Charles Oxnard

Evolutionary radiations and convergences in the structural organization of mammalian brains

The sizes of mammalian brain components seem to be mostly related to the sizes of the whole brain (and body), suggesting a one-dimensional scale of encephalization. Previous multivariate study of such data concludes that evolutionary selection for enlargement of any one brain part is constrained to selection for a concerted enlargement of the whole brain. However, interactions between structurally related pairs of brain parts confirm reports of differential change in brain nuclei, and imply mosaic rather than concerted evolution. Here we analyse a large number of variables simultaneously using multi-dimensional methods. We show that the relative proportions of different systems of functionally integrated brain structures vary independently between different mammalian orders, demonstrating separate evolutionary radiations in mammalian brain organization. Within each major order we identify clusters of unrelated species that occupy similar behavioural niches and have convergently evolved similar brain proportions. We conclude that within orders, mosaic brain organization is caused by selective adaptation, whereas between orders it suggests an interplay between selection and constraints.

Assessing mechanical function of the zygomatic region in macaques: validation and sensitivity testing of finite element models.

Crucial to the interpretation of the results of any finite element analysis of a skeletal system is a test of the validity of the results and an assessment of the sensitivity of the model parameters. We have therefore developed finite element models of two crania of Macaca fascicularis and investigated their sensitivity to variations in bone material properties, the zygomatico‐temporal suture and the loading regimen applied to the zygomatic arch. Maximum principal strains were validated against data derived from ex vivo strain gauge experiments using non‐physiological loads applied to the macaque zygomatic arch. Elastic properties of the zygomatic arch bone and the zygomatico‐temporal suture obtained by nanoindentation resulted in a high degree of congruence between experimental and simulated strains. The findings also indicated that the presence of a zygomatico‐temporal suture in the model produced strains more similar to experimental values than a completely separated or fused arch. Strains were distinctly higher when the load was applied through the modelled superficial masseter compared with loading an array of nodes on the arch. This study demonstrates the importance of the accurate selection of the material properties involved in predicting strains in a finite element model. Furthermore, our findings strongly highlight the influence of the presence of craniofacial sutures on strains experienced in the face. This has implications when investigating craniofacial growth and masticatory function but should generally be taken into account in functional analyses of the craniofacial system of both extant and extinct species.

Sexual Dimorphism in the Subadult Mandible: Quantification Using Geometric Morphometrics

ABSTRACT: There have been numerous attempts, with varying degrees of success, to differentiate males from females on the basis of the immature skeleton. We investigate here whether the mandible can discriminate immature individuals by sex; the techniques we apply are from the field of geometric morphometrics. The application of these methods in forensic anthropology is still relatively new; thus, an important aspect of this research is that it demonstrates potential applications in this discipline. The sample comprises 96 known age and sex subadult individuals; the three‐dimensional coordinates of 38 landmarks are analyzed using the shape analysis software morphologika. Multivariate regressions indicated no significant sexual dimorphism in the subadult sample; this result is supported by poor cross‐validated classification accuracy (59%). Our results suggest that the subadult mandible is not dimorphic (to the extent that dimorphism is not evident within the sample we studied); thus, sex determination using previously described criteria is likely to yield poor results.

Masticatory loading and bone adaptation in the supraorbital torus of developing macaques

Research on the evolution and adaptive significance of primate craniofacial morphologies has focused on adult, fully developed individuals. Here, we investigate the possible relationship between the local stress environment arising from masticatory loadings and the emergence of the supraorbital torus in the developing face of the crab-eating macaque Macaca fascicularis. By using finite element analysis (FEA), we are able to evaluate the hypothesis that strain energy density (SED) magnitudes are high in subadult individuals with resulting bone growth in the supraorbital torus. We developed three micro-CT-based FEA models of M. fascicularis skulls ranging in dental age from deciduous to permanent dentitions and validated them against published experimental data. Applied masticatory muscle forces were estimated from physiological cross-sectional areas of macaque cadaveric specimens. The models were sequentially constrained at each working side tooth to simulate the variation of the bite point applied during masticatory function. Custom FEA software was used to solve the voxel-based models and SED and principal strains were computed. A physiological superposition SED map throughout the face was created by allocating to each element the maximum SED value from each of the load cases. SED values were found to be low in the supraorbital torus region throughout ontogeny, while they were consistently high in the zygomatic arch and infraorbital region. Thus, if the supraorbital torus arises to resist masticatory loads, it is either already adapted in each of our subadult models so that we do not observe high SED or a lower site-specific bone deposition threshold must apply.

Are the small human-like fossils found on Flores human endemic cretins?

Fossils from Liang Bua (LB) on Flores, Indonesia, including a nearly complete skeleton (LB1) dated to 18 kyr BP, were assigned to a new species, Homo floresiensis. We hypothesize that these individuals are myxoedematous endemic (ME) cretins, part of an inland population of (mostly unaffected) Homo sapiens. ME cretins are born without a functioning thyroid; their congenital hypothyroidism leads to severe dwarfism and reduced brain size, but less severe mental retardation and motor disability than neurological endemic cretins. We show that the fossils display many signs of congenital hypothyroidism, including enlarged pituitary fossa, and that distinctive primitive features of LB1 such as the double rooted lower premolar and the primitive wrist morphology are consistent with the hypothesis. We find that the null hypothesis (that LB1 is not a cretin) is rejected by the pituitary fossa size of LB1, and by multivariate analyses of cranial measures. We show that critical environmental factors were potentially present on Flores, how remains of cretins but not of unaffected individuals could be preserved in caves, and that extant oral traditions may provide a record of cretinism.

Animal lifestyles and anatomies : the case of the prosimian primates

An ambitious and original attempt to quantify, analyze, and compare the environment, behavior, and morphology of prosimian primates on a global scale. A species-by-species presentation, along with superb drawings of the animals and graphs showing complex combinations of variables, is put to the serv

Discriminant function sexing of the mandible of Indigenous South Africans

South Africa currently has a high homicide rate. This results in a large number of unidentified bodies being recovered each year, many of which are referred to the forensic examiner. This situation has resulted in considerable growth of forensic anthropological research devoted to devising standards for specific application in South African medico-legal investigations. The standards suitable for Black South Africans now encompass a wide variety of skeletal elements (e.g. cranium, humerus, pelvis, femur, patella, talus, calcaneus), each with differing degrees of accuracy. Apart from a preliminary investigation of the Zulu local population, however, we note that there appears to be no established metric mandible discriminant function standards for sex determination in this population. The purpose of the present study is to undertake a comprehensive analysis of sexual dimorphism in the mandible of Black South Africans, incorporating individuals from a selection of the larger local population groupings; the primary aim is to produce a series of metrical standards for the determination of sex. The sample analyzed comprises 225 non-pathological mandibles of Black South African individuals drawn from the R.A. Dart Collection. Nine linear measurements, obtained from mathematically transformed three-dimensional landmark data, are analyzed using basic univariate statistics and discriminant function analyses. All of the measurements examined are found to be sexually dimorphic; the dimensions of the ramus and corpus lengths are most dimorphic. The sex classification accuracy of the discriminant functions ranged from 70.7 to 77.3% for the univariate method, 81.8% for the stepwise method, and 63.6 to 84% for the direct method. We conclude that the mandible is a very useful element for sex determination in this population.

Bone and bones, architecture and stress, fossils and osteoporosis

The combined use of architectural and stress technologies in osteological studies is starting to provide the basic biomechanical underpinnings to both evolutionary and applied medical investigations of bone. The architectural investigations, though tested using invasive methods, are aimed at non-invasive ways of obtaining information from radiographs of bones, fossils and people. They include optical (Fourier) data analysis (ODA) and computational Fourier transformations (FFT). The stress studies, though initially involving older techniques such as photoelastic stress analysis, now employ finite element analysis (FEA) and, most recently, fast Lagrangian analysis of continua (FLAC). Taken together, these methods are capable of providing more detailed knowledge of bone form and function that is important (a) in revealing functional adaptation in evolutionary studies of fossils and (b) for making early diagnosis and understanding pathological fractures in the late stages of osteoporosis.

determination of sex in south african blacks by discriminant function analysis of mandibular linear dimensions

The determination of sex is a critical component in forensic anthropological investigation. The literature attests to numerous metrical standards, each utilizing diffetent skeletal elements, for sex determination in South A frican Blacks. Metrical standards are popular because they provide a high degree of expected accuracy and are less error-prone than subjective nonmetric visual techniques. We note, however, that there appears to be no established metric mandible discriminant function standards for sex determination in this population.We report here on a preliminary investigation designed to evaluate whether the mandible is a practical element for sex determination in South African Blacks. The sample analyzed comprises 40 nonpathological Zulu individuals drawn from the R.A. Dart Collection. Ten linear measurements, obtained from mathematically trans-formed three-dimensional landmark data, are analyzed using basic univariate statistics and discriminant function analyses. Seven of the 10 measurements examined are found to be sexually dimorphic; the dimensions of the ramus are most dimorphic. The sex classification accuracy of the discriminant functions ranged from 72.5 to 87.5% for the univariate method, 92.5% for the stepwise method, and 57.5 to 95% for the direct method. We conclude that the mandible is an extremely useful element for sex determination in this population.

Post-cranial skeletons of hypothyroid cretins show a similar anatomical mosaic as Homo floresiensis.

Human remains, some as recent as 15 thousand years, from Liang Bua (LB) on the Indonesian island of Flores have been attributed to a new species, Homo floresiensis. The definition includes a mosaic of features, some like modern humans (hence derived: genus Homo), some like modern apes and australopithecines (hence primitive: not species sapiens), and some unique (hence new species: floresiensis). Conversely, because only modern humans (H. sapiens) are known in this region in the last 40 thousand years, these individuals have also been suggested to be genetic human dwarfs. Such dwarfs resemble small humans and do not show the mosaic combination of the most complete individuals, LB1 and LB6, so this idea has been largely dismissed. We have previously shown that some features of the cranium of hypothyroid cretins are like those of LB1. Here we examine cretin postcrania to see if they show anatomical mosaics like H. floresiensis. We find that hypothyroid cretins share at least 10 postcranial features with Homo floresiensis and unaffected humans not found in apes (or australopithecines when materials permit). They share with H. floresiensis, modern apes and australopithecines at least 11 postcranial features not found in unaffected humans. They share with H. floresiensis, at least 8 features not found in apes, australopithecines or unaffected humans. Sixteen features can be rendered metrically and multivariate analyses demonstrate that H. floresiensis co-locates with cretins, both being markedly separate from humans and chimpanzees (P<0.001: from analysis of similarity (ANOSIM) over all variables, ANOSIM, global R>0.999). We therefore conclude that LB1 and LB6, at least, are, most likely, endemic cretins from a population of unaffected Homo sapiens. This is consistent with recent hypothyroid endemic cretinism throughout Indonesia, including the nearby island of Bali.