Variation in limb loading magnitude and timing in tetrapods.

Comparative analyses of locomotion in tetrapods reveal two patterns of stride cycle variability. Tachymetabolic tetrapods (birds and mammals) have lower inter-cycle variation in stride duration than bradymetabolic tetrapods (amphibians, lizards, turtles and crocodilians). This pattern has been linked to the fact that birds and mammals share enlarged cerebella, relatively enlarged and heavily myelinated Ia afferents, and γ-motoneurons to their muscle spindles. Both tachymetabolic tetrapod lineages also possess an encapsulated Golgi tendon morphology, thought to provide more spatially precise information on muscle tension. The functional consequence of this derived Golgi tendon morphology has never been tested. We hypothesized that one advantage of precise information on muscle tension would be lower and more predictable limb bone stresses, achieved in tachymetabolic tetrapods by having less variable substrate reaction forces than bradymetabolic tetrapods. To test this hypothesis, we analyzed hindlimb substrate reaction forces during locomotion of 55 tetrapod species in a phylogenetic comparative framework. Variation in species means of limb loading magnitude and timing confirm that, for most of the variables analyzed, variance in hindlimb loading and timing is significantly lower in species with encapsulated versus unencapsulated Golgi tendon organs. These findings suggest that maintaining predictable limb loading provides a selective advantage for birds and mammals by allowing energy savings during locomotion, lower limb bone safety factors and quicker recovery from perturbations. The importance of variation in other biomechanical variables in explaining these patterns, such as posture, effective mechanical advantage and center-of-mass mechanics, remains to be clarified.

Limb phase flexibility in walking: a test case in the squirrel monkey (Saimiri sciureus).

BACKGROUND: Previous analyses of factors influencing footfall timings and gait selection in quadrupeds have focused on the implications for energetic cost or gait mechanics separately. Here we present a model for symmetrical walking gaits in quadrupedal mammals that combines both factors, and aims to predict the substrate contexts in which animals will select certain ranges of footfall timings that (1) minimize energetic cost, (2) minimize rolling and pitching moments, or (3) balance the two. We hypothesize that energy recovery will be a priority on all surfaces, and will be the dominant factor determining footfall timings on flat, ground-like surfaces. The ability to resist pitch and roll, however, will play a larger role in determining footfall choice on narrower and more complex branch-like substrates. As a preliminary test of the expectations of the model, we collected sample data on footfall timings in a primate with relatively high flexibility in footfall timings - the squirrel monkey (Saimiri sciureus) - walking on a flat surface, straight pole, and a pole with laterally-projecting branches to simulate simplified ground and branch substrates. We compare limb phase values on these supports to the expectations of the model. RESULTS: As predicted, walking steps on the flat surface tended towards limb phase values that promote energy exchange. Both pole substrates induced limb phase values predicted to favor reduced pitching and rolling moments. CONCLUSIONS: These data provide novel insight into the ways in which animals may choose to adjust their behavior in response to movement on flat versus complex substrates and the competing selective factors that influence footfall timing in mammals. These data further suggest a pathway for future investigations using this perspective.

Functional and evolutionary aspects of axial stability in euarchontans and other mammals.

The presence of a stable thoracolumbar region, found in many arboreal mammals, is considered advantageous for bridging and cantilevering between discontinuous branches. However, no study has directly explored the link between osteological features cited as enhancing axial stability and the frequency of cantilevering and bridging behaviors in a terminal branch environment. To fill this gap, we collected metric data on costal and vertebral morphology of primate and nonprimate mammals known to cantilever and bridge frequently and those that do not. We also quantified the frequency and duration of cantilevering and bridging behaviors using experimental setups for species that have been reported to show differences in use of small branches and back anatomy (Caluromys philander, Loris tardigradus, Monodelphis domestica, and Cheirogaleus medius). Phylogenetically corrected principal component analysis reveals that taxa employing frequent bridging and cantilevering (C. philander and lorises) also exhibit reduced intervertebral and intercostal spaces, which can serve to increase thoracolumbar stability, when compared to closely related species (M. domestica and C. medius). We observed C. philander cantilevering and bridging significantly more often than M. domestica, which never cantilevered or crossed any arboreal gaps. Although no difference in the frequency of cantilevering was observed between L. tardigradus and C. medius, the duration of cantilevering bouts was significantly greater in L. tardigradus. These data suggest that osteological features promoting axial rigidity may be part of a morpho-behavioral complex that increases stability in mammals moving and foraging in a terminal branch environment.

Comparative and quantitative myology of the forearm and hand of prosimian primates.

Associations between the relative development of muscles of the forearm and hand of prosimian primates and locomotor behavior, body size, and grasping specializations have been made on the basis of qualitative observations. These associations are here tested through comparative and quantitative analyses of muscle mass data for a broad sample of prosimian species (i.e., strepsirrhines and tarsiers). The musculature of the forearm and hand of 17 fresh-frozen specimens representing six families and 12 species was dissected and weighed. Muscle weights were scaled relative to body mass of individual specimens using regression and compared by limb compartment and functional group. Forearm and digital flexor muscle masses are highly correlated with body mass (r = 0.97 and r = 0.96, respectively) and scale isometrically. As a general trend within the prosimian sample, the relative mass of the flexor compartment increases with body size. Interspecific comparisons of functional groups of muscles did not identify any association between larger muscle mass and locomotor adaptations or grasping specializations of the hand. However, compared to other prosimians, the adductor pollicis muscle of Nycticebus is more developed (52% of the intrinsic hand musculature sampled) and the flexor digitorum profundus muscle has two well-separated heads with more individualized tendons, with the pollex and digit IV receiving tendinous contributions from both muscle heads. These differences that characterize the forearm and hand of Nycticebus correlate with the extreme thumb divergence and pincer-like grips of lorises.

Teaching cricothyrotomy: a multisensory surgical education approach for final-year medical students.

OBJECTIVE: This study sought to evaluate the efficacy of a multisensory teaching approach in imparting the necessary knowledge, technical skills, and confidence to perform a cricothyrotomy to a cohort of fourth-year medical students. METHODS: One hundred twenty students were recruited into the study. Subjects commenced by viewing an upper airway obstruction vodcast. Students subsequently observed an expert demonstration of cricothyrotomy on human cadavers. Following this, students were able to practice on cadaveric simulators guided by experts. The students' confidence was assessed with pre- and post-session confidence-level questionnaires. Thirty randomly selected students were then assessed with a post-session objective skill assessment test. Finally, a multiple choice examination was administered to test knowledge. RESULTS: Ninety-five percent of student subjects achieved a score of at least 80% on the knowledge examination. All sampled students were considered competent to perform cricothyrotomy as judged by 2independent expert observers using the objective skill assessment test instrument. There was a statistically significant improvement between the pre- and post-CLQ scores (p<0.001). In keeping with current education theories on multisensory learning, qualitative feedback suggested a student preference for this teaching approach. CONCLUSION: This study provides further evidence that a multisensory teaching intervention effectively improves the knowledge, skill, and confidence of fourth-year medical students in performing cricothyrotomy.

Tail architecture and function of Cebupithecia sarmientoi, a Middle Miocene platyrrhine from La Venta, Colombia.

Cebupithecia sarmientoi, an early pitheciine from the Middle Miocene of La Venta, Colombia, preserves an almost complete caudal vertebral sequence (18 vertebrae). Behavioral reconstructions for this taxon based on appendicular elements suggest a locomotor profile similar to that of Pithecia for which vertical clinging postures and leaping behavior are frequently adopted. General tail morphology suggests some similarity with prehensile-tailed Cebus in the proximal tail region, although overwhelming similarity with nonprehensile-tailed Pithecia is evident in the distal tail region. Indices of caudal muscle attachment sites show marked similarities to nonprehensile-tailed platyrrhines, especially Pithecia. However, the cortices of Cebupithecia caudal vertebral bodies are thicker than those of most other nonprehensile-tailed New World primates. Mechanically, this would provide high resistance to bending and torsional stresses, falling within the range exhibited by prehensile-tailed monkeys. These results suggest that Cebupithecia may have employed its tail differently than most nonprehensile-tailed platyrrhines living today, behaviors that possibly involved tail-bracing or twisting during hindlimb (pedal grasping) suspensory behaviors. Such behaviors may serve as a preadaptive model for the full-fledged evolution of below-branch tail suspension and prehensility seen in other New World primates.

The effect of substrate size on the locomotion and gait patterns of the kinkajou (Potos flavus).

Diagonal-sequence (DS) gaits, which are very rare among mammals, are common and well documented in primates and some arboreal marsupials. DS walking gaits have been reported in the kinkajou (Potos flavus), which shows ecological similarities with primates and arboreal opossums but lacks prehensile specializations of the hindfoot. Nevertheless, the actual frequency of DS gaits and the functional context in which these gaits occur in this highly arboreal mammal remain unknown. We examined the effect of substrate size on the locomotion and gait patterns of kinkajous by recording gaits in two individuals walking and running on poles of two different diameters and on a runway. Diagonality and limb duty factors were calculated for 534 gait cycles. Kinkajous relied mostly on DS gaits and trots during walking, and increased the diagonality of their gait patterns on thinner substrates. The proposed functional link between locomotion on thin branches and the presence of a grasping, primate-like hindfoot is not supported by these data. However, further analysis of kinkajou gait cycles showed that DS gaits may have advantages overlooked earlier. DS gaits, during walking, minimize the distance between two ipsilateral feet during short periods of unilateral bipedality, and per corollary maximize the distance between two contralateral feet during the much longer periods of diagonal bipedality. Such foot positioning during the gait cycle could be beneficial in walking on a relatively thin substrate and could explain why kinkajous adopt DS walking gaits, especially on thinner poles, despite lacking prehensile specializations of the hindfoot.

Brief communication: Forelimb compliance in arboreal and terrestrial opossums.

Primates display high forelimb compliance (increased elbow joint yield) compared to most other mammals. Forelimb compliance, which is especially marked among arboreal primates, moderates vertical oscillations of the body and peak vertical forces and may represent a basal adaptation of primates for locomotion on thin, flexible branches. However, Larney and Larson (Am J Phys Anthropol 125 [2004] 42-50) reported that marsupials have forelimb compliance comparable to or greater than that of most primates, but did not distinguish between arboreal and terrestrial marsupials. If forelimb compliance is functionally linked to locomotion on thin branches, then elbow yield should be highest in marsupials relying on arboreal substrates more often. To test this hypothesis, we compared forelimb compliance between two didelphid marsupials, Caluromys philander (an arboreal opossum relying heavily on thin branches) and Monodelphis domestica (an opossum that spends most of its time on the ground). Animals were videorecorded while walking on a runway or a horizontal 7-mm pole. Caluromys showed higher elbow yield (greater changes in degrees of elbow flexion) on both substrates, similar to that reported for arboreal primates. Monodelphis was characterized by lower elbow yield that was intermediate between the values reported by Larney and Larson (Am J Phys Anthropol 125 [2004] 42-50) for more terrestrial primates and rodents. This finding adds evidence to a model suggesting a functional link between arboreality--particularly locomotion on thin, flexible branches--and forelimb compliance. These data add another convergent trait between arboreal primates, Caluromys, and other arboreal marsupials and support the argument that all primates evolved from a common ancestor that was a fine-branch arborealist.

Intrinsic hand proportions of euarchontans and other mammals: implications for the locomotor behavior of plesiadapiforms.

Arboreal primates have distinctive intrinsic hand proportions compared with many other mammals. Within Euarchonta, platyrrhines and strepsirrhines have longer manual proximal phalanges relative to metacarpal length than colugos and terrestrial tree shrews. This trait is part of a complex of features allowing primates to grasp small-diameter arboreal substrates. In addition to many living and Eocene primates, relative elongation of proximal manual phalanges is also present in most plesiadapiforms. In order to evaluate the functional and evolutionary implications of manual similarities between crown primates and plesiadapiforms, we measured the lengths of the metacarpal, proximal phalanx, and intermediate phalanx of manual ray III for 132 extant mammal species (n=702 individuals). These data were compared with measurements of hands in six plesiadapiform species using ternary diagrams and phalangeal indices. Our analyses reveal that many arboreal mammals (including some tree shrews, rodents, marsupials, and carnivorans) have manual ray III proportions similar to those of various arboreal primates. By contrast, terrestrial tree shrews have hand proportions most similar to those of other terrestrial mammals, and colugos are highly derived in having relatively long intermediate phalanges. Phalangeal indices of arboreal species are significantly greater than those of the terrestrial species in our sample, reflecting the utility of having relatively long digits in an arboreal context. Although mammals known to be capable of prehensile grips demonstrate long digits relative to palm length, this feature is not uniquely associated with manual prehension and should be interpreted with caution in fossil taxa. Among plesiadapiforms, Carpolestes, Nannodectes, Ignacius, and Dryomomys have manual ray III proportions that are unlike those of most terrestrial species and most similar to those of various arboreal species of primates, tree shrews, and rodents. Within Euarchonta, Ignacius and Carpolestes have intrinsic hand proportions most comparable to those of living arboreal primates, while Nannodectes is very similar to the arboreal tree shrew Tupaia minor. These results provide additional evidence that plesiadapiforms were arboreal and support the hypothesis that Euarchonta originated in an arboreal milieu.

New hand bones of Hadropithecus stenognathus: implications for the paleobiology of the Archaeolemuridae.

A partial, associated skeleton of Hadropithecus stenognathus (AHA-I) was discovered in 2003 at Andrahomana Cave in southeastern Madagascar. Among the postcranial elements found were the first hand bones (right scaphoid, right hamate, left first metacarpal, and right and left fifth metacarpals) attributed to this rare subfossil lemur. These hand bones were compared to those of extant strepsirrhines and catarrhines in order to infer the positional adaptations of Hadropithecus, and they were compared to those of Archaeolemur in order to assess variation in hand morphology among archaeolemurids. The scaphoid tubercle does not project palmarly as in suspensory and climbing taxa, and the hamate has no hook at all (just a small tubercle), which also points to a poorly developed carpal tunnel. There is a distinctive, radioulnarly directed "spiral" facet for articulation with the triquetrum that is most similar in orientation to that of more terrestrial primates (i.e., Lemur catta, Papio, and Gorilla). The first metacarpal is very reduced and represents only 48% of the length of metacarpal V, as in Archaeolemur, which suggests that pollical grasping of arboreal supports was not important. Compared to Archaeolemur, the shaft of metacarpal V is gracile, and the head has no dorsal ridge and lacks characteristics functionally associated with digitigrade, extended metacarpophalangeal joint postures. Proximally, the articular facet for the hamate is oriented more dorsally. Thus, the carpometacarpal joint V appears to have a distinctive hyperextended set, which has no analog among living or extinct primates. The carpals of Hadropithecus are diagnostic of a pronograde, arboreal and terrestrial (although not digitigrade) locomotor repertoire that typifies Lemur catta and some Old World monkeys. No clinging, suspensory, or climbing specializations that characterize indriids or lorises can be found in the hand of this subfossil lemur. The hand of Hadropithecus likely had similar ranges of movement at the radiocarpal and midcarpal joints as of those of pronograde primates, such as lemurids, for which the hand is held in a more extended, pronated, and neutral (i.e., showing less ulnar deviation) position during locomotion in comparison to that of vertical clingers or slow climbers. Although highly autapomorphic, the hand of Hadropithecus resembles that of its sister taxon, Archaeolemur, in having a very reduced pollex and an articular facet on the scaphoid for a sizeable prepollex. These unusual hand features reinforce the monophyly of the Archaeolemuridae.

Comparative morphometrics of the primate apical tuft.

The relationship between the structure and function of the primate apical tuft is poorly understood. This study addresses several hypotheses about apical tuft morphology using a large modern primate comparative sample. Two indices of tuft size are employed: expansion and robusticity. First, comparisons of relative apical tuft size were drawn among extant nonhuman primate groups in terms of locomotion and phylogenetic category. Both of these factors appear to play a role in apical tuft size among nonhuman primates. Suspensory primates and all platyrrhines had the smallest apical tufts, while terrestrial quadrupeds and all strepsirrhines (regardless of locomotor category) had the largest tufts. Similarly, hypotheses regarding the apical tufts of hominins, especially the large tufts of Neandertals were addressed using a comparison of modern warm- and cold-adapted humans. The results showed that cold-adapted populations possessed smaller apical tufts than did warm-adapted groups. Therefore, the cold-adaptation hypothesis for Neandertal distal phalangeal morphology is not supported. Also, early modern and Early Upper Paleolithic humans had apical tufts that were significantly less expanded and less robust than those of Neandertals. The hypothesis that a large apical tuft serves as support for an expanded digital pulp is supported by radiographic analysis of modern humans in that a significant correlation was discovered between the width of the apical tuft and the width of the pulp. The implications of these findings for hypotheses about the association of apical tuft size and tool making in the hominin fossil record are discussed.

Body size and scaling of the hands and feet of prosimian primates.

The hands and feet of primates fulfill a variety of biological roles linked with food acquisition and positional behavior. Current explanations of shape differences in cheiridial morphology among prosimians are closely tied to body size differences. Although numerous studies have examined the relationships between body mass and limb morphology in prosimians, no scaling analysis has specifically considered hand and foot dimensions and intrinsic proportions. In this study, we present such an analysis for a sample of 270 skeletal specimens distributed over eight prosimian families. The degree of association between size and shape was assessed using nonparametric correlational techniques, while the relationship between each ray element length and body mass (from published data and a body mass surrogate) was tested for allometric scaling. Since tarsiers and strepsirrhines encompass many taxa of varying degrees of phylogenetic relatedness, effective degrees of freedom were calculated, and comparisons between families were performed to partially address the problem of statistical nonindependence and "phylogenetic inertia." Correlational analyses indicate negative allometry between relative phalangeal length (as reflected by phalangeal indices) and body mass, except for the pollex and hallux. Thus, as size increases, there is a significant decrease in the relative length of the digits when considering all prosimian taxa sampled. Regression analyses show that while the digital portion of the rays scales isometrically with body mass, the palmar/plantar portion of the rays often scales with positive allometry. Some but not all of these broadly interspecific allometric patterns remain statistically significant when effective degrees of freedom are taken into account. As is often the case in interspecific scaling, comparisons within families show different scaling trends in the cheiridia than those seen across families (i.e., lorisids, indriids, and lemurids exhibit rather different allometries). The interspecific pattern of positive allometry that appears to best characterize the metapodials of prosimians, especially those of the foot, parallels differences found in the morphology of the volar skin. Indeed, relatively longer metapodials appear to covary with flatter and more coalesced volar pads, which in turn slightly improve frictional force for animals that are at a comparative disadvantage while climbing because of their larger mass. Despite the essentially isometric relationship found between digit length and body mass across prosimians, examination of the residual variation reveals that tarsiers and Daubentonia possess, relative to their body sizes, remarkably long fingers. Such marked departures between body size and finger length observed in these particular primates are closely linked with specialized modes of prey acquisition and manipulation involving the hands.

Adaptive value of ambling gaits in primates and other mammals.

At speeds between the walk and the gallop, most mammals trot. Primates almost never trot, and it has been claimed that they transition directly from a walk to a gallop without any distinctive mid-speed running gait. If true, this would be another characteristic difference between the locomotion of primates and that of most other quadrupedal mammals. Presently, however, few data exist concerning the actual presence or absence of intermediate-speed gaits (i.e. gaits that are used between a walk and a gallop) in primates. Video records of running in twelve primate species reveal that, unlike most other mammals, all the primates studied almost exclusively adopt an 'amble'--an intermediate-speed running gait with no whole-body aerial phase--rather than trot. Ambling is also common in elephants and some horses, raising the question of why ambling is preferred over trotting in these diverse groups of animals. Mathematical analyses presented here show that ambling ensures continuous contact of the body with the substrate while dramatically reducing vertical oscillations of the center of mass. This may explain why ambling appears to be preferable to trotting for extremely large terrestrial mammals such as elephants and for arboreal mammals like primates that move on unstable branches. These findings allow us to better understand the mechanics of these unusual running gaits and shed new light on primate locomotor evolution.

Role of the prehensile tail during ateline locomotion: experimental and osteological evidence.

The dynamic role of the prehensile tail of atelines during locomotion is poorly understood. While some have viewed the tail of Ateles simply as a safety mechanism, others have suggested that the prehensile tail plays an active role by adjusting pendulum length or controlling lateral sway during bimanual suspensory locomotion. This study examines the bony and muscular anatomy of the prehensile tail as well as the kinematics of tail use during tail-assisted brachiation in two primates, Ateles and Lagothrix. These two platyrrhines differ in anatomy and in the frequency and kinematics of suspensory locomotion. Lagothrix is stockier, has shorter forelimbs, and spends more time traveling quadrupedally and less time using bimanual suspensory locomotion than does Ateles. In addition, previous studies showed that Ateles exhibits greater hyperextension of the tail, uses its tail to grip only on alternate handholds, and has a larger abductor caudae medialis muscle compared to Lagothrix. In order to investigate the relationship between anatomy and behavior concerning the prehensile tail, osteological data and kinematic data were collected for Ateles fusciceps and Lagothrix lagothricha. The results demonstrate that Ateles has more numerous and smaller caudal elements, particularly in the proximal tail region. In addition, transverse processes are relatively wider, and sacro-caudal articulation is more acute in Ateles compared to Lagothrix. These differences reflect the larger abductor muscle mass and greater hyperextension in Ateles. In addition, Ateles shows fewer side-to-side movements during tail-assisted brachiation than does Lagothrix. These data support the notion that the prehensile tail represents a critical dynamic element in the tail-assisted brachiation of Ateles, and may be useful in developing inferences concerning behavior in fossil primates.

Locomotor mechanics of the slender loris (Loris tardigradus).

The quadrupedal walking gaits of most primates can be distinguished from those of most other mammals by the presence of diagonal-sequence (DS) footfall patterns and higher peak vertical forces on the hindlimbs compared to the forelimbs. The walking gait of the woolly opossum (Caluromys philander), a highly arboreal marsupial, is also characterized by diagonal-sequence footfalls and relatively low peak forelimb forces. Among primates, three species--Callithrix, Nycticebus, and Loris--have been reported to frequently use lateral-sequence (LS) gaits and experience relatively higher peak vertical forces on the forelimbs. These patterns among primates and other mammals suggest a strong association between footfall patterns and force distribution on the limbs. However, current data for lorises are limited and the frequency of DS vs. LS walking gaits in Loris is still ambiguous. To test the hypothesis that patterns of footfalls and force distribution on the limbs are functionally linked, kinematic and kinetic data were collected simultaneously for three adult slender lorises (Loris tardigradus) walking on a 1.25 cm horizontal pole. All subjects in this study consistently used diagonal-sequence walking gaits and always had higher peak vertical forces on their forelimbs relative to their hindlimbs. These results call into question the hypothesis that a functional link exists between the presence of diagonal-sequence walking gaits and relatively higher peak vertical forces on the hindlimbs. In addition, this study tested models that explain patterns of force distribution based on limb protraction angle or limb compliance. None of the Loris subjects examined showed kinematic patterns that would support current models proposing that weight distribution can be adjusted by actively shifting weight posteriorly or by changing limb stiffness. These data reveal the complexity of adaptations to arboreal locomotion in primates and indicate that diagonal-sequence walking gaits and relatively low forelimb forces could have evolved independently.

Seasonal variation in body mass and locomotor kinetics of the fat-tailed dwarf lemur (Cheirogaleus medius).

The fat-tailed dwarf lemur (Cheirogaleus medius) is unusual among primates in storing large amounts of fat subcutaneously prior to hibernating during the winter months. In doing so, it increases its body mass by more than 50%, with a substantial weight gain in the tail. This seasonal increase in mass provides a unique natural experiment to examine how changes in body mass affect substrate reaction forces during locomotion. As body mass increases, it is expected that the limbs of the fat-tailed dwarf lemur will be subjected to greater peak vertical substrate reaction forces during quadrupedal walking. However, whether or not these peak substrate reaction forces will increase proportionally across forelimbs and hindlimbs as body mass increases is unknown. Substrate reaction forces were collected on four adult C. medius walking quadrupedally on a 28-mm pole attached to a force platform. Peak vertical substrate reaction forces (Vpk) (N) were analyzed and compared for a cross-sectional sample of different body masses (180-300 g). Forelimb and hindlimb Vpk were positively correlated with body mass, with hindlimb Vpk always higher than forelimb Vpk. However, the rate at which Vpk increased relative to body mass was higher for the hindlimb than the forelimb. This disproportion in weight distribution between the forelimbs and hindlimbs as body mass increases appears to be linked to the accumulation of fat in the tail. It is likely that storing fat in the tail region may shift the center of mass more caudally, from a more cranial position when the tail is thinner. Such a caudal shift of the center of mass-either morphological or dynamic-is believed to have played an important role in the functional differentiation of the limbs and the evolution of locomotor modes of several tetrapod groups, including dinosaurs and primates.

Hand and body position during locomotor behavior in the aye-aye (Daubentonia madagascariensis).

Aye-ayes (Daubentonia madagascariensis) have unique hands among primates, with extraordinarily long fingers in relation to body size. These long digits may be vulnerable to damage from forces during locomotion, particularly during head-first descent-a locomotor mode that the aye-aye utilizes frequently. Previous behavioral studies of aye-aye locomotion reported that Daubentonia must curl its fingers during horizontal quadrupedalism and/or descent to reduce potential stresses on its long fingers. To test this hypothesis, we examined hand and body position in three captive adult aye-ayes while they walked quadrupedally on horizontal and oblique branches. Substantial variation in hand position was observed among individuals for each substrate orientation. While hand postures with curled fingers were preferred by one individual during descent, they were not preferred by the other two individuals, contrary to our expectations. Differences in body position were more consistent among all three individuals. The angle of the body relative to the substrate was significantly reduced during descent (8.4 degrees ) compared to horizontal locomotion (16.9 degrees ). These results suggest that changes in body position, rather than hand position, may help reduce stresses on the digits. A biomechanical model is proposed that demonstrates how a reduction in the body angle in relation to substrate may act to move the center of mass more caudally. This mechanism of moderating loads by altering body position, rather than hand position, may represent an important functional aspect of arboreal locomotion in aye-ayes and other primates.

Origins of primate locomotion: gait mechanics of the woolly opossum.

The locomotion of primates differs from that of other mammals in three fundamental ways. During quadrupedal walking, primates use diagonal sequence gaits, protract their arms more at forelimb touchdown, and experience lower vertical substrate reaction forces on their forelimbs relative to their hindlimbs. It is widely held that the unusual walking gaits of primates represent a basal adaptation for movement on thin, flexible branches and reflect a major change in the functional role of the forelimb. However, little data on nonprimate arboreal mammals exist to test this notion. To that end, we examined the gait mechanics of the woolly opossum (Caluromys philander), a marsupial convergent with small-bodied prosimians in ecology, behavior, and morphology. Data on the footfall sequence, relative arm protraction, and peak vertical substrate reaction forces were obtained from videotapes and force records for three adult woolly opossums walking quadrupedally on a wooden runway and a thin pole. For all steps recorded on both substrates, woolly opossums always used diagonal sequence walking gaits, protracted their arms beyond 90 degrees relative to horizontal body axis, and experienced peak vertical substrate reaction forces on forelimbs that were significantly lower than on hindlimbs. The woolly opossum is the first nonprimate mammal to show locomotor mechanics that are identical to those of primates. This case of convergence between primates and a committed fine-branch, arboreal marsupial strongly implies that the earliest primates evolved gait specializations for fine-branch locomotion, which reflect important changes in forelimb function.