Comparative analysis of the shape and size of the middle ear cavity of turtles reveals no correlation with habitat ecology.

The middle ear of turtles differs from other reptiles in being separated into two distinct compartments. Several ideas have been proposed as to why the middle ear is compartmentalized in turtles, most suggesting a relationship with underwater hearing. Extant turtle species span fully marine to strictly terrestrial habitats, and ecomorphological hypotheses of turtle hearing predict that this should correlate with variation in the structure of the middle ear due to differences in the fluid properties of water and air. We investigate the shape and size of the air-filled middle ear cavity of 56 extant turtles using 3D data and phylogenetic comparative analysis to test for correlations between habitat preferences and the shape and size of the middle ear cavity. Only weak correlations are found between middle ear cavity size and ecology, with aquatic taxa having proportionally smaller cavity volumes. The middle ear cavity of turtles exhibits high shape diversity among species, but we found no relationship between this shape variation and ecology. Surprisingly, the estimated acoustic transformer ratio, a key functional parameter of impedance-matching ears in vertebrates, also shows no relation to habitat preferences (aquatic/terrestrial) in turtles. We suggest that middle ear cavity shape may be controlled by factors unrelated to hearing, such as the spatial demands of surrounding cranial structures. A review of the fossil record suggests that the modern turtle ear evolved during the Early to Middle Jurassic in stem turtles broadly adapted to freshwater and terrestrial settings. This, combined with our finding that evolutionary transitions between habitats caused only weak evolutionary changes in middle ear structure, suggests that tympanic hearing in turtles evolved as a compromise between subaerial and underwater hearing.

Modeling neck mobility in fossil turtles.

Turtles have the unparalleled ability to retract their heads and necks within their shell but little is known about the evolution of this trait. Extensive analysis of neck mobility in turtles using radiographs, CT scans, and morphometry reveals that basal turtles possessed less mobility in the neck relative to their extant relatives, although the anatomical prerequisites for modern mobility were already established. Many extant turtles are able to achieve hypermobility by dislocating the central articulations, which raises cautions about reconstructing the mobility of fossil vertebrates. A 3D-model of the Late Triassic turtle Proganochelys quenstedti reveals that this early stem turtle was able to retract its head by tucking it sideways below the shell. The simple ventrolateral bend seen in this stem turtle, however, contrasts with the complex double-bend of extant turtles. The initial evolution of neck retraction therefore occurred in a near-synchrony with the origin of the turtle shell as a place to hide the unprotected neck. In this early, simplified retraction mode, the conical osteoderms on the neck provided further protection.

Hand to mouth in a neandertal: right-handedness in Regourdou 1.

We describe and analyze a Neandertal postcranial skeleton and dentition, which together show unambiguous signs of right-handedness. Asymmetries between the left and right upper arm in Regourdou 1 were identified nearly 20 years ago, then confirmed by more detailed analyses of the inner bone structure for the clavicle, humerus, radius and ulna. The total pattern of all bones in the shoulder and arm reveals that Regourdou 1 was a right-hander. Confirmatory evidence comes from the mandibular incisors, which display a distinct pattern of right oblique scratches, typical of right-handed manipulations performed at the front of the mouth. Regourdou's right handedness is consistent with the strong pattern of manual lateralization in Neandertals and further confirms a modern pattern of left brain dominance, presumably signally linguistic competence. These observations along with cultural, genetic and morphological evidence indicate language competence in Neandertals and their European precursors.

Vertebral microanatomy in squamates: structure, growth and ecological correlates.

The histological study of vertebrae in extant squamates shows that the internal vertebral structure in this group differs from that of other tetrapods. Squamate vertebrae are lightly built and basically composed of two roughly concentric osseous tubes--one surrounding the neural canal and the other constituting the peripheral cortex of the vertebra--connected by few thin trabeculae. This structure, which characteristically evokes that of a tubular bone, results from a peculiar remodelling process characterised by an imbalance between local bone resorption and redeposition; in both periosteal and endosteo-endochondral territories, bone is extensively resorbed but not reconstructed in the same proportion by secondary deposits. This process is particularly intense in the deep region of the centrum, where originally compact cortices are made cancellous, and where the endochondral spongiosa is very loose. This remodelling process starts at an early stage of development and remains active throughout subsequent growth. The growth of squamate centra is also strongly asymmetrical, with the posterior (condylar) part growing much faster than the anterior (cotylar) part. Preliminary analyses testing for associations between vertebral structure and habitat use suggest that vertebrae of fossorial taxa are denser than those of terrestrial taxa, those in aquatic taxa being of intermediate density. However, phylogenetically informed analyses do not corroborate these findings, thus suggesting a strong phylogenetic signal in the data. As our analyses demonstrate that vertebrae in snakes are generally denser than those of lizards sensu stricto, this may drive the presence of a phylogenetic signal in the data. More comprehensive sampling of fossorial and aquatic lizards is clearly needed to more rigorously evaluate these patterns.

Technical note: Morphometric maps of long bone shafts and dental roots for imaging topographic thickness variation.

Qualitative and quantitative characterization through functional imaging of mineralized tissues is of potential value in the study of the odontoskeletal remains. This technique, widely developed in the medical field, allows the bi-dimensional, planar representation of some local morphometric properties, i.e., topographic thickness variation, of a three-dimensional object, such as a long bone shaft. Nonetheless, the use of morphometric maps is still limited in (paleo)anthropology, and their feasibility has not been adequately tested on fossil specimens. Using high-resolution microtomographic images, here we apply bi-dimensional virtual "unrolling" and synthetic thickness mapping techniques to compare cortical bone topographic variation across the shaft in a modern and a fossil human adult femur (the Magdalenian from Chancelade). We also test, for the first time, the possibility to virtually unroll and assess for dentine thickness variation in modern and fossil (the Neanderthal child from Roc de Marsal) human deciduous tooth roots. The analyses demonstrate the feasibility of using two-dimensional morphometric maps for the synthetic functional imaging and comparative biomechanical interpretation of cortical bone thickness variation in extant and fossil specimens and show the interest of using this technique also for the subtle characterization of root architecture and dentine topography. More specifically, our preliminary results support the use of virtual cartography as a tool for assessing to what extent internal root morphology is capable of responding to loading and directional stresses and strains in a predictable way.

Textural characteristics of the iliac-femoral trabecular pattern in a bipedally trained Japanese macaque.

Previous research has revealed that Japanese macaques (Macaca fuscata) trained in bipedal performance (Saru-mawashi) display a number of functionally related external skeletal changes, as well as some site-specific endostructural cortical and cancellous adaptations. Through radiography assisted by digital image processing, we investigated the trabecular pattern of the ilium and femoral neck of Sansuke, a macaque habitually bipedally trained for 8 years. A set of eight regions of interest on Sansuke were recorded for subtle structural characterization of the cancellous network and compared to a sample of 5 ilia and 23 femurs from 26 adult wild Japanese macaques. The measured variables include trabecular thickness, trabecular bone volume, and degree of anisotropy. As a whole, the site-specific textural characteristics of the cancellous network detected on the ilium and proximal femur of the bipedally trained macaque can be interpreted as functional adaptive responses to more compressive loads dissipated along the axis from the sacro-iliac to the coxo-femoral joint, and back.