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Creators/Authors contains: "Bhullar, Bhart‐Anjan S."

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  1. Free, publicly-accessible full text available July 9, 2026
  2. Free, publicly-accessible full text available January 2, 2026
  3. Abstract The trigeminus nerve (cranial nerve V) is a large and significant conduit of sensory information from the face to the brain, with its three branches extending over the head to innervate a wide variety of integumentary sensory receptors, primarily tactile. The paths of the maxillary (V2) and mandibular (V3) divisions of the trigeminus frequently transit through dedicated canals within the bones of the upper and lower jaws, thus allowing this neuroanatomy to be captured in the fossil record and be available to interpretations of sensory ability in extinct taxa. Here, we use microCT and synchrotron scans from 38 extant and fossil species spanning a wide phylogenetic sample across tetrapods to investigate whether maxillary and mandibular canal morphology can be informative of sensory biology in the synapsid lineage. We found that in comparison to an amphibian and sauropsid outgroup, synapsids demonstrate a distinctive evolutionary pattern of change from canals that are highly ramified near the rostral tip of the jaws to canals with increasingly simplified morphology. This pattern is especially evident in the maxillary canal, which came to feature a shortened infraorbital canal terminating in a single large infraorbital foramen that serves as the outlet for branches of V2that then enter the soft tissues of the face. A comparison with modern analogues supports the hypothesis that this morphological change correlates to an evolutionary history of synapsid‐specific innovations in facial touch. We interpret the highly ramified transitional form found in early nonmammalian synapsids as indicative of enhanced tactile sensitivity of the rostrum via direct or proximal contact, similar to tactile specialists such as probing shorebirds and alligators that possess similar proliferative ramifications of the maxillary and mandibular canals. The transition toward a simplified derived form that emerged among Mid‐Triassic prozostrodont cynodonts and is retained among modern mammals is a unique configuration correlated with an equally unique and novel tactile sensory apparatus: mobile mystacial whiskers. Our survey of maxillary and mandibular canals across a phylogenetic and ecological variety of tetrapods highlights the morphological diversity of these structures, but also the need to establish robust form‐function relationships for future interpretations of osteological correlates for sensory biology. 
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    Free, publicly-accessible full text available November 24, 2025
  4. Abstract Our knowledge of vertebrate functional evolution depends on inferences about joint function in extinct taxa. Without rigorous criteria for evaluating joint articulation, however, such analyses risk misleading reconstructions of vertebrate animal motion. Here we propose an approach for synthesizing raycast-based measurements of 3-D articular overlap, symmetry, and congruence into a quantitative “articulation score” for any non-interpenetrating six-degree-of-freedom joint configuration. We apply our methodology to bicondylar hindlimb joints of two extant dinosaurs (guineafowl, emu) and, through comparison with in vivo kinematics, find that locomotor joint poses consistently have high articulation scores. We then exploit this relationship to constrain reconstruction of a pedal walking stride cycle for the extinct dinosaurDeinonychus antirrhopus, demonstrating the utility of our approach. As joint articulation is investigated in more living animals, the framework we establish here can be expanded to accommodate additional joints and clades, facilitating improved understanding of vertebrate animal motion and its evolution. 
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  5. Abstract Squamata is the most diverse clade of terrestrial vertebrates. Although the origin of pan-squamates lies in the Triassic, the oldest undisputed members of extant clades known from nearly complete, uncrushed material come from the Cretaceous. Here, we describe three-dimensionally preserved partial skulls of two new crown lizards from the Late Jurassic of North America. Both species are placed at the base of the skink, girdled, and night lizard clade Pan-Scincoidea, which consistently occupies a position deep inside the squamate crown in both morphological and molecular phylogenies. The new lizards show that several features uniting pan-scincoids with another major lizard clade, the pan-lacertoids, in trees using morphology were convergently acquired as predicted by molecular analyses. Further, the palate of one new lizard bears a handful of ancestral saurian characteristics lost in nearly all extant squamates, revealing an underappreciated degree of complex morphological evolution in the early squamate crown. We find strong evidence for close relationships between the two new species and Cretaceous taxa from Eurasia. Together, these results suggest that early crown squamates had a wide geographic distribution and experienced complicated morphological evolution even while the Rhynchocephalia, now solely represented by the tuatara, was the dominant clade of lepidosaurs. 
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  6. Abstract The rapid radiation and dispersal of crown reptiles following the end‐Permian mass extinction characterizes the earliest phase of the Mesozoic. Phylogenetically, this early radiation is difficult to interpret, with polytomies near the crown node, long ghost lineages, and enigmatic origins for crown group clades. Better understanding of poorly known taxa from this time can aid in our understanding of this radiation and Permo‐Triassic ecology. Here, we describe an Early Triassic specimen of the diapsidPalacrodonfrom the Fremouw Formation of Antarctica. WhilePalacrodonis known throughout the Triassic and exhibits a cosmopolitan geographic range, little is known of its evolutionary relationships. We recoverPalacrodonoutside of crown reptiles (Sauria) but more crownward thanYoungina capensisand other late Permian diapsids. Furthermore,Palacrodonpossesses anatomical features that add clarity to the evolution of the stapes within the reptilian lineage, as well as incipient adaptations for arboreality and herbivory during the earliest phases of the Permo–Triassic recovery. 
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  7. Ichthyornishas long been recognized as a pivotally important fossil taxon for understanding the latest stages of the dinosaur–bird transition, but little significant new postcranial material has been brought to light since initial descriptions of partial skeletons in the 19thCentury. Here, we present new information on the postcranial morphology ofIchthyornisfrom 40 previously undescribed specimens, providing the most complete morphological assessment of the postcranial skeleton ofIchthyornisto date. The new material includes four partially complete skeletons and numerous well-preserved isolated elements, enabling new anatomical observations such as muscle attachments previously undescribed for Mesozoic euornitheans. Among the elements that were previously unknown or poorly represented forIchthyornis, the new specimens include an almost-complete axial series, a hypocleideum-bearing furcula, radial carpal bones, fibulae, a complete tarsometatarsus bearing a rudimentary hypotarsus, and one of the first-known nearly complete three-dimensional sterna from a Mesozoic avialan. Several pedal phalanges are preserved, revealing a remarkably enlarged pes presumably related to foot-propelled swimming. Although diagnosable asIchthyornis, the new specimens exhibit a substantial degree of morphological variation, some of which may relate to ontogenetic changes. Phylogenetic analyses incorporating our new data and employing alternative morphological datasets recoverIchthyornisstemward of Hesperornithes andIaceornis, in line with some recent hypotheses regarding the topology of the crownward-most portion of the avian stem group, and we establish phylogenetically-defined clade names for relevant avialan subclades to help facilitate consistent discourse in future work. The new information provided by these specimens improves our understanding of morphological evolution among the crownward-most non-neornithine avialans immediately preceding the origin of crown group birds. 
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