosaurs remain a subject of debate. Here we report on a new avian-line archosaur from the early Middle Triassic (Anisian) of Tanzania. Phylogenetic analysis places Asilisaurus kongwe gen. et sp. nov. as an avian-line archosaur and a member of the Silesauridae, which is here considered the sister taxon to Dinosauria. Silesaurids were diverse and had a wide distribution by the Late Triassic, with a novel ornithodiran bauplan including leaf-shaped teeth, a beak-like lower jaw, long, gracile limbs, and a quadrupedal stance. Our analysis suggests that the dentition and diet of silesaurids, ornithischians and sauropodomorphs evolved independently from a plesiomorphic carnivorous form. As the oldest avian-line archosaur, Asilisaurus demonstrates the antiquity of both Ornithodira and the dinosaurian lineage. The initial diversification of Archosauria, previously documented by crocodilian-line archosaurs in the Anisian, can now be shown to include a contemporaneous avian-line radiation. The unparalleled taxonomic diversity of the Manda archosaur assemblage indicates that archosaur diversification was well underway by the Middle Triassic or earlier.

A Complete Skeleton of a Late Triassic Saurischian and the Early Evolution of Dinosaurs

Early Dinosaur Discovery Our understanding of the evolution of early dinosaurs is hampered by limited material, especially compared to the many Jurassic and Cretaceous samples. Nesbitt et al. (p. 1530) provide a complete view of a Late Triassic theropod based on several nearly complete skeletons from New Mexico. The dinosaur elucidates the likely relationships between early theropods and shows that some prominent features were already derived by this time. Comparison among Triassic dinosaur fauna and other early species suggests that Triassic North American fauna were diverse but not endemic, perhaps arising from earlier migrants from South America. A complete theropod from New Mexico implies that early dinosaurs dispersed widely, perhaps originating from South America. Characterizing the evolutionary history of early dinosaurs is central to understanding their rise and diversification in the Late Triassic. However, fossils from basal lineages are rare. A new theropod dinosaur from New Mexico is a representative of the early North American diversification. Known from several nearly complete skeletons, it reveals a mosaic of plesiomorphic and derived features that clarify early saurischian dinosaur evolution and provide evidence for the antiquity of novel avian character systems including skeletal pneumaticity. The taxon further reveals latitudinal differences among saurischian assemblages during the Late Triassic, demonstrates that the theropod fauna from the Late Triassic of North America was not endemic, and suggests that intercontinental dispersal was prevalent during this time.

Early ornithischian dinosaurs: the Triassic record

Abstract Ornithischian dinosaurs are one of the most taxonomically diverse dinosaur clades during the Mesozoic, yet their origin and early diversification remain virtually unknown. In recent years, several new Triassic ornithischian taxa have been proposed, mostly based upon isolated teeth. New discoveries of skeletal material of some of these tooth taxa indicate that these teeth can no longer be assigned to the Ornithischia using unambiguous synapomorphies. The Triassic record of ornithischian dinosaurs now comprises only three probable occurrences: Pisanosaurus and an unnamed heterodontosaurid from Argentina, and an unnamed specimen from the uppermost Triassic of South Africa. This revised Triassic record suggests that ornithischians were not very diverse or abundant through the Triassic, and there are large gaps in the Triassic ornithischian fossil record. Moreover, traditional living analogues for interpreting the feeding ecology of early ornithischians from their tooth morphology are generally inappropriate, and “herbivorous” archosaur teeth such as those found in early ornithischians are not necessarily diagnostic of herbivorous feeding.

A critical re‐evaluation of the Late Triassic dinosaur taxa of North America

Synopsis The North American Triassic dinosaur record has been repeatedly cited as one of the most complete early dinosaur assemblages. The discovery of Silesaurus from Poland and the recognition that Herrerasaurus and Eoraptor may not be theropods have forced a re‐evaluation of saurischian and theropod synapomorphies. Here, we re‐evaluate each purported Triassic dinosaur from North America on a specimen by specimen basis using an apomorphy‐based approach. We attempt to assign specimens to the most exclusive taxon possible. Our revision of purported Late Triassic dinosaur material from North America indicates that dinosaurs were rarer and less diverse in these strata than previously thought. This analysis concludes that non‐dinosaurian dinosauriforms were present in North America in the Late Triassic. Most of the proposed theropod specimens are fragmentary and/or indistinguishable from corresponding elements in the only well‐known Triassic theropod of North America, Coelophysis bauri. No Triassic material from North America can be assigned to Sauropodomorpha, because none of the purported ‘prosauropod’ material is diagnostic. Recent discovery of the skull and skeleton of Revueltosaurus callenderi from Arizona shows that it is a pseudosuchian archosaur, not an ornithischian dinosaur. As a result, other purported North American ornithischian teeth cannot be assigned to the Ornithischia and therefore, there are no confirmed North American Triassic ornithischians. Non‐tetanuran theropods and possible basal saurischians are the only identifiable dinosaurs recognised in North America until the beginning of the Jurassic Period.

Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda)

Living archosaurs comprise birds (dinosaurs) and crocodylians (suchians). The morphological diversity of birds and stem group dinosaurs is tremendous and well-documented. Suchia, the archosaurian group including crocodylians, is generally considered more conservative. Here, we report a new Late Triassic suchian archosaur with unusual, highly specialized features that are convergent with ornithomimid dinosaurs. Several derived features of the skull and postcranial skeleton are identical to conditions in ornithomimids. Such cases of extreme convergence in multiple regions of the skeleton in two distantly related vertebrate taxa are rare. This suggests that these archosaurs show iterative patterns of morphological evolution. It also suggests that this group of suchians occupied the adaptive zone that was occupied by ornithomimosaurs later in the Mesozoic.

Osteology of the Middle Triassic pseudosuchian archosaur Arizonasaurus babbitti

Arizonasaurus babbitti is rediagnosed from a well-preserved partial skeleton from the early Middle Triassic Moenkopi Formation of northern Arizona. The skull, excluding the braincase, is described and compared to other “rauisuchians” and basal archosaurs. Two possible autapomorphies include a deep fossa hidden from lateral and medial views on the posteroventral edge of the ascending process of the maxilla and a tongue (pubic peduncle of the ilium) and groove (proximal portion of the pubis) articulation between the pubis and the ilium. The postcrania is described in detail. Tall neural spines form a sail that is similar to those of other basal archosaurs with unknown affinities (Ctenosauriscus, Lotosaurus, Bromsgroveia, “Hypselorhachis”). Additional characters in the pelvic girdle in Arizonasaurus unite this taxon with Ctenosauriscus, Lotosaurus, Bromsgroveia, and Hypselorhachus to form a group, the ctenosauriscids. Pelvic characters of the ctenosauriscids are also found in Poposaurus and “Chatterjeea”, suggesting that ctenosauriscids are closely related. A possibly monophyletic group containing Arizonasaurus, other ctenosauriscids, Poposaurus, Sillosuchus, and “Chatterjeea” the exclusions of most other rauisuchians is proposed based on a new understanding of pseudosuchian characters.

The Late Triassic pseudosuchian Revueltosaurus callenderi and its implications for the diversity of early ornithischian dinosaurs

A new discovery of skeletons of Revueltosaurus callenderi from the Upper Triassic Chinle Formation of Petrified Forest National Park, Arizona clearly shows that Revueltosaurus is not an ornithischian dinosaur as previously supposed. Features such as the presence of a postfrontal, crocodile-normal ankle and paramedian osteoderms with anterior bars place R. callenderi within the Pseudosuchia, closer to crocodylomorphs than to dinosaurs. Therefore, dental characters previously used to place Revueltosaurus within the Ornithischia evolved convergently among other archosaur taxa, and cannot be used to diagnose ornithischian dinosaur teeth. As a result, all other putative North American Late Triassic ornithischians, which are all based exclusively on teeth, are cast into doubt. The only reasonably well-confirmed Late Triassic ornithischians worldwide are Pisanosaurus mertii and an unnamed heterodontosaurid from Argentina. This considerably changes the understanding of early dinosaur diversity, distribution and evolution in the Late Triassic.

Provincialization of terrestrial faunas following the end-Permian mass extinction

In addition to their devastating effects on global biodiversity, mass extinctions have had a long-term influence on the history of life by eliminating dominant lineages that suppressed ecological change. Here, we test whether the end-Permian mass extinction (252.3 Ma) affected the distribution of tetrapod faunas within the southern hemisphere and apply quantitative methods to analyze four components of biogeographic structure: connectedness, clustering, range size, and endemism. For all four components, we detected increased provincialism between our Permian and Triassic datasets. In southern Pangea, a more homogeneous and broadly distributed fauna in the Late Permian (Wuchiapingian, ∼257 Ma) was replaced by a provincial and biogeographically fragmented fauna by Middle Triassic times (Anisian, ∼242 Ma). Importantly in the Triassic, lower latitude basins in Tanzania and Zambia included dinosaur predecessors and other archosaurs unknown elsewhere. The recognition of heterogeneous tetrapod communities in the Triassic implies that the end-Permian mass extinction afforded ecologically marginalized lineages the ecospace to diversify, and that biotic controls (i.e., evolutionary incumbency) were fundamentally reset. Archosaurs, which began diversifying in the Early Triassic, were likely beneficiaries of this ecological release and remained dominant for much of the later Mesozoic.