Mathieu Gabriel Faure-Brac

The inner microstructure of aquatic amniote ribs can reveal adaptations to physiology and locomotion, but are relatively poorly known for ichthyosaurs, especially ophthalmosaurid taxa whose body shape converges to modern odontocetes. In this study, we describe the microstructure of ribs and gastralia from two Late Jurassic ichthyosaurs († Palvennia hoybergeti and † Keilhauia sp.) and compare them to extant odontocetes ( Delphinapterus leucas and Phocoena phocoena ). Ribs were sampled at proximal, midshaft, and distal locations to assess the potential for reliable growth marks, and quantitative microanatomical values were collected to enable a comparison to other ichthyosaurs. The ichthyosaur ribs preserve alternating vascular and avascular bands resulting from localized growth variation, but no unambiguous growth marks, whereas gastralia seem more promising for skeletochronology. The toothed whale ribs preserve no growth marks. Ichthyosaur skeletal elements often have a spongy internal structure without a thick cortex, but the ophthalmosaurid ribs in this study have relatively compact bone throughout, compared to the cetaceans, something that is possibly an adaptation to counter ballast the buoyancy of the lungs.

Les effets de la thermophysiologie sur l'évolution des Pseudosuchiens (Archosauria): Il est aujourd’hui communément admis que les archosaures sont ancestralement endothermes, soit qu’ils présentent des mécanismes de thermogenèse non frissonnante. Parmi les archosaures, les crocodiliens actuels sont pourtant des ectothermes, mais présentent des caractéristiques anatomiques et moléculaires traditionnellement associées à l’endothermie : un cœur à quatre cavité ; plusieurs acquisitions liées à une respiration efficace, comme un muscle diaphragmatique et un flux d’air unidirectionnel ; la formation de tissus osseux liés à des taux de croissance élevés ; un taux d’évolution mitochondrial important. Des études sur des pseudosuchiens éteints ont conclus que la plupart des espèces du Trias étaient endothermes. La perte de l’endothermie au sein du clade doit cependant toujours être définie. Durant cette thèse, j’ai utilisé la géochimie des isotopes stables et l’ostéohistologie quantitative pour inférer la température corporelle et le taux métabolique au repos, deux proxies de l’endothermie, de plusieurs métasuchiens. J’ai conclus que le clade Metasuchia était ancestralement ectotherme et, à l’aide d’une reconstruction ancestrale des caractères, que l’endothermie a probablement été perdue au nœud Crocodylomorpha. J’en déduit l’hypothèse suivante : l’endothermie des pseudosuchiens était probablement à un stade plus primitif que celle des dinosaures. Contrairement à ces derniers, les pseudosuchiens endothermes n’ont pu survivre à l’extinction de masse de la fin du Trias, ne laissant pour seul survivant que les crocodylomorphes ectothermes.

Pseudosuchia, one of the two main clades of Archosauria, is today only represented by some 20 extant species, the crocodilians, representing only a fraction of its extinct diversity. Extant crocodilians are ectotherms but present morphological and anatomical features usually associated with endothermy. In 2004, it was proposed that pseudosuchians were ancestrally endothermic and the features observed in extant crocodilians are the remains of this lost legacy. This contribution has two parts: the first part covers 20 years of studies on this subject, first exploring the evidence for a loss of endothermy in extant crocodilians, before covering the variety of proxies used to infer the thermophymetabolic regime of extinct pseudosuchians. In the second part, the quantitative results of these previous studies are integrated into a comprehensive ancestral state reconstruction to discuss a potential scenario for the evolution of thermometabolism. Pseudosuchian endothermy would then have been lost close to the node Crocodylomorpha. The end‐Triassic mass extinction is proposed to have played the role of a filter, leading to the extinction of endothermic pseudosuchians and the survival of ectothermic ones. This difference in survival in Pseudosuchia is compared to those of dinosaurs, and difference in their metabolism is also considered. Pseudosuchian endothermy might have been of a different level than the dinosaurian one and more studies are expected to clarify this question.

Paleophysiology is an emergent discipline. Organismic (integrative) approaches seem more appropriate than studies focusing on the variation of specific features because traits are tightly related in actual organisms. Here, we used such an organismic approach (including lifestyle, thermometabolism, and hunting behavior) to understand the paleobiology of the lower Jurassic (Toarcian) thalattosuchian metriorhynchoid Pelagosaurus typus. First, we show that the lifestyle (aquatic, amphibious, terrestrial) has an effect on the femoral compactness profiles in amniotes. The profile of Pelagosaurus indicates that it was amphibious, with a foraging activity in shallow marine environments (as suggested by the presence of salt glands) and thermoregulatory basking behavior in land (as suggested by the presence of osteoderms with highly developed ornamentation). As for the thermometabolism, we show that the mass‐independent resting metabolic rate of Pelagosaurus is relatively high compared to the sample of extant ectothermic amniotes, but analysis of vascular canal diameter and inferences of red blood cell size refute the hypothesis suggesting incipient endothermy. Finally, the foraging behavior was inferred using two proxies. Pelagosaurus had a mass‐independent maximum metabolic rate and an aerobic scope higher than those measured in the almost motionless Iguana iguana, similar to those measured in the sit‐and‐wait predator Crocodylus porosus but lower than those quantified in the active hunter Varanus gouldii. These results suggest that Pelagosaurus may have had a hunting behavior involving a slow sustained swimming or a patient waiting in shallow waters, and may have caught preys like gharials, using fast sideways sweeping motions of the head.

Notosuchia is a group of mostly terrestrial crocodyliforms. The presence of a prominent crest overhanging the acetabulum, slender straight-shafted long bones with muscular insertions close to the joints, and a stable knee joint suggests that they had an erect posture. This stance has been proposed to be linked to endothermy, because it is present in mammals and birds and contributes to the efficiency of their respiratory systems. However, a bone paleohistological study unexpectedly suggested that Notosuchia were ectothermic organisms. The thermophysiological status of Notosuchia deserves further analysis, because the methodology of the previous study can be improved. First, it was based on a relationship between red blood cell size and bone vascular canal diameter tested using 14 extant tetrapod species. Here we present evidence for this relationship using a more comprehensive sample of extant tetrapods (31 species). Moreover, contrary to previous results, bone cross-sectional area appears to be a significant explanatory variable (in addition to vascular canal diameter). Second, red blood cell size estimations were performed using phylogenetic eigenvector maps, and this method excludes a fraction of the phylogenetic information. This is because it generates a high number of eigenvectors requiring a selection procedure to compile a subset of them to avoid model overfitting. Here we inferred the thermophysiology of Notosuchia using phylogenetic logistic regressions, a method that overcomes this problem by including all of the phylogenetic information and a sample of 46 tetrapods. These analyses suggest that Araripesuchus wegeneri , Armadillosuchus arrudai , Baurusuchus sp., Iberosuchus macrodon , and Stratiotosuchus maxhechti were ectothermic organisms.

Sang chaud ou sang froid ? Ce qu’en disent les os was published in Paléontologie d'aujourd'hui on page 190.

The evolution of thermometabolism in pseudosuchians (Late Triassic to the present) remains a partly unsolved issue: extant taxa (crocodilians) are ectothermic, but the clade was inferred ancestrally endothermic. Here we inferred the thermometabolic regime of two neosuchian groups, Goniopholididae (Early Jurassic to Late Cretaceous) and Dyrosauridae (middle Cretaceous to late Eocene), close relatives of extant crocodilians, in order to elucidate the evolutionary pattern across Metasuchia (Early Jurassic to the present), a clade comprising Neosuchia (Early Jurassic to the present) and Notosuchia (Middle Jurassic until the late Miocene). We propose a new integrative approach combining geochemical analyses to infer body temperature from the stable oxygen isotope composition of tooth phosphate and paleohistology and phylogenetic comparative methods to infer resting metabolic rates and red blood cell dimensions. † Dyrosaurus and † Goniopholis share with extant crocodilians similar lifestyles, body forms, bone tissue organization, body temperatures, metabolic rates, and red blood cell dimensions. Consistently, we infer ectothermy for † Dyrosaurus and † Goniopholis with the parsimonious implication of neosuchians and metasuchians being primitively ectothermic.

In 1995, Pleijel raised an issue on the coding of complex (multistate) characters: to find a unique coding approach satisfying both informational and semantical criteria. Following Pleijel’s work, this study aims to propose a new answer to this problematic. We proposed here to use hierarchical characters instead of classical partitional ones as hierarchy allows to deal effectively with the different uncertainties met by phylogeneticists since the beginning of phylogenetic data matrices: missing data, inapplicable data and polymorphism. We translated all previous proposed approaches into hierarchical ones and add three new approaches, only coded using hierarchy. Using phylogenetic and show with different metric how one of these new approaches displaces the other ones. The results from this study then supports the idea than phylogenetic characters should be coded using only hierarchies, as they offer a better management of uncertainties and propose new approaches more informative.

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.