Inferring the lifestyles of extinct Crocodyliformes using osteoderm ornamentation.

Osteoderms are bony plates formed within the dermis of diverse vertebrate groups. They are present in all crocodylomorphs but Metriorhynchidae. Most of them show typical bone ornamentation consisting of pits and ridges on their outer surface. The most widely discussed functional hypothesis suggests that the ornamentation of osteoderms influences heat exchange with the environment through the adjacent vascular network, facilitating the absorption of solar radiation. This process allows semiaquatic crocodiles to compensate for heat loss resulting from the high thermal conductivity of surrounding water. In order to test this assertion, we conducted a phylogenetic logistic regression analysis to evaluate the relationship between osteoderm relative area of pits (RAP) and lifestyle (terrestrial versus aquatic) in a sample of crocodyliforms. Our results revealed that lifestyle is significantly explained by RAP: the lower the degree of ornamentation (RAP), the higher the probability of a terrestrial lifestyle. We used this model to infer the lifestyle of two extinct taxa, Peirosaurus torminni and Microsuchus schilleri. We concluded that terrestrial notosuchians may have lost osteoderm ornamentation due to the lower thermal conductivity of air and reduced heat loss in a terrestrial environment compared to what happens in water. Among these notosuchians, we hypothesize that large terrestrial baurusuchids maintained a stable body temperature due to thermal inertia, whereas small notosuchians took advantage of the early morning sun exposure to warm up and stayed in terrestrial burrows during periods of intense solar radiation. Finally, unlike the almost motionless behavior of freshwater crocodiles, fully marine Metriorhynchidae probably lost osteoderms because they constantly swim, generating heat by muscular contraction, so osteoderms with a thermoregulatory function for heat absorption were no longer positively selected.