New Patagonian baurusuchids (Crocodylomorpha; Notosuchia) from the Bajo de la Carpa Formation (Upper Cretaceous; Neuquén,Argentina): New evidences of the early sebecosuchian diversification in Gondwana

The Bajo de la Carpa Formation (Santonian) of the Neuquén basin (southwestern Argentina) has one of the most complete crocodyliform records of South America, together with the Adamantina and Marília formations of Brazil. In this contribution we report two fragmentary individuals from two different localities of the Bajo de la Carpa Formation: a middle-sized individual (MLP 26-IV-30-2), represented by postcranial remains from the Boca del Sapo locality and a large-bodied individual (MACN Pv-RN 1150), consisting only of snout elements from Paso Córdoba locality. Despite the remains of both specimens are fragmentary, they display anatomical characters of a specialized notosuchian carnivore clade: Baurusuchidae. Our phylogenetic analysis recovers these new individuals as non-baurusuchine baurusuchids, although they take multiple phylogenetic positions on the different most parsimonious trees. These new remains highlight the relevance of the Bajo de la Carpa Formation for understanding the rise of notosuchians during the Late Cretaceous.     

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

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 new Sphagesaurus (Mesoeucrocodylia: Notosuchia) from the Upper Cretaceous of Monte Alto City (Bauru Group,Brazil), and a revision of the Sphagesauridae

Oxygen consumption and ammonia excretion rates were assessed for Terebratulina retusa (L.) held under 3 different regimes of temperature and food availability. These were: 5.6?C, no food (cold, starved); 5.8?C, food present (cold, fed) and 10.7?C food present (warm, fed), which simulated winter conditions, summer conditions and an intermediate treatment. Regressions of oxygen consumption on ash‐free dry weight (AFDW) had slopes which were not significantly different from each other and ranged from 0.953 to 0.999. A common slope of 0.976 was calculated and intercepts based on the common slope used to compare oxygen consumption in each treatment. The rise from cold, starved conditions to warm, fed was 24.5 per cent and this was significant (P < 0.05). Other differences were not significant (P > 0.05) but the cold, fed result was 12.6 per cent higher than cold, starved. Therefore feeding and temperature probably account for equivalent proportions of the rise in metabolism from winter to summer. Ammonia production data were much more variable. Excretion rates of a 50 mg AFDW individual (in ng‐at NH₃‐N.h^‐1) were as follows: cold, starved: 30.2 cold, fed: 7.1; and warm, fed: 22.9. Oxygen to nitrogen (O:N) ratios reflected these results. Mean O:N ratios were: cold, starved: 8.0; cold, fed: 42.4; warm, fed: 16.3. This shows that the simulated winter group relied heavily on protein to fuel their metabolism, the simulated summer group were less dependent on protein and the intermediate group probably used lipids and carbohydrates to fuel metabolic demands. This possibly reflected a trade off between food supply and increased metabolism from treatment to treatment, demonstrating a flexibility which could have been a contributing factor in the ecological tolerance and geological longevity of some brachiopods.     

Environmental drivers of crocodyliform extinction across the Jurassic/Cretaceous transition

Crocodyliforms have a much richer evolutionary history than represented by their extant descendants, including several independent marine and terrestrial radiations during the Mesozoic. However, heterogeneous sampling of their fossil record has obscured their macroevolutionary dynamics, and obfuscated attempts to reconcile external drivers of these patterns. Here, we present a comprehensive analysis of crocodyliform biodiversity through the Jurassic/Cretaceous (J/K) transition using subsampling and phylogenetic approaches and apply maximum-likelihood methods to fit models of extrinsic variables to assess what mediated these patterns. A combination of fluctuations in sea-level and episodic perturbations to the carbon and sulfur cycles was primarily responsible for both a marine and non-marine crocodyliform biodiversity decline through the J/K boundary, primarily documented in Europe. This was tracked by high extinction rates at the boundary and suppressed origination rates throughout the Early Cretaceous. The diversification of Eusuchia and Notosuchia likely emanated from the easing of ecological pressure resulting from the biodiversity decline, which also culminated in the extinction of the marine thalattosuchians in the late Early Cretaceous. Through application of rigorous techniques for estimating biodiversity, our results demonstrate that it is possible to tease apart the complex array of controls on diversification patterns in major archosaur clades.     

Phylogenetic structure of the extinction and biotic factors explaining differential survival of terrestrial notosuchians at the Cretaceous–Palaeogene crisis

Although the clade Crocodylomorpha is represented by few extant species (Crocodylia), it has a rich fossil record. Hundreds of species, adapted to terrestrial, semi-aquatic and marine environments, have existed over more than 200 million years. Numerous studies have attempted to characterize the factors driving the diversification and extinction events of Crocodylomorpha, resulting in ambiguous and even contradictory conclusions, which points to the need for phylogenetically and temporally smaller-scaled studies. Here, we focus on differential survival at the Cretaceous–Palaeogene (K–Pg) crisis of Notosuchia, a diverse clade of mostly terrestrial Crocodylomorpha that achieved great diversity during the Cretaceous. More precisely, we tested the effect of body size and palaeotemperatures on notosuchian survival probability during the K–Pg crisis as well as the effect of diet on the evolution of their body size. We find that Notosuchia showed an evolutionary trend towards larger body sizes through time, associated with a shift from an omnivorous to a carnivorous diet. This may explain why sebecids were the only notosuchians to survive the K–Pg crisis. We also corroborate the conclusions of previous studies that detected a Lagerstätten effect occurring in the Adamantina Formation (Upper Cretaceous, Brazil, Bauru Group). This work confirms the value of more finely-scaled macroevolutionary studies for understanding the history of a rich and complex group such as Crocodylomorpha.