Mosasaurids Bare the Teeth: An Extraordinary Ecological Disparity in the Phosphates of Morocco Just Prior to the K/Pg Crisis

Abstract

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.