The architecture of the shoulder in some mammals

A series of nine features of the shoulder girdle, chosen as having functional significance in relation to the movements of the shoulder in arboreal locomotion, have been studied in 1188 specimens of 194 genera of mammals. The features were defined metrically and examined by means of a multivariate statistical technique: viz. canonical analysis. The study has shown that those mammals which are nonarboreal differ considerably among themselves and form the arboreal forms. But the myriad shapes of the shoulder girdle in a wide range of mammals (e.g. some marsupials, edentates, rodents, carnivores and primates) which climb or forage in trees, can be summarized mathematically by a very small number of similar canonical variates. This information correlates well with that of a previous series of studies carried out on the primates alone. The biological information that was postulated as being reflected by the individual canonical variates for the primates is also apparent for the arboreal mammals. The different variates separate the forms in ways which are consonant with what is known about the function of the shoulder in locomotion. Aspects of the shape of the shoulder defined by the analysis appear to be discernible from an examination of the contribution of the original variables to each individual canonical variate. This seems to confirm that the shape of the shoulder girdle within a very wide range of mammals is limited by a very small number of underlying factors of biological significance. One interpretation of the results suggests that the genetic model of the mammalian shoulder may have been sufficiently fixed at an early stage in the evolution of the class as to place considerable constraints upon the subsequent evolution of the shoulder in the different Orders.     

Glenohumeral mobility in primates

This study refutes the traditional idea that the glenohumeral joint of hominoids is more mobile than that of other primates, a belief that forms a basis for the two prominent theories of hominoid evolution. According to the brachiation theory, many anatomical features of the hominoid shoulder (including those of the glenohumeral joint) increase shoulder mobility and are interpreted as adaptations for brachiation. The slow climbing theory explains the same set of features as adaptations for slow climbing. The slow-climbing primates should therefore also possess these features, and their glenohumeral mobility should be the same as that of hominoids and be higher than that of other primates. This study presents three-dimensional glenohumeral mobility data, measured using a single video camera method on fresh specimens. The results show that the hominoid glenohumeral joint is actually less mobile than those of non-hominoid primates, including the habitually slow-climbing lorines, but it is characterized by a smooth excursion in the scapulocranial direction.     

Morphometric affinities of the human shoulder

To analyze differences between apes and monkeys and the affinities of man, we have studied the shoulder girdle of 327 specimens of anthropoid primates. The scapula, clavicle and humerus are viewed as an integrated functional complex on the basis of 18 measurements. Several varieties of multivariate analysis show that man is clearly closer to other hominoids than to the included monkey taxa (whether terrestrial or arboreal, Old World or New World). The marked shoulder differences between apes and monkeys and similarities between apes and man correlate with the muscular anatomy, which in hominoids allows the motions involved in their locomotion and feeding behavior. As the hominid-pongid correspondence in shoulder morphology is especially detailed regarding the functionally important joint surfaces, it is consistent with a fairly recent period of common ancestry and behavior. No hypothetical evolutionary pathway or ancestral form of the human shoulder need look far beyond the model afforded by extant pongids. In contrast with previous studies on the primate shoulder, these results agree with information accumulating from other systems—comparative anatomy, primate behavior, and molecular biology — in suggesting very close relationship between man and extant African pongids.     

The form of the talus in some higher primates: a multivariate study

Sixteen measurements of the talus have been taken on 334 tali of a total of eleven primate groups and several additional single individual specimens. Multivariate morphometric (canonical and generalized distance) analyses of these data in extant higher primates are presented and used to define the relative morphological positions of fossils of the genera Proconsul and Limnopithecus, of individual specimens from Kromdraai, Olduvai and Kiik-Koba (Homo neanderthalensis), and a group of specimens of Bronze Age man from Jericho. Following preliminary studies the ultimate analysis suggests that the various extant arboreal primates examined fall within an envelope that is defined by Macaca together with various other Old World monkeys and extending in different directions to the extreme genera (a) Pongo, (b) Hylobates and (c) Ateles. This separation is thus one which defines generally quadrupedal monkeys and separates the various extreme arboreal locomotor modes of (a) acrobatic climbing and hanging, (b) richochetal brachiation and (c) prehensile-tailed arm-swinging and hanging, respectively. Man and the African apes are well separated both from each other and from this spectrum of arboreally adapted genera. Bronze Age man from Jericho and Neandertal man from Kiik-Koba lie relatively close to the position for modern man although significantly separated from him. Limnopithecus, Proconsul, and the specimens from Kromdraai and Olduvai all lie within the envelope of arboreal species and specifically rather close to, although significantly different from, the orang-utan; they differ markedly from both man and the African apes. The possibility exists that the resemblances of Proconsul and Limnopithecus relate to arboreal habitus in these species. The findings for the specimens from Kromdraai and Olduvai suggest either that the morphological resemblances to arboreal forms relate to a previous arboreal history for these species, or that bipedality is much less advanced or uniquely different from that displayed by Homo. It is not inconceivable that both conditions might apply.     

Functional aspects of primate pelvic structure: A multivariate approach

This study aims to clarify the relationship of primate bony pelvic structure to locomotor habit. As with most of the postcranial skeleton, the pelvic bones of species within the Ceboidea and the Cercopithecoidea are remarkably similar visually except for variations in size. Yet there are substantial differences in locomotor pattern between the species in these taxa. I performed canonical analyses on a sample of 17 pelvic variables describing 22 primate species of the Ceboidea, the Cercopithecoidea, and the Hominoidea to discover which variables were significant in separating them into groups. In both analyses there was good separation of major taxa and additional separation of groups that differed in locomotor habit. The separation of colobine from cercopithecine monkeys was particularly consistent. In the analysis, including all 22 species, the variables given particular weight by the canonical analysis were the same as those traditionally used by anatomists for the same purpose. Specifically, breadth of the ischial tuberosity (reflecting presence or absence of ischial callosities) separated the Old from the New World monkeys. Breadth of the iliac tuberosity, in which man and to some extent other hominoids differ from other primates, and ilium height, in which man differs from other primates, were significant. Sagittal diameter of the pelvis was also substantially weighted. Having established that the technique would select variables of anatomical significance, the same method was applied to a study of monkeys only where the characteristics that differ between groups are not well established. Breadth of the ischial tuberosity was again important in separating the Ceboidea from the Cercopithecoidea. Discrimination of locomotor groups within these large divisions was brought about mainly by ischial length and the sagittal diameter of the pelvis. In studying these variables and their relationship to size in greater detail, it was found that among cercopithecoid monkeys, the colobines showed relatively lower values than did cercopithecines for both these dimensions. Atelines showed low values for ischial length but high values for the sagittal pelvic diameter. Biomechanical explanations of these observations are suggested.     

Muscle mass scaling in primates: an energetic and ecological perspective

Body composition is known to vary dramatically among mammals, even in closely related species, yet this issue has never been systematically investigated. Here, we examine differences in muscle mass scaling among mammals, and explore how primate body composition compares to that of nonprimate mammals. We use a literature-based sample of eutherian and metatherian mammals, and combine this with new dissection-based data on muscularity in a variety of strepsirrhine primates and the haplorhine, Tarsius syrichta. Our results indicate an isometric scaling relationship between total muscle mass and total body mass across mammals. However, we documented substantial variation in muscularity in mammals (21-61% of total body mass), which can be seen both within and between taxonomic groups. We also found that primates are under-muscled when compared to other mammals. This difference in body composition may in part reflect the functional consequences of arboreality, as arboreal species have significantly lower levels of muscularity than terrestrial species.     

Kinematic and EMG analysis of horizontal bimanual climbing in humans

Climbing is an increasingly popular recreational and competitive behavior, engaged in a variety of environments and styles. However, injury rates are high in climbing populations, especially in the upper extremity and shoulder. Despite likely arising from an arboreal, climbing ancestor and being closely related to primates that are highly proficient climbers, the modern human shoulder has devolved a capacity for climbing. Limited biomechanical research exists on manual climbing performance. This study assessed kinematic and muscular demands during a bimanual climbing task that mimicked previous work on climbing primates. Thirty participants were recruited – 15 experienced and 15 inexperienced climbers. Motion capture and electromyography (EMG) measured elbow, thoracohumeral and trunk angles, and activity of twelve shoulder muscles, respectively, of the right-side while participants traversed across a horizontal climbing apparatus. Statistical parametric mapping was used to detect differences between groups in kinematics and muscle activity. Experienced climbers presented different joint motions that more closely mimicked the kinematics of climbing primates, including more elbow flexion (p = 0.0045) and internal rotation (p = 0.021), and less thoracohumeral elevation (p = 0.046). Similarly, like climbing primates, experienced climbers generally activated the shoulder musculature at a lower percentage of maximum, particularly during the exchange from support to swing and swing to support phase. However, high muscle activity was recorded in all muscles in both participant groups. Climbing experience coincided with a positive training effect, but not enough to overcome the high muscular workload of bimanual climbing. Owing to the evolved primary usage of the upper extremity for low-force, below shoulder-height tasks, bimanual climbing may induce high risk of fatigue-related musculoskeletal disorders.     

Morphometric correlates of karyotype and host plant in genus Euceraphis (Hemiptera: Aphididae)

Abstract Samples of birch- and alder-feeding aphids of genus Euceraphis were measured and analysed using multiple discriminant analysis (canonical variates) to find out if morphological variation correlated with previously reported differences of karyotype and host association. The dataset comprised groups of specimens defined solely by locality and collection date. Mean scores on the first two canonical variates clustered the samples fully in accordance with their karyotypes and host plants, confirming the existence of a number of morphologically similar but distinct host-specific taxa within the E. betulae group. Three new species are described: Euceraphis borealis Blackman, sp.n. on Betula glandulosa and B. nana , Euceraphis papyrifericola Blackman, sp.n. on B. papyrifera , and Euceraphis quednaui Blackman, sp.n. on B. occidentalis . A key is provided to alate viviparae of the genus.     

The functional morphology of the primate shoulder as revealed by comparative anatomical,osteometric and discriminant function techniques

A background to assessments of the ecological adaptations of fossil primates is the relationship of the detailed shape of bones in living forms to their known locomotor patterns, and this has been here attempted for the shoulder. Within the locomotion of the primates the function of the shoulder varies according to the extent to which the trunk is suspended by the arms. An analysis of the differences in the shoulder muscles has shown that much of their quantitative variation is mechanically in phase with these functional differences. A series of features of the shoulder bones, chosen because of their association with the mechanically meaningful features of the musculature, have been found to vary (a) in association with the known contrasts in locomotion and (b) in such a way as to render more efficient mechanically the associated muscular structure. Investigation of bony dimensions “residual” to such a study has shown that they are not highly correlated with primate locomotion but are, in contrast, associated with the commonly accepted taxonomic grouping of the order. The combination by discriminant functions of such sets of “locomotor” and “residual” dimensions reveals unsuspected information for living primates and might well allow more precise definition of the functional and taxonomic status of a fossil. The experimental testing of functional inferences from morphology is a necessary part of such studies, and preliminary reports of experimental stress analysis utilising the photoelastic technique confirm and reinforce their validity.     

Occurrence of human-pathogenic Enterocytozoon bieneusi,Giardia duodenalis and Cryptosporidium genotypes in laboratory macaques in Guangxi,China

Captive nonhuman primates have been identified as common hosts of Enterocytozoon bieneusi, Giardia duodenalis, Cryptosporidium hominis, and Cyclospora spp., thus are potential reservoirs of some enteric parasites in humans. However, few studies have examined the source and human-infective potential of enteric parasites in laboratory nonhuman primates. In the present work, 205 fecal specimens were collected from three groups of captive Macaca fascicularis kept in different densities in a laboratory animal facility in Guangxi, China, and examined by PCR for E. bieneusi, G. duodenalis, Cryptosporidium spp., and Cyclospora spp. The infection rates of E. bieneusi and G. duodenalis were 11.3% and 1.2% in Group 1 (young animals kept individually; n = 168), 72.2% and 11.1% in Group 2 (young animals kept in groups; n = 18), and 31.6% and 5.3% in Group 3 (adults kept in groups; n = 19), respectively. Sequence analysis of PCR products showed the presence of five E. bieneusi genotypes, with genotype D (in 16/36 genotyped specimens) and a new genotype (in 15/36 genotyped specimens) as the dominant genotypes. All five E. bieneusi genotypes belonged to the zoonotic group (Group 1). The G. duodenalis genotypes (assemblages AII and B) in five specimens and C. hominis subtype (IdA14) in one specimen were also known human-pathogens, although the Cyclospora seen in one animal appeared to be unique to macaque monkeys. The higher infection rate in younger animals reared in groups and common occurrence of zoonotic genotypes indicated that human-pathogenic E. bieneusi could be transmitted efficiently in captive nonhuman primates, and group-housing was a risk factor for transmission of zoonotic pathogens in young nonhuman primates in research facilities.     

Multivariate analysis of vegetational variation in different habitats at Omayed,Egypt

The application of indicator species analysis has proved useful in classifying stands into groupings coinciding with topographic variations. 41 indicator pseudo-species are identified in the vegetation along a phytosociological gradient: some are characteristic of specific habitats, and the others indicate transition between groups of habitats.The use of qualitative estimations (transformed density records) in multivariate analysis in the present study indicates that they may be more preferable than quantitative estimations, because of their easy and fast recording in the field, and the less comprehensive computations, while yielding precise results.The X-axis of the reciprocal averaging ordination is related to the salinity and fertility gradients, while the Y-axis reflects soil texture.The phytosociological gradients of the canonical variate axes reflect essentially contrasts between groups of species, each correlated with one or the other of the environmental gradients.Comparing the results obtained by the two ordination methods (reciprocal averaging and canonical variates) we find that the first method detects the overall phytosociological changes along strong environmental gradients which would be helpful in studying large surveys. On the other hand, the second technique, which is mainly predictive, is more sensitive to changes within both the phytosociological and environmental gradients and can detect the impact of these changes on the overall variance of each gradient.Nomenclature follows Täckholm (1964).The authors are grateful to Dr J. Jeffers and Mr and Mrs P. Howard of the Institute of Terrestrial Ecology, England, for their help with applying the multivariate techniques to the vegetation data.     

Expression of Myosin Heavy Chain Isoforms in the Supraspinatus Muscle of Different Primate Species: Implications for the Study of the Adaptation of Primate Shoulder Muscles to Different Locomotor Modes

The supraspinatus muscle is a key component of the soft tissues of the shoulder. In pronograde primates, its main function, in combination with the other rotator cuff muscles (subscapularis, infraspinatus, and teres minor), is to stabilize the glenohumeral joint, whereas in orthograde primates it functions together with the deltoid, to elevate the upper extremity in the scapular plane. To determine whether these functional differences are also reflected in the molecular biochemistry of the supraspinatus muscles involved in these different locomotor modes, we used real-time polymerase chain reaction (RT-PCR) to analyze the expression of the myosin heavy chain (MHC) isoforms in supraspinatus muscles from modern humans and 12 species of pronograde and orthograde primates. The MHC expression pattern in the supraspinatus muscle of pronograde primates was consistent with its function as a tonic and postural muscle, whereas the MHC expression pattern observed in the supraspinatus muscle of nonhuman orthograde primates was that of a muscle that emphasizes speed, strength, and less resistance to fatigue. These findings are consistent with the role of the supraspinatus in the posture and locomotor modes of these groups of nonhuman primates. The humans included in the study had an expression pattern similar to that of the nonhuman orthograde primates. In conclusion, molecular analysis of skeletal muscles via RT-PCR can contribute to a better understanding of the morphological and functional characteristics of the primate musculoskeletal system.     

A morphometric study of limb proportions in leaping prosimians

Study of individual limb proportions—most notably, for instance, of the intermembral index—distinguishes from most other primates those various prosimians capable of leaping. The present investigation analyzes 24 measures of limb proportions (taken by F.-K. Jouffroy and J. Lessertisseur) from 161 specimens representing 12 species of prosimians, using the multivariate statistical method of multiple discriminant analysis (carried out by R. German and C.E. Oxnard). The results indicate that there are three major gradients: (1) indriids, (2) cheirogaleines, and (3) galagines plus Tarsius, radiating from a fourth centrally located group of species, Lemur, Lepilemur, and Hapalemur. It is likely that the morphometric relationships displayed here can be associated with behavioral (locomotor) parallels among these various forms. Though leaping is a most important adaptation among prosimians, it seems clear that it has evolved separately a number of times, and it seems at least likely that the biomechanical nature of the adaptation differs in different groups. The results, therefore, present certain new hypotheses about prosimian locomotion that can only be tested with further field and laboratory studies. This study also emphasizes the fact that anatomical adaptation is complex. Though the analysis of each anatomical region separately provides a similar qualitative picture of the variation among genera, it is only in the analysis of all measures taken together that the complete pattern described above appears. Though no new variables have been added in that analysis, new information is obviously contained within the results. Since this information stems from the interactions of the variables, it must be due to associations between these anatomical areas.     

An evaluation of the systematic value of skull morphology in the Trimeresurus radiation (Serpentes: Viperidae: Crotalinae) of Asian pitvipers

Trimeresurus (in its widest sense) is a very diverse and widespread radiation of Asian pitvipers, which has been subject to numerous taxonomic revisions, some of which have been based on characteristics of the skull. In order to evaluate the taxonomic utility of such characters, we conducted a comparison of the skulls of 57 specimens representing nine genera, and two currently unassigned species that are very closely related to each other. A canonical variate analysis reveals three distinct phenetic groups: the Protobothrops group, a group containing Ovophis monticola , ' Ovophis ' okinavensis , and ' Trimeresurus ' gracilis , and finally a group comprising the remaining species and characterized by considerable overlap between most genera with the exception of the monotypic Peltopelor and Himalayophis . Agreement between phenetic similarity based on skull characteristics and phylogenetic relationships based on molecular evidence varies between different groups: the morphological similarity of the skull of Protobothrops sieversorum to the other Protobothrops species is congruent with their recent synonymization while the phenetic similarity among the species within the second group does not reflect current molecular phylogenetic relationships and indicates that convergent or parallel evolution may be responsible for at least some of the phenetic similarity detected among skulls of the Asian pit vipers examined. A test of phylogenetic independence, however, indicates that there is still a significant phylogenetic signal that can be recovered from several skull characteristics. Thus, we conclude that skull morphology can contribute to an overall understanding of pitviper taxonomy, but that it would be unwise to rely on skull characteristics alone.     

New characters for the functional interpretation of primate scapulae and proximal humeri

The study of muscle function in nonhuman primates through the technique of electromyography (EMG) has facilitated the identification of specific functional roles for muscles in particular behaviors. This has led to a more complete understanding of the biomechanics of certain regions of the musculoskeletal system, and should facilitate our ability to identify morphological features useful in the functional interpretation of fossil material. The current paper represents one such investigation of a new set of morphometric characters of the scapula and proximal humerus suggested by EMG analyses of shoulder muscle function. A set of new metric variables were examined on the scapulae and proximal humeri of 25 species of extant anthropoid primates, as well as on casts of scapulae and humeri of three fossil primate taxa. The variables are primarily related to the line of action and attachments of the rotator cuff muscles. The position of the scapular spine, the degree of lateral expansion of the subscapular fossa, the size and shape of the subscapularis insertion facet on the lesser tubercle, and the orientation of the infraspinatus insertion facet on the greater tubercle all appear to successfully sort the extant taxa into locomotor groups. Their appearance on the fossil specimens generally supports previous functional interpretations of each taxon's locomotor abilities based on a variety of other characters, suggesting that these traits are equally applicable to fossil material. © Wiley-Liss, Inc.     

Tempo and mode of evolution of a primate-specific retrotransposon belonging to the LINE 1 family

L1P_MA2 is a primate-specific subfamily of L1 retrotransposons. The consensus sequence of this element differs from the canonical L1 consensus by the presence of a 3800-bp region in 5' (L1M1_5). Part of this region has been proposed to be involved in a dystrophin mutation affecting the correct splicing of the gene and causing an X-linked dilated cardiomyopathy. In consideration of the potential involvement in splicing regulation of this element and also because of its atypical structure, we investigated its evolutionary history by analyzing the inter- and intraspecific divergence of L1P_MA2 sequences in various species of primates. The resulting phylogenetic trees show long terminal branches and short basal internodes, as expected for a rapid event of diversification that occurred in the past. The phylogenetic analysis and the intraspecific divergence estimates revealed a pattern of evolution for this element similar in all primates with the exception of lemurs, thus suggesting that the major wave of expansion of L1P_MA2 in primate genomes occurred after the divergence between Prosimiae and Anthropoidea. These results clearly indicate that a phylogenetic approach is more appropriate than methods based on sequence data from a single species, when investigating time and mode of evolution of retro-elements.     

Environmental factors influence on chemical polymorphism of the essential oils of Lychnophora ericoides

Lychnophora ericoides is a Brazilian medicinal plant used in folk medicine as an anti-inflammatory and analgesic agent. The essential oils from leaves of two populations with and without scent, collected at 2-month intervals during an 1-year period, were analysed by GC-MS. The results were submitted to principal component and cluster analysis which allowed two groups of essential oils to be distinguished with respect to sampling site and scent: cluster I (Vianópolis site, with specimens exhibiting an aromatic scent) containing a high percentage of alpha-bisabolol (44.7-76.4%) and alpha-cadinol (10.9-23.5%), and cluster II (Cristalina site, with specimens without scent) characterised by a high content of (E)-nerolidol (31.3-47.1%) and ar-dihydro-turmerone (4.8-15.4%). The canonical discriminant analysis showed that using the data set of the seven sampling months and (E)-nerolidol and alpha-bisabolol as predictable variables, it was possible to distinguish between the samples harvested according to Cerrado seasons, dry winter (May-September) and humid summer (November-March). In addition, canonical correlation analysis between the soil sampling sites and the populations revealed a significant relationship between oil components and edaphic factors. Oxygenated sesquiterpenes and potential acidity, Al saturation, cationic exchange capacity, silt, and sand load as the first canonical variate were fairly strongly related to samples collected in Vianópolis site. On the other hand, monoterpenes and sesquiterpene hydrocarbons were strongly related to chemical balance in soils (organic matter, P and base saturation), which is related to samples at the Cristalina site. The chemovariation observed appears to be environmentally determined.     

Predicting the shapes of bones at a joint: application to the shoulder

This paper presents a novel method to explore the intrinsic morphological correlation between the bones of a shoulder joint (humerus and scapula). To model this correlation, canonical correlation analysis (CCA) is used. We also propose a technique to predict a three-dimensional (3D) bone shape from its adjoining segment at a joint based on partial least squares regression (PLS). The high dimensional 3D surface information of a bone is represented by a few variables using principal component analysis, which also captures the pattern of variability of the shapes in our datasets. Our results show that the humerus set and scapula set have highly linear morphological relationship and that the correlation information can be used as a classifier. In this study, primate shoulder bone datasets were categorised into two clusters: great apes (including humans) and monkeys. A leave one out experiment was performed to test the robustness of this prediction method. The prediction behaviour using this method shows statistically significantly better results than using the mean shape from the training set.     

Canonical correlation analysis of marine macrobenthos survey data

Canonical correlation analysis is applied to measurements of environmental variables and species distributions made during a survey of macrobenthos around a sewage-treatment farm drain. The implications of data reduction, necessary to enable the method to proceed, are discussed. The amount of data was reduced by discarding the rarest species, discarding species occurring at fewest stations, and including only those species and environmental variables which correlated highly with the greatest number of other variables. Only the third data-reduction scheme gave ecologically sensible results. Use of station scores on the first two canonical variates (CV1 and CV2) enabled the sampling grid to be divided into a group of nearshore stations, a group of intermediate depth, and a group of deep offshore stations. Loadings of environmental variables on the canonical variates were found to be unstable but correlations between these variables and canonical variates enabled the variates to be interpreted: CV1 as a gradient of depth and associated changes in sediment characteristics, CV2 with depth- and nutrient-related components, and CV3 as patchiness in sediment characteristics different from that normally expected with depth. Use of correlations between species and canonical variates enables definition of two major species groups, one confined to nearshore environments and a second offshore. These groups (and their sub-groups) related well to groups defined previously by hierarchical classification. It is concluded that, with careful attention to the method of data reduction, canonical correlation analysis can be an effective tool in the analysis of marine benthic survey data.