A rich fossil record chronicles the distant origins of mammals, but the evolution of defining soft tissue characters of extant mammals, such as mammary glands and hairs is difficult to interpret because soft tissue does not readily fossilize. As many soft tissue features are derived from dermic structures, their evolution is linked to that of the nervous syutem, and palaeoneurology offers opportunities to find bony correlates of these soft tissue features. Here, a CT scan study of 29 fossil skulls shows that non-mammaliaform Prozostrodontia display a retracted, fully ossified, and non-ramified infraorbital canal for the infraorbital nerve, unlike more basal therapsids. The presence of a true infraorbital canal in Prozostrodontia suggests that a motile rhinarium and maxillary vibrissae were present. Also the complete ossification of the parietal fontanelle (resulting in the loss of the parietal foramen) and the development of the cerebellum in Probainognathia may be pleiotropically linked to the appearance of mammary glands and having body hair coverage since these traits are all controlled by the same homeogene, Msx2, in mice. These suggest that defining soft tissue characters of mammals were already present in their forerunners some 240 to 246 mya.

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Dinocephalian therapsids are renowned for their massive, pachyostotic and ornamented skulls adapted for head-to-head fighting during intraspecific combat. Synchrotron scanning of the tapinocephalid Moschops capensis reveals, for the first time, numerous anatomical adaptations of the central nervous system related to this combative behaviour. Many neural structures (such as the brain, inner ear and ophthalmic branch of the trigeminal nerve) were completely enclosed and protected by bones, which is unusual for non-mammaliaform therapsids. The nearly complete ossification of the braincase enables precise determination of the brain cavity volume and encephalization quotient, which appears greater than expected for such a large and early herbivore. The practice of head butting is often associated with complex social behaviours and gregariousness in extant species, which are known to influence brain size evolution. Additionally, the plane of the lateral (horizontal) semicircular canal of the bony labyrinth is oriented nearly vertically if the skull is held horizontally, which suggests that the natural position of the head was inclined about 60–65°to the horizontal. This is consistent with the fighting position inferred from osteology, as well as ground-level browsing. Finally, the unusually large parietal tube may have been filled with thick conjunctive tissue to protect the delicate pineal eye from injury sustained during head butting.

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We provide the first detailed description of the inner ear of the oldest artiodactyl, Diacodexis, based on a three-dimensional reconstruction extracted from computed tomography imagery of a skull of Diacodexis ilicis of earliest Wasatchian age (ca. 55 Ma). This description provides new anatomical data for the earliest artiodactyls, and reveals that the bony labyrinth of Diacodexis differs greatly from that of modern artiodactyls described so far. The bony labyrinth of Diacodexis presents a weakly coiled cochlea (720°), a secondary common crus, a dorsal extension of the anterior semicircular canal more pronounced than that of the posterior one, and a small angle between the basal turn of the bony cochlear canal and the lateral semicircular canal. This suite of characters also occurs in basal eutherian mammals. Diacodexis strongly resembles small living tragulid ruminants in its overall body shape and hindlimb proportions. Comparison of the bony labyrinth of Diacodexis to that of the tragulid Moschiola meminna (Indian mouse deer) reveals great morphological difference in cochlear shape and semicircular canal disposition. The shape of the cochlea suggests that Diacodexis was a high-frequency hearing specialist, with a high low-frequency hearing limit (543 Hz at 60 dB). By comparison, the estimated low-frequency limit of Moschiola meminna is much lower (186.0 Hz at 60 dB). We also assess the locomotor agility of Diacodexis based on measurements of the semicircular canals. Locomotor agility estimates for Diacodexis range between 3.62 and 3.93, and suggest a degree of agility compatible with a nimble, fast running to jumping animal. These results are congruent with the postcranial functional analysis for this extinct taxon.

Anomodontia was the most successful herbivorous clade of the mammalian stem lineage (non-mammalian synapsids) during the late Permian and Early Triassic. Among anomodonts, Dicynodontia stands apart because of the presence of an osseous beak that shows evidence of the insertion of a cornified sheath, the ramphotheca. In this study, fourteen anomodont specimens were microCT-scanned and their trigeminal canals reconstructed digitally to understand the origin and evolution of trigeminal nerve innervation of the ramphotheca. We show that the pattern of innervation of the anomodont "beak" is more similar to that in chelonians (the nasopalatine branch is enlarged and innervates the premaxillary part of the ramphotheca) than in birds (where the nasopalatine and maxillary branches play minor roles). The nasopalatine branch is noticeably enlarged in the beak-less basal anomodont Patranomodon, suggesting that this could be an anomodont or chainosaur synapomorphy. Our analyses suggest that the presence or absence of tusks and postcanine teeth are often accompanied by corresponding variations of the rami innervating the caniniform process and the alveolar region, respectively. The degree of ossification of the canal for the nasal ramus of the ophthalmic branch also appears to correlate with the presence of a nasal boss. The nasopalatine canal is absent from the premaxilla in the Bidentalia as they uniquely show a large plexus formed by the internal nasal branch of the maxillary canal instead. The elongated shape of this plexus in Lystrosaurus supports the hypothesis that the rostrum evolved as an elongation of the subnarial region of the snout. Finally, the atrophied and variable aspect of the trigeminal canals in Myosaurus supports the hypothesis that this genus had a reduced upper ramphotheca.

The origin and evolution of the mammalian brain has long been the focus of scientific enquiry. Conversely, little research has focused on the palaeoneurology of the stem group of Mammaliaformes, the Permian and Triassic non-mammaliaform Therapsida (NMT). This is because the majority of the NMT have a non-ossified braincase, making the study of their endocranial cast (sometimes called the “fossil brain”) problematic. Thus, descriptions of the morphology and size of NMT endocranial casts have been based largely on approximations rather than reliable determination. Accordingly, here we use micro-CT scans of the skulls of 1 Dinocephalia and 3 Biarmosuchia, which are NMT with a fully ossified braincase and thus a complete endocast. For the first time, our work enables the accurate determination of endocranial shape and size in NMT. This study suggests that NMT brain size falls in the upper range of the reptilian and amphibian variation. Brain size in the dicynodont Kawingasaurus is equivalent to that of early Mammaliaformes, whereas the Dinocephalia show evidence of a secondary reduction of brain size. In addition, unlike other NMT in which the endocast has a tubular shape and its parts are arranged in a linear manner, the biarmosuchian endocast is strongly flexed at the level of the midbrain, creating a near right angle between the fore- and hindbrain. These data highlight an unexpected diversity of endocranial size and morphology in NMT, features that are usually considered conservative in this group.

Choerosaurus dejageri, a non-mammalian eutheriodont therapsid from the South African late Permian (~259 Ma), has conspicuous hemispheric cranial bosses on the maxilla and the mandible. These bosses, the earliest of this nature in a eutheriodont, potentially make C. dejageri a key species for understanding the evolutionary origins of sexually selective behaviours (intraspecific competition, ritualized sexual and intimidation displays) associated with cranial outgrowths at the root of the clade that eventually led to extant mammals. Comparison with the tapinocephalid dinocephalian Moschops capensis, a therapsid in which head butting is strongly supported, shows that the delicate structure of the cranial bosses and the gracile structure of the skull of Choerosaurus would be more suitable for display and low energy combat than vigorous head butting. Thus, despite the fact that Choerosaurus is represented by only one skull (which makes it impossible to address the question of sexual dimorphism), its cranial bosses are better interpreted as structures involved in intraspecific selection, i.e. low-energy fighting or display. Display structures, such as enlarged canines and cranial bosses, are widespread among basal therapsid clades and are also present in the putative basal therapsid Tetraceratops insignis. This suggests that sexual selection may have played a more important role in the distant origin and evolution of mammals earlier than previously thought. Sexual selection may explain the subsequent independent evolution of cranial outgrowths and pachyostosis in different therapsid lineages (Biarmosuchia, Dinocephalia, Gorgonopsia and Dicynodontia).