Terrestrial Quadrupeds Research Articles

Metacarpophalangeal Joint Orientation in Anthropoid Manual Phalanges

The proximal articular surface angle of orientation (AO) of proximal phalanges of the hand and foot has been used to infer the locomotor profile of extinct Miocene catarrhines and early hominins. Previous work has found that joint orientation distinguishes quadrupedal from suspensory anthropoids. The purpose of this study is to expand on previous research by examining this feature within and across several primate clades, allowing us to investigate the potential influences of locomotion, substrate usage, hand posture, and phylogeny. We also report AO measurements in human proximal hand phalanges, allowing us to examine human skeletal variation within a wide comparative context. The angle of orientation was measured on manual proximal third phalanges of 21 extant anthropoid species using a Microscribe digitizer. Comparisons were made between locomotor groups within hominoids, platyrrhines, and cercopithecoids. Proximal phalanges of quadrupedal species were characterized by greater dorsal orientation than those of suspensory taxa in hominoids and atelids. In addition, arboreal quadrupeds had greater AO values than terrestrial quadrupeds within the Cercopithecoidea. However, within the terrestrial locomotor group, mean AO values did not differ between palmigrade and digitigrade taxa. Thus, while there appears to be a functional signal related to substrate usage, differences in use of hand postures when moving on the ground were not reflected in proximal joint orientation of the proximal phalanx. Finally, we measured relatively low AO values in human phalanges, which might be related to integration with serially homologous pedal phalanges that are under strong selective pressure related to bipedalism.

A universal scaling relationship between body mass and proximal limb bone dimensions in quadrupedal terrestrial tetrapods.

BackgroundBody size is intimately related to the physiology and ecology of an organism. Therefore, accurate and consistent body mass estimates are essential for inferring numerous aspects of paleobiology in extinct taxa, and investigating large-scale evolutionary and ecological patterns in the history of life. Scaling relationships between skeletal measurements and body mass in birds and mammals are commonly used to predict body mass in extinct members of these crown clades, but the applicability of these models for predicting mass in more distantly related stem taxa, such as non-avian dinosaurs and non-mammalian synapsids, has been criticized on biomechanical grounds. Here we test the major criticisms of scaling methods for estimating body mass using an extensive dataset of mammalian and non-avian reptilian species derived from individual skeletons with live weights.ResultsSignificant differences in the limb scaling of mammals and reptiles are noted in comparisons of limb proportions and limb length to body mass. Remarkably, however, the relationship between proximal (stylopodial) limb bone circumference and body mass is highly conserved in extant terrestrial mammals and reptiles, in spite of their disparate limb postures, gaits, and phylogenetic histories. As a result, we are able to conclusively reject the main criticisms of scaling methods that question the applicability of a universal scaling equation for estimating body mass in distantly related taxa.ConclusionsThe conserved nature of the relationship between stylopodial circumference and body mass suggests that the minimum diaphyseal circumference of the major weight-bearing bones is only weakly influenced by the varied forces exerted on the limbs (that is, compression or torsion) and most strongly related to the mass of the animal. Our results, therefore, provide a much-needed, robust, phylogenetically corrected framework for accurate and consistent estimation of body mass in extinct terrestrial quadrupeds, which is important for a wide range of paleobiological studies (including growth rates, metabolism, and energetics) and meta-analyses of body size evolution.

A Cassowary Casuarius casuarius (Linnaeus, 1758) Record from Alexandria, Egypt, in 20 B.C. (Aves, Ratitae, Casuaridae)

The reverse side of the Artemidorus Papyrus, which was latest created early in the first century A.D. in Alexandria, features 47 drawings of animals by the same illustrator. In most cases, the Greek name of the animal is given. According to an Aristotelian heading, the papyrus shows terrestrial quadrupeds, birds, fish and whales. The taxa vary: one jellyfish, one mantis shrimp, five fishes s. l., six reptiles s. l., eleven birds and seventeen mammals. The work fits into the Hellenistic tradition of realistic animal illustrations. The papyrus was obviously produced and used as a pattern book. All the animals depicted are from Africa or the Mediterranean, except for eleven which can be said with certainty to come from India and four others which occur in both Africa and Asia. The Indian animals were presented to Princeps Augustus (r. 31 B.C. - 14 A.D.) in the summer of 20 B.C. in Daphne, Antioch and in the winter of 20/19 B.C. on the island of Samos by a delegation sent by King Poros of India (ruler of 600 kings), a Gujarati monarch hoping to establish trade relations with the Roman Empire. The delegation made its way to Rome via Antioch where it split for Samos and Athens accompanying Augustus, and via Alexandria, where a number of its animals were recorded on the Artemidorus Papyrus. Some of the species portrayed are also attested to by Strabo fide Nikolaos of Damascus. Others of the same exotic origin to be depicted in Alexandria include the four-horned antelope and the cassowary examined below. The complexity of the animal depictions on the reverse of this papyrus and the numerous details pinning it to historical events are enough to put paid to the notion that the Artemidorus Papyrus is a forgery. An asiatic bird named cornica which is described in an apocryph Plinius edition cited by medieval authors, unmistakeably is a cassowary, probably the same specimen.

Topsy-turvy locomotion: biomechanical specializations of the elbow in suspended quadrupeds reflect inverted gravitational constraints

Some tetrapods hang upside down from tree branches when moving horizontally. The ability to walk in quadrupedal suspension has been acquired independently in at least 14 mammalian lineages. During the stance (supportive) phase of quadrupedal suspension, the elbow joint flexor muscles (not the extensors as in upright vertebrates moving overground) are expected to contract to maintain the flexed limb posture. Therefore muscular control in inverted, suspended quadrupeds may require changes of muscle control, and even morphologies, to conditions opposite to those in upright animals. However, the relationships between musculoskeletal morphologies and elbow joint postures during the stance phase in suspended quadrupeds have not been investigated. Our analysis comparing postures and skeletal morphologies in Choloepus (Pilosa), Pteropus (Chiroptera), Nycticebus (Primates) and Cynocephalus (Dermoptera) revealed that the elbow joints of these animals were kept at flexed angles of 70-100 ° during the stance phase of quadrupedal suspension. At these joint angles the moment arms of the elbow joint flexors were roughly maximized, optimizing that component of antigravity support. Our additional measurements from various mammalian species show that suspended quadrupeds have relatively small extensor/flexor ratios in both muscle masses and maximum moment arms. Thus, in contrast to the pattern in normal terrestrial quadrupeds, suspended quadrupeds emphasize flexor over extensor muscles for body support. This condition has evolved independently multiple times, attendant with a loss or reduction of the ability to move in normal upright postures.

Effects of grade and mass distribution on the mechanics of trotting in dogs

Quadrupedal running on grades requires balancing of pitch moments about the center of mass (COM) while supplying sufficient impulse to maintain a steady uphill or downhill velocity. Here, trotting mechanics on a 15 deg grade were characterized by the distribution of impulse between the limbs and the angle of resultant impulse at each limb. Anterior-posterior manipulation of COM position has previously been shown to influence limb mechanics during level trotting of dogs; hence, the combined effects of grade and COM manipulations were explored by adding 10% body mass at the COM, shoulder or pelvis. Whole body and individual limb ground reaction forces, as well as spatiotemporal step parameters, were measured during downhill and uphill trotting. Deviations from steady-speed locomotion were determined by the net impulse angle and accounted for in the statistical model. The limbs exerted only propulsive force during uphill trotting and, with the exception of slight hindlimb propulsion in late stance, only braking force during downhill trotting. Ratios of forelimb impulse to total impulse were computed for normal and shear components. Normal impulse ratios were more different from level values during uphill than downhill trotting, indicating that the limbs act more as levers on the incline. Differential limb function was evident in the extreme divergence of forelimb and hindlimb impulse angles, amplifying forelimb braking and hindlimb propulsive biases observed during level trotting. In both downhill and uphill trotting, added mass at the up-slope limb resulted in fore-hind distributions of normal impulse more similar to those of level trotting and more equal fore-hind distributions of shear impulse. The latter result suggests a functional trade-off in quadruped design: a COM closer to the hindlimbs would distribute downhill braking more equally, whereas a COM closer to the forelimbs would distribute uphill propulsion more equally. Because muscles exert less force when actively shortening than when lengthening, it would be advantageous for the forelimb and hindlimb muscles to share the propulsive burden more equally during uphill trotting. This functional advantage is consistent with the anterior COM position of most terrestrial quadrupeds.

Gait dynamics of Cebus apella during quadrupedalism on different substrates

Primates are distinguished from many mammals by emphasizing arboreal lifestyles. Primate arboreal adaptations include specializations for enhancing balance and manipulative skills. Compliant gait and diagonal sequence (DS) footfalls are hypothesized mechanisms for improving balance during arboreal quadrupedalism (AQ), while simultaneously permitting vertical peak force reductions sustained by limbs, particularly forelimbs (FLs). Capuchin monkeys (Cebus apella) are arboreally-adapted quadrupeds that use both lateral sequence (LS) and DS footfalls. As tool-users, capuchins experience selective pressures for FL manipulative capabilities, which seemingly conflict with encountering substantial locomotor stresses. We evaluate kinetic and 3-D kinematic data from 172 limb contacts of two adult males on terrestrial and arboreal substrates to address questions about C. apella gait compliancy, kinematics of LS and DS footfalls during quadrupedalism on different substrates, and whether capuchins reduce FL vertical peak forces relative to hind limb (HL) forces more than other primates that use tools or those that do not. Lower vertical peak forces during AQ are consistent with compliant gait, but mixed kinematic results obscure how the reduction occurs. Forearm adduction angle is one consistent kinematic difference between terrestrial and arboreal quadrupedalism, which may implicate frontal plane movements in gait compliancy. Major differences between DS and LS gaits were not observed in kinetic or kinematic comparisons. Capuchins exhibit low FL/HL vertical peak force ratios like several anthropoids, including tool-users (e.g., chimpanzees), and species not considered tool-users in free-ranging conditions (e.g., spider monkeys). Additional selective pressures besides simply tool use appear responsible for the relative reduction in primate forelimb forces.

Optimal running speed and the evolution of hominin hunting strategies

Recent discussion of the selective pressures leading to the evolution of modern human postcranial morphology, seen as early as Homo erectus, has focused on the relative importance of walking versus running. Specifically, these conversations have centered on which gait may have been used by early Homo to acquire prey. An element of the debate is the widespread belief that quadrupeds are constrained to run at optimally efficient speeds within each gait, whereas humans are equally efficient at all running speeds. The belief in the lack of optimal running speeds in humans is based, however, on a number of early studies with experimental designs inadequate for the purpose of evaluating optimality. Here we measured the energetic cost of human running ( n = 9) at six different speeds for five minutes at each speed, with careful replicates and controls. We then compared the fit of linear versus curvilinear models to the data within each subject. We found that individual humans do, in fact, have speeds at which running is significantly less costly than at other speeds (i.e., an optimal running speed). In addition, we demonstrate that the use of persistence hunting methods to gain access to prey at any running speed, even the optimum, would be extremely costly energetically, more so than a persistence hunt at optimal walking speed. We argue that neither extinct nor extant hominin populations are as flexible in the chosen speeds of persistence hunting pursuits as other researchers have suggested. Variations in the efficiency of human locomotion appear to be similar to those of terrestrial quadrupeds.

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.

Mediolateral forces associated with quadrupedal gaits of lemurids

Substrate reaction forces (SRFs) were quantified for 502 forelimb and hindlimb contacts in two primate species, Eulemur fulvus and Lemur catta. Three individuals of each taxon were encouraged to traverse a force transducer. Reaction forces were evaluated for linear locomotion on runway and simulated arboreal substrates. Preferred gaits differed between substrates: galloping on the runway and walking on simulated arboreal substrates. Unlike vertical and fore–aft (FA) forces, mediolateral (ML) forces during linear locomotion were relatively independent of speed, which suggests balancing the centre of body mass in the coronal plane was not speed related. Since peak ML forces occasionally exceeded peak FA forces, ML forces were not of trivial importance during primate quadrupedal locomotion. Both lemurids regularly experienced lower ML peak forces during ‘arboreal’ than ‘terrestrial’ quadrupedalism, probably to minimize side-to-side fluctuations of their centre of mass over the simulated arboreal branch supports. Eulemur fulvus encountered greater ML peak forces in their forelimbs, while L. catta encountered greater hindlimb peak ML forces. Lemurids most frequently exerted medially directed peak side-to-side forces during arboreal quadrupedalism and laterally directed peak side-to-side forces during terrestrial quadrupedalism. The direction of hindlimb ML peak force, however, was less consistent than forelimb ML peak force regardless of substrate. The lemurs experienced oscillations in the direction of the forelimb and hindlimb ML force over the duration of support phase, suggesting that both pairs of limbs contributed to balance. More frequent changes in ML force direction of the hindlimb support the predominance of these limbs in keeping side-to-side balance. The results of our study suggest that ML forces warrant consideration in locomotion studies, especially when evaluating joint moments and bending regimes of long bones.

How does the white-headed langur (Trachypithecus leucocephalus) adapt locomotor behavior to its unique limestone hill habitat?

Limestone hill habitats pose unique challenges to langurs. One of the characteristics of this habitat is its cliffs, which account for about 10-20% of the total area. We have never observed langurs falling from the cliffs during our 10-year field study. Five patterns of locomotion were exhibited by the white-headed langur: (1) arboreal ascent and descent, (2) arboreal quadupedalism, (3) terrestrial quadrupadelism, (4) moving on cliffs and (5) leaping on cliffs. Locomotor patterns varied according to the substrate, but terrestrial quadrupedalism accounted for more than 50% of locomotion time. Moving on cliffs and leaping on cliffs may be modes of locomotion unique to the white-headed langur, at least in terms of frequency. White-headed langurs have an intermembral index of 76 and, compared to langurs with a similar intermembral index, are more terrestrial. Further analysis indicates that greater terrestrialism may be the result of adaptation to their limestone habitat. Interestingly, white-headed langurs select caves on the cliff as their sleeping sites, and they exhibit special behaviors for exiting and entering the cave very early in the morning and late in the evening.

Comparative postcranial body shape and locomotion inChlorocebus aethiops andCercopithecus mitis

Body weight and length, chest girth, and seven postcranial limb segment lengths are compared between two guenon species, Chlorocebus (Cercopithecus) aethiops (vervets) and Cercopithecus mitis (blue monkeys), exhibiting different habitual locomotor preferences. The subjects, all adults, were wild caught for a non-related research project (Turner et al. [1986] Genetic and morphological studies on two species of Kenyan monkeys, C. aethiops and C. mitis. In: Else JG, Lee PC, editors. Primate evolution, proceedings of the Xth International Congress of Primatology, Cambridge. London). The morphological results are interpreted within the context of previously published observations of primate locomotion and social organization. The sample is unique in that the body weight of each individual is known, allowing the effects of body-size scaling to be assessed in interspecific and intersexual comparisons. C. mitis has a significantly (P < 0.05) greater body weight and trunk length than C. aethiops. A shorter trunk may function to reduce spinal flexibility for ground-running in the latter. Proximal limb segments (arm and thigh) are significantly greater in C. mitis, reflecting known adaptations to committed arboreal quadrupedal locomotion. By contrast, relative distal limb segments (forearm, crus, and foot) are significantly longer in C. aethiops, concordant with a locomotor repertoire that includes substantial terrestrial quadrupedalism, in addition to arboreal agility, and also the requisite transition between ground and canopy. Although normally associated with arboreal monkeys, greater relative tail length occurs in the more terrestrial vervets. However, because vervets exploit both arboreal and terrestrial habitats, a longer tail may compensate for diminished balance during arboreal quadrupedalism resulting from the greater "brachial" and "crural" indices that enhance their ground quadrupedalism. Most interspecific differences in body proportions are explicable by differences in locomotor modalities. Some results, however, contradict commonly held "tenets" that relate body size and morphology exclusively to locomotion. Generally associated with terrestriality, sexual dimorphism (male/female) is greater in the more arboreal blue monkeys. A more intense, seasonal mating competition may account for this incongruity.

Functional anatomy of the olecranon process in hominoids and plio-pleistocene hominins.

This study examines the functional morphology of the olecranon process in hominoids and fossil hominins. The length of the bony lever of the triceps brachii muscle (TBM) is measured as the distance between the trochlear articular center and the most distant insertion site of the TBM, and olecranon orientation is measured as the angle that this bony lever makes with the long axis of the ulna. Results show that Homo, Pan, Gorilla, most monkeys, and the Australopithecus fossils studied have similar relative olecranon lengths. Suspensory hominoids and Ateles have shorter olecranons, suggesting, in some instances, selection for greater speed in extension. The orientation that the lever arm of the TBM makes with the long axis of the ulna varies with preferred locomotor mode. Terrestrial primates have olecranons that are more posteriorly oriented as body size increases, fitting general models of terrestrial mammalian posture. Arboreal quadrupeds have more proximally oriented lever arms than any terrestrial quadrupeds, which suggests use of the TBM with the elbow in a more flexed position. Olecranon orientation is not consistent in suspensory hominoids, although they are all characterized by orientations that are either similar or more posterior than those observed in quadrupeds. Homo and the fossils have olecranons that are clearly more proximally oriented than expected for a quadruped of their size. This suggests that Homo and Australopithecus used their TBM in a flexed position, a position most consistent with manipulatory activities.

Fiber architecture of the intrinsic muscles of the shoulder and arm in semiterrestrial and arboreal guenons.

The internal organization of myofibers and connective tissues has important physiologic implications for muscle function and for naturalistic behavior. In this study of forelimb muscle morphology and primate locomotion, fiber architecture is examined in the intrinsic muscles of the shoulder (musculi deltoideus, infraspinatus, supraspinatus, subscapularis, teres major, and t. minor) and arm (m. coracobrachialis, biceps brachii, brachialis, and triceps brachii) in the semiterrestrial vervets (Chlorocebus aethiops) and arboreal red-tailed guenons (Cercopithecus ascanius). Wet weights and lengths of whole muscles, lengths of fasciculi and their associated proximal and distal tendons, and angles of pinnation were measured to estimate morphologic correlates of physiologic properties of individual muscles: force, velocity/excursion, energy expense, and relative isometric or isotonic contraction. Neither mean total-shoulder:total-arm ratios for muscle mass nor total reduced physiological cross-sectional area exhibited significant (P < 0.05) interspecific differences, thus emphasizing the importance of fine-tuning musculoskeletal analyses by the data collected here. The results generally support those previously published for quadriceps femoris and triceps surae of the hind limb in these species (Anapol and Barry [1996] Am. J. Phys. Anthropol. 99:429-447). The fiber architecture of the semiterrestrial vervets is largely suited for higher velocity while running on the ground. By contrast, the architectural configuration of red-tailed monkeys implies relatively isometric muscle contraction and passive storage of elastic strain energy for exploitation of the compliant canopy, where substrate components are situated beneath the sagittal plane of the animal. With respect to relative distribution of maximum potential force output among muscles of either shoulder or arm groups in these otherwise hind limb-dominated quadrupedal primates, statistically significant interspecific differences are best interpreted in light of braking, climbing, and, for vervets, the transition between ground and canopy. The interspecific differences shown here for the intrinsic muscles of the shoulder and arm underscore the significance of intramuscular morphology in reconciling structure and function with regard to locomotor behavior. Its analysis and interpretation lend support to consideration of "semiterrestrial" as a bona fide locomotor category uniquely different from what is practiced by dedicated arboreal and terrestrial quadrupeds that occasionally visit the habitat of one another. Data from a more committed terrestrial species would clarify this enigma.

Mediolateral reaction forces and forelimb anatomy in quadrupedal primates: implications for interpreting locomotor behavior in fossil primates.

The forelimb joints of terrestrial primate quadrupeds appear better able to resist mediolateral (ML) shear forces than those of arboreal quadrupedal monkeys. These differences in forelimb morphology have been used extensively to infer locomotor behavior in extinct primate quadrupeds. However, the nature of ML substrate reaction forces (SRF) during arboreal and terrestrial quadrupedalism in primates is not known. This study documents ML-SRF magnitude and orientation and forelimb joint angles in six quadrupedal anthropoid species walking across a force platform attached to terrestrial (wooden runway) and arboreal supports (raised horizontal poles). On the ground all subjects applied a lateral force in more than 50% of the steps collected. On horizontal poles, in contrast, all subjects applied a medially directed force to the substrate in more than 75% of the steps collected. In addition, all subjects on arboreal supports combined a lower magnitude peak ML-SRF with a change in the timing of the ML-SRF peak force. As a result, during quadrupedalism on the poles the overall SRF resultant was relatively lower than it was on the runway. Most subjects in this study adduct their humerus while on the poles. The kinetic and kinematic variables combine to minimize the tendency to collapse or translate forelimbs joints in an ML plane in primarily arboreal quadrupedal primates compared to primarily terrestrial quadrupedal ones. These data allow for a more complete understanding of the anatomy of the forelimb in terrestrial vs. arboreal quadrupedal primates. A better understanding of the mechanical basis of morphological differences allows greater confidence in inferences concerning the locomotion of extinct primate quadrupeds.

Vestibular evidence for the evolution of aquatic behaviour in early cetaceans

Early cetaceans evolved from terrestrial quadrupeds to obligate swimmers, a change that is traditionally studied by functional analysis of the postcranial skeleton. Here we assess the evolution of cetacean locomotor behaviour from an independent perspective by looking at the semicircular canal system, one of the main sense organs involved in neural control of locomotion. Extant cetaceans are found to be unique in that their canal arc size, corrected for body mass, is approximately three times smaller than in other mammals. This reduces the sensitivity of the canal system, most plausibly to match the fast body rotations that characterize cetacean behaviour. Eocene fossils show that the new sensory regime, incompatible with terrestrial competence, developed quickly and early in cetacean evolution, as soon as the taxa are associated with marine environments. Dedicated agile swimming of cetaceans thus appeared to have originated as a rapid and fundamental shift in locomotion rather than as the gradual transition suggested by postcranial evidence. We hypothesize that the unparalleled modification of the semicircular canal system represented a key 'point of no return' event in early cetacean evolution, leading to full independence from life on land.

Origin of human bipedalism: The knuckle-walking hypothesis revisited

Some of the most long-standing questions in paleoanthropology concern how and why human bipedalism evolved. Over the last century, many hypotheses have been offered on the mode of locomotion from which bipedalism originated. Candidate ancestral adaptations include monkey-like arboreal or terrestrial quadrupedalism, gibbon- or orangutan-like (or other forms of) climbing and suspension, and knuckle-walking. This paper reviews the history of these hypotheses, outlines their predictions, and assesses them in light of current phylogenetic, comparative anatomical, and fossil evidence. The functional significance of characteristics of the shoulder and arm, elbow, wrist, and hand shared by African apes and humans, including their fossil relatives, most strongly supports the knuckle-walking hypothesis, which reconstructs the ancestor as being adapted to knuckle-walking and arboreal climbing. Future fossil discoveries, and a clear understanding of anthropoid locomotor anatomy, are required to ultimately test these hypotheses. If knuckle-walking was an important component of the behavioral repertoire of the prebipedal human ancestor, then we can reject scenarios on the origin of bipedalism that rely on a strictly arboreal ancestor. Moreover, paleoenvironmental data associated with the earliest hominins, and their close relatives, contradict hypotheses that place the agents of selection for bipedality in open savanna habitats. Existing hypotheses must explain why bipedalism would evolve from an ancestor that was already partly terrestrial. Many food acquisition and carrying hypotheses remain tenable in light of current evidence. Yrbk Phys Anthropol 44:70–105, 2001. © 2001 Wiley-Liss, Inc.

Origin of human bipedalism: The knuckle-walking hypothesis revisited.

Some of the most long-standing questions in paleoanthropology concern how and why human bipedalism evolved. Over the last century, many hypotheses have been offered on the mode of locomotion from which bipedalism originated. Candidate ancestral adaptations include monkey-like arboreal or terrestrial quadrupedalism, gibbon- or orangutan-like (or other forms of) climbing and suspension, and knuckle-walking. This paper reviews the history of these hypotheses, outlines their predictions, and assesses them in light of current phylogenetic, comparative anatomical, and fossil evidence. The functional significance of characteristics of the shoulder and arm, elbow, wrist, and hand shared by African apes and humans, including their fossil relatives, most strongly supports the knuckle-walking hypothesis, which reconstructs the ancestor as being adapted to knuckle-walking and arboreal climbing. Future fossil discoveries, and a clear understanding of anthropoid locomotor anatomy, are required to ultimately test these hypotheses. If knuckle-walking was an important component of the behavioral repertoire of the prebipedal human ancestor, then we can reject scenarios on the origin of bipedalism that rely on a strictly arboreal ancestor. Moreover, paleoenvironmental data associated with the earliest hominins, and their close relatives, contradict hypotheses that place the agents of selection for bipedality in open savanna habitats. Existing hypotheses must explain why bipedalism would evolve from an ancestor that was already partly terrestrial. Many food acquisition and carrying hypotheses remain tenable in light of current evidence.

Climbing, brachiation, and terrestrial quadrupedalism: historical precursors of hominid bipedalism.

The vertical-climbing account of the evolution of locomotor behavior and morphology in hominid ancestry is reexamined in light of recent behavioral, anatomical, and paleontological findings and a more firmly established phylogeny for the living apes. The behavioral record shows that African apes, when arboreal, are good vertical climbers, and that locomotion during traveling best separates the living apes into brachiators (gibbons), scrambling/ climbing/brachiators (orangutans), and terrestrial quadrupeds (gorillas and chimpanzees). The paleontological record documents frequent climbing as an ancestral catarrhine ability, while a reassessment of the morphology of the torso and forelimb in living apes and Atelini suggests that their shared unique morphological pattern is best explained by brachiation and forelimb suspensory positional behavior. Further, evidence from the hand and foot points to a terrestrial quadrupedal phase in hominoid evolution prior to the adoption of bipedalism. The evolution of positional behavior from early hominoids to hominids appears to have begun with an arboreal quadrupedal-climbing phase and proceeded though an orthograde, brachiating, forelimb-suspensory phase, which was in turn followed by arboreal and terrestrial quadrupedal phases prior to the advent of hominid bipedality. The thesis that protohominids climbed down from the trees to become terrestrial bipeds needs to be reexamined in light of a potentially long history of terrestriality in the ancestral protohominid.

Morphometry and allometry of the primate humerus

The primate distal humerus has been used both in phylogenetic reconstruction and in assessing locomotor and postural adaptations. This study uses an allometric approach to predict locomotor patterns of extant primates regardless of phylogenetic position. By showing the relationship between form and function in living primate taxa it will be possible to use this data set to predict locomotor behavior of extinct primates. Several linear measurements were taken from the distal humerus of 71 extant primate species (anthropoids and prosimians). Allometric regressions of each measurement were performed with mandibular M2 area as a surrogate for body size. These measurements were used to determine if significant differences in distal humerus morphology exist among locomotor groups. The results were then used to test several hypotheses about the relationship between humeral form and function. For example, the hypothesis that suspensory primates have a large medial epicondyle is confirmed; the hypothesis that terrestrial quadrupeds have a deep olecranon fossa could not be confirmed with quantitative data. In addition to this hypothesis testing, the residuals from the allometric regressions of the humeral measurements were used in a discriminant functions analysis to estimate locomotor behavior from distal humerus morphology. The discriminant functions analysis correctly reclassified 64/71 (90%) species.

Limb skeleton and locomotor adaptations of apidium phiomense, an Oligocene anthropoid from Egypt

Apidium phiomense is the most common primate from the early Oligocene deposits of Fayum, Egypt. It is known from hundreds of dental remains and dozens of skeletal remains, including numerous representatives of the long bones of the forelimb and hindlimb. Apidium phiomense was a small (1,600 g) arboreal quadruped. The forelimb bones of this species show features characteristic of arboreal quadrupeds and lack characteristic features found in the forelimb bones of vertial clingers, terrestrial quadrupeds, or suspensory species. The pelvis and hindlimb bones show numerous adaptations for leaping from a quadrupedal position. In general, Apidium lacks characteristic features of either cercopithecoid monkeys or hominoid apes. Overall, the skeleton shows greatest similarities to the same elements of small platyrrhines such as Saimiri and is also very similar to the hypothetical morphotype for ancestral platyrrhine. The skeleton of Apidium phiomense is the most primitive anthropoid postcranial skeleton known.