Anneke H. van Heteren

Functional morphology of the cave bear (Ursus spelaeus) mandible: a 3D geometric morphometric analysis

The diet of the fossil cave bears (Ursus spelaeus group) has been debated extensively. Thought traditionally to be herbivorous, more recent studies have proposed more meat in the cave bear diet. To test this, the mandibular morphology of cave bears was analysed using 3D geometric morphometrics and compared to that of extant Ursidae. Landmarks for 3D digitisation of the mandible were chosen to reflect functional morphology relating to the temporalis and masseter muscles. Extant and extinct Pleistocene Ursidae were digitised with a MicroScribe G2. Generalised Procrustes superimposition was performed, and data were allometrically and phylogenetically corrected. Principal component analysis (PCA), two-block partial least squares analysis (2B-PLS), regression analysis and discriminant function analysis were performed. PCA and 2B-PLS differentiate between known dietary niches in extant Ursidae. The lineage of the cave bear runs parallel to that of the panda (Ailuropoda melanoleuca) in morphospace, implying the development of morphological adaptations for eating foliage. A regression of shape onto foliage content in the diet and a discriminant function analysis also indicate that the cave bear diet consisted primarily of foliage.

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.

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

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.

A review of the dodo and its ecosystem: insights from a vertebrate concentration Lagerstätte in Mauritius

ABSTRACT The dodo Raphus cucullatus Linnaeus, 1758, an extinct and flightless, giant pigeon endemic to Mauritius, has fascinated people since its discovery, yet has remained surprisingly poorly known. Until the mid-19th century, almost all that was known about the dodo was based on illustrations and written accounts by 17th century mariners, often of questionable accuracy. Furthermore, only a few fragmentary remains of dodos collected prior to the birds extinction exist. Our understanding of the dodos anatomy was substantially enhanced by the discovery in 1865 of subfossil bones in a marsh called the Mare aux Songes, situated in southeastern Mauritius. However, no contextual information was recorded during early excavation efforts, and the majority of excavated material comprised larger dodo bones, almost all of which were unassociated. Here we present a modern interdisciplinary analysis of the Mare aux Songes, a 4200-year-old multitaxic vertebrate concentration Lagerstätte. Our analysis of the deposits at this site provides the first detailed overview of the ecosystem inhabited by the dodo. The interplay of climatic and geological conditions led to the exceptional preservation of the animal and associated plant remains at the Mare aux Songes and provides a window into the past ecosystem of Mauritius. This interdisciplinary research approach provides an ecological framework for the dodo, complementing insights on its anatomy derived from the only associated dodo skeletons known, both of which were collected by Etienne Thirioux and are the primary subject of this memoir. Citation for this article: Rijsdijk, K. F., J. P. Hume, P. G. B. de Louw, H. J. M. Meijer, A. Janoo, E. J. de Boer, L. Steel, J. de Vos, L. G. van der Sluis, H. Hooghiemstra, F. B. V. Florens, C. Baider, T. J. J. Vernimmen, P. Baas, A. H. van Heteren, V. Rupear, G. Beebeejaun, A. Grihault, J. van der Plicht, M. Besselink, J. K. Lubeek, M. Jansen, S. J. Kluiving, H. Hollund, B. Shapiro, M. Collins, M. Buckley, R. M. Jayasena, N. Porch, R. Floore, F. Bunnik, A. Biedlingmaier, J. Leavitt, G. Monfette, A. Kimelblatt, A. Randall, P. Floore, and L. P. A. M. Claessens. 2015. A review of the dodo and its ecosystem: insights from a vertebrate concentration Lagerstätte in Mauritius; pp. 3–20 in L. P. A. M. Claessens, H. J. M. Meijer, J. P. Hume, and K. F. Rijsdijk (eds.), Anatomy of the Dodo (Raphus cucullatus L., 1758): An Osteological Study of the Thirioux Specimens. Society of Vertebrate Paleontology Memoir 15. Journal of Vertebrate Paleontology 35(6, Supplement).

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.

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

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.

Cranial and mandibular morphology of Middle Pleistocene cave bears (Ursus deningeri): implications for diet and evolution

ABSTRACT Deninger’s bears (Ursus deningeri) have been studied less frequently than Ursus spelaeus s.l. Our objective is to present, for the first time, an analysis of the skull shape of U. deningeri. Bear crania and mandibles were digitised with a Microscribe or CT-scanned and the surface models subsequently landmarked. The landmarks were chosen based on a compromise between functional morphology and sample size. Results show that U. deningeri and U. spelaeus mandibles display very similar morphologies and allometric trajectories, both to each other and to Ailuropoda melanoleuca. It is inferred that masticatory adaptations to a herbivorous diet were already present in the Middle Pleistocene. U. deningeri displays a cranial morphology that is similar to that of U. spelaeus when comparing all species, but U. deningeri has a relatively narrower and dorsoventrally lower zygomatic arch than U. spelaeus, although the masticatory signal is less strong in the skull. We observe intraspecific differences between different populations of U. deningeri, which could parallel the genetic diversity found in U. spelaeus. The intraspecific differences found within U. deningeri may be temporal and/or geographical in nature and could be related to the evolution of the Late Pleistocene cave bear, but this hypothesis remains to be tested.

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

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.

A three-dimensional analysis of tooth-root morphology in living bears and implications for feeding behaviour in the extinct cave bear

ABSTRACT The morphology of both crowns and tooth-roots reflects dietary specialisation in mammalian carnivores. In this article, we analyse the tooth-root morphology of maxillary teeth from CT scans of living bears (Ursus arctos, Ursus americanus, Ursus maritimus, Ursus thibetanus, Melursus ursinus, Helarctos malayanus, Tremarctos ornatus and Ailuropoda melanoleuca) in order to make inferences about the diet and feeding behaviour of the extinct cave bear (Ursus spelaeus sensu lato). Specifically, we investigate two major mitochondrial clades of extinct cave bears recognized by previous authors: Ursus ingressus and Ursus spelaeus (U. spelaeus spelaeus, U. spelaeus ladinicus, U. spelaeus eremus). Our results indicate a close association between tooth-root surface area and feeding behaviour in all living bear species. Tooth-root surface area values of cave bears suggest that they relied more on vegetative matter than living brown bears (Ursus arctos) but subtle differences between these species/subspecies could also indicate different feeding strategies among the members of cave bear complex.

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.