The semicircular canals (SCs), part of the vestibular apparatus of the inner ear, are directly involved in the detection of angular motion of the head for maintaining balance, and exhibit adaptive patterns for locomotor behaviour. Consequently, they are generally believed to show low levels of intraspecific morphological variation, but few studies have investigated this assumption. On the basis of high-resolution computed tomography, we present here, to our knowledge, the first comprehensive study of the pattern of variation of the inner ear with a focus on Xenarthra. Our study demonstrates that extant three-toed sloths show a high level of morphological variation of the bony labyrinth of the inner ear. Especially, the variation in shape, relative size and angles of their SCs greatly differ from those of other, faster-moving taxa within Xenarthra and Placentalia in general. The unique pattern of variation in three-toed sloths suggests that a release of selection and/or constraints on their organ of balance is associated with the observed wide range of phenotypes. This release is coincident with their slow and infrequent locomotion and may be related, among other possible factors, to a reduced functional demand for a precise sensitivity to movement.

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Aim To determine the evolutionary history, relationships and distinctiveness of allopatric populations of Hispaniolan solenodon (Solenodon paradoxus), a highly threatened Caribbean 'relict' mammal, to understand spatio-temporal patterns of gene flow and the distribution of diversity across complex large island landscapes and inform spatial conservation prioritization.Location Historical and modern-day solenodon specimens were analysed from sampling localities across Hispaniola, representing this geotectonically complex island's distinct northern, south-eastern and south-western biogeographic provinces.Methods We successfully amplified mitochondrial cytochrome b and control region sequences from 34 solenodon samples. Bayesian phylogenetic analyses were applied to assess the relationship between these sequence data, and coalescent simulation and approximate Bayesian computation were used to identify which model of solenodon intra-island demographic history best explains the observed patterns of variation. We also conducted morphometric analysis of 110 solenodon specimens to investigate whether allopatric Hispaniolan populations can be differentiated using craniodental characteristics.Results Unique haplotypes were identified in solenodon samples from each biogeographic region, with no haplotype sharing between regions. Higher marginal posterior probability values were found for a three-population model comprising allopatric northern, south-eastern and south-western Hispaniolan populations, with exceptionally low migration rates inferred between all populations, indicating that they are genetically isolated. Modal estimates of long-term effective female population size are extremely low for south-western and south-eastern populations. Morphometric differentiation is observed between all three populations.Main conclusions Evolutionary differentiation of Hispaniolan solenodons into three distinct populations is congruent with phylogenetic patterns observed in several other Hispaniolan species, with population isolation possibly associated with past marine transgression. We interpret these populations as distinct subspecies, with the two genetically impoverished southern subspecies particularly vulnerable to environmental change. Our improved understanding of Hispaniolan solenodon evolutionary history provides an important baseline for identifying wider patterns of intra-island diversification and prioritizing conservation attention for evolutionarily significant populations.

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Virtually reconstructed and natural endocranial casts are used in the study of brain evolution through geological time. We here present work investigating the paleoneurological evolution of afrotherian mammals. Using microCT-generated endocasts we show that, with the exception of the subfamilies Macroscelidinae and Tenrecoidea, most Afroinsectiphilia display a more or less gyrencephalic and ventrally expanded neopallium, two derived features that are unexpected for these insectivore-grade afrotherians. This implies that the endocranial cast morphology at the root of the afrotherian clade may have been more advanced than previously thought. The reconstructed endocranial morphology of the Afrotheria's last common ancestor reaches the level of complexity of some early Cenozoic archaic ungulates. Our result gives support to the hypothesis of an ungulate-like ancestral body plan for Afrotheria. It also implies that the a priori ‘primitive' suite of traits evident in the brain of Afroinsectivora, especially in the tenrecs, may have been secondarily acquired. Implications on the overestimation of the divergence age of Afrotheria are discussed.

We investigated if and how the inner ear region undergoes similar adaptations in small, fossorial, insectivoran-grade mammals, and found a variety of inner ear phenotypes. In our sample, afrotherian moles (Chrysochloridae) and the marsupial Notoryctes differ from most other burrowing mammals in their relatively short radii of semicircular canal curvature; chrysochlorids and fossorial talpids share a relatively long interampullar width. Chrysochlorids are unique in showing a highly coiled cochlea with nearly four turns. Extensive cochlear coiling may reflect their greater ecological dependence on low frequency auditory cues compared to talpids, tenrecids, and the marsupial Notoryctes. Correspondingly, the lack of such extensive coiling in the inner ear of other fossorial species may indicate a greater reliance on other senses to enable their fossorial lifestyle, such as tactile sensation from vibrissae and Eimer's organs. The reliance of chrysochlorids on sound is evident in the high degree of coiling and in the diversity of its mallear types, and may help explain the lack of any semiaquatic members of that group. The simplest mallear types among chrysochlorids are not present in the basal-most members of that clade, but all extant chrysochlorids investigated to date exhibit extensive cochlear coiling. The chrysochlorid ear region thus exhibits mosaic evolution; our data suggest that extensive coiling evolved in chrysochlorids prior to and independently of diversification in middle ear ossicle size and shape.

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Cranial sutures are fibrocellular joints between the skull bones that are progressively replaced with bone throughout ontogeny, facilitating growth and cranial shape change. This transition from soft tissue to bone is reflected in the biomechanical properties of the craniofacial complex. However, the mechanical significance of cranial sutures has only been explored at a few localised areas within the mammalian skull, and as such our understanding of suture function in overall skull biomechanics is still limited. Here, we sought to determine how the overall strain environment is affected by the complex network of cranial sutures in the mammal skull.We combined two computational biomechanical methods, multibody dynamics analysis and finite element analysis, to simulate biting in a rat skull and compared models with and without cranial sutures. Our results show that including complex sutures in the rat model does not substantially change overall strain gradients across the cranium, particularly strain magnitudes in the bones overlying the brain.However, local variations in strain magnitudes and patterns can be observed in areas close to the sutures. These results show that, during feeding, sutures may be more important in some regions than others. Sutures should therefore be included in models that require accurate local strain magnitudes and patterns of cranial strain, particularly if models are developed for analysis of specific regions, such as the temporomandibular joint or zygomatic arch. Our results suggest that, for mammalian skulls, cranial sutures might be more important for allowing brain expansion during growth than redistributing biting loads across the cranium in adults.