The dicynodonts are an emblematic group of herbivorous therapsids that survived the Permo-Triassic (P-Tr) crisis. Laotian dicynodonts from stratigraphically constrained beds, recently dated using the U-Pb zircon method, yield new insights into terrestrial faunas of Southeast Asia during the latest Permian and earliest Triassic. Summarily described, they were originally attributed to the genus Dicynodon. We provide a new phylogenetic analysis for Laotian dicynodonts, based on three well-preserved skulls, indicating that they belong to two new taxa: Counillonia superoculis, gen. et sp. nov., and Repelinosaurus robustus, gen. et sp. nov. Our phylogenetic analysis of Dicynodontia indicates that (1) Counillonia is closely related to some ‘Dicynodon’-grade taxa and (2) Repelinosaurus is a kannemeyeriiform. The phylogenetic affinities of these new Laotian dicynodonts allow discussion of the survivorship of multiple lineages (Kannemeyeriiformes and ‘Dicynodon’-grade dicynodontoids) across the P-Tr crisis. The Laotian dicynodonts also shed new light on the paleobiogeography of Southeast Asia from the late Paleozoic to the early Mesozoic, particularly the timing of collisions between the Indochina, the South China, and the North China blocks. The presence of dicynodonts in Laos, most likely in the Early Triassic, thus implies that the connection between the Indochina Block and the South China Block occurred no later than the latest Permian or earliest Triassic (i.e., when the dicynodonts provide direct evidence for a connection).

This paper is aimed at constraining the phylogenetic frame of the acquisition of endothermy by Archosauromorpha. We analyzed the bone histology of Azendohsaurus laaroussii . Stylopodial and zeugopodial bones show three tissue types: (1) avascular lamellar zonal bone formed at low growth rates; (2) a scaffold of parallel-fibered bone containing either small primary osteons or simple vascular canals; and (3) fibrolamellar bone formed at high growth rates. We used quantitative histology to infer the thermometabolic regime of this taxon. We define endothermy as the presence of any mechanism of nonshivering thermogenesis that increases both body temperature and resting metabolic rate. Thus, estimating the resting metabolic rate of an extinct organism may be a good proxy to infer its thermometabolic regime (endothermy vs. ectothermy). High resting metabolic rates have been shown to be primitive for the clade Prolacerta –Archosauriformes. Therefore, we inferred the resting metabolic rates of A. laaroussii , a sister group of this clade, and of 14 extinct related taxa, using phylogenetic eigenvector maps. All the inferences obtained are included in the range of variation of resting metabolic rates measured in mammals and birds, so we can reasonably assume that all these taxa (including Azendohsaurus ) were endotherms. A parsimony optimization of the presence of endothermy on a phylogenetic tree of tetrapods shows that this derived character state was acquired by the last common ancestor of the clade Azendohsaurus –Archosauriformes and that there is a reversion in Crocodylia.

Mosasaurid teeth are abundant in the fossil record and often diagnostic to low taxonomic levels, allowing to document the taxonomic diversity and ecological disparity through time and with fewer biases than in other marine reptiles. The upper Maastrichtian Phosphates of Morocco, with at least fifteen coeval species representing a wide range of sizes and morphologies, undoubtedly represent the richest outcrop in the world for this clade of iconic Mesozoic squamates and one of the richest known marine tetrapod assemblages. Until now, the methods used to link tooth morphology to diets in marine amniotes were mainly qualitative in nature. Here, using the dental morphology of mosasaurids from Morocco, we combine two complementary approaches—a thorough comparative anatomical description and 2D/3D geometric morphometry—to quantitatively categorize the main functions of these teeth during feeding processes and infer diet preferences and niche-partitioning of these apex predators. Our results from combining these two approaches show the following: (1) Mosasaurids from the upper Maastrichtian Phosphates of Morocco occupy the majority of dental guilds ever colonized by Mesozoic marine reptiles. (2) As seen elsewhere in the Maastrichtian, mosasaurines dominate the regional mosasaurid assemblage, exhibiting the greatest taxonomic diversity (two-thirds of the species) and the largest range of morphologies, body sizes (2 m to more than 10 m) and ecological disparities (participating in nearly all predatory ecological guilds); strikingly, mosasaurines did not developed flesh piercers and, conversely, are the only ones to include durophagous species. (3) Halisaurines, though known by species of very different sizes (small versus large) and cranial morphologies (gracile versus robust), maintain a single tooth shape (piercer). (4) Plioplatecarpines were medium-size cutters and piercers, known by very morphologically diverging species. (5) Tylosaurines currently remain scarce, represented by a very large generalist species; they were largely replaced by mosasaurines as apex predators over the course of the Maastrichtian, as observed elsewhere. Also, when comparing tooth shapes with body sizes, the largest taxa (>8 m long) occupied a restricted area of tooth shapes (generalist, durophagous), whereas small and medium-sized species (<8 m long) range across all of them (generalists, durophagous, cutters, piercers). In other words, and probably related to the specificities and advantages of biomechanical resistance, apex predators are never dedicated piercers, micro-predators are conversely never generalists, and meso-predators show the widest range of dental adaptations. These diversities and disparities strongly suggest that Tethyan mosasaurids evolved strong niche-partitioning in the shallow marine environment of the upper Maastrichtian Phosphates of Morocco. Such a high diversity sensu lato just prior to the K/Pg biological crisis suggests that their extinction was rather sudden, though the exact causes of their extinction remain unknown. Finally, Gavialimimus Strong et al., 2020 is systematically reassigned to Gavialimimus ptychodon (Arambourg, 1952), and an emended diagnosis (for teeth and dentition) is proposed for this species.

Mosasaurids, a clade of specialized marine squamates, saw a major adaptive radiation in the Late Cretaceous, evolving a wide range of body sizes, shapes, and specialized tooth morphologies. The most diverse known mosasaurid faunas come from the late Maastrichtian phosphates of Morocco. Here, we report an unusual new mosasaurid, Stelladens mysteriosus, based on a partial jaw and associated tooth crowns from lower Couche III phosphatic deposits at Sidi Chennane, Oulad Abdoun Basin, Morocco. Stelladens is characterized by short, triangular tooth crowns with a series of strong, elaborate, and serrated ridges on the lingual surface of the tooth, functioning as accessory carinae. Morphology of the teeth and associated jaw fragment suggest affinities with Mosasaurinae. No close analogues to the unique tooth morphology of Stelladens are known, either extant or extinct. It may have had an unusual and highly specialized diet, a specialized prey-capture strategy, or both. The diversity of mosasaurid teeth is much higher than that of plesiosaurs, ichthyosaurs, or extant marine mammals, and likely reflects both the ecological diversity of mosasaurids and complex developmental mechanisms responsible for tooth formation in mosasaurines. Mosasaurid diversity continued to increase up to the Cretaceous–Paleogene boundary.