The effects of locomotion on the structural characteristics of avian limb bones

Despite the wide range of locomotor adaptations in birds, little detailed attention has been given to the relationships between the quantitative structural characteristics of avian limb bones and bird behaviour. Possible differences in forelimb relative to hindlimb strength across species have been especially neglected. We generated cross‐sectional, geometric data from peripheral quantitative computed tomography scans of the humerus and femur of 127 avian skeletons, representing 15 species of extant birds in 13 families. The sample includes terrestrial runners, arboreal perchers, hindlimb‐propelled divers, forelimb‐propelled divers and dynamic soarers. The hindlimb‐propelled diving class includes a recently flightless island form. Our results demonstrate that locomotor dynamics can be differentiated in most cases based on cross‐sectional properties, and that structural proportions are often more informative than bone length proportions for determining behaviour and locomotion. Recently flightless forms, for example, are more easily distinguished using structural ratios than using length ratios. A proper phylogenetic context is important for correctly interpreting structural characteristics, especially for recently flightless forms. Some of the most extreme adaptations to mechanical loading are seen in aquatic forms. Penguins have forelimbs adapted to very high loads. Aquatic species differ from non‐aquatic species on the basis of relative cortical thickness. The combination of bone structural strength and relative cortical area of the humerus successfully differentiates all of our locomotor groups. The methods used in this study are highly applicable to fossil taxa, for which morphology is known but behaviour is not. The use of bone structural characteristics is particularly useful in palaeontology not only because it generates strong signals for many locomotor guilds, but also because analysing such traits does not require knowledge of body mass, which can be difficult to estimate reliably for fossil taxa. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 601–624.     

Biomechanics and allometric scaling in primate locomotion and morphology

Body size has a dominant influence on locomotor performance and the morphology of the locomotor apparatus. In locomotion under the influence of gravity, body mass acts as weight force and is a mechanical variable. Accordingly, the application of biomechanical principles and methods allows a functional understanding of scaling effects in locomotion. This is demonstrated here using leaping primates as an example. With increasing body size, the decreasing ratio of muscle force available for acceleration during takeoff to the body mass that has to be accelerated dictates both the movement pattern and the proportions of the hindlimbs. In an arm-swinging movement, the long, heavy arms of the large-bodied leapers are effectively used to gain additional momentum. A new perspective on decreasing size identifies the absolutely small acceleration distance and time available for propulsion as factors limiting leaping distance and extensively determining locomotor behavior and body proportions. As the mechanical constraints differ according to body size for a given mode of locomotion, a typological approach to morphology in relation to locomotor category is ruled out. Across locomotor categories, dynamic similarity (sensu Alexander) can be expected if the propulsive mechanisms as well as the selective pressures acting upon locomotion are the same.     

Influences of limb proportions and body size on locomotor kinematics in terrestrial primates and fossil hominins

During locomotion, mammalian limb postures are influenced by many factors including the animal's limb length and body mass. Polk (2002) compared the gait of similar-sized cercopithecine monkeys that differed limb proportions and found that longer-limbed monkeys usually adopt more extended joint postures than shorter-limbed monkeys in order to moderate their joint moments. Studies of primates as well as non-primate mammals that vary in body mass have demonstrated that larger animals use more extended limb postures than smaller animals. Such extended postures in larger animals increase the extensor muscle mechanical advantage and allow postures to be maintained with relatively less muscular effort (Polk, 2002; Biewener 1989). The results of these previous studies are used here to address two anthropological questions. The first concerns the postural effects of body mass and limb proportion differences between australopithecines and members of the genus Homo. That is, H. erectus and later hominins all have larger body mass and longer legs than australopithecines, and these anatomical differences suggest that Homo probably used more extended postures and probably required relatively less muscular force to resist gravity than the smaller and shorter-limbed australopithecines. The second question investigates how animals with similar size but different limb proportions differ in locomotor performance. The effects of limb proportions on gait are relevant to inferring postural and locomotor differences between Neanderthals and modern Homo sapiens which differ in their crural indices and relative limb length. This study demonstrates that primates with relatively long limbs achieve higher walking speeds while using lower stride frequencies and lower angular excursions than shorter-limbed monkeys, and these kinematic differences may allow longer-limbed taxa to locomote more efficiently than shorter-limbed species of similar mass. Such differences may also have characterized the gait of Homo sapiens in comparison to Neanderthals, but more experimental data on humans that vary in limb proportions are necessary in order to evaluate this question more thoroughly.     

The correspondence between proximal phalanx morphology and locomotion: implications for inferring the locomotor behavior of fossil catarrhines

Phalanges are considered to be highly informative in the reconstruction of extinct primate locomotor behavior since these skeletal elements directly interact with the substrate during locomotion. Variation in shaft curvature and relative phalangeal length has been linked to differences in the degree of suspension and overall arboreal locomotor activities. Building on previous work, this study investigated these two skeletal characters in a comparative context to analyze function, while taking evolutionary relationships into account. This study examined the correspondence between proportions of suspension and overall substrate usage observed in 17 extant taxa and included angle of curvature and relative phalangeal length. Predictive models based on these traits are reported. Published proportions of different locomotor behaviors were regressed against each phalangeal measurement and a size proxy. The relationship between each behavior and skeletal trait was investigated using ordinary least-squares, phylogenetic generalized least-squares (pGLS), and two pGLS transformation methods to determine the model of best-fit. Phalangeal curvature and relative length had significant positive relationships with both suspension and overall arboreal locomotion. Cross-validation analyses demonstrated that relative length and curvature provide accurate predictions of relative suspensory behavior and substrate usage in a range of extant species when used together in predictive models. These regression equations provide a refined method to assess the amount of suspensory and overall arboreal locomotion characterizing species in the catarrhine fossil record.     

Long bone articular and diaphyseal structure in old world monkeys and apes. I: locomotor effects

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.     

Semicircular canal system in early primates

Mammals with more rapid and agile locomotion have larger semicircular canals relative to body mass than species that move more slowly. Measurements of semicircular canals in extant mammals with known locomotor behaviours can provide a basis for testing hypotheses about locomotion in fossil primates that is independent of postcranial remains, and a means of reconstructing locomotor behaviour in species known only from cranial material. Semicircular canal radii were measured using ultra high resolution X-ray CT data for 9 stem primates (“plesiadapiforms”; n = 11), 7 adapoids (n = 12), 4 omomyoids (n = 5), and the possible omomyoid Rooneyia viejaensis (n = 1). These were compared with a modern sample (210 species including 91 primates) with known locomotor behaviours. The predicted locomotor agilities for extinct primates generally follow expectations based on known postcrania for those taxa. “Plesiadapiforms” and adapids have relatively small semicircular canals, suggesting they practiced less agile locomotion than other fossil primates in the sample, which is consistent with reconstructions of them as less specialized for leaping. The derived notharctid adapoids (excluding Cantius) and all omomyoids sampled have relatively larger semicircular canals, suggesting that they were more agile, with Microchoerus in particular being reconstructed as having had very jerky locomotion with relatively high magnitude accelerations of the head. Rooneyia viejaensis is reconstructed as having been similarly agile to omomyids and derived notharctid adapoids, which suggests that when postcranial material is found for this species it will exhibit features for some leaping behaviour, or for a locomotor mode requiring a similar degree of agility.     

Swimmers,Diggers, Climbers and More,a Study of Integration Across the Mustelids’ Locomotor Apparatus (Carnivora: Mustelidae)

Phenotypic integration, defined as the coordinated co-variation of parts of an organism can be an important constraint on phenotypic diversification. Functional factors, by having an heterogeneous impact across the animal body, may reinforce the integration of some parts while causing a perturbation of the integration among other parts. The integration across the locomotor apparatus should thus reflect to a certain extent the locomotor ecology of the animal. Using the mustelids as study group, we track changes in the patterns of co-variation in species belonging to four different locomotor ecologies (terrestrial, semi-arboreal, semi-fossorial, and semi-aquatic). Our results highlight the strong overall integration in mustelid long bones. The main shape changes associated with co-variations between skeletal elements are the bone robustness and proportions of the epiphyses. The pattern of co-variation is, however, only slightly impacted by allometry. Changes in co-variation between species mostly scale with phylogenetic divergence time, except for the (Mustela putorius, M. eversmanni, M. lutreola) clade which, despite a short divergence time, presents strong differences in co-variation. Co-variation patterns differ between locomotor ecologies, but few of these variations match the hypothesis of a reduction of integration due to functional specialization. This may reflect our lack of knowledge on the functional modules in species with locomotor ecologies that differ from terrestrial locomotion rather than invalidate our a priori hypotheses.     

Locomotion in ornithischian dinosaurs: an assessment using three‐dimensional computational modelling

Ornithischian dinosaurs were primitively bipedal with forelimbs modified for grasping, but quadrupedalism evolved in the clade on at least three occasions independently. Outside of Ornithischia, quadrupedality from bipedal ancestors has only evolved on two other occasions, making this one of the rarest locomotory transitions in tetrapod evolutionary history. The osteological and myological changes associated with these transitions have only recently been documented, and the biomechanical consequences of these changes remain to be examined. Here, we review previous approaches to understanding locomotion in extinct animals, which can be broadly split into form–function approaches using analogy based on extant animals, limb‐bone scaling, and computational approaches. We then carry out the first systematic attempt to quantify changes in locomotor muscle function in bipedal and quadrupedal ornithischian dinosaurs. Using three‐dimensional computational modelling of the major pelvic locomotor muscle moment arms, we examine similarities and differences among individual taxa, between quadrupedal and bipedal taxa, and among taxa representing the three major ornithischian lineages (Thyreophora, Ornithopoda, Marginocephalia). Our results suggest that the ceratopsid Chasmosaurus and the ornithopod Hypsilophodon have relatively low moment arms for most muscles and most functions, perhaps suggesting poor locomotor performance in these taxa. Quadrupeds have higher abductor moment arms than bipeds, which we suggest is due to the overall wider bodies of the quadrupeds modelled. A peak in extensor moment arms at more extended hip angles and lower medial rotator moment arms in quadrupeds than in bipeds may be due to a more columnar hindlimb and loss of medial rotation as a form of lateral limb support in quadrupeds. We are not able to identify trends in moment arm evolution across Ornithischia as a whole, suggesting that the bipedal ancestry of ornithischians did not constrain the development of quadrupedal locomotion via a limited number of functional pathways. Functional anatomy appears to have had a greater effect on moment arms than phylogeny, and the differences identified between individual taxa and individual clades may relate to differences in locomotor performance required for living in different environments or for clade‐specific behaviours.     

Frog tendon structure and its relationship with locomotor modes

Tendon collagen fibrils are the basic force‐transmitting units of the tendon. Yet, surprisingly little is known about the diversity in tendon anatomy and ultrastructure, and the possible relationships between this diversity and locomotor modes utilized. Our main objectives were to investigate: (a) the ultra‐structural anatomy of the tendons in the digits of frogs; (b) the diversity of collagen fibril diameters across frogs with different locomotor modes; (c) the relationship between morphology, as expressed by the morphology of collagen fibrils and tendons, and locomotor modes. To assess the relationship between morphology and the locomotor modes of the sampled taxa we performed a principal component analysis considering body length, fibrillar cross sectional area (CSA) and tendon CSA. A MANOVA showed that differences between species with different locomotor modes were significant with collagen fibril diameter being the discriminating factor. Overall, our data related the greatest collagen fibril diameter to the most demanding locomotor modes, conversely, the smallest collagen fibril CSA and the highest tendon CSA were observed in animals showing a hopping locomotion requiring likely little absorption of landing forces given the short jump distances.     

The Calcaneum—On the Heels of Marsupial Locomotion

The potential for making functional interpretations from a single postcranial element for marsupials was investigated through morphometric analysis of the calcanea of 61 extant species from Australia and New Guinea. Extant species were grouped into locomotor categories and a canonical variates analysis was carried out on measurements of their calcanea. A relationship between measurements of the calcanea and the locomotor behavior of species was found, allowing for prediction of locomotor behavior from calcaneum morphometrics. This was applied to fossil marsupial taxa, from early–late Miocene/?Pliocene deposits at Riversleigh, in an attempt to determine their locomotor behavior. Hopping (saltatorial) taxa are distinguished from quadruped terrestrial taxa and taxa capable of climbing (arboreal and scansorial) by their relatively longer tuber calcis and wider calcaneal head, by their dorso-ventrally thicker calcaneal head, and by their calcaneocuboid facet being less steeply angled antero-posteriorly. Taxa capable of climbing are distinguished from quadruped terrestrial taxa by their shorter tuber calcis relative to the calcaneal head and by their smaller calcaneo-astragalar facet. The locomotor categories distinguished in this study (arboreal/scansorial, quadruped terrestrial, and saltatorial) highlight differences between species in their use of available substrates and thus are informative with regards to the structural components of their habitat. The results of this analysis can be used, in combination with other data, to make inferences about the habitats of paleocommunities at Riversleigh through the Miocene. The calcaneum is a dense and very robust element and, therefore, has a good chance of being preserved. This method provides a quick and easy way of inferring locomotion and has a wide potential for application to many fossil deposits because it requires only a single element.     

Neither season nor sex affects the cost of terrestrial locomotion in a circumpolar diving duck: the common eider (Somateria mollissima)

Locomotion accounts for a significant proportion of the energy budget in birds, and selection is likely to act on its economy, particularly where energy conservation is essential for survival. Birds are capable of different forms of locomotion, such as walking/running, swimming, diving and flying, and adaptations for these affect the energetic cost [cost of locomotion (CoL)] and kinematics of terrestrial locomotion. Furthermore, seasonal changes in climate and photoperiod elicit physiological and behavioural adaptations for survival and reproduction, which also influence energy budget. However, little is understood about how this might affect the CoL. Birds are also known to exhibit sex differences in size, behaviour and physiology; however, sex differences in terrestrial locomotion have only been studied in two cursorially adapted galliform species in which males achieved higher maximum speeds, and in one case had a lower mass-specific CoL than females. Here, using respirometry and high-speed video recordings, we sought to determine whether season and sex would affect the CoL and kinematics of a principally aquatic diving bird: the circumpolar common eider (Somateria mollissima). We demonstrate that eiders are only capable of a walking gait and exhibit no seasonal or sex differences in mass-specific CoL or maximum speed. Despite sharing identical limb morphometrics, the birds exhibited subtle sex differences in kinematic parameters linked to the greater body mass of the males. We suggest that their principally aquatic lifestyle accounts for the observed patterns in their locomotor performance. Furthermore, sex differences in the CoL may only be found in birds in which terrestrial locomotion directly influences male reproductive success.     

Comparative aspects of gait, scaling and mechanics in mammals.

In phylogenetically based systematics, Mammalia is the nomenclatural term which designates the clade stemming from the most recent common ancestry of monotremes and theria [, Sys. Biol. 43 (1994) 497]. Considering that locomotor performance is a prevalent function to provide the necessary conditions to survive and transmit genes, it may be questioned if the diverse types of locomotion exhibited by extant mammals could have played a role in their evolution, or have only followed it. We may look after the structural and behavioural features which are involved in mammal locomotion compared to other tetrapods and test if they fit with the proposed phylogeny. Several factors may be checked: scaling effect in relation to gravitational constraints; geometrical distribution of masses in the body, and relative mechanical role of the limbs in the production of the external forces necessary to forward motion. Classically, it was thought that the fastest gaits used by terrestrial mammals were based upon a unique kind of limb motion co-ordination, called asymmetrical gaits, which in turn may be thought to be related to a peculiar neuronal wiring. Kinematic analysis brings an insight to this topic. Is the search for an ancestral mammalian locomotor pattern judicious? Notice the small size of many of the first mammals and their probable locomotor plasticity. (relation between grain size of the elements within the substrate and the organism scale). At a small size, the gravitational constraint is less important, and the distinction between terrestrial and arboreal has probably no sense when the limbs are the principal motor elements. There remains the importance of the geometrical distribution of body elements, the proportions of the limbs and of the head-neck complex, the tail merely as an appendix, a set of factors which may have generated the frame of constraints within which diverse locomotor modes have evolved.     

Limb growth in captive Galago senegalensis: getting in shape to be an adult

Since primate infants are not simply miniature adults, adult shape results from differential growth patterns of individual body segments. Initially an infant relies on its mother for transportation, and later begins independent locomotion. Skeletal growth patterns must meet the functional demands of independent locomotion. In this study we sought to determine whether Galago senegalensis braccatus follow the general primate pattern of decreasing intermembral index (IMI) throughout ontogeny. We also asked whether ontogenetic attainment of adult limb proportions coincides with attainment of independent locomotion, i.e., do infants reach adult limb proportions near the time they begin independent locomotion (approximately 7 weeks of age)? Mixed‐longitudinal data were taken from a sample of 10 captive‐born Galago senegalensis. Linear lengths of the trunk, arm, forearm, thigh, and leg were measured in the animals from birth until they were approximately 500 days old. The IMI and the ratio of each limb segment to both trunk length and the cube root of body mass were calculated. The results of a Mann‐Whitney Wilcoxon rank‐sum test for unmatched samples indicate that G. senegalensis do exhibit the primate pattern of decreasing IMI throughout ontogeny, and that the IMIs of infants at the time of initial locomotor independence are significantly higher than those of adult IMIs. Some (but not all) measures of relative limb lengths differed between neonates or 7‐week‐old infants and adults. Therefore, the hypothesis that infants acquire adult limb proportions by the time they begin independent locomotion is not supported by this study. The current results indicate that ontogenetic shape changes in galagos are a complex process and apparently cannot be explained by simple initial locomotor competency. Am. J. Primatol. 69:103–111, 2007. © 2006 Wiley‐Liss, Inc.     

Interlimb coordination,gait, and neural control of quadrupedalism in chimpanzees

Interlimb coordination is directly relevant to the understanding of the neural control of locomotion, but few studies addressing this topic for nonhuman primates are available, and no data exist for any hominoid other than humans. As a follow-up to Jungers and Anapol's ([1985] Am. J. Phys. Anthropol. 67:89–97) analysis on a lemur and talapoin monkey, we describe here the patterns of interlimb coordination in two chimpanzees as revealed by electromyography. Like the lemur and talapoin monkey, ipsilateral limb coupling in chimpanzees is characterized by variability about preferred modes within individual gaits. During symmetrical gaits, limb coupling patterns in the chimpanzee are also influenced by kinematic differences in hindlimb placement (“overstriding”). These observations reflect the neurological constraints placed on locomotion but also emphasize the overall flexibility of locomotor neural mechanisms. Interlimb coordination patterns are also species-specific, exhibiting significant differences among primate taxa and between primates and cats. Interspecific differences may be suggestive of phylogenetic divergence in the basic mechanisms for neural control of locomotion, but do not preclude morphological explanations for observed differences in interlimb coordination across species. Am J Phys Anthropol 102:177–186, 1997 © 1997 Wiley-Liss, Inc.     

The semicircular canal system and locomotion: The case of extinct lemuroids and lorisoids

Paleontologists reconstruct the locomotor and postural behavior of extinct species by analogy with living forms and biomechanical analyses. In rare cases, behavioral evidence such as footprints can be used to confirm fossil‐based reconstructions for predominantly terrestrial orders of mammals. For instance, the chalicothere prints from Laetoli show that these perissodactyls supported their body weight on the metacarpals, as previously reconstructed. 1 Unfortunately, primates are mostly arboreal and rarely leave footprints. The cercopithecid and hominin prints at Laetoli are a rare exception. We have recently shown that the semicircular canal system can be used to test and augment locomotor reconstructions based on postcranial material or to provide first estimations of locomotor behavior for taxa not known from the postcranium. Using a sample of modern primates, we have been able to demonstrate that the radii of curvature of the semicircular canals are significantly correlated with both body mass and agility of locomotion. 2 This paper reviews those results and examines the relationship between semicircular canal morphology and other evidence in efforts to reconstruct locomotor behavior in subfossil lemurs from the Holocene of Madagascar and fossil lorisoids from the Miocene of Africa.     

Femoral/humeral strength in early African Homo erectus

Lower-to-upper limb-bone proportions give valuable clues to locomotor behavior in fossil taxa. However, to date only external linear dimensions have been included in such analyses of early hominins. In this study, cross-sectional measures of femoral and humeral diaphyseal strength are determined for the two most complete early Homo erectus (or ergaster) associated skeletons--the juvenile KNM-WT 15000 and the adult KNM-ER 1808. Modern comparative samples include an adult human skeletal sample representative of diverse body shapes, a human longitudinal growth series, and an adult chimpanzee sample. When compared to appropriately age-matched samples, both H. erectus specimens fall very close to modern human mean proportions and far from chimpanzee proportions (which do not overlap with those of humans). This implies very similar mechanical load-sharing between the lower and upper limbs, and by implication, similar locomotor behavior in early H. erectus and modern humans. Thus, by the earliest Pleistocene (1.7 Ma), completely modern patterns of bipedal behavior were fully established in at least one early hominin taxon.     

Locomotor Behavior and Body Mass of Paramys delicatus (Ischyromyidae,Rodentia) and Commentary on Other Early North American Paramyines

Paramyine ischyromyids are one of the first ancestral rodent groups to appear in North America. Studying ecological indicators of these extinct animals enables us to better understand how they integrated into North American mammalian communities. In this study we reassess the locomotor behavior of a nearly complete skeleton of a paramyine, Paramys delicatus (AMNH FM 12506), using functional limb indices and living squirrels as extant analogues. We then used the results of the functional limb index study to select an appropriate locomotor group for body mass estimations of Paramys delicatus and other early North American (Wasatchian-Bridgerian) paramyines. This was done because body mass is strongly tied to locomotor patterns and more reliable body mass estimates can be generated from an extant sample that functionally resembles the fossils being studied. Functional limb indices were calculated for three locomotor groups (arboreal, semifossorial, and gliding) of living sciurids. Comparisons among arboreal, semifossorial, and gliding sciurids show that the functional indices related to mechanical advantage of muscles and limb robusticity enable distinction among locomotor groups; however, there is considerable overlap between arboreal and semifossorial taxa. Paramys delicatus was found to have generally greater mechanical advantages and limb robusticity than most living squirrels, including semifossorial taxa. As these traits are associated with semifossorial squirrels that frequently use scratch-digging, this suggests that Paramys delicatus and perhaps other early paramyines were likely proficient scratch-diggers. However, indices reflecting limb proportions of paramyines suggest that these early rodents may have used more hind limb dominated locomotion than do living squirrels. Body mass estimations for early paramyines were therefore derived from a semifossorial squirrel sample. Statistical comparisons suggest that many of the most reliable estimators for body mass in Paramys delicatus and other paramyines are those derived from humeral dimensions, with the most reliable estimate being humeral head superoinferior breadth. Using these estimators, individual body mass estimates of early paramyines range from 3391 to 4005 g for Paramys delicatus, 1137–1329 g for Paramys copei, 1291 g for Paramys taurus, and 3357 g for Notoparamys costilloi. All body mass estimations derived from postcranial elements are substantially larger than previously published estimates derived from the dentition, which may be because postcranial elements play a larger role in supporting body weight.

     

The bony labyrinth of the early platyrrhine primate Chilecebus

We document the morphology of the bony labyrinth of Chilecebus carrascoensis, one of the best preserved early platyrrhines known, based on high resolution CT scanning and 3D digital reconstruction. The cochlea is low and conical in form, as in other anthropoids, but has only 2.5 spiral turns. When the allometric relationship with body mass is considered, cochlear size is similar to that in extant primates. The relative size of the semicircular canals, which is well within the range of other primates, indicates that Chilecebus carrascoensis was probably not as agile in its locomotion as other small-bodied platyrrhines such as Leontopithecus rosalia, Saguinus oedipus, and Callithrix jacchus, but it probably was not a suspensory acrobat or a slow climber. The proportion, shape, and orientation of the semicircular canals in Chilecebus carrascoensis also mirror that typically seen in extant primates. However, no single variable can be used for predicting the locomotor pattern in Chilecebus carrascoensis. Based on Principle Component Analysis (PCA) scores we calculated rescaled Euclidean distances for various taxa; primates with similar locomotor patterns tend to share shorter distances. Results for Chilecebus carrascoensis underscore its general resemblance to living quadrupedal primate taxa, but it is not positioned especially near any single living taxon.     

Hindlimb proportions, allometry, and biomechanics in Old World monkeys (primates, Cercopithecidae).

The traditional focus on morphological rather than mechanical units has obscured some significant functional differences in the hindlimbs of primates. This paper examines the allometric and biomechanical basis for some distinctive proportional differences among pairs of morphological units in the hindlimb, and especially the foot, of cercopithecid primates. Five major conclusions are reached. First, many hindlimb dimensions scale allometrically with body mass to maintain mechanical similarity within taxonomic and locomotor groups. Therefore, the majority of traditional indices which describe the shape of the foot within cercopithecids reveal differences which are primarily a function of size. Second, the hindlimb segments in colobines, and especially in Presbytis, are relatively long, probably to enhance leaping. Third, the major distinction of terrestrial cercopithecines among the features analysed is reduction in the length of the phalanges, due to the reduced importance of grasping during locomotion and the assumption of digitigrady. Fourth, Theropithecus and male Erythrocebus have high crural indices, relative to their body masses, which can facilitate curosoriality. Female E. patas already has a high crural index as a function of its body mass. Fifth, macaques form a distinctive group among cercopithecines, characterized by relatively short hindlimbs. Relatively very short hindlimbs in Macaca fuscata and M. thibetana suggest that climatic conditions can have an added effect on the lengths of the hindlimb segments. In summary, this analysis of the lengths of the hindlimb segments relative to body size reveals taxonomic differences which are due in part to phylogeny, to differences in locomotor behavior, and to substrate use.     

Locomotion in some small to medium-sized mammals: a geometric morphometric analysis of the penultimate lumbar vertebra,pelvis and hindlimbs

We assessed the influence of a variety of aspects of locomotion and ecology including gait and locomotor types, maximal running speed, home range, and body size on postcranial shape variation in small to medium-sized mammals, employing geometric morphometric analysis and phylogenetic comparative methods. The four views analyzed, i.e., dorsal view of the penultimate lumbar vertebra, lateral view of the pelvis, posterior view of the proximal femur and proximal view of the tibia, showed clear phylogenetic signal and interesting patterns of association with movement. Variation in home range size was related to some tibia shape changes, while speed was associated with lumbar vertebra, pelvis and tibia shape changes. Femur shape was not related to any locomotor variables. In both locomotor type and high-speed gait analyses, locomotor groups were distinguished in both pelvis and tibia shape analyses. These results suggest that adaptations to both typical and high-speed gaits could explain a considerable portion of the shape of those elements. In addition, lumbar vertebra and tibia showed non-significant relationships with body mass, which suggests that they might be used in morpho-functional analyses and locomotor inferences on fossil taxa, with little or no bias for body size. Lastly, we observed morpho-functional convergences among several mammalian taxa and detected some taxa that achieve similar locomotor features following different morphological paths.