Predatory synapsid ecomorphology signals growing dynamism of late Palaeozoic terrestrial ecosystems.

Terrestrial ecosystems evolved substantially through the Palaeozoic, especially the Permian, gaining much new complexity, especially among predators. Key among these predators were non-mammalian synapsids. Predator ecomorphology reflect interactions with prey and competitors, which are key controls on carnivore diversity and ecology. Therefore, carnivorous synapsids may offer insight on wider ecological evolution as the first complex, tetrapod-dominated, terrestrial ecosystems formed through the late Palaeozoic. Using morphometric and phylogenetic comparative methods, we chart carnivorous synapsid trophic morphology from the latest Carboniferous to the earliest Triassic (307-251.2 Ma). We find a major morphofunctional shift in synapsid carnivory between the early and middle Permian, via the addition of new feeding modes increasingly specialised for greater biting power or speed that captures the growing antagonism and dynamism of terrestrial tetrapod predator-prey interactions. The further evolution of new hypo- and hypercarnivorous synapsids highlight the nascent intrinsic pressures and complexification of terrestrial ecosystems across the mid-late Permian.

Locomotion and the early Mesozoic success of Archosauromorpha.

The Triassic was a time of ecological upheaval as life recovered from the Permian-Triassic mass extinction. Archosauromorphs were a key component of the recovery, diversifying substantially during the Triassic and encompassing the origins of dinosaurs, pterosaurs and crocodylomorphs. Here, we explore the evolution of locomotion in Archosauromorpha to test whether dinosaurs show any distinctive locomotory features that might explain their success. We implement geometric morphometrics on limb bone shapes and use limb ratios to calculate bipedality and cursoriality metrics. We find that the Avemetatarsalia (dinosaurs, pterosaurs and relatives) exhibit more variable limb form and limb ratios than any other group, indicating a wider range of locomotory modes. The earliest avemetatarsalians were bipedal and cursorial, and their range of form increased through the Triassic with notable diversification shifts following extinction events. This is especially true of dinosaurs, even though these changes cannot be discriminated from a stochastic process. By contrast, the Pseudosuchia (crocodilians and relatives) were more restricted in limb form and locomotor mode with disparity decreasing through time, suggesting more limited locomotor adaptation and vulnerability to extinction. Perhaps the greater locomotor plasticity of dinosaurs gave them a competitive advantage in the changing climates of the Late Triassic.

A Carboniferous synapsid with caniniform teeth and a reappraisal of mandibular size-shape heterodonty in the origin of mammals.

Heterodonty is a hallmark of early mammal evolution that originated among the non-mammalian therapsids by the Middle Permian. Nonetheless, the early evolution of heterodonty in basal synapsids is poorly understood, especially in the mandibular dentition. Here, we describe a new synapsid, Shashajaia bermani gen. et sp. nov., based on a well-preserved dentary and jaw fragments from the Carboniferous-Permian Halgaito Formation of southern Utah. Shashajaia shares with some sphenacodontids enlarged (canine-like) anterior dentary teeth, a dorsoventrally deep symphysis and low-crowned, subthecodont postcanines having festooned plicidentine. A phylogenetic analysis of 20 taxa and 154 characters places Shashajaia near the evolutionary divergence of Sphenacodontidae and Therapsida (Sphenacodontoidea). To investigate the ecomorphological context of Palaeozoic sphenacodontoid dentitions, we performed a principal component analysis based on two-dimensional geometric morphometrics of the mandibular dentition in 65 synapsids. Results emphasize the increasing terrestrialization of predator-prey interactions as a driver of synapsid heterodonty; enhanced raptorial biting (puncture/gripping) aided prey capture, but this behaviour was probably an evolutionary antecedent to more complex processing (shearing/tearing) of larger herbivore prey by the late Early to Middle Permian. The record of Shashajaia supports the notion that the predatory feeding ecology of sphenacodontoids emerged in palaeotropical western Pangea by late Carboniferous times.

Niche partitioning shaped herbivore macroevolution through the early Mesozoic.

The Triassic (252-201 Ma) marks a major punctuation in Earth history, when ecosystems rebuilt themselves following the devastating Permian-Triassic mass extinction. Herbivory evolved independently several times as ecosystems comprising diverse assemblages of therapsids, parareptiles and archosauromorphs rose and fell, leading to a world dominated by dinosaurs. It was assumed that dinosaurs prevailed either through long-term competitive replacement of the incumbent clades or rapidly and opportunistically following one or more extinction events. Here we use functional morphology and ecology to explore herbivore morphospace through the Triassic and Early Jurassic. We identify five main herbivore guilds (ingestion generalists, prehension specialists, durophagous specialists, shearing pulpers, and heavy oral processors), and find that herbivore clades generally avoided competition by almost exclusively occupying different guilds. Major ecosystem remodelling was triggered multiple times by external environmental challenges, and previously dominant herbivores were marginalised by newly emerging forms. Dinosaur dominance was a mix of opportunity following disaster, combined with competitive advantage in their new world.