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Upper Cretaceous coastal marine deposits are widespread in the Southern Urals with a number of marine vertebrates previously reported from this region. However, previous studies on the vertebrate faunas in this region often lack detailed taxonomic descriptions and illustrations, rendering comparisons to other faunal assemblages difficult. A new diverse vertebrate assemblage comprising cartilaginous and bony fishes, as well as marine reptiles, is described here from the Orenburg region near Akkermanovka (Southern Urals, Russia). Thirty five taxa are identified, including three holocephalans ( Elasmodus sp., Ischyodus yanschini , Chimaeroid indet.), two hybodontiform sharks ( Meristodonoides sp., cf. Polyacrodus sp.), 17 neoselachians ( Paraorthacodus cf. andersoni, Paraorthacodus sp., Synechodus sp., Cederstroemia nilsi, Acrolamna acuminata, Archaeolamna ex gr. kopingensis, Cretalamna sarcoportheta, Cretoxyrhina mantelli, Eostriatolamia segedini, E. venusta, Hispidaspis horridus, H . cf. gigas, Pseudocorax laevis, Pseudoscapanorhynchus compressidens, Scapanorhynchus rhaphiodon, Squalicorax kaupi, Ptychodus rugosus ), a holostean (Lepisosteidae indet.), nine teleosts ( Protosphyraena sp., Saurodontidae indet., cf. Pachyrhizodus sp., Pachyrhizodontidae indet., Enchodus petrosus, E. ferox, E . cf. gladiolus, E . spp., Alepisauroidei indet.), two plesiosaurs (Polycotylidae indet., Plesiosauria indet.), and one mosasaurid (Tylosaurinae indet.). Based on the faunal assemblage, a Santonian-?early Campanian age is proposed. Lamniform sharks are the best represented group in terms of taxic diversity and relative abundance, probably reflecting the peak in diversity this group experienced following the Cenomanian radiation in the Late Cretaceous. The faunal assemblage of Akkermanovka exhibits significant taxonomic overlaps with assemblages reported from Asia and North America, but not from Southern Hemisphere continents, indicating east-west dispersal of several marine taxa during the Late Cretaceous.

Mosasaurid squamates were the dominant amniote predators in marine ecosystems during most of the Late Cretaceous. Here, we use a suite of biomechanically rooted, functionally descriptive ratios in a framework adapted from population ecology to investigate how the morphofunctional disparity of mosasaurids evolved prior to the Cretaceous-Palaeogene (K/Pg) mass extinction. Our results suggest that taxonomic turnover in mosasaurid community composition from Campanian to Maastrichtian is reflected by a notable global increase in morphofunctional disparity, especially driving the North American record. Ecomorphospace occupation becomes polarized during the Late Maastrichtian, with morphofunctional disparity plateauing in the Southern Hemisphere and decreasing in the Northern Hemisphere. We show that these changes are not strongly associated with mosasaurid size, but rather with the functional capacities of their skulls. Our novel approach indicates that mosasaurid morphofunctional disparity was in decline in multiple provincial communities before the K/Pg mass extinction, highlighting region-specific patterns of disparity evolution and the importance of assessing vertebrate extinctions both globally and locally. Ecomorphological differentiation in mosasaurid communities, coupled with declines in other formerly abundant marine reptile groups, indicates widespread restructuring of higher trophic levels in marine food webs was well underway when the K/Pg mass extinction took place.

Mosasaurs were large, globally distributed aquatic lizards that lived during the Late Cretaceous. Despite numerous specimens of varying maturity, a detailed growth series has not been proposed for any mosasaur taxon. Two taxa- Tylosaurus proriger and T. kansasensis/nepaeolicus -have robust fossil records with specimens spanning a wide range of sizes and are thus ideal for studying mosasaur ontogeny. Tylosaurus is a genus of particularly large mosasaurs with long, edentulous anterior extensions of the premaxilla and dentary that lived in Europe and North America during the Late Cretaceous. An analysis of growth in Tylosaurus provides an opportunity to test hypotheses of the synonymy of T. kansasensis with T. nepaeolicus , sexual dimorphism, anagenesis, and heterochrony. Fifty-nine hypothetical growth characters were identified, including size-dependent, size-independent, and phylogenetic characters, and quantitative cladistic analysis was used to recover growth series for the two taxa. The results supported the synonymy of T. kansasensis with T. nepaeolicus and that T. kansasensis represent juveniles of T. nepaeolicus . A Spearman rank-order correlation test resulted in a significant correlation between two measures of size (total skull length and quadrate height) and maturity. Eleven growth changes were shared across both species, neither of the ontogram topologies showed evidence of skeletal sexual dimorphism, and a previous hypothesis of paedomorphy in T. proriger was not rejected. Finally, a novel hypothesis of anagenesis in Western Interior Seaway Tylosaurus species, driven by peramorphy, is proposed here., Competing Interests: The author declares that they have no competing interests., (© 2020 Zietlow.)

Squamates present a unique challenge to our understanding of dental evolution in amniotes because they are the only extant tooth-bearing group for which a ligamentous tooth attachment is considered to be absent. This has led to the assumption that mammals and crocodilians have convergently evolved a ligamentous tooth attachment, composed of root cementum, periodontal ligament, and alveolar bone, whereas squamates are thought to possess a single bone of attachment tissue that fuses teeth to the jaws. The identity and homology of tooth attachment tissues between squamates, crocodilians, and mammals have thus been a focal point of debate for decades. We provide a novel interpretation of the mineralized attachment tissues in two focal taxa in this debate, mosasaurids and snakes, and compare dental tissue histology with that of the extant crocodilian Caiman sclerops. We identify a periodontal ligament in these squamates that usually exists temporarily as a soft connective tissue anchoring each tooth to the alveolar bone. We also identify two instances where complete calcification of the periodontal ligament does not occur: in a durophagous mosasaur, and in the hinged teeth of fossil and modern snakes. We propose that the periodontal ligament rapidly calcifies in the majority of mosasaurids and snakes, ankylosing the tooth to the jaw. This gives the appearance of a single, bone-like tissue fusing the tooth to the jaw in ankylosed teeth, but is simply the end stage of dental tissue ontogeny in most snakes and mosasaurids., (© 2017 Anatomical Society.)

Living in water imposes severe constraints on the evolution of the vertebrate body. As a result of these constraints, numerous extant and extinct aquatic vertebrate groups evolved convergent osteological and soft-tissue adaptations. However, one important suite of adaptations is still poorly understood: dermal cover morphologies and how they influence surface fluid dynamics. This is especially true for fossil aquatic vertebrates where the soft tissue of the dermis is rarely preserved. Recent studies have suggested that the keeled scales of mosasaurids (pelagic lizards that lived during the Late Cretaceous) aided in surface frictional drag reduction in a manner analogous to the riblets on shark placoid scales. However, here we demonstrate that mosasaurid scales were over an order of magnitude too large to have this effect. More likely they increased the frictional drag of the body and may have played a role in controlling flow separation by acting as surface roughness that turbulated the boundary layer. Such a role could have reduced pressure drag and enhanced manoeuvrability. We caution those studying fossil aquatic vertebrates from positing the presence of surface drag reducing morphologies, because as we show herein, to be effective such features need to have a spacing of approximately 0.1 mm or less.