Understanding the origin, expansion and loss of biodiversity is fundamental to evolutionary biology. The approximately 26 living species of crocodylomorphs (crocodiles, caimans, alligators and gharials) represent just a snapshot of the group's rich 230-million-year history, whereas the fossil record reveals a hidden past of great diversity and innovation, including ocean and land-dwelling forms, herbivores, omnivores and apex predators. In this macroevolutionary study of skull and jaw shape disparity, we show that crocodylomorph ecomorphological variation peaked in the Cretaceous, before declining in the Cenozoic, and the rise and fall of disparity was associated with great heterogeneity in evolutionary rates. Taxonomically diverse and ecologically divergent Mesozoic crocodylomorphs, like marine thalattosuchians and terrestrial notosuchians, rapidly evolved novel skull and jaw morphologies to fill specialized adaptive zones. Disparity in semi-aquatic predatory crocodylians, the only living crocodylomorph representatives, accumulated steadily, and they evolved more slowly for most of the last 80 million years, but despite their conservatism there is no evidence for long-term evolutionary stagnation. These complex evolutionary dynamics reflect ecological opportunities, that were readily exploited by some Mesozoic crocodylomorphs but more limited in Cenozoic crocodylians.

https://royalsocietypublishing.org/doi/pdf/10.1098/rspb.2021.0069

Ecological opportunity and the rise and fall of crocodylomorph evolutionary innovation.

Climatic constraints on the biogeographic history of Mesozoic dinosaurs

Notosuchia is a clade of crocodyliforms that was highly successful and diverse in the Cretaceous of Gondwana. Araripesuchus gomesii is a small notosuchian from the Early Cretaceous of Brazil that belongs to Uruguaysuchidae, one of the subgroups of notosuchians that first radiated, during the Aptian-Albian. Here we present a finite element analysis of A. gomesii based on a model reconstructed from CT scans and performed using published bone properties for crocodiles. The adductor musculature and their respective attachment areas were reconstructed based on Extant Phylogenetic Bracket. Different functional scenarios were tested applying an estimated 158 N bite force: unilateral bite, bilateral bite, pullback, head-shake, and head-twist. The results obtained were compared with those of Alligator mississippiensis, one of its closest living relatives. In the different simulations, the skull and lower jaws of Araripesuchus suffers more stress in the head-shake movement, followed by the unilateral and pullback bites with stress focalized in the premaxillary region. In contrast, the head-twist is the one with smaller stress values. Araripesuchus possess an oreinirostral skull that may provide greater overall resistance in the different scenarios on average, unlike Alligator that has a platyrostral skull with less resistance to dorsoventral mechanical loads. Previous hypotheses that considered A. gomesii as omnivorous coupled with our results, its small size, and likely limited bite force, suggest this taxon probably fed on small prey and other trophic items that could catch and handle entirely with its mouth, such as insects and small vertebrates.

Biomechanical performance of the cranio-mandibular complex of the small notosuchian Araripesuchus gomesii (Notosuchia, Uruguaysuchidae)

New data on the palaeosteohistology and growth dynamic of the notosuchian Araripesuchus Price, 1959

Osteoderms of eight extant and extinct species of crocodylomorphs are studied histologically and morphologically. Most osteoderms display the typical “crocodilian” structure with a woven‐fibered matrix surrounded by an upper and a lower parallel fibered matrix. The dorsal ornamentation of those specimens consists of a pit‐and‐ridge structure, with corresponding remodeling mechanisms. However, an osteoderm of Iberosuchus, studied here for the first time, differs in being nearly devoid of ornamentation; moreover, it shows strong bundles of straight Sharpey's fibers perpendicular to the surface in its lateral and dorsal walls, along with a rough plywood‐like structure in its basal plate. This suggests that this osteoderm was more deeply anchored within the dermis than the other osteoderms studied hitherto. This peculiar structure might have been linked to a terrestrial ecology and a specific thermoregulation strategy. Some other notosuchians in our sample do not exhibit ornamentation on their osteoderms, as opposed to neosuchians. Considering current interpretations of osteoderm function(s) in crocodilians, our observations are discussed in reference to possible ecophysiological peculiarities of Notosuchia in general, and Iberosuchus in particular.

A survey of osteoderm histology and ornamentation among Crocodylomorpha: A new proxy to infer lifestyle?

\n Most Notosuchia were active terrestrial predators. A few were semi-aquatic, or were insectivorous, omnivorous or herbivorous. A question relative to their thermometabolism remains to be answered: were Notosuchia warm-blooded? Here we use quantitative bone palaeohistology to answer this question. Two variables were used as proxies to infer thermometabolism: resting metabolic rate and red blood cell dimensions. Resting metabolic rate was inferred using relative primary osteon area and osteocyte size, shape and density. Blood cell dimensions were inferred using harmonic mean canal diameter and minimum canal diameter. All inferences were performed using phylogenetic eigenvector maps. Both sets of analyses suggest that the seven species of Notosuchia sampled in this study were ectotherms. Given that extant Neosuchia (their sister group) are also ectotherms, and that archosaurs were primitively endotherms, parsimony suggests that endothermy may have been lost at the node Metasuchia (Notosuchia–Neosuchia) by the Early Jurassic. Semi-aquatic taxa such as Pepesuchus may have had thermoregulatory strategies similar to those of recent crocodylians, whereas the terrestrial taxa (Araripesuchus, Armadillosuchus, Iberosuchus, Mariliasuchus, Stratiotosuchus) may have been thermoregulators similar to active predatory varanids. Thermal inertia may have contributed to maintaining a stable temperature in large notosuchians such as Baurusuchus.

Guillaume Houee

Papers

Were Notosuchia (Pseudosuchia: Crocodylomorpha) warm-blooded? A palaeohistological analysis suggests ectothermy

Expression of 20 SCPP genes during tooth and bone mineralization in Senegal bichir.