Major evolutionary transitions, in which animals develop new body plans and adapt to dramatically new habitats and lifestyles, have punctuated the history of life. The origin of cetaceans from land-living mammals is among the most famous of these events. Much earlier, during the Mesozoic Era, many reptile groups also moved from land to water, but these transitions are more poorly understood. We use computed tomography to study changes in the inner ear vestibular system, involved in sensing balance and equilibrium, as one of these groups, extinct crocodile relatives called thalattosuchians, transitioned from terrestrial ancestors into pelagic (open ocean) swimmers. We find that the morphology of the vestibular system corresponds to habitat, with pelagic thalattosuchians exhibiting a more compact labyrinth with wider semicircular canal diameters and an enlarged vestibule, reminiscent of modified and miniaturized labyrinths of other marine reptiles and cetaceans. Pelagic thalattosuchians with modified inner ears were the culmination of an evolutionary trend with a long semiaquatic phase, and their pelagic vestibular systems appeared after the first changes to the postcranial skeleton that enhanced their ability to swim. This is strikingly different from cetaceans, which miniaturized their labyrinths soon after entering the water, without a prolonged semiaquatic stage. Thus, thalattosuchians and cetaceans became secondarily aquatic in different ways and at different paces, showing that there are different routes for the same type of transition.

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Shartegosuchids are a poorly known, early-branching group of Asian and North American crocodylomorphs. Shartegosuchids have been hypothesized to have eusuchian-type secondary palates, but a paucity of described material makes assessing this difficult. Our fieldwork in western Mongolia recovered cranial material of a new Shartegosuchus specimen from the Ulan Malgait Formation, which we CT-scanned and digitally reconstructed to investigate its palatal morphology. We then incorporated this new anatomical information into a revised phylogenetic dataset to assess its affinities. Our study confirms that Shartegosuchus has a posteriorly placed choana that is fully enclosed by the pterygoids, but differs from Eusuchia in possessing a secondary palatal fenestra and reduced palatine bones. Shartegosuchus, together with Adzhosuchus, Fruitachampsa, and Nominosuchus, forms the monophyletic group Shartegosuchidae. Shartegosuchidae is nested within a larger clade Shartegosuchoidea, and this clade is an earlierdiverging lineage than Eusuchia, showing that a eusuchian-type secondary palate evolved multiple times in crocodylomorphs, including very early in the group's evolutionary history. The co-occurrence of Nominosuchus in the Ulan Malgait Formation and the Shishigou Formation allows us to assign an early Oxfordian age to Shartegosuchus. The independent evolution of a eusuchian-type secondary palate in an oreinorostral group suggests that the link between platyrostry and a closed secondary palate has been overstated.

The Massospondylus Assemblage Zone is the youngest tetrapod biozone in the Karoo Basin (upper Stormberg Group, Karoo Supergroup) and records one of the oldest dinosaur dominated ecosystems in southern Gondwana. Recent qualitative and quantitative investigations into the biostratigraphy of the lower and upper Elliot formations (lEF, uEF) and Clarens Formation in the main Karoo Basin resulted in the first biostratigraphic review of this stratigraphic interval in nearly four decades, allowing us to introduce a new biostratigraphic scheme, the Massospondylus Assemblage Zone (MAZ). The MAZ expands upon the Massospondylus Range Zone by including the crocodylomorph Protosuchus haughtoni and the ornithischian Lesothosaurus diagnosticus as two co-occurring index taxa alongside the main index taxon, the sauropodomorph Massospondylus carinatus. With a maximum thickness of ~320 m in the southeastern portion of the basin, our new biozone is contained within the uEF and Clarens formations (upper Stormberg Group), however, based on vertebrate ichnofossils evidence, it may potentially extend into the sedimentary units of the lowermost Drakensberg Group. We do not propose any further subdivisions, and do not consider the Tritylodon Acme Zone (TAZ) as a temporal biostratigraphic marker within the MAZ. The MAZ is currently accepted to range in age between the Hettangian and Pliensbachian, however a faunal turnover, which observes an increase in the diversity of dinosaur clades, crocodylomorph, and mammaliaform taxa in the lower uEF, could reflect effects of the end-Triassic extinction event (ETE).