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Homo floresiensis is an endemic hominin species that occupied Liang Bua, a limestone cave on Flores in eastern Indonesia, during the Late Pleistocene epoch. The skeleton of the type specimen (LB1) of H. floresiensis includes a relatively complete left foot and parts of the right foot. These feet provide insights into the evolution of bipedalism and, together with the rest of the skeleton, have implications for hominin dispersal events into Asia. Here we show that LB1's foot is exceptionally long relative to the femur and tibia, proportions never before documented in hominins but seen in some African apes. Although the metatarsal robusticity sequence is human-like and the hallux is fully adducted, other intrinsic proportions and pedal features are more ape-like. The postcranial anatomy of H. floresiensis is that of a biped, but the unique lower-limb proportions and surprising combination of derived and primitive pedal morphologies suggest kinematic and biomechanical differences from modern human gait. Therefore, LB1 offers the most complete glimpse of a bipedal hominin foot that lacks the full suite of derived features characteristic of modern humans and whose mosaic design may be primitive for the genus Homo. These new findings raise the possibility that the ancestor of H. floresiensis was not Homo erectus but instead some other, more primitive, hominin whose dispersal into southeast Asia is still undocumented.

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Bones of the lower extremity have been recovered for up to nine different individuals of Homo floresiensis - LB1, LB4, LB6, LB8, LB9, LB10, LB11, LB13, and LB14. LB1 is represented by a bony pelvis (damaged but now repaired), femora, tibiae, fibulae, patellae, and numerous foot bones. LB4/2 is an immature right tibia lacking epiphyses. LB6 includes a fragmentary metatarsal and two pedal phalanges. LB8 is a nearly complete right tibia (shorter than that of LB1). LB9 is a fragment of a hominin femoral diaphysis. LB10 is a proximal hallucal phalanx. LB11 includes pelvic fragments and a fragmentary metatarsal. LB13 is a patellar fragment, and LB14 is a fragment of an acetabulum. All skeletal remains recovered from Liang Bua were extremely fragile, and some were badly damaged when they were removed temporarily from Jakarta. At present, virtually all fossil materials have been returned, stabilized, and hardened. These skeletal remains are described and illustrated photographically. The lower limb skeleton exhibits a uniquely mosaic pattern, with many primitive-like morphologies; we have been unable to find this combination of ancient and derived (more human-like) features in either healthy or pathological modern humans, regardless of body size. Bilateral asymmetries are slight in the postcranium, and muscle markings are clearly delineated on all bones. The long bones are robust, and the thickness of their cortices is well within the ranges seen in healthy modern humans. LB1 is most probably a female based on the shape of her greater sciatic notch, and the marked degree of lateral iliac flaring recalls that seen in australopithecines such as "Lucy" (AL 288-1). The metatarsus has a human-like robusticity formula, but the proximal pedal phalanges are relatively long and robust (and slightly curved). The hallux is fully adducted, but we suspect that a medial longitudinal arch was absent.

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Several bones of the upper extremity were recovered during excavations of Late Pleistocene deposits at Liang Bua, Flores, and these have been attributed to Homo floresiensis. At present, these upper limb remains have been assigned to six different individuals - LB1, LB2, LB3, LB4, LB5, and LB6. Several of these bones are complete or nearly so, but some are quite fragmentary. All skeletal remains recovered from Liang Bua were extremely fragile, but have now been stabilized and hardened in the laboratory in Jakarta. They are now curated in museum-quality containers at the National Research and Development Centre for Archaeology in Jakarta, Indonesia. These skeletal remains are described and illustrated photographically. The upper limb presents a unique mosaic of derived (human-like) and primitive morphologies, the combination of which is never found in either healthy or pathological modern humans.

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The strain environment of the tibial midshaft of two female macaques was evaluated through in vivo bone strain experiments using three rosette gauges around the circumference of the bones. Strains were collected for a total of 123 walking and galloping steps as well as several climbing cycles. Principal strains and the angle of the maximum (tensile) principal strain with the long axis of the bone were calculated for each gauge site. In addition, the normal strain distribution throughout the cross section was determined from the longitudinal normal strains (strains in the direction of the long axis of the bone) at each of the three gauge sites, and at the corresponding cross-sectional geometry of the bone. This strain distribution was compared with the cross-sectional properties (area moments) of the midshaft. For both animals, the predominant loading regime was found to be bending about an oblique axis running from anterolateral to posteromedial. The anterior and part of the medial cortex are in tension; the posterior and part of the lateral cortex are in compression. The axis of bending does not coincide with the maximum principal axis of the cross section, which runs mediolaterally. The bones are not especially buttressed in the plane of bending, but offer the greatest strength anteroposteriorly. The cross-sectional geometry therefore does not minimize strain or bone tissue. Peak tibial strains are slightly higher than the peak ulnar strains reported earlier for the same animals (Demes et al. [1998] Am J Phys Anthropol 106:87-100). Peak strains for both the tibia and the ulna are moderate in comparison to strains recorded during walking and galloping activities in nonprimate mammals.

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Extant apes are similar to one another, and different from monkeys, in features granting them greater range of forearm rotation and greater size of the muscles that produce this motion. Although these traits may have been independently acquired by the various apes, the possibility arises that such features reflect adaptation to the stem behavior of the hominoid lineage. Anticipating that knowledge of forearm rotatory muscle recruitment during brachiation, vertical climbing, arm-hanging during feeding, and voluntary reaching might point to this stem behavior, we undertook telemetered electromyographic experiments on the supinator, pronator quadratus, ulnar head of pronator teres, and a variety of other upper limb muscles in two gibbons and four chimpanzees. The primary rotator muscles of the hominoid forearm were recruited at high levels in a variety of behaviors. As had been suspected by previous researchers, the supinator is usually active during the support phase of armswinging, but we observed numerous instances of this behavior during which the muscle was inactive. No other muscle took over its role. Kinetic analyses are required to determine how apes can execute body rotation of armswinging without active muscular effort. The one behavior that is common to most extant apes, is rare in monkeys, and which places a consistently great demand on the primary forearm rotatory muscles, is hang-feeding. The muscles of the supporting limb are essential to properly position the body; those of the free limb are essential for grasping food. Since the greater range of forearm rotation characterizing apes is also best explained by adaptation to this behavior, we join previous authors who assert that it lies at the very origin of the Hominoidea.

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Among the characteristics that are thought to set primate quadrupedal locomotion apart from that of nonprimate mammals are a more protracted limb posture and larger limb angular excursion. However, kinematic aspects of primate or nonprimate quadrupedal locomotion have been documented in only a handful of species, and more widely for the hind than the forelimb. This study presents data on arm (humerus) and forelimb posture during walking for 102 species of mammals, including 53 nonhuman primates and 49 nonprimate mammals. The results demonstrate that primates uniformly display a more protracted arm and forelimb at hand touchdown of a step than nearly all other mammals. Although primates tend to end a step with a less retracted humerus, their total humeral or forelimb angular excursion exceeds that of other mammals. It is suggested that these features are components of functional adaptations to locomotion in an arboreal habitat, using clawless, grasping extremities.

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Using telemetered electromyography and immunocytochemical fibre typing (of both fresh frozen and preserved specimens), the present paper demonstrates clearly that at the elbow, knee, and ankle joints, the rhesus monkey (Macaca mulatta) is endowed with one extensor-muscle head specialized for posture. These postural heads are distinguished by (a) recruitment at low levels to maintain joint position against the effect of gravity, (b) recruitment near maximum levels during walking, and (c) high content and relatively large size of slow, fatigue-resistant (type I) muscle fibres. The nonpostural heads of the investigated muscles were recruited at levels correlated to the strenuousness of the effort and are notable by the small percentage and size of slow muscle fibres. The postmortem stability of the structural properties of myosin makes immunocytochemical fibre typing suitable for the study of preserved cadavers.

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In vivo bone strain experiments were performed on the ulnae of three female rhesus macaques to test how the bone deforms during locomotion. The null hypothesis was that, in an animal moving its limbs predominantly in sagittal planes, the ulna experiences anteroposterior bending. Three rosette strain gauges were attached around the circumference of the bone slightly distal to midshaft. They permit a complete characterization of the ulna's loading environment. Strains were recorded during walking and galloping activities. Principal strains and strain directions relative to the long axis of the bone were calculated for each gauge site. In all three animals, the lateral cortex experienced higher tensile than compressive principal strains during the stance phase of walking. Compressive strains predominated at the medial cortex of two animals (the gauge on this cortex of the third animal did not function). The posterior cortex was subject to lower strains; the nature of the strain was highly dependent on precise gauge position. The greater principal strains were aligned closely with the long axis of the bone in two animals, whereas they deviated up to 45 degrees from the long axis in the third animal. A gait change from walk to gallop was recorded for one animal. It was not accompanied by an incremental change in strain magnitudes. Strains are at the low end of the range of strain magnitudes recorded for walking gaits of nonprimate mammals. The measured distribution of strains in the rhesus monkey ulna indicates that mediolateral bending, rather than anteroposterior bending, is the predominant loading regime, with the neutral axis of bending running from anterior and slightly medial to posterior and slightly lateral. A variable degree of torsion was superimposed over this bending regime. Ulnar mediolateral bending is apparently caused by a ground reaction force vector that passes medial to the forearm. The macaque ulna is not reinforced in the plane of bending. The lack of buttressing in the loaded plane and the somewhat counterintuitive bending direction recommend caution with regard to conventional interpretations of long bone cross-sectional geometry.

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

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In this report we provide detailed data on the patterns and frequency of heel contact with terrestrial and arboreal supports in primates. These data can help resolve the question of whether African apes and humans are uniquely "plantigrade" (Gebo [1992] Am. J. Phys. Anthropol. 89:29-58; Gebo [1993a] Am. J. Phys. Anthropol. 91:382-385; Gebo [1993b] Postcranial Adaptation in Nonhuman Primates), or if plantigrady is common in other primates (Meldrum [1993] Am. J. Phys. Anthropol. 91:379-381). Using biplanar and uniplanar videotapes, we recorded the frequency and timing of heel contact for a variety of primates (32 species) walking on the ground and on simulated arboreal supports at a range of natural speeds. Our results indicate that Pongo as well as the African apes exhibit a "heel-strike" at the end of swing phase. Ateles and Hylobates make heel contact on all supports shortly after mid-foot contact, although spider monkeys do so only at slow or moderate speeds. Data available from uniplanar videotapes suggest that this pattern occurs in Alouatta and Lagothrix as well. No other New or Old World monkey or prosimian in this study made heel contact during quadrupedalism on any substrate. Thus, heel contact occurs in all apes and atelines, but only the great apes exhibit a heel-strike. We suggest that heel contact with the substrate is a by-product of an active posterior weight-shift mechanism involving highly protracted hindlimbs at touchdown. Force plate studies indicate that this mechanism is most extreme in arboreally adapted primate quadrupeds walking on arboreal supports. Although heel contact and heel-strike may have no evolutionary link, it is possible that both patterns are the result of a similar weight shift mechanism. Therefore, the regular occurrence of heel contact in a variety of arboreal primates, and the absence of a true biomechanical link between limb elongation, heel contact, and terrestriality, calls into question the claim that hominid foot posture was necessarily derived from a quadrupedal terrestrial ancestor.

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Unlike all other primates, the digastric muscle of the orangutan lacks an anterior belly; the posterior belly, while present, inserts directly onto the mandible. To understand the functional consequences of this morphologic novelty, the EMG activity patterns of the digastric muscle and other potential mandibular depressors were studied in a gibbon and an orangutan. The results suggest a significant degree of functional differentiation between the two digastric bellies. In the gibbon, the recruitment pattern of the posterior digastric during mastication is typically biphasic. It is an important mandibular depressor, active in this role during mastication and wide opening. It also acts with the anterior suprahyoid muscles to move the hyoid prior to jaw opening during mastication. The recruitment patterns of the anterior digastric suggest that it is functionally allied to the geniohyoid and mylohyoid. For example, although it transmits the force of the posterior digastric during mandibular depression, it functions independent of the posterior digastric during swallowing. Of the muscles studied, the posterior digastric was the only muscle to exhibit major differences in recruitment pattern between the two species. The posterior digastric retains its function as a mandibular depressor in orangutans, but is never recruited biphasically, and is not active prior to opening. The unique anatomy of the digastric muscle in orangutans results in decoupling of the mechanisms for hyoid movement and mandibular depression, and during unilateral activity it potentially contributes to substantial transverse movements of the mandible. Hypotheses to explain the loss of the anterior digastric should incorporate these functional conclusions.

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Differences in the degree of projection of the greater tubercle above the level of the humeral head in primate proximal humeri have been associated with differing leverage requirements for supraspinatus during arboreal vs. terrestrial quadrupedal locomotion. Since most workers have assumed that supraspinatus acts as a humeral protractor, interpretations of the variation in greater tubercle height have focused on the need for powerful vs. rapid humeral protraction during the swing phase of quadrupedal locomotion. However, in an EMG study on the activity patterns of supraspinatus in the vervet monkey, Larson and Stern (Am. J. Phys. Anthropol. 79:369-377, 1989) reported that although supraspinatus is active during arm elevations against gravity, it is silent during the swing phase of quadrupedal locomotion, and instead acts as a joint stabilizer during support phase. They suggested that the pattern of activity for supraspinatus observed in the vervet was common for all quadrupedal primates, and that differences in greater tubercle projection could be related to the degree of mobility of the shoulder. In the current study, we present additional EMG data on a baboon and three macaques supporting the suggestions offered by Larson and Stern (1989).

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The importance of arm-raising has been a major consideration in the functional interpretation of differences in shoulder morphology among species of nonhuman primates. Among the characters that have been associated with enhancement of the arm-raising mechanism in hominoid primates are the relative enlargement of cranial trapezius and caudal serratus anterior, as the main scapular rotators, as well as changes in scapular morphology associated with their improved leverage for scapular rotation. Yet in an EMG study of cranial trapezius and caudal serratus anterior function in the great apes, Tuttle and Basmajian (Yrbk. Phys. Anthropol. 20:491-497, 1977) found these muscles to be essentially inactive during arm-raising. Although Tuttle and Basmajian suggest that the cranial orientation of the glenoid fossa in apes has reduced the demand for scapular rotation during arm-raising, subsequent EMG studies on other primate species suggest that these muscles do play a significant role in arm motion during active locomotion. This paper presents a reexamination of muscle recruitment patterns for trapezius and caudal serratus anterior in the chimpanzee. All but the lowest parts of caudal serratus anterior were found to be highly active during arm-raising motions, justifying earlier morphological interpretations of differences in caudal serratus anterior development. The lowest digitations of this muscle, while inactive during arm-raising, displayed significant activity during suspensory postures and locomotion, presumably to control the tendency of the scapula to shift cranially relative to the rib cage. Cranial trapezius did not appear to be involved in arm-raising; instead, its recruitment was closely tied to head position.

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In comparative anatomical studies of the shoulder, the humeral retractors are often grouped together as propulsive muscles, which are important in the propulsive stroke of the forelimb during quadrupedal locomotion. Electromyographic (EMG) analyses of these muscles in opossums, cats, and dogs in general have confirmed such conclusions. An EMG study of chimpanzee shoulder muscles during knuckle-walking found, however, that the humeral retractors are either inactive or perform a function unrelated to propulsion (Larson & Stern, 1987). This contrast in muscle recruitment patterns between chimpanzees and more "typical" mammalian quadrupeds was attributed to the derived morphology of the chimpanzee shoulder. The present study examines the activity patterns of the humeral retractors in the vervet monkey, a primate more closely resembling nonprimate mammals in its shoulder morphology. The results of this EMG analysis show that despite the significant differences in anatomy between chimpanzees and vervets, the two species display very similar muscle recruitment patterns during quadrupedalism, and there is evidence for this same pattern in other species of primates. These differences in muscle activity patterns between primates and nonprimate mammals may be related to changes in the neurological control of locomotion in primates due to the evolutionary development of manipulative abilities in the primate forelimb.

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Various researchers have noted that terrestrial and arboreal monkeys often differ in the degree to which the greater tubercle of the humerus projects proximally beyond the profile of the humeral head. These differences have been associated with differing leverage requirements for supraspinatus during quadrupedal locomotion. Although Jolly (The Baboon in Medical Research, Vol. II., pp. 23-50, 1967) suggested that a projecting tubercle can be related to the ability of supraspinatus to control more exactly the passive humeral retraction that occurs during support phase of quadrupedalism, most workers emphasize a link between this bony trait and the need for powerful or rapid protraction of the forelimb during swing phase of terrestrial quadrupedalism. We report on an EMG analysis of supraspinatus function showing that the muscle does not act as a brachial protractor during quadrupedalism, even in the relatively cursorial vervet monkey. We suggest that differences in greater tubercle projection can be related to the degree of mobility of the shoulder, and that supraspinatus size is determined by the interaction of greater tubercle height, adaptive importance of brachial elevation, and body size.

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Current views on the function of the deltoid and rotator cuff muscles emphasize their roles in arm-raising as participants in a scapulohumeral force "couple." The acceptance of such a mechanism is based primarily on a 1944 EMG study of human shoulder muscle action. More recently, it has been suggested that shoulder joint stabilization constitutes a second and equally important function of the cuff musculature, especially in nonhuman primates which habitually use their forelimbs in overhead postural and locomotor activities. Few comparative data exist, however, on the actual recruitment patterns of these muscles in different species. In order to assess the general applicability of a scapulohumeral force couple model, and the functional significance of the differential development of the scapulohumeral musculature among primate species, we have undertaken a detailed study of shoulder muscle activity patterns in nonhuman primates employing telemetered electromyography, which permits examination of unfettered natural behaviors and locomotion. The results of our research on the chimpanzee, Pan troglodytes, on voluntary reaching and two forms of "arboreal" locomotion reveal four ways in which previous perceptions of the function of the scapulohumeral muscles must be revised: 1) the posterior deltoid is completely different in function from the middle and anterior regions of this muscle; 2) the integrity of the glenohumeral joint during suspensory postures is not maintained solely by osseoligamentous structures; 3) the function of teres minor is entirely different from that of the other rotator cuff muscles and is more similar to the posterior deltoid and teres major; and 4) each remaining member of the rotator cuff plays a distinct, and often unique, role during natural behaviors. These results clearly refute the view that the muscles of the rotator cuff act as a single functional unit in any way, and an alternative to the force couple model is proposed.

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The importance of allometry as an analytic tool is well recognized in the literature of primate morphology. However, a number of recent studies have illustrated how interpretive difficulties can arise when researchers confound different types of allometric data. Such confusion is due less to carelessness than to uncertainty about how different types of allometry are related. The present study examines the relationship between two types--ontogenetic and interspecific allometry--in the case of organ weight scaling in six species of Old World monkeys. Accepting the interpretation of interspecific allometry as a reflection of functional scaling constraints, the results of this analysis indicate how ontogenetic patterns have been modified in different-sized species to maintain compliance with these constraints. Specifically, for the heart and lungs it appears that vertical transpositions of individual species' ontogenies are dictated by isometric interspecific allometry, while in the case of the kidneys and liver, the relation of negative allometry across species entails alteration of the relative growth coefficients of the individual species. While these conclusions can at present only be applied to organ weight scaling, the approach of examining interspecific patterns in light of developmental differences between species should prove very helpful in our efforts to understand the phenomena of size and scaling.

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Heart weight scaling among members of the primate tribe Papionini is analyzed in relation to the proposal that evolutionary size change may sometimes come about in such a way that geometric similarity is maintained across species. The results indicate that in terms of relative heart weight, the six species examined can be divided into two groups of geometric variants. Although these groupings do not conform to any hypothesized phylogenetic relationships, there is some evidence to support the associations based on shared patterns of ontogenetic development. An account is proposed for the differences between the two groups in terms of derived versus primitive patterns of relative heart weight growth.

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The allometric scaling of nine internal organs was examined for Macaca arctoides. Significant organ weight-body weight regressions were obtained for heart, lungs, kidneys, pancreas, thyroid, liver, and testes. The spleen and adrenal glands exhibited strong variability and were only loosely correlated to body weight. Using allometry as a criterion of subtraction, observed sex differences in mean organ weights were seen to be primarily the result of differences in average body weight. It is postulated that analysis of observed differences in organ weights between this species and Macaca mulatta would yield similar conclusions. Comparison of intraspecific slope values obtained in the present study with interspecific values reported in the literature reveals a pattern paralleling the brain-body weight relation. A discussion of the relationship between intra- and interspecific slopes is presented.

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