Femoral morphology of sciuromorph rodents in light of scaling and locomotor ecology.

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.

Homeotic transformations reflect departure from the mammalian 'rule of seven' cervical vertebrae in sloths: inferences on the Hox code and morphological modularity of the mammalian neck.

BACKGROUND: Sloths 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. RESULTS: Shape 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. CONCLUSION: Although 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.

Trabecular architecture in the forelimb epiphyses of extant xenarthrans (Mammalia).

BACKGROUND: Bone 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. RESULTS: Several 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. CONCLUSIONS: Most 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.

Covariation of proximal finger and toe phalanges in Homo sapiens: A novel approach to assess covariation of serially corresponding structures.

OBJECTIVES: As hands and feet are serially repeated corresponding structures in tetrapods, the morphology of fingers and toes is expected to covary due to a shared developmental origin. The present study focuses on the covariation of the shape of proximal finger and toe phalanges of adult Homo sapiens to determine whether covariation is different in the first ray relative to the others, as its morphology is also different. MATERIAL AND METHODS: Proximal phalanges of 76 individuals of unknown sex (Muséum national d'Histoire naturelle, Paris, and the Natural History Museum, London) were digitized using a surface scanner. Landmarks were positioned on 3D surface models of the phalanges. Generalized Procrustes analysis and two-block partial least squares (PLS) analyses were conducted. A novel landmark-based geometric morphometric approach focusing on covariation is based on a PCoA of the angles between PLS axes in morphospace. The results can be statistically evaluated. RESULTS: The difference in PCo scores between the first and the other rays indicates that the integration between the thumb and the big toe is different from that between the lateral rays of the hand and foot. DISCUSSION: We speculate that the results are possibly the evolutionary consequence of differential selection pressure on the big toe relative to the other toes related to the rise of bipedalism, which is proposed to have emerged very early in the hominin clade. In contrast, thumb morphology and its precision grip never ceased undergoing changes, suggesting less acute selection pressures related to the evolution of the precision grip.

Ancient DNA of the pygmy marmoset type specimen Cebuella pygmaea (Spix, 1823) resolves a taxonomic conundrum.

The pygmy marmoset, the smallest of the anthropoid primates, has a broad distribution in Western Amazonia. Recent studies using molecular and morphological data have identified two distinct species separated by the Napo and Solimões-Amazonas rivers. However, reconciling this new biological evidence with current taxonomy, i.e., two subspecies, Cebuella pygmaea pygmaea (Spix, 1823) and Cebuella pygmaea niveiventris (Lönnberg, 1940), was problematic given the uncertainty as to whether Spix's pygmy marmoset ( Cebuella pygmaea pygmaea) was collected north or south of the Napo and Solimões-Amazonas rivers, making it unclear to which of the two newly revealed species the name pygmaea would apply. Here, we present the first molecular data from Spix's type specimen of Cebuella pygmaea, as well as novel mitochondrial genomes from modern pygmy marmosets sampled near the type locality (Tabatinga) on both sides of the river. With these data, we can confirm the correct names of the two species identified, i.e., C. pygmaea for animals north of the Napo and Solimões-Amazonas rivers and C. niveiventris for animals south of these two rivers. Phylogenetic analyses of the novel genetic data placed into the context of cytochrome b gene sequences from across the range of pygmy marmosets further led us to re-evaluate the geographical distribution for the two Cebuella species. We dated the split of these two species to 2.54 million years ago. We discuss additional, more recent, subdivisions within each lineage, as well as potential contact zones between the two species in the headwaters of these rivers.

Scythes, sickles and other blades: defining the diversity of pectoral fin morphotypes in Pachycormiformes.

The traditional terminology of 'scythe' or 'sickle' shaped is observed to be flawed as an effective descriptor for pectoral fin shape in pachycormids. The diversity of pachycormid pectoral fin shapes is assessed across the 14 recognised genera that preserve complete pectoral fins, and improved terms are defined to more effectively describe their form, supported by anatomical observation and aspect ratio analysis of individual fins, and corroborated by landmark analysis. Three clear and distinct pectoral fin structural morphotypes emerge (falceform, gladiform, falcataform), reflecting a diversity of pachycormid lifestyles throughout the Mesozoic, from agile pursuit predator to slow-cruising suspension feeder.

Trabecular architecture in the sciuromorph femoral head: allometry and functional adaptation.

BACKGROUND: Sciuromorpha (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. METHODS: Based 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. RESULTS: We 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. CONCLUSION: This 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.

Neither slim nor fat: estimating the mass of the dodo (Raphus cucullatus, Aves, Columbiformes) based on the largest sample of dodo bones to date.

The dodo (Raphus cucullatus) might be the most enigmatic bird of all times. It is, therefore, highly remarkable that no consensus has yet been reached on its body mass; previous scientific estimates of its mass vary by more than 100%. Until now, the vast amount of bones stored at the Natural History Museum in Mauritius has not yet been studied morphometrically nor in relation to body mass. Here, a new estimate of the dodo's mass is presented based on the largest sample of dodo femora ever measured (n = 174). In order to do this, we have used the regression method and chosen our variables based on biological, mathematical and physical arguments. The results indicate that the mean mass of the dodo was circa 12 kg, which is approximately five times as heavy as the largest living Columbidae (pigeons and doves), the clade to which the dodo belongs.

Correction: Neither slim nor fat: estimating the mass of the dodo (Raphus cucullatus, Aves, Columbiformes) based on the largest sample of dodo bones to date.

[This corrects the article DOI: 10.7717/peerj.4110.].

Geometric morphometric analysis of intratrackway variability: a case study on theropod and ornithopod dinosaur trackways from Münchehagen (Lower Cretaceous, Germany).

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.

The Quadratojugal of Eryops studied by computed tomography and the morphological variability of foramina and canals in the quadratojugal of basal tetrapods.

With respect to its large size and abundance, Eryops is an important representative of Permo-Carboniferous basal tetrapods and one of the best-known large temnospondyl amphibians of this period. This taxon forms a significant component of the Early Permian tetrapod fauna of Texas and New Mexico and here we describe a new record of skull remains, the first one from Brushy Creek (30 km northeast of Seymour) in Texas (Petrolia Formation, Wichita Group; Lower Permian - lower Artinskian). Our material, found in 2015, consists of a left nasal, a jaw fragment (premaxilla or maxilla), left quadratojugal fragments, and a partial left mandible. We used computed tomography methods (micro-CT) for imaging both internal and external structures, for the first time, for Eryops. The quadratojugal presented here is exceptional compared to all known basal tetrapods in having four different internal foramina. CT data show that these foramina are interconnected by canals within the bone. This indicates that the morphology of the foramina and the course of the canals in the quadratojugal of basal tetrapods are more variable than hitherto thought. Anat Rec, 299:1073-1079, 2016. © 2016 Wiley Periodicals, Inc.

Is LB1 diseased or modern? A review of the proposed pathologies.

The fossil remains of Homo floresiensis have been debated extensively over the past few years. This paper gives a review of the various pathologies ascribed to LB1, the type specimen of H. floresiensis, and associated individuals. This paper will assess the arguments for growth anomalies, microcephaly, Laron syndrome and cretinism. Additionally, some of the analyses done by proponents of the pathology theory will be methodologically evaluated. Subsequently, a brief overview of the alternative hypotheses regarding the origin of H. floresiensis will be given.