Endocranial anatomy and phylogenetic position of the crocodylian Eosuchus lerichei from the late Paleocene of northwestern Europe and potential adaptations for transoceanic dispersal in gavialoids.

Eosuchus lerichei is a gavialoid crocodylian from late Paleocene marine deposits of northwestern Europe, known from a skull and lower jaws, as well as postcrania. Its sister taxon relationship with the approximately contemporaneous species Eosuchus minor from the east coast of the USA has been explained through transoceanic dispersal, indicating a capability for salt excretion that is absent in extant gavialoids. However, there is currently no anatomical evidence to support marine adaptation in extinct gavialoids. Furthermore, the placement of Eosuchus within Gavialoidea is labile, with some analyses supporting affinities with the Late Cretaceous to early Paleogene "thoracosaurs." Here we present novel data on the internal and external anatomy of the skull of E. lerichei that enables a revised diagnosis, with 6 autapormorphies identified for the genus and 10 features that enable differentiation of the species from Eosuchus minor. Our phylogenetic analyses recover Eosuchus as an early diverging gavialid gavialoid that is not part of the "thoracosaur" group. In addition to thickened semi-circular canal walls of the endosseous labyrinth and paratympanic sinus reduction, we identify potential osteological correlates for salt glands in the internal surface of the prefrontal and lacrimal bones of E. lerichei. These salt glands potentially provide anatomical evidence for the capability of transoceanic dispersal within Eosuchus, and we also identify them in the Late Cretaceous "thoracosaur" Portugalosuchus. Given that the earliest diverging and stratigraphically oldest gavialoids either have evidence for a nasal salt gland and/or have been recovered from marine deposits, this suggests the capacity for salt excretion might be ancestral for Gavialoidea. Mapping osteological and geological evidence for marine adaptation onto a phylogeny indicates that there was probably more than one independent loss/reduction in the capacity for salt excretion in gavialoids.

Neuroanatomy of the crocodylian Tomistoma dowsoni from the Miocene of North Africa provides insights into the evolutionary history of gavialoids.

The interrelationships of the extant crocodylians Gavialis gangeticus and Tomistoma schlegelii have been historically disputed. Whereas molecular analyses indicate a sister taxon relationship between these two gavialoid species, morphological datasets typically place Gavialis as the outgroup to all other extant crocodylians. Recent morphological-based phylogenetic analyses have begun to resolve this discrepancy, recovering Gavialis as the closest living relative of Tomistoma; however, several stratigraphically early fossil taxa are recovered as closer to Gavialis than Tomistoma, resulting in anomalously early divergence timings. As such, additional morphological data might be required to resolve these remaining discrepancies. 'Tomistoma' dowsoni is an extinct species of gavialoid from the Miocene of North Africa. Utilising CT scans of a near-complete, referred skull, we reconstruct the neuroanatomy and neurosensory apparatus of 'Tomistoma' dowsoni. Based on qualitative and quantitative morphometric comparisons with other crocodyliforms, the neuroanatomy of 'Tomistoma' dowsoni is characterised by an intermediate morphology between the two extant gavialoids, more closely resembling Gavialis. This mirrors the results of recent studies based on the external anatomy of these three species and other fossil gavialoids. Several neuroanatomical features of these species appear to reflect ecological and/or phylogenetic signals. For example, the 'simple' morphology of their neurosensory apparatus is broadly similar to that of other long and narrow-snouted (longirostrine), aquatic crocodyliforms. A dorsoventrally short, anteroposteriorly long endosseous labyrinth is also associated with longirostry. These features indicate that snout and skull morphology, which are themselves partly constrained by ecology, exert an influence on neuroanatomical morphology, as has also been recognised in birds and turtles. Conversely, the presence of a pterygoid bulla in Gavialis and several extinct gavialoids, and its absence in Tomistoma schlegelii, could be interpreted as a phylogenetic signal of crocodylians more closely related to Gavialis than to Tomistoma. Evaluation of additional fossil gavialoids will be needed to further test whether these and other neuroanatomical features primarily reflect a phylogenetic or ecological signal. By incorporating such previously inaccessible information of extinct and extant gavialoids into phylogenetic and macroecological studies, we can potentially further constrain the clade's interrelationships, as well as evaluate the timing and ecological association of the evolution of these neuroanatomical features. Finally, our study supports recent phylogenetic analyses that place 'Tomistoma' dowsoni as being phylogenetically closer to Gavialis gangeticus than to Tomistoma schlegelii, indicating the necessity of a taxonomic revision of this fossil species.