Geographic variation in gorilla limb bones.

Gorilla systematics has received increased attention over recent decades from primatologists, conservationists, and paleontologists. Studies of geographic variation in DNA, skulls, and teeth have led to new taxonomic proposals, such as recognition of two gorilla species, Gorilla gorilla (western gorilla) and Gorilla beringei (eastern gorilla). Postcranial differences between mountain gorillas (G. beringei beringei) and western lowland gorillas (G. g. gorilla) have a long history of study, but differences between the limb bones of the eastern and western species have not yet been examined with an emphasis on geographic variation within each species. In addition, proposals for recognition of the Cross River gorilla as Gorilla gorilla diehli and gorillas from Tshiaberimu and Kahuzi as G. b. rex-pymaeorum have not been evaluated in the context of geographic variation in the forelimb and hindlimb skeletons. Forty-three linear measurements were collected from limb bones of 266 adult gorillas representing populations of G. b. beringei, Gorilla beringei graueri, G. g. gorilla, and G. g. diehli in order to investigate geographic diversity. Skeletal elements included the humerus, radius, third metacarpal, third proximal hand phalanx, femur, tibia, calcaneus, first metatarsal, third metatarsal, and third proximal foot phalanx. Comparisons of means and principal components analyses clearly differentiate eastern and western gorillas, indicating that eastern gorillas have absolutely and relatively smaller hands and feet, among other differences. Gorilla subspecies and populations cluster consistently by species, although G. g. diehli may be similar to the eastern gorillas in having small hands and feet. The subspecies of G. beringei are distinguished less strongly and by different variables than the two gorilla species. Populations of G. b. graueri are variable, and Kahuzi and Tshiaberimu specimens do not cluster together. Results support the possible influence of higher-altitude Pleistocene refugia on patterns of geographic variation in gorillas.

Scaling effect on the mid-diaphysis properties of long bones-the case of the Cervidae (deer).

How skeletal elements scale to size is a fundamental question in biology. While the external shape of long bones was intensively studied, an important component of their organization is also found in their less accessible inner structure. Here, we studied mid-diaphyseal properties of limb long bones, characterizing notably the thickness of their cortices (bone walls), in order to test whether body size directly influences bone inner organization. Previous examinations of scaling in long bones used broad samplings to encompass a wide range of body sizes. To account for the effect of confounding factors related to different lifestyles, we focused our comprehensive sampling on a mammalian clade that comprises various body sizes but a relatively uniform lifestyle, the Cervidae. Positive allometry was found in femoral cross-sectional shape, indicating greater directional bending rigidity in large-sized taxa. None of the compactness parameters scaled allometrically in any of their bones. The cortices of sampled zeugopodial bones (tibia and radius) were found as significantly thicker than those of stylopodial bones (femur and humerus). Furthermore, while the mean relative cortical thickness values for both stylopodial and zeugopodial bones are close to mass-saving optima, the variance for the stylopodial bones is significantly lower. This suggests that mass saving is less intensively selected in zeugopodial bones. Finally, the long-legged Elk (Alces) and the short-legged dwarf Cretan deer (Candiacervus) featured rather thin and thick cortices, respectively, suggesting that the acquisition of a different limb proportion is accompanied by a modification of the relative mid-diaphyseal cortical thickness.

A three-dimensional analysis of the morphological evolution and locomotor behaviour of the carnivoran hind limb.

BACKGROUND: The shape of the appendicular bones in mammals usually reflects adaptations towards different locomotor abilities. However, other aspects such as body size and phylogeny also play an important role in shaping bone design.We used 3D landmark-based geometric morphometrics to analyse the shape of the hind limb bones (i.e., femur, tibia, and pelvic girdle bones) of living and extinct terrestrial carnivorans (Mammalia, Carnivora) to quantitatively investigate the influence of body size, phylogeny, and locomotor behaviour in shaping the morphology of these bones. We also investigated the main patterns of morphological variation within a phylogenetic context. RESULTS: Size and phylogeny strongly influence the shape of the hind limb bones. In contrast, adaptations towards different modes of locomotion seem to have little influence. Principal Components Analysis and the study of phylomorphospaces suggest that the main source of variation in bone shape is a gradient of slenderness-robustness. CONCLUSION: The shape of the hind limb bones is strongly influenced by body size and phylogeny, but not to a similar degree by locomotor behaviour. The slender-robust "morphological bipolarity" found in bone shape variability is probably related to a trade-off between maintaining energetic efficiency and withstanding resistance to stresses. The balance involved in this trade-off impedes the evolution of high phenotypic variability. In fact, both morphological extremes (slender/robust) are adaptive in different selective contexts and lead to a convergence in shape among taxa with extremely different ecologies but with similar biomechanical demands. Strikingly, this "one-to-many mapping" pattern of evolution between morphology and ecology in hind limb bones is in complete contrast to the "many-to-one mapping" pattern found in the evolution of carnivoran skull shape. The results suggest that there are more constraints in the evolution of the shape of the appendicular skeleton than in that of skull shape because of the strong biomechanical constraints imposed by terrestrial locomotion.

A genome-wide association study of limb bone length using a Large White × Minzhu intercross population.

BACKGROUND: In pig, limb bone length influences ham yield and body height to a great extent and has important economic implications for pig industry. In this study, an intercross population was constructed between the indigenous Chinese Minzhu pig breed and the western commercial Large White pig breed to examine the genetic basis for variation in limb bone length. The aim of this study was to detect potential genetic variants associated with porcine limb bone length. METHODS: A total of 571 F2 individuals from a Large White and Minzhu intercross population were genotyped using the Illumina PorcineSNP60K Beadchip, and phenotyped for femur length (FL), humerus length (HL), hipbone length (HIPL), scapula length (SL), tibia length (TL), and ulna length (UL). A genome-wide association study was performed by applying the previously reported approach of genome-wide rapid association using mixed model and regression. Statistical significance of the associations was based on Bonferroni-corrected P-values. RESULTS: A total of 39 significant SNPs were mapped to a 11.93 Mb long region on pig chromosome 7 (SSC7). Linkage analysis of these significant SNPs revealed three haplotype blocks of 495 kb, 376 kb and 492 kb, respectively, in the 11.93 Mb region. Annotation based on the pig reference genome identified 15 genes that were located near or contained the significant SNPs in these linkage disequilibrium intervals. Conditioned analysis revealed that four SNPs, one on SSC2 and three on SSC4, showed significant associations with SL and HL, respectively. CONCLUSIONS: Analysis of the 15 annotated genes that were identified in these three haplotype blocks indicated that HMGA1 and PPARD, which are expressed in limbs and influence chondrocyte cell growth and differentiation, could be considered as relevant biological candidates for limb bone length in pig, with potential applications in breeding programs. Our results may also be useful for the study of the mechanisms that underlie human limb length and body height.

Sexual dimorphism and ancestral variation in the pectoral and pelvic girdles of modern humans.

Discussions of the evolution of sexual dimorphism in torso shape and the pectoral region assume that this dimorphism exists independently of body size. We test this assumption in two human populations and further examine what is needed to understand sexual dimorphism in the pectoral region. Modern human males have broad shoulders and narrow hips relative to females, lending males a more triangular torso. The wider female pelvis is commonly attributed to obstetric pressures while the broader male pectoral girdle has been argued to be an adaptation that improves hunting or intrasexual competition. While sexual dimorphism in the pelvic girdle is known to exist after adjusting for body size across human populations, most studies of sexual dimorphism in the pectoral girdle have not adjusted the data to account for sexual size dimorphism or compared different ancestral groups. The aforementioned hypotheses explaining sexual dimorphism in the clavicle and scapula as products of natural selection are predicated on the untested assumption that sex differences do not scale with body size. This study tests this assumption by comparing various measurements of the pectoral girdle, the pelvic girdle, and six pectoral-pelvic indices of black and white South Africans of known sex and height to test whether the sexes and ancestral groups will differ in these values after adjusting for differences in body size. Comparisons of ancestral groups reveal that white South Africans have larger pectoral and pelvic dimensions than black South Africans, but that blacks have larger index values than whites. Regardless of differences in ancestry and body size, males have significantly broader pectoral regions as indicated by comparisons of both individual pectoral measurements and pectoral-pelvic indices. This pattern of sexual dimorphism is reversed in the pelvic region where females have larger skeletal elements. In addition to finding both absolute and relative differences in mean values for the pectoral and pelvic skeleton, females and males and blacks and whites differ in the scaling relationship of these traits, suggesting different allometric trajectories for these bones that may be explained by their distinct evolutionary functions, their adaptations to specific environments, or by changes in lengths due to age. These results suggest that sexual dimorphism in the pectoral region is not a product of scaling and that differences in this region reflect adaptive forces acting in unique ways on each sex, consistent with the assumptions of earlier evolutionary explanations.

Specialized connective tissue: bone, the structural framework of the upper extremity.

Bone is a connective tissue containing cells, fibers, and ground substance. There are many functions in the body in which the bone participates, such as storing minerals, providing internal support, protecting vital organs, enabling movement, and providing attachment sites for muscles and tendons. Bone is unique because its collagen framework absorbs energy, whereas the mineral encased within the matrix allows bone to resist deformation. This article provides an overview of the structure and function of bone tissue from a macroscopic to microscopic level and discusses the physiological processes contributing to upper extremity bone health. It concludes by discussing common conditions influencing upper extremity bone health.

Neandertal postcranial remains from the Sima de las Palomas del Cabezo Gordo, Murcia, southeastern Spain.

The Sima de las Palomas, southeastern Spain, has yielded a series of Neandertal postcranial remains, including immature and mature isolated elements and the fragmentary partial skeleton of a young adult (Palomas 92). The remains largely conform to the general late archaic/Neandertal morphological pattern in terms of humeral diaphyseal shape, pectoralis major tuberosity size and pillar thickness, ulnar coronoid process height, manual middle phalangeal epiphyseal breadth, manual distal phalangeal tuberosity shape and breadth, femoral diaphyseal shape, and probably body proportions. Palomas 92 contrasts with the Neandertals in having variably gracile hand remains, a more sellar trapezial metacarpal 1 facet, more anteroposteriorly expanded mid-proximal femoral diaphysis, and less robust pedal proximal phalanges. The Palomas Neandertals contrast with more northern European Neandertals particularly in various reflections of overall body size.

Enthesopathies as occupational stress markers: evidence from the upper limb.

Enthesopathies--that is, "musculo-skeletal stress markers"--are frequently used to reconstruct past lifestyles and activity patterns. Relatively little attention has been paid in physical anthropology to methodological gaps implicit in this approach: almost all methods previously employed neglect current medical insights into enthesopathies and the distinction between healthy and pathological aspects has been arbitrary. This study presents a new visual method of studying fibrocartilaginous enthesopathies of the upper limb (modified from Villotte: Bull Mém Soc Anthropol Paris n.s. 18 (2006) 65-85), and application of this method to 367 males who died between the 18th and 20th centuries, from four European identified skeletal collections: the Christ Church Spitalfields Collection, the identified skeletal collection of the anthropological museum of the University of Coimbra, and the Sassari and Bologna collections of the museum of Anthropology, University of Bologna. The analysis, using generalized estimating equations to model repeated binary outcome variables, has established a strong link between enthesopathies and physical activity: men with occupations involving heavy manual tasks have significantly (P-value < 0.001) more lesions of the upper limbs than nonmanual and light manual workers. Probability of the presence of an enthesopathy also increases with age and is higher for the right side compared with the left. Our study failed to distinguish significant differences between the collections when adjusted for the other effects. It appears that enthesopathies can be used to reconstruct past lifestyles of populations if physical anthropologists: 1) pay attention to the choice of entheses in their studies and 2) use appropriate methods.

Investigating the impact of captivity and domestication on limb bone cortical morphology: an experimental approach using a wild boar model.

The lack of bone morphological markers associated with the human control of wild animals has prevented the documentation of incipient animal domestication in archaeology. Here, we assess whether direct environmental changes (i.e. mobility reduction) could immediately affect ontogenetic changes in long bone structure, providing a skeletal marker of early domestication. We relied on a wild boar experimental model, analysing 24 wild-born specimens raised in captivity from 6 months to 2 years old. The shaft cortical thickness of their humerus was measured using a 3D morphometric mapping approach and compared with 23 free-ranging wild boars and 22 pigs from different breeds, taking into account sex, mass and muscle force differences. In wild boars we found that captivity induced an increase in cortical bone volume and muscle force, and a topographic change of cortical thickness associated with muscular expression along a phenotypic trajectory that differed from the divergence induced by selective breeding. These results provide an experimental proof of concept that changes in locomotor behaviour and selective breeding might be inferred from long bones morphology in the fossil and archaeological record. These trends need to be explored in the archaeological record and further studies are required to explore the developmental changes behind these plastic responses.

Descriptions of the upper limb skeleton of Homo floresiensis.

Several bones of the upper extremity were recovered during excavations of Late Pleistocene deposits at Liang Bua, Flores, and these have been attributed to Homo floresiensis. At present, these upper limb remains have been assigned to six different individuals - LB1, LB2, LB3, LB4, LB5, and LB6. Several of these bones are complete or nearly so, but some are quite fragmentary. All skeletal remains recovered from Liang Bua were extremely fragile, but have now been stabilized and hardened in the laboratory in Jakarta. They are now curated in museum-quality containers at the National Research and Development Centre for Archaeology in Jakarta, Indonesia. These skeletal remains are described and illustrated photographically. The upper limb presents a unique mosaic of derived (human-like) and primitive morphologies, the combination of which is never found in either healthy or pathological modern humans.

Australopithecus garhi: a new species of early hominid from Ethiopia.

The lack of an adequate hominid fossil record in eastern Africa between 2 and 3 million years ago (Ma) has hampered investigations of early hominid phylogeny. Discovery of 2.5 Ma hominid cranial and dental remains from the Hata beds of Ethiopia's Middle Awash allows recognition of a new species of Australopithecus. This species is descended from Australopithecus afarensis and is a candidate ancestor for early Homo. Contemporary postcranial remains feature a derived humanlike humeral/femoral ratio and an apelike upper arm-to-lower arm ratio.

A transformation method to estimate muscle attachments based on three bony landmarks.

In order to create musculoskeletal models that can be scalable to different subject specificities the calculation of the exact locations of muscle attachment is required. For this purpose, a scaling method is presented that estimates muscle attachment locations in homologous segments using three bony landmarks per segment. A data-set of 17 muscles' attachment lines from the shoulders of seven cadavers was used to assess the estimation quality of the scaling method. By knowing from the cadaver data the measured location of the muscles' attachment lines it is possible to assess the quality of the estimated ones. The scaling results showed an overall mean RMSE for the scapula and humerus muscles of 7.6 and 11.1mm, respectively. These results were then analyzed with an upper extremity model, in order to compute the influence of the RMSE in glenohumeral elevation muscle moment arms in the scapular plane. The results presented were considered to be satisfactory. Among other error contributors, the inter- and intra-subject variability should be further investigated, along with the sensitivity of a biomechanical model to these error variations.

Formation and ossification of limb elements in Trachemys scripta and a discussion of autopodial elements in turtles.

Though sequences of formation and ossification of bony elements have been described for many taxa, controversy surrounds the formation of limb elements in turtles. Three hypotheses for patterns of formation of autopodial elements have been proposed, differing primarily in the origin of Distal Carpal/Tarsal 3, the digital arch, and Centrale 4. Patterns of formation and ossification of limb elements are described for Trachemys scripta. These patterns are compared to similar data for representatives of four families of turtles (Cheloniidae, Chelydridae, Emydidae, and Trionychidae). Hypotheses of limb formation are compared in the context of new and published data. Three species (Trachemys scripta, Chrysemys picta, and Chelydra serpentina) suggest that Distal Carpal 3 forms by branching from the ulnare, whereas Distal Carpal 3 may branch from Distal Carpal 4 in Macrochelys temminckii and Chelonia mydas; data from Graptemys nigrinoda, Apalone spinifera, and Eretmochelys imbricata did not provide evidence for the origin of Distal Carpal 3. Centrale 4 was not observed to branch from the ulnare and apparently arises by de-novo condensation. Distal Carpal 4 did not branch from Centrale 4 in any species. Until the developmental origins of Distal Carpal 3 and Centrale 4 are understood, interspecific variation in the origin of these elements remains, and may explain some of the observed differences. Trends of ossification in the fore- and hind limb autopodium also are summarized. Homology of elements in pedal Digit V is discussed, and we suggest that the hooked proximal element of this digit be recognized as Distal Tarsal 5.

New evidence from China for the nature of the pterosaur evolutionary transition.

Pterosaurs are extinct flying reptiles, the first vertebrates to achieve powered flight. Our understanding of the evolutionary transition between basal, predominantly long-tailed forms to derived short-tailed pterodactyloids remained poor until the discovery of Wukongopterus and Darwinopterus in western Liaoning, China. In this paper we report on a new genus and species, Douzhanopterus zhengi, that has a reduced tail, 173% the length of the humerus, and a reduced fifth pedal digit, whose first phalange is ca. 20% the length of metatarsal III, both unique characters to Monofenestra. The morphological comparisons and phylogenetic analysis presented in this paper demonstrate that Douzhanopterus is the sister group to the 'Painten pro-pterodactyloid' and the Pterodactyloidea, reducing the evolutionary gap between long- and short-tailed pterosaurs.

Postcranial skeleton of Shinisaurus crocodilurus (Squamata: Anguimorpha).

The postcranial skeleton is poorly known for Shinisaurus crocodilurus, the Chinese crocodile lizard. Discrepancies exist between published accounts of Shinisaurus; moreover, comparisons with complete specimens show important differences from the published descriptions. Contrary to some publications, the axial skeleton variably consists of 26 or 27 presacral vertebrae, including eight cervical vertebrae. Humeral entepicondylar and ectepicondylar foramina are present, as are an epipubis and hypoischium, and the post-tubercular portion of the pubis is subequal in length to the proximal portion. Sesamoids are present in the knee, elbow, and between the penultimate phalanges and unguals. A cartilaginous strut joins the clavicles and interclavicle. Comparative investigation of extant anguimorphs add context to these observations and support the conclusion that Mosasauroidea possessed eight or more (rather than seven) cervical vertebrae. Overall, the postcranium of Shinisaurus is relatively unspecialized for Anguimorpha, although it may be diagnosed accurately by a combination of character states.

Patterns of morphological integration in the appendicular skeleton of mammalian carnivores.

We investigated patterns of evolutionary integration in the appendicular skeleton of mammalian carnivores. The findings are discussed in relation to performance selection in terms of organismal function as a potential mechanism underlying integration. Interspecific shape covariation was quantified by two-block partial least-squares (2B-PLS) analysis of 3D landmark data within a phylogenetic context. Specifically, we compared pairs of anatomically connected bones (within-limbs) and pairs of both serially homologous and functional equivalent bones (between-limbs). The statistical results of all the comparisons suggest that the carnivoran appendicular skeleton is highly integrated. Strikingly, the main shape covariation relates to bone robustness in all cases. A bootstrap test was used to compare the degree of integration between specialized cursorial taxa (i.e., those whose forelimbs are primarily involved in locomotion) and noncursorial species (i.e., those whose forelimbs are involved in more functions than their hindlimb) showed that cursors have a more integrated appendicular skeleton than noncursors. The findings demonstrate that natural selection can influence the pattern and degree of morphological integration by increasing the degree of bone shape covariation in parallel to ecological specialization.

Perinatal Specimens of Saurolophus angustirostris (Dinosauria: Hadrosauridae), from the Upper Cretaceous of Mongolia.

BACKGROUND: The Late Cretaceous Nemegt Formation, Gobi Desert, Mongolia has already yielded abundant and complete skeletons of the hadrosaur Saurolophus angustirostris, from half-grown to adult individuals. METHODOLOGY/PRINCIPAL FINDINGS: Herein we describe perinatal specimens of Saurolophus angustirostris, associated with fragmentary eggshell fragments. The skull length of these babies is around 5% that of the largest known S. angustirostris specimens, so these specimens document the earliest development stages of this giant hadrosaur and bridge a large hiatus in our knowledge of the ontogeny of S. angustirostris. CONCLUSIONS/SIGNIFICANCE: The studied specimens are likely part of a nest originally located on a riverbank point bar. The perinatal specimens were buried by sediment carried by the river current presumably during the wet summer season. Perinatal bones already displayed diagnostic characters for Saurolophus angustirostris, including premaxillae with a strongly reflected oral margin and upturned premaxillary body in lateral aspect. The absence of a supracranial crest and unfused halves of the cervical neural arches characterize the earliest stages in the ontogeny of S. angustirostris. The eggshell fragments associated with the perinatal individuals can be referred to the Spheroolithus oogenus and closely resemble those found in older formations (e.g. Barun Goyot Fm in Mongolia) or associated with more basal hadrosauroids (Bactrosaurus-Gilmoreosaurus in the Iren Dabasu Fm, Inner Mongolia, China). This observation suggests that the egg microstructure was similar in basal hadrosauroids and more advanced saurolophines. COMPETING INTERESTS: One of the authors (FE) is employed by the commercial organization Eldonia. Eldonia provided support in the form of a salary for FE, but did not have any additional role or influence in the study design, data collection and analysis, decision to publish, or preparation of the manuscript and it does not alter the authors' adherence to all the PLoS ONE policies on sharing data and materials.

Thoracic Limb Morphology of the Ring-tailed Lemur (Lemur catta) Evidenced by Gross Osteology and Radiography.

There is limited information available on the morphology of the thoracic limb of the ring-tailed lemur (Lemur catta). This study describes the morphology of the thoracic limb of captive ring-tailed lemurs evidenced by gross osteology and radiography as a guide for clinical use. Radiographic findings of 12 captive ring-tailed lemurs are correlated with bone specimens of three adult animals. The clavicle is well developed. The scapula has a large area for the origin of the m. teres major. The coracoid and hamate processes are well developed. The lateral supracondylar crest and medial epicondyle are prominent. The metacarpal bones are widely spread, and the radial tuberosity is prominent. These features indicate the presence of strong flexor muscles and flexibility of thoracic limb joints, which are important in arboreal quadrupedal locomotion. Furthermore, an ovoid ossicle is always seen at the inter-phalangeal joint of the first digit. Areas of increased soft tissue opacity are superimposed over the proximal half of the humerus and distal half of the antebrachium in male animals as a result of the scent gland. Knowledge of the morphology of the thoracic limb of individual species is important for accurate interpretation and diagnosis of musculoskeletal diseases.

Characterization of the integrity of three-dimensional trabecular bone microstructure by connectivity and shape analysis using high-resolution magnetic resonance imaging in vivo.

Bone mineral density and bone structure are the main determinants of bone strength in osteoporosis. In this study we used high-resolution magnetic resonance imaging to visualize the bone microstructure in the finger phalanges in vivo and to assess the topological three-dimensional connectivity of the trabecular network and the shape of the trabeculae as measures of bone quality. We visualized the phalanges of young and elderly healthy volunteers in vivo with a spatial resolution of 152 microm x 152 microm x 280 microm. Image processing software to quantify three measures of connectedness was developed and tested: connectivity, global connectivity density, and local connectivity density. Global three-dimensional connectivity ranged from 904 to 1,607 connections. Global connectivity density ranged from 2.9 to 4.7 connections per mm with large intersubject differences. We found a decrease of local connectivity density with growing distance from the joint ranging from 5.1 to 0.2 connections per mm. These preliminary results represent a quantitative description of the well-known rarefication of the trabecular network when moving from epiphysis to the diaphysis. Three-dimensional visualization showed a dense network consisting mostly of rod-like trabeculae at the epiphysis changing to a less dense network of a few plate-like structures near the medullary canal. An algorithm for the quantitative classification of trabecular architecture with regard to plate or rod-like shape was tested for feasibility. We conclude that in vivo assessment of three-dimensional properties of the trabecular network is possible in human phalanges. Determination of connectivity and shape will allow quantification of structural aspects of osteoporotic changes and may improve assessment of fracture risk.

Estimating muscle attachment contours by transforming geometrical bone models.

For individualization of a biomechanical model, it is necessary to estimate the muscle attachments of the person to whom it is to be adapted. One of the methods to estimate muscle attachments is to use model transformations to transform a model with known muscle attachments to the bones of a person. We hypothesize that the location and shape of muscle attachment sites correlate with the shape of the bones they are attached to. If this hypothesis holds, it is possible to predict the location of muscle attachments when the shape of the bones is known. To validate this hypothesis, geometric models of three sets of shoulder bones were built. These models consist of 3-D surface models of the scapula, clavicle, and humerus, with the muscle attachment contours connected to them. By means of geometric transformations, the models were transformed, so the muscle attachments of the different data sets could be compared. Using these techniques, 50 per cent of the muscle attachment contours could be predicted with high accuracy. The muscle attachment contours that could not be predicted were all influenced by measurement errors. For 30 per cent of the muscle attachment contours, it was not possible to distinguish the interindividual differences from the inaccuracies of the method used. From this study, we concluded that most muscle attachment contours can be predicted by means of geometric models of the bones.