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Abstract The stem lineage of Archosauria is populated by a diverse fossil record that remains notably understudied relative to the crown clade. Prominent among these specimens is a beautifully preserved skull of the early mid-Triassic rhynchosaur Mesosuchus browni [Iziko South African Museum (SAM) 6536], whose phylogenetic position has considerable influence on patterns of pan-archosaurian cranial evolution. We used high-resolution, micro-computed tomography to re-examine the anatomy of this specimen, building on previous studies that were either limited to external observations or restricted to the braincase. A digital segmentation of the cranial elements and primary neurovascular canals of SAM-PK-6536 allows for expanded character scoring and constitutes a foundation for future comparative insights. Our data support the phylogenetically oldest instance of a pneumatized maxilla in a pan-archosaur, bringing the record of antorbital pneumatization into closer alignment with that of the neurocranium. The nasal cavity and primary palate of Mesosuchus includes a complex septomaxilla, a novel element anterior to the vomer, and is likely to have supported a well-developed vomeronasal system. The evolution of this system is discussed in terms of both phylogenetic pattern and how the skeletal architecture of Mesosuchus and other fossils could inform the signalling dynamics that pattern the vomeronasal system during development. 

Abstract Resolving the phylogenetic relationships of early amniotes, in particular stem reptiles, remains a difficult problem. Three‐dimensional morphological analysis of well‐preserved stem‐reptile specimens can reveal important anatomical data and clarify regions of phylogeny. Here, we present the first thorough description of the unusual early Permian stem reptileBolosaurus major, including the first comprehensive description of a bolosaurid braincase. We describe previously obscured details of the palate, allowing for insight into bolosaurid feeding mechanics. Aspects of the rostrum, palate, mandible, and neurocranium suggest thatB. majorhad a particularly strong bite. We additionally foundB. majorhas a surprisingly slender stapes, similar to that of the middle Permian stem reptileMacroleter poezicus, which may suggest enhanced hearing abilities compared to other Paleozoic amniotes (e.g., captorhinids). We incorporated our new anatomical information into a large phylogenetic matrix (150 OTUs, 590 characters) to explore the relationship of Bolosauridae among stem reptiles. Our analyses generally recovered a paraphyletic “Parareptilia,” and found Bolosauridae to diverge after Captorhinidae + Araeoscelidia. We also includedB. majorwithin a smaller matrix (10 OTUs, 27 characters) designed to explore the interrelationships of Bolosauridae and found all species ofBolosaurusto be monophyletic. While reptile relationships still require further investigation, our phylogeny suggests repeated evolution of impedance‐matching ears in Paleozoic stem reptiles.