Sauropodomorph evolution across the Triassic-Jurassic boundary: body size, locomotion, and their influence on morphological disparity.

Sauropodomorph dinosaurs were the dominant medium to large-sized herbivores of most Mesozoic continental ecosystems, being characterized by their long necks and reaching a size unparalleled by other terrestrial animals (> 60 tonnes). Our study of morphological disparity across the entire skeleton shows that during the Late Triassic the oldest known sauropodomorphs occupied a small region of morphospace, subsequently diversifying both taxonomically and ecologically, and shifting to a different and broader region of the morphospace. After the Triassic-Jurassic boundary event, there are no substancial changes in sauropodomorph morphospace occupation. Almost all Jurassic sauropodomorph clades stem from ghost lineages that cross the Triassic-Jurassic boundary, indicating that variations after the extinction were more related to changes of pre-existing lineages (massospondylids, non-gravisaurian sauropodiforms) rather than the emergence of distinct clades or body plans. Modifications in the locomotion (bipedal to quadrupedal) and the successive increase in body mass seem to be the main attributes driving sauropodomorph morphospace distribution during the Late Triassic and earliest Jurassic. The extinction of all non-sauropod sauropodomorphs by the Toarcian and the subsequent diversification of gravisaurian sauropods represent a second expansion of the sauropodomorph morphospace, representing the onset of the flourishing of these megaherbivores that subsequently dominated in Middle and Late Jurassic terrestrial assemblages.

An early trend towards gigantism in Triassic sauropodomorph dinosaurs.

Dinosaurs dominated the terrestrial ecosystems for more than 140 Myr during the Mesozoic era, and among them were sauropodomorphs, the largest land animals recorded in the history of life. Early sauropodomorphs were small bipeds, and it was long believed that acquisition of giant body size in this clade (over 10 tonnes) occurred during the Jurassic and was linked to numerous skeletal modifications present in Eusauropoda. Although the origin of gigantism in sauropodomorphs was a pivotal stage in the history of dinosaurs, an incomplete fossil record obscures details of this crucial evolutionary change. Here, we describe a new sauropodomorph from the Late Triassic of Argentina nested within a clade of other non-eusauropods from southwest Pangaea. Members of this clade attained large body size while maintaining a plesiomorphic cyclical growth pattern, displaying many features of the body plan of basal sauropodomorphs and lacking most anatomical traits previously regarded as adaptations to gigantism. This novel strategy highlights a highly accelerated growth rate, an improved avian-style respiratory system, and modifications of the vertebral epaxial musculature and hindlimbs as critical to the evolution of gigantism. This reveals that the first pulse towards gigantism in dinosaurs occurred over 30 Myr before the appearance of the first eusauropods.

Novel insight into the origin of the growth dynamics of sauropod dinosaurs.

Sauropod dinosaurs include the largest terrestrial animals and are considered to have uninterrupted rapid rates of growth, which differs from their more basal relatives, which have a slower cyclical growth. Here we examine the bone microstructure of several sauropodomorph dinosaurs, including basal taxa, as well as the more derived sauropods. Although our results agree that the plesiomorphic condition for Sauropodomorpha is cyclical growth dynamics, we found that the hypothesized dichotomy between the growth patterns of basal and more derived sauropodomorphs is not supported. Here, we show that sauropod-like growth dynamics of uninterrupted rapid growth also occurred in some basal sauropodomorphs, and that some basal sauropods retained the plesiomorphic cyclical growth patterns. Among the sauropodomorpha it appears that the basal taxa exploited different growth strategies, but the more derived Eusauropoda successfully utilized rapid, uninterrupted growth strategies.

A new sphenodontian (Lepidosauria: Rhynchocephalia) from the Late Triassic of Argentina and the early origin of the herbivore opisthodontians.

Sphenodontians were a successful group of rhynchocephalian reptiles that dominated the fossil record of Lepidosauria during the Triassic and Jurassic. Although evidence of extinction is seen at the end of the Laurasian Early Cretaceous, they appeared to remain numerically abundant in South America until the end of the period. Most of the known Late Cretaceous record in South America is composed of opisthodontians, the herbivorous branch of Sphenodontia, whose oldest members were until recently reported to be from the Kimmeridgian-Tithonian (Late Jurassic). Here, we report a new sphenodontian, Sphenotitan leyesi gen. et sp. nov., collected from the Upper Triassic Quebrada del Barro Formation of northwestern Argentina. Phylogenetic analysis identifies Sphenotitan as a basal member of Opisthodontia, extending the known record of opisthodontians and the origin of herbivory in this group by 50 Myr.

A new basal sauropodomorph (Dinosauria: Saurischia) from Quebrada del Barro Formation (Marayes-El Carrizal Basin), northwestern Argentina.

BACKGROUND: Argentinean basal sauropodomorphs are known by several specimens from different basins; Ischigualasto, El Tranquilo, and Mogna. The Argentinean record is diverse and includes some of the most primitive known sauropodomorphs such as Panphagia and Chromogisaurus, as well as more derived forms, including several massospondylids. Until now, the Massospondylidae were the group of basal sauropodomorphs most widely spread around Pangea with a record in almost all continents, mostly from the southern hemisphere, including the only record from Antarctica. METHODOLOGY/PRINCIPAL FINDING: We describe here a new basal sauropodomorph, Leyesaurus marayensis gen. et sp. nov., from the Quebrada del Barro Formation, an Upper Triassic-Lower Jurassic unit that crops out in northwestern Argentina. The new taxon is represented by a partial articulated skeleton that includes the skull, vertebral column, scapular and pelvic girdles, and hindlimb. Leyesaurus is diagnosed by a set of unique features, such as a sharply acute angle (50 degrees) formed by the ascending process of the maxilla and the alveolar margin, a straight ascending process of the maxilla with a longitudinal ridge on its lateral surface, noticeably bulging labial side of the maxillary teeth, greatly elongated cervical vertebrae, and proximal articular surface of metatarsal III that is shelf-like and medially deflected. Phylogenetic analysis recovers Leyesaurus as a basal sauropodomorph, sister taxon of Adeopapposaurus within the Massospondylidae. Moreover, the results suggest that massospondylids achieved a higher diversity than previously thought. CONCLUSIONS/SIGNIFICANCE: Our phylogenetic results differ with respect to previous analyses by rejecting the massospondylid affinities of some taxa from the northern hemisphere (e.g., Seitaad, Sarahsaurus). As a result, the new taxon Leyesaurus, coupled with other recent discoveries, suggests that the diversity of massospondylids in the southern hemisphere was higher than in other regions of Pangea. Finally, the close affinities of Leyesaurus with the Lower Jurassic Massospondylus suggest a younger age for the Quebrada del Barro Formation than previously postulated.