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Primate shoulder morphology has been linked with locomotor habits, oftentimes irrespective of phylogenetic heritage. Among hominoids, juvenile African apes are known to climb more frequently than adults, while orangutans and gibbons maintain an arboreal lifestyle throughout ontogeny. This study examined if these ontogenetic locomotor differences carry a morphological signal, which should be evident in the scapulae of chimpanzees and gorillas but absent in taxa that do not display ontogenetic behavioral shifts. The scapular morphology of five hominoid primates and one catarrhine outgroup was examined throughout ontogeny to evaluate if scapular traits linked with arboreal activities are modified in response to ontogenetic behavioral shifts away from climbing. Specifically, the following questions were addressed: 1) which scapular characteristics distinguish taxa with different locomotor habits; and 2) do these traits show associated changes during development in taxa known to modify their behavioral patterns? Several traits characterized suspensory taxa from nonsuspensory forms, such as cranially oriented glenohumeral joints, obliquely oriented scapular spines, relatively narrow infraspinous fossae, and inferolaterally expanded subscapularis fossae. The relative shape of the dorsal scapular fossae changed in Pan, Gorilla, and also Macaca in line with predictions based on reported ontogenetic changes in locomotor behavior. These morphological changes were mostly distinct from those seen in Pongo, Hylobates, and Homo and imply a unique developmental pattern, possibly related to ontogenetic locomotor shifts. Accordingly, features that sorted taxa by locomotor habits and changed in concert with ontogenetic behavioral patterns should be particularly useful for reconstructing the locomotor habits of fossil forms. Am J Phys Anthropol 152:239–260, 2013. © 2013 Wiley Periodicals, Inc.  相似文献   

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Objectives

Several studies have investigated potential functional signals in the trabecular structure of the primate proximal humerus but with varied success. Here, we apply for the first time a “whole‐epiphyses” approach to analysing trabecular bone in the humeral head with the aim of providing a more holistic interpretation of trabecular variation in relation to habitual locomotor or manipulative behaviors in several extant primates and Australopithecus africanus.

Materials and methods

We use a “whole‐epiphysis” methodology in comparison to the traditional volume of interest (VOI) approach to investigate variation in trabecular structure and joint loading in the proximal humerus of extant hominoids, Ateles and A. africanus (StW 328).

Results

There are important differences in the quantification of trabecular parameters using a “whole‐epiphysis” versus a VOI‐based approach. Variation in trabecular structure across knuckle‐walking African apes, suspensory taxa, and modern humans was generally consistent with predictions of load magnitude and inferred joint posture during habitual behaviors. Higher relative trabecular bone volume and more isotropic trabeculae in StW 328 suggest A. africanus may have still used its forelimbs for arboreal locomotion.

Discussion

A whole‐epiphysis approach to analysing trabecular structure of the proximal humerus can help distinguish functional signals of joint loading across extant primates and can provide novel insight into habitual behaviors of fossil hominins.
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Observational data were collected on the positional behavior of habituated adult female orangutans in the rain forest of the Kutai National Park, East Kalimantan, Indonesia. Results revealed the following about locomotion during travel: movement was concentrated in the understory and lower main canopy; and brachiation (without grasping by the feet) accounted for 11% of travel distance, quadrupedalism for 12%, vertical climbing for 18%, tree-swaying for 7%, and clambering for 51%. In climbing and clambering, the animal was orthograde and employed forelimb suspension with a mixture of hindlimb suspension and support. Thus suspension by the forelimbs occurred in at least 80% of travel. Locomotion in feeding trees resembled that during travel but with more climbing and less brachiation. Feeding was distributed more evenly among canopy levels than was travel, and use of postures (by time) included sitting 50%, suspension with the body vertical 11%, and suspension by hand and foot with the body horizontal 36%. The traditional explanation of the evolution of the distinctive hominoid postcranium stresses brachiation. More recently it has been proposed that climbing, broadly defined and partly equivalent to clambering in this study, is the most significant behavior selecting for morphology. The biomechanical similarity of brachiation and the orthograde clambering of orangutans precludes the present results from resolving the issue for the evolution of Pongo. The orangutan is by far the largest mammal that travels in forest canopy, and a consideration of the ways that its positional behavior solves problems posed by habitat structure, particularly the tapering of branches and gaps between trees, indicates that suspensory capacities have been essential in permitting the evolution and maintenance of its great body size.  相似文献   

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Lever systems within a skeleton transmit force with a capacity determined by the mechanical advantage, A. A is the distance from input force to a joint, divided by the distance from the joint to the output force. A lever with a relatively high A in static equilibrium has a great capacity to generate force but moves a load over a small distance. Therefore, the geometry of a skeletal lever presents a trade-off between force and speed under quasi-static conditions. The present study considers skeletal dynamics that do not assume static equilibrium by modelling kicking by a locust leg, which is powered by stored elastic energy. This model predicts that the output force of this lever is proportional to A, but its maximum speed is independent of A. Therefore, no trade-off between force and velocity exists in a lever system with spring-mass dynamics. This demonstrates that the motion of a skeleton depends on the major forces that govern its dynamics and cannot be inferred from skeletal geometry alone.  相似文献   

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Contiguous high resolution computed tomography images were obtained at a 1.5 mm slice thickness perpendicular to the neck axis from the base of the femoral head to the trochanteric line in a sample of 10 specimens each of Homo sapiens, Pan troglodytes, and Gorilla gorilla, plus five specimens of Pan paniscus. Superior, inferior, anterior, and posterior cortical thicknesses were automatically measured directly from these digital images. Throughout the femoral neck H. sapiens displays thin superior cortical bone and inferior cortical bone that thickens distally. In marked contrast, cortical bone in the femoral neck of African apes is more uniformly thick in all directions, with even greater thickening of the superior cortical bone distally. Because the femoral neck acts as a cantilevered beam, its anchorage at the neck-shaft junction is subjected to the highest bending stresses and is the most biomechanically relevant region to inspect for response to strain. As evinced by A.L. 128-1, A.L. 211-1 and MAK-VP-1/1, Australopithecus afarensis is indistinguishable from H. sapiens, but markedly different from African apes in cortical bone distribution at the femoral neck-shaft junction. Cortical distribution in the African ape indicates much greater variation in loading conditions consistent with their more varied locomotor repertoire. Cortical distribution in hominids is a response to the more stereotypic loading pattern imposed by habitual bipedality, and thin superior cortex in A. afarensis confirms the absence of a significant arboreal component in its locomotor repertoire. Am J Phys Anthropol 104:117–131, 1997. © 1997 Wiley-Liss, Inc.  相似文献   

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The relationship between locomotor behavior and long bone structural proportions is examined in 179 individuals and 13 species of hominoids and cercopithecoids. Articular surface areas, estimated from linear caliper measurements, and diaphyseal section moduli (strengths), determined from CT scans, were obtained for the femur, tibia, humerus, radius, and ulna. Both within-bone (articular to shaft) and between-bone (forelimb to hindlimb) proportions were calculated and compared between taxa. It was hypothesized that: 1) species emphasizing slow, cautious movement and/or more varied limb positioning (i.e., greater joint excursion) would exhibit larger articular to cross-sectional shaft proportions, and 2) species with more forelimb suspensory behavior would have relatively stronger/larger forelimbs, while those with more leaping would have relatively stronger/larger hindlimbs. The results of the analysis generally confirm both hypotheses. Several partial exceptions can be explained on the basis of more detailed structural-functional considerations. Associations between locomotion and structural proportions can be demonstrated both across major groupings (hominoids and cercopithecoids) and between relatively closely related taxa, e.g., mountain and lowland gorillas, siamangs and gibbons, and Trachypithecus and other colobines. Furthermore, structure and function do not always covary with taxonomy. For example, compared to cercopithecoids, mountain gorillas have relatively larger joints, like other hominoids, but do not have relatively stronger forelimbs, unlike other hominoids. This is consistent with a locomotor repertoire emphasizing relatively slow movement but with very little forelimb suspension. Proportions of Proconsul nyanzae, Proconsul heseloni, Morotopithecus bishopi, and Theropithecus oswaldi are compared with modern distributions to illustrate the application of the techniques to fossil taxa.  相似文献   

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Abstract:  For some decades, a major focus of research has been on how locomotor modes changed in some archosaurian reptiles from a more or less 'sprawling' to an 'erect' posture, whether there were discrete intermediate stages, and how many times 'erect' posture evolved. The classic paradigm for the evolution of stance and gait in archosaurs, a three-stage transition from sprawling to 'semi-erect' to erect posture, has been replaced by a subtler understanding of a continuum of changing limb joint angles. We suggest a further separation of terminology related to stance vs. gait so as not to entail different processes: 'sprawling' and 'erect' should refer to continua of stance; 'rotatory' and 'parasagittal' are more appropriate ends of a continuum that describes the motions of gait. We show that the Triassic trackway Apatopus best fits the anatomy and proportions of phytosaurs, based on a new reconstruction of their foot skeleton; it is less likely to have been made by another pseudosuchian or non-archosaurian archosauromorph. Moreover, the trackmaker was performing the high walk. A phytosaurian trackmaker would imply that the common ancestor of pseudosuchians, and therefore archosaurs could approximate the high walk (depending on phylogeny), and if so, erect stance and parasagittal gait did not evolve independently in pseudosuchians and ornithosuchians, although the kinematic mechanisms differed in the two groups. It remains to be seen how far outside Archosauria, if at all, more or less erect posture and parasagittal gait may have evolved.  相似文献   

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Differences in scapular morphology between modern humans and the African and lesser apes are associated with the distinct locomotor habits of these groups. However, several traits, particularly aspects of the supraspinous fossa, are convergent between Homo and Pongo—an unexpected result given their divergent locomotor habits. Many morphological assessments of the scapula rely on the limited number of static landmarks available, and traditional approaches like these tend to oversimplify scapular shape. Here, we present the results of two geometric morphometric (GM) analyses of hominoid supraspinous fossa shape—one employing five homologous landmarks and another with 83 sliding semilandmarks—alongside those of traditional methods to evaluate if three-dimensional considerations of fossa shape afford more comprehensive insights into scapular shape and functional morphology. Traditional measures aligned Pongo and Homo with narrow and transversely oriented supraspinous fossae, whereas African ape and Hylobates fossae are broader and more obliquely situated. However, our GM results highlight that much of the convergence between Homo and Pongo is reflective of their more medially positioned superior angles. These approaches offered a more complete assessment of supraspinous shape and revealed that the Homo fossa, with an intermediate superior angle position and moderate superoinferior expansion, is actually reminiscent of the African ape shape. Additionally, both Pongo and Hylobates were shown to have more compressed fossae, something that has not previously been identified through traditional analyses. Thus, the total morphological pattern of the Pongo supraspinous fossa is unique among hominoids, and possibly indicative of its distinctive locomotor habits. Am J Phys Anthropol 156:498–510, 2015. © 2015 Wiley Periodicals, Inc.  相似文献   

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Previous research on subfossil lemurs has revealed much about the positional behavior of these extinct strepsirrhines, but a thorough quantitative analysis of their vertebral form and function has not been performed. In this study, 156 lumbar vertebrae of Pachylemur, Archaeolemur, Megaladapis, Mesopropithecus, Babakotia, and Palaeopropithecus (11 species in all) were compared to those of 26 species of extant strepsirrhines and haplorhines. Lumbar shape was compared among species, using a principal components analysis (PCA) in conjunction with selected vertebral indices. The first principal component revealed strong separation between Palaeopropithecus at one extreme, and Archaeolemur/Pachylemur at the other, with Babakotia, Mesopropithecus, and Megaladapis in an intermediate position. Palaeopropithecus has markedly shorter spinous processes and wider laminae than do the other subfossil taxa, consistent with sloth-like, inverted suspensory postures. The moderately reduced lumbar spinous processes of Babakotia, Mesopropithecus, and Megaladapis are convergent with those of lorisids and Pongo, reflecting antipronogrady, but a less specialized adaptation than that of Palaeopropithecus. Archaeolemur and Pachylemur share relatively elongated spinous processes, in conjunction with other features (e.g., transverse process orientation and relatively short vertebral bodies) indicative of pronograde, quadrupedal locomotion characterized by reduced agility. All subfossil taxa exhibit adaptations emphasizing lumbar spinal stability (e.g., relatively short vertebral bodies, and transverse processes that are not oriented ventrally); we believe this probably reflects convergent mechanical demands connected to large body size, irrespective of specific locomotor mode. Reconstructions of positional behavior in subfossil lemurs based on lumbar vertebrae are largely consistent with those based on other aspects of the postcrania.  相似文献   

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The relationship between form and function in the lumbar vertebral column has been well documented among platyrrhines and especially catarrhines, while functional studies of postcranial morphology among strepsirrhines have concentrated predominantly on the limbs. This morphometric study investigates biomechanically relevant attributes of the lumbar vertebral morphology of 20 species of extant strepsirrhines. With this extensive sample, our goal is to address the influence of positional behavior on lumbar vertebral form while also assessing the effects of body size and phylogenetic history. The results reveal distinctions in lumbar vertebral morphology among strepsirrhines in functional association with their habitual postures and primary locomotor behaviors. In general, strepsirrhines that emphasize pronograde posture and quadrupedal locomotion combined with leaping (from a pronograde position) have the relatively longest lumbar regions and lumbar vertebral bodies, features promoting sagittal spinal flexibility. Indrids and galagonids that rely primarily on vertical clinging and leaping with orthograde posture share a relatively short (i.e., stable and resistant to bending) lumbar region, although the length of individual lumbar vertebral bodies varies phylogenetically and possibly allometrically. The other two vertical clingers and leapers, Hapalemur and Lepilemur, more closely resemble the pronograde, quadrupedal taxa. The specialized, suspensory lorids have relatively short lumbar regions as well, but the lengths of their lumbar regions are influenced by body size, and Arctocebus has dramatically longer vertebral bodies than do the other lorids. Lumbar morphology among galagonids appears to reflect a strong phylogenetic signal superimposed on a functional one. In general, relative length of the spinous processes follows a positively allometric trend, although lorids (especially the larger-bodied forms) have relatively short spinous processes for their body size, in accordance with their positional repertoire. The results of the study broaden our understanding of postcranial adaptation in primates, while providing an extensive comparative database for interpreting vertebral morphology in fossil primates.  相似文献   

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During posthatching development the fins of fishes undergo striking changes in both structure and function. In this article we examine the development of the pectoral fins from larval through adult life history stages in the zebrafish (Danio rerio), describing in detail their pectoral muscle morphology. We explore the development of muscle structure as a way to interpret the fins' role in locomotion. Genetic approaches in the zebrafish model are providing new tools for examining fin development and we take advantage of transgenic lines in which fluorescent protein is expressed in specific tissues to perform detailed three-dimensional, in vivo fin imaging. The fin musculature of larval zebrafish is organized into two thin sheets of fibers, an abductor and adductor, one on each side of an endoskeletal disk. Through the juvenile stage the number of muscle fibers increases and muscle sheets cleave into distinct muscle subdivisions as fibers orient to the developing fin skeleton. By the end of the juvenile period the pectoral girdle and fin muscles have reoriented to take on the adult organization. We find that this change in morphology is associated with a switch of fin function from activity during axial locomotion in larvae to use in swim initiation and maneuvering in adults. The examination of pectoral fins of the zebrafish highlights the yet to be explored diversity of fin structure and function in subadult developmental stages. J. Morphol. (c) 2005 Wiley-Liss, Inc.  相似文献   

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Divergent morphologies among related species are often correlated with distinct behaviors and habitat uses. Considerable morphological and behavioral differences are found between two major clades within the polychaete family Opheliidae. For instance, Thoracophelia mucronata burrows by peristalsis, whereas Armandia brevis exhibits undulatory burrowing. We investigate the anatomical differences that allow for these distinct burrowing behaviors, then interpret these differences in an evolutionary context using broader phylogenetic (DNA‐based) and morphological analyses of Opheliidae and taxa, such as Scalibregmatidae and Polygordiidae. Histological three‐dimensional‐reconstruction of A. brevis reveals bilateral longitudinal muscle bands as the prominent musculature of the body. Circular muscles are absent; instead oblique muscles act with unilateral contraction of longitudinal muscles to bend the body during undulation. The angle of helical fibers in the cuticle is consistent with the fibers supporting turgidity of the body rather than resisting radial expansion from longitudinal muscle contraction. Circular muscles are present in the anterior of T. mucronata, and they branch away from the body wall to form oblique muscles. Helical fibers in the cuticle are more axially oriented than those in undulatory burrowers, facilitating radial expansion during peristalsis. A transition in musculature accompanies the change in external morphology from the thorax to the abdomen, which has oblique muscles similar to A. brevis. Muscles in the muscular septum, which extends posteriorly to form the injector organ, act in synchrony with the body wall musculature during peristalsis: they contract to push fluid anteriorly and expand the head region following a direct peristaltic wave of the body wall muscles. The septum of A. brevis is much thinner and is presumably used for eversion of a nonmuscular pharynx. Mapping of morphological characters onto the molecular‐based phylogeny shows close links between musculature and behavior, but less correlation with habitat. J. Morphol. 275:548–571, 2014. © 2014 Wiley Periodicals, Inc.  相似文献   

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