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BackgroundSciuromorpha (squirrels and close relatives) are diverse in terms of body size and locomotor behavior. Individual species are specialized to perform climbing, gliding or digging behavior, the latter being the result of multiple independent evolutionary acquisitions. Each lifestyle involves characteristic loading patterns acting on the bones of sciuromorphs. Trabecular bone, as part of the bone inner structure, adapts to such loading patterns. This network of thin bony struts is subject to bone modeling, and therefore reflects habitual loading throughout lifetime. The present study investigates the effect of body size and lifestyle on trabecular structure in Sciuromorpha.MethodsBased upon high-resolution computed tomography scans, the femoral head 3D inner microstructure of 69 sciuromorph species was analyzed. Species were assigned to one of the following lifestyle categories: arboreal, aerial, fossorial and semifossorial. A cubic volume of interest was selected in the center of each femoral head and analyzed by extraction of various parameters that characterize trabecular architecture (degree of anisotropy, bone volume fraction, connectivity density, trabecular thickness, trabecular separation, bone surface density and main trabecular orientation). Our analysis included evaluation of the allometric signals and lifestyle-related adaptation in the trabecular parameters.ResultsWe show that bone surface density, bone volume fraction, and connectivity density are subject to positive allometry, and degree of anisotropy, trabecular thickness, and trabecular separation to negative allometry. The parameters connectivity density, bone surface density, trabecular thickness, and trabecular separation show functional signals which are related to locomotor behavior. Aerial species are distinguished from fossorial ones by a higher trabecular thickness, lower connectivity density and lower bone surface density. Arboreal species are distinguished from semifossorial ones by a higher trabecular separation.ConclusionThis study on sciuromorph trabeculae supplements the few non-primate studies on lifestyle-related functional adaptation of trabecular bone. We show that the architecture of the femoral head trabeculae in Sciuromorpha correlates with body mass and locomotor habits. Our findings provide a new basis for experimental research focused on functional significance of bone inner microstructure.Electronic supplementary materialThe online version of this article (10.1186/s40851-018-0093-z) contains supplementary material, which is available to authorized users.

BackgroundBone structure has a crucial role in the functional adaptations that allow vertebrates to conduct their diverse lifestyles. Much has been documented regarding the diaphyseal structure of long bones of tetrapods. However, the architecture of trabecular bone, which is for instance found within the epiphyses of long bones, and which has been shown experimentally to be extremely plastic, has received little attention in the context of lifestyle adaptations (virtually only in primates). We therefore investigated the forelimb epiphyses of extant xenarthrans, the placental mammals including the sloths, anteaters, and armadillos. They are characterised by several lifestyles and degrees of fossoriality involving distinct uses of their forelimb. We used micro computed tomography data to acquire 3D trabecular parameters at regions of interest (ROIs) for all extant genera of xenarthrans (with replicates). Traditional, spherical, and phylogenetically informed statistics (including the consideration of size effects) were used to characterise the functional signal of these parameters.ResultsSeveral trabecular parameters yielded functional distinctions. The main direction of the trabeculae distinguished lifestyle categories for one ROI (the radial trochlea). Among the other trabecular parameters, it is the degree of anisotropy (i.e., a preferential alignment of the trabeculae) that yielded the clearest functional signal. For all ROIs, the armadillos, which represent the fully terrestrial and fossorial category, were found as characterised by a greater degree of anisotropy (i.e., more aligned trabeculae). Furthermore, the trabeculae of the humeral head of the most fossorial armadillos were also found to be more anisotropic than in the less fossorial species.ConclusionsMost parameters were marked by an important intraspecific variability and by a size effect, which could, at least partly, be masking the functional signal. But for some parameters, the degree of anisotropy in particular, a clear functional distinction was recovered. Along with data on primates, our findings suggest that a trabecular architecture characterised by a greater degree of anisotropy is to be expected in species in which the relevant epiphyses withstand a restricted range of load directions. Trabecular architecture therefore is a promising research avenue for the reconstruction of lifestyles in extinct or cryptic species.Electronic supplementary materialThe online version of this article (10.1186/s12983-017-0241-x) contains supplementary material, which is available to authorized users.

Sciuromorph rodents are a monophyletic group comprising about 300 species with a body mass range spanning three orders of magnitude and various locomotor behaviors that we categorized into arboreal, fossorial and aerial. The purpose of this study was to investigate how the interplay of locomotor ecology and body mass affects the morphology of the sciuromorph locomotor apparatus. The most proximal skeletal element of the hind limb, i.e. the femur, was selected, because it was shown to reflect a functional signal in various mammalian taxa. We analyzed univariate traits (effective femoral length, various robustness variables and the in‐levers of the muscles attaching to the greater, third and lesser trochanters) as well as femoral shape, representing a multivariate trait. An ordinary least‐squares regression including 177 species was used to test for a significant interaction effect between body mass and locomotor ecology on the variables. Specifically, it tested whether the scaling patterns of the fossorial and aerial groups differ when compared with the arboreal, because the latter was identified as the ancestral sciuromorph condition via stochastic character mapping. We expected aerial species to display the highest trait values for a given body mass as well as the steepest slopes, followed by the arboreal and fossorial species along this order. An Ornstein–Uhlenbeck regression fitted to a phylogenetically pruned dataset of 140 species revealed the phylogenetic inertia to be very low in the univariate traits, hence justifying the utilization of standard regressions. These variables generally scaled close to isometry, suggesting that scaling adjustments might not have played a major role for most of the femoral features. Nevertheless, the low phylogenetic inertia indicates that the observed scaling patterns needed to be maintained during sciuromorph evolution. Significant interaction effects were discovered in the femoral length, the centroid size of the condyles, and the in‐levers of the greater and third trochanters. Additionally, adjustments in various femoral traits reflect the acquisitions of fossorial and aerial behaviors from arboreal ancestors. Using sciuromorphs as a focal clade, our findings exemplify the importance of statistically accounting for potential interaction effects of different environmental factors in studies relating morphology to ecology.

A profound understanding of the influence of trackmaker anatomy, foot movements and substrate properties is crucial for any interpretation of fossil tracks. In this case study we analyze variability of footprint shape within one large theropod (T3), one medium-sized theropod (T2) and one ornithopod (I1) trackway from the Lower Cretaceous of Münchehagen (Lower Saxony, Germany) in order to determine the informativeness of individual features and measurements for ichnotaxonomy, trackmaker identification, and the discrimination between left and right footprints. Landmark analysis is employed based on interpretative outline drawings derived from photogrammetric data, allowing for the location of variability within the footprint and the assessment of covariation of separate footprint parts. Objective methods to define the margins of a footprint are tested and shown to be sufficiently accurate to reproduce the most important results. The lateral hypex and the heel are the most variable regions in the two theropod trackways. As indicated by principal component analysis, a posterior shift of the lateral hypex is correlated with an anterior shift of the margin of the heel. This pattern is less pronounced in the ornithopod trackway, indicating that variation patterns can differ in separate trackways. In all trackways, hypices vary independently from each other, suggesting that their relative position a questionable feature for ichnotaxonomic purposes. Most criteria commonly employed to differentiate between left and right footprints assigned to theropods are found to be reasonably reliable. The described ornithopod footprints are asymmetrical, again allowing for a left–right differentiation. Strikingly, 12 out of 19 measured footprints of the T2 trackway are stepped over the trackway midline, rendering the trackway pattern a misleading left–right criterion for this trackway. Traditional measurements were unable to differentiate between the theropod and the ornithopod trackways. Geometric morphometric analysis reveals potential for improvement of existing discriminant methods.

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A profound understanding of the influence of trackmaker anatomy, foot movements and substrate properties is crucial for any interpretation of fossil tracks. In this case study we analyze variability of footprint shape within one large theropod (T3), one medium-sized theropod (T2) and one ornithopod (I1) trackway from the Lower Cretaceous of Münchehagen (Lower Saxony, Germany) in order to determine the informativeness of individual features and measurements for ichnotaxonomy, trackmaker identification, and the discrimination between left and right footprints. Landmark analysis is employed based on interpretative outline drawings derived from photogrammetric data, allowing for the location of variability within the footprint and the assessment of covariation of separate footprint parts. Objective methods to define the margins of a footprint are tested and shown to be sufficiently accurate to reproduce the most important results.The lateral hypex and the heel are the most variable regions in the two theropod trackways. As indicated by principal component analysis, a posterior shift of the lateral hypex is correlated with an anterior shift of the margin of the heel. This pattern is less pronounced in the ornithopod trackway, indicating that variation patterns can differ in separate trackways. In all trackways, hypices vary independently from each other, rendering their relative position a questionable feature for ichnotaxonomic purposes. Most criteria commonly employed to differentiate between left and right footprints assigned to theropods are found to be reasonably reliable. The described ornithopod footprints are asymmetrical, again allowing for a left-right differentiation. Strikingly, 12 out of 19 measured footprints of the T2 trackway are stepped over the trackway midline, rendering the trackway pattern a misleading left-right criterion for this trackway. Traditional measurements were unable to differentiate between the theropod and the ornithopod trackways. Geometric morphometric analysis reveals potential for improvement of existing discriminant methods. A profound understanding of the influence of trackmaker anatomy, foot movements and substrate properties is crucial for any interpretation of fossil tracks. In this case study we analyze variability of footprint shape within one large theropod (T3), one medium-sized theropod (T2) and one ornithopod (I1) trackway from the Lower Cretaceous of Münchehagen (Lower Saxony, Germany) in order to determine the informativeness of individual features and measurements for ichnotaxonomy, trackmaker identification, and the discrimination between left and right footprints. Landmark analysis is employed based on interpretative outline drawings derived from photogrammetric data, allowing for the location of variability within the footprint and the assessment of covariation of separate footprint parts. Objective methods to define the margins of a footprint are tested and shown to be sufficiently accurate to reproduce the most important results. The lateral hypex and the heel ...

BackgroundSloths are one of only two exceptions to the mammalian ‘rule of seven’ vertebrae in the neck. As a striking case of breaking the evolutionary constraint, the explanation for the exceptional number of cervical vertebrae in sloths is still under debate. Two diverging hypotheses, both ultimately linked to the low metabolic rate of sloths, have been proposed: hypothesis 1 involves morphological transformation of vertebrae due to changes in the Hox gene expression pattern and hypothesis 2 assumes that the Hox gene expression pattern is not altered and the identity of the vertebrae is not changed. Direct evidence supporting either hypothesis would involve knowledge of the vertebral Hox code in sloths, but the realization of such studies is extremely limited. Here, on the basis of the previously established correlation between anterior Hox gene expression and the quantifiable vertebral shape, we present the morphological regionalization of the neck in three different species of sloths with aberrant cervical count providing indirect insight into the vertebral Hox code.ResultsShape differences within the cervical vertebral column suggest a mouse-like Hox code in the neck of sloths. We infer an anterior shift of HoxC-6 expression in association with the first thoracic vertebra in short-necked sloths with decreased cervical count, and a posterior shift of HoxC-5 and HoxC-6 expression in long-necked sloths with increased cervical count.ConclusionAlthough only future developmental analyses in non-model organisms, such as sloths, will yield direct evidence for the evolutionary mechanism responsible for the aberrant number of cervical vertebrae, our observations lend support to hypothesis 1 indicating that the number of modules is retained but their boundaries are displaced. Our approach based on quantified morphological differences also provides a reliable basis for further research including fossil taxa such as extinct ‘ground sloths’ in order to trace the pattern and the underlying genetic mechanisms in the evolution of the vertebral column in mammals.Electronic supplementary materialThe online version of this article (10.1186/s12862-018-1202-5) contains supplementary material, which is available to authorized users.