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Lagomorpha is a clade of herbivorous mammals nested within Euarchontoglires, one of the major placental groups represented today. It comprises two extant families with markedly different body plans: the long-eared and long-limbed Leporidae (hares and rabbits) and the short-eared and short-limbed Ochotonidae (pikas). These two lagomorph lineages diverged probably during the latest Eocene/early Oligocene, but it is unclear whether the last common ancestor of crown lagomorphs was more leporid- or more ochotonid-like in morphology. Palaeolagus, an early lagomorph dominant in western North America from the late Eocene to Oligocene is of particular importance for addressing this controversy. Here, we present new and comprehensive data on the cranial anatomy of Palaeolagus haydeni, the type species for the genus, based on micro-computed tomography (μCT). Our μCT data allow us to confirm, revise and score for the very first time the states of several leporid-like and ochotonid-like characters in the skull of Palaeolagus. This mixed cranial architecture differentiates Palaeolagus from the crown groups of Lagomorpha and supports its phylogenetic status as a stem taxon.

Synapsids are unique in having developed multirooted teeth and complex occlusions. These innovations evolved in at least two lineages of mammaliamorphs (Tritylodontidae and Mammaliaformes). Triassic fossils demonstrate that close to the origins of mammals, mammaliaform precursors were "experimenting" with tooth structure and function, resulting in novel patterns of occlusion. One of the most surprising examples of such adaptations is present in the haramiyidan Glade, which differed from contemporary mammaliaforms in having two rows of cusps on molariform crowns adapted to omnivorous/herbivorous feeding. However, the origin of the multicusped tooth pattern present in haramiyidans has remained enigmatic. Here we describe the earliest-known mandibular fossil of a mammaliaform with double molariform roots and a crown with two rows of cusps from the Late Triassic of Greenland. The crown morphology is intermediate between that of morganucodontans and haramiyidans and suggests the derivation of the multicusped molariforms of haramiyidans from the triconodont molar pattern seen in morganucodontids. Although it is remarkably well documented in the fossil record, the significance of tooth root division in mammaliaforms remains enigmatic. The results of our biomechanical analyses (finite element analysis [FEA]) indicate that teeth with two roots can better withstand stronger mechanical stresses like those resulting from tooth occlusion, than teeth with a single root.

The Middle Jurassic was a key interval of mammalian evolutionary history that witnessed the diversification of the therian stem group. Great Britain has yielded a significant record of mammalian fossils from this interval, represented by numerous isolated jaws and teeth from the Bathonian of Oxfordshire and the Isle of Skye. This record captures a key period in early cladotherian evolution, with amphitheriids, peramurans and ‘stem zatherians’ displaying intermediate talonid morphologies that document the evolutionary assembly of tribosphenic molars. We present a mandible with near-complete dentition from the late Bathonian (c. 167.4–166.5 Ma) Kilmaluag Formation, near Elgol, Skye, representing the amphitheriid Palaeoxonodon ooliticus, previously known only from isolated teeth. The specimen sheds new light on the taxonomic diversity of British Middle Jurassic stem therians, as the morphological variation within the preserved tooth row encompasses that previously ascribed to three distinct species within two genera: Palaeoxonodon ooliticus, P. freemani and Kennetheridium leesi. Thus, both P. freemani and K. leesi are subjective junior synonyms of P. ooliticus. The dental formula of P. ooliticus (i4:c1:p5:m5) is intermediate between the primitively larger postcanine count (p5:m6–7) of Amphitherium and the reduced number in peramurans and tribosphenidans (p5:m3). Phylogenetic analyses of P. ooliticus generally confirm a close affinity with Amphitherium, but highlight the lack of strong empirical support for hypothesized patterns of divergences among early cladotherians.