Locomotor Repertoire Research Articles

Cutting corners: the dynamics of turning behaviors in two primate species

In an attempt to characterize more fully the variation in substrate reaction forces in the locomotor repertoire of primates, we recorded the forces involved in directional changes for two species. These are the first records of turning forces for vertebrate quadrupeds, much less primates. Three ring-tailed lemurs and two patas monkeys performed turns of approximately 30 degrees as they crossed a force platform. The ring-tailed lemurs also turned on a horizontal branch-like support with a segment attached to the force transducer. Mediolateral forces of up to 40% body weight were recorded. These are considerably higher than during linear locomotion. Pivot limbs in ground turns and turns on the branch differed in the lemurs, suggesting that substrate influences turning strategies. Limbs encountered both medial and lateral reaction forces, and as a result, they may be exposed to variable bending regimes in the frontal plane. The stereotypy in bending regimes suggested by in vivo bone strain studies, therefore, may characterize linear locomotion only. The lemurs showed hindlimb dominance in turns, both in terms of frequency used as well as force magnitude (hindlimb steering). Hindlimb dominance in weight support characterizes both species (and primates in general), but it is more pronounced in the lemurs. In the patas monkeys, forces were more evenly distributed among the two pairs of limbs. The mediolateral turning forces therefore seem to track the amount of weight to be shifted sideways. Overall variance in mediolateral forces was greater in the arboreal and versatile lemurs than in the terrestrial and cursorial patas monkeys.

Locomotor network modeling based on identified zebrafish neurons

The larval zebrafish generates a discrete set of locomotor maneuvers, each with distinctive bending patterns and tail-beat frequencies (TBFs). It is not known how these locomotor patterns are generated. We had previously shown that aspects of the locomotor repertoire could be modeled with a simple 2-cell segmental oscillator replicated in series to simulate the 30 segment larval spinal cord. This model, however, conflicted with known features of the spinal circuitry and was not able to produce the natural whole-cord activity patterns. We present here three new more realistic CPG models which incorporate anatomical and neurotransmitter features of identified zebrafish spinal interneurons. These whole-cord models were able to produce oscillatory rhythms across the range of natural TBFs in ways that the simpler model could not.

Postcranial Remains of the Extinct Monkeys of the Greater Antilles, with Evidence for Semiterrestriality in Paralouatta1

This paper describes postcranial remains pertaining to the endemic xenotrichin callicebines of the Greater Antilles, all of which are extinct: Xenothrix mcgregori (Jamaica), Antillothrix bernensis (Hispaniola), and Paralouatta varonai and P. marianae (Cuba). These monkeys differed considerably in body size and inferred locomotor behavior. Xenothrix and Antillothrix are estimated to have weighed 2–5 kg, which is well within the middle range of body sizes found in extant South American monkeys, but Paralouatta (~9–10 kg) would have been nearly as large as the largest living platyrrhines. In line with previous studies, we interpret Xenothrix mcgregori as a rather short-limbed, slow-moving arboreal quadruped possessing some unusual features not otherwise seen in platyrrhines (e.g., adductor process or “fourth trochanter” of the femur). Its closest locomotor analog among living primates remains uncertain. Paralouatta varonai also exhibits features not seen in other platyrrhines, but in this case there are intriguing resemblances to certain Old World monkeys (e.g., retroflexed medial epicondyle and narrow trochlea on humerus, stabilization features of talocrural joint, short digital rays), especially so-called semiterrestrial cercopithecines whose locomotor repertoire includes a significant amount of movement on the ground (e.g., Cercopithecus lhoesti). At the same time, the Cuban monkey conspicuously lacks most features uniquely connected with suspensory activities, otherwise seen in all living platyrrhines of large body size. The locomotor and postural repertoire of Antillothrix is unresolved, as the only element currently available for analysis is a distal tibia. The tibia of the Hispaniolan monkey is not very informative from a functional standpoint, although it exhibits less emphasis on talocrural stabilization than does the equivalent element in Paralouatta (e.g., size of medial malleolus). The diverse postcranial specializations exhibited by xenotrichins are consistent with their long isolation (at least since Oligocene) on land masses in the Caribbean Sea.

The cross-sectional geometry of the hand and foot bones of the Hominoidea and its relationship to locomotor behavior

Cheiridia are valuable indicators of positional behavior, as they directly contact the substrate, but systematic comparison of the structural properties of both metacarpals and metatarsals has never been carried out. Differences in locomotor behavior among the great apes (knuckle-walking vs. quadrumanous climbing) can produce biomechanical differences that may be elucidated by the parallel study of cross-sectional characteristics of metacarpals and metatarsals. The aim of this work is to study the cross-sectional geometric properties of these bones and their correlation with locomotor behavior in large-bodied hominoids. The comparisons between bending moments of metacarpals and metatarsals of the same ray furnished interesting results. Metacarpals III and especially IV of the knuckle-walking African apes were relatively stronger than those of humans and orangutans, and metatarsal V of humans was relatively stronger than those of the great apes. Interestingly, the relative robusticity of the metacarpal IV of the quadrumanous orangutan was between that of the African apes and that of humans. The main conclusions of the study are: 1) cross-sectional dimensions of metacarpals and metatarsals are influenced by locomotor modes in great apes and humans; 2) interlimb comparisons of cross-sectional properties of metacarpals and metatarsals are good indicators of locomotor modes in great apes and humans; and 3) the results of this study are in accord with those of previous analyses of plantar pressure and morphofunctional traits of the same bones, and with behavioral studies. These results provide a data base from which it will be possible to compare the morphology of the fossils in order to gain insight into the locomotor repertoires of extinct taxa.

Movement control strategies during jumping in a lizard ( Anolis valencienni)

Whereas maximal performance is subjected to specific control criteria, sub-maximal movements theoretically allow for an infinite number of control strategies. Yet sub-maximal movements are predominant in the locomotor repertoire of most organisms and often little understood. Previous data on sub-maximal vertical jumping in humans has suggested that a movement effectiveness criterion might best explain the observed control strategy employed. Here we test the generality of this criterion in jumping by inducing lizards to jump both at a range of distances as well as a range of take-off angles. Our results show that while movement effectiveness appears to best explain jumping for different take-off angles, a ‘push harder’ strategy (i.e. mostly increasing the force output of the system), is used in the control of distance jumping. Thus, our data support the generality of the movement effectiveness criterion for vertical jumping, but not for distance jumping. Sub-maximal distance jumping in the lizard Anolis valencienni appears to be governed by a relatively simple control strategy that allows a rapid response. This accords well to the ecological circumstances in which long jumps are typically used (escape from predators).

The Oreopithecus thumb: a strange case in hominoid evolution

Pliopithecoids are a diverse group of Miocene catarrhine primates from Eurasia. Their positional behavior is still unknown, and many species are known exclusively from dentognathic remains. Here, we describe a proximal radius (IPS66267) from the late Miocene of Castell de Barberà (Vallès-Penedès Basin, NE Iberian Peninsula) that represents the first postcranial specimen of the pliopithecoid Barberapithecus huerzeleri. A body mass estimate based on the radius is compared with dental estimates, and its morphology is compared with that of extant and fossil anthropoids by qualitative means as well as by landmark-based three-dimensional geometric morphometrics. The estimated body mass of ∼5 kg for IPS66267 closely matches the dental estimates for the (female) holotype, thereby discounting an alternative attribution to the large-bodied hominoid recorded at Castell de Barberà. In multiple features (oval and moderately tilted head with a pronounced lateral lip and a restricted articular area for the capitulum; proximodistally expanded proximal radioulnar joint; and short, robust, and anteroposteriorly compressed neck), the specimen differs from hominoids and resembles instead extant nonateline monkeys and stem catarrhines. The results of the morphometric analysis further indicate that the Barberapithecus proximal radius shows closer similarities with nonsuspensory arboreal cercopithecoids and the dendropithecid Simiolus. From a locomotor viewpoint, the radius of Barberapithecus lacks most of the features functionally related to climbing and/or suspensory behaviors and displays instead a proximal radioulnar joint that would have been particularly stable under pronation. On the other hand, the Barberapithecus radius differs from other stem catarrhines in the less anteroposteriorly compressed and less tilted radial head with a deeper capitular fovea, suggesting a somewhat enhanced mobility at the elbow joint. We conclude that pronograde arboreal quadrupedalism was the main component of the locomotor repertoire of Barberapithecus but that, similar to other crouzeliids, it might have displayed better climbing abilities than pliopithecids.

The hands and feet of Archaeolemur: metrical affinities and their functional significance

Recent expeditions to Madagascar have recovered abundant skeletal remains of Archaeolemur, one of the so-called “monkey lemurs” known from Holocene deposits scattered across the island. These new skeletons are sufficiently complete to permit reassembly of entire hands and feet—postcranial elements crucial to drawing inferences about substrate preferences and positional behavior. Univariate and multivariate analysis of intrinsic hand and foot proportions, phalangeal indices, relative pollex and hallux lengths, phalangeal curvature, and distal phalangeal shape reveal a highly derived and unique morphology for an extinct strepsirrhine that diverges dramatically from that of living lemurs and converges in some respects on that of Old World monkeys (e.g., mandrills, but not baboons or geladas). The hands and feet of Archaeolemur are relatively short (extremely so relative to body size); the carpus and tarsus are both “long” relative to total hand and foot lengths, respectively; phalangeal indices of both the hands and feet are low; both pollex and hallux are reduced; the apical tufts of the distal phalanges are very broad; and the proximal phalanges are slightly curved (but more so than in baboons). Overall grasping capabilities may have been compromised to some extent, and dexterous handling of small objects seems improbable. Deliberate and noncursorial quadrupedalism was most likely practiced on both the ground and in the trees. A flexible locomotor repertoire in conjunction with a eurytopic trophic adaptation allowed Archaeolemur to inhabit much of Madagascar and may explain why it was one of the latest surviving subfossil lemurs.

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.

The Tarsal Complex of Afro-Malagasy Tenrecoidea: A Search for Phylogenetically Meaningful Characters

Morphological characters should be assessed in an ecological and evolutionary framework before their use in phylogenetic analyses. We have attempted such an assessment for the tarsal complex of the Tenrecoidea. The 30+/− extant species of these small mammals live in a diverse range of microhabitats. They exhibit markedly different positional behaviors, encompassing four basic locomotor repertoires: terrestrial running and walking, scansorial climbing, digging, and swimming. Articular surfaces from the upper, middle, and lower ankle joints in 10 Malagasy tenrecid, 1 potamogalid, 1 solenodontid, and 1 macroscelidid species were compared. The results were tested against published data on the correlation between function and morphology in other therian locomotor specialists. Descriptive accounts, supported by quantitative analyses, demonstrate significant differences in many aspects of tarsal morphology that may be explained by function-based hypotheses within the context of the tenrecoid heritage. Three closely related tenrecines (Hemicentetes, Setifer, and Echinops) show divergences in form that are consistent with their respective semifossorial, terrestrial, and scansorial/arboreal modes of locomotion. In addition to functional–adaptive considerations, we propose several synapomorphies for the tenrecine and oryzorictine clades that appear to corroborate their monophyly. There are apparent convergences between the habitual diggers from different subfamilies (Hemicentetes and Oryzorictes), and there are adaptive differences within subfamilies (e.g. arboreal Echinops vs. terrestrial Setifer). The few likenesses between Potamogale and Limnogale cannot be supported as homologies, and the proposal of a recent phylogenetic affiliation between them is therefore rejected here. Manifest differences in tarsal form between Geogale and the oryzorictines support recognition of the subfamily Geogalinae. As expected, the tarsal complex of Solenodon is fundamentally unlike that of the tenrecoids. Finally, the similarities in detail between Macroscelididae and Potamogalidae reflect the stabilization of UAJ during plantarflexion, and therefore such attributes are rejected as synapomorphies. Traits with clear species-specific adaptations are a potential interference in cladistic analyses and cannot be meaningfully used without ecology-based character assessment. While this practice may ultimately reduce the size of a database, it will nonetheless result in taxonomic properties with lasting value against which phylogenetic hypotheses may be tested with confidence.

Tail loss in Proconsul heseloni

The middle Miocene stem kenyapithecine Nacholapithecus kerioi (16–15 Ma; Nachola, Kenya) is represented by a large number of isolated fossil remains and one of the most complete skeletons in the hominoid fossil record (KNM-BG 35250). Multiple fieldwork seasons performed by Japanese–Kenyan teams during the last part of the 20th century resulted in the discovery of a large sample of Nacholapithecus fossils. Here, we describe the new femoral remains of Nacholapithecus. In well-preserved specimens, we evaluate sex differences and within-species variation using both qualitative and quantitative traits. We use these data to determine whether these specimens are morphologically similar to the species holotype KNM-BG 35250 (which shows some plastic deformation) and to compare Nacholapithecus with other Miocene hominoids and extant anthropoids to evaluate the distinctiveness of its femur. The new fossil evidence reaffirms previously reported descriptions of some distal femoral traits, namely the morphology of the patellar groove. However, results also show that relative femoral head size in Nacholapithecus is smaller, relative neck length is longer, and neck–shaft angle is lower than previously reported for KNM-BG 35250. These traits have a strong functional signal related to the hip joint kinematics, suggesting that the morphology of the proximal femur in Nacholapithecus might be functionally related to quadrupedal-like behaviors instead of more derived antipronograde locomotor modes. Results further demonstrate that other African Miocene apes (with the exception of Turkanapithecus kalakolensis) generally fall within the Nacholapithecus range of variation, whose overall femoral shape resembles that of Ekembo spp. and Equatorius africanus. Our results accord with the previously inferred locomotor repertoire of Nacholapithecus, indicating a combination of generalized arboreal quadrupedalism combined with other antipronograde behaviors (e.g., vertical climbing).

Function of pectoral fins in rainbow trout: behavioral repertoire and hydrodynamic forces.

Salmonid fishes (trout, salmon and relatives) have served as a model system for study of the mechanics of aquatic animal locomotion, yet little is known about the function of non-axial propulsors in this major taxonomic group. In this study we examine the behavioral and hydromechanical repertoire of the paired pectoral fins of rainbow trout Oncorhynchus mykiss, performing both steady rectilinear swimming and unsteady maneuvering locomotion. A combination of kinematic analysis and quantitative flow visualization (using digital particle image velocimetry) enables identification of the propulsive roles played by pectoral fin motions. During constant-speed swimming (0.5 and 1.0 body length s(-1)), the pectoral fins remain adducted against the body. These fins are actively recruited, however, for a variety of maneuvering behaviors, including station holding in still water (hovering), low-speed (i.e. non-fast-start) turning, and rapid deceleration of the body during braking. Despite having a shallow pectoral-fin base orientation (the plesiomorphic teleost condition), trout are capable of rotating the fin base over 30 degrees during maneuvering, which affords the fin an impressive degree of kinematic versatility. When hovering, the pectoral fins are depressed beneath the body and twisted along their long axes to allow anteroposterior sculling. During turning and braking, the fins undergo spanwise rotation in the opposite direction and exhibit mediolateral and dorsoventral excursions. Water velocity fields and calculated momentum flows in the wake of the pectoral fins reveal that positive thrust is not generated during maneuvering, except during the retraction half-stroke of hovering. Relatively large laterally directed fluid force (mean 2.7 mN) is developed during turning, whose reaction powers yawing rotation of the body (4-41 degrees s(-1)). During deceleration, the wake-force line of action falls below the center of mass of the body, and this result supports a long-standing mechanical model of braking by fishes with ventrally positioned paired fins. Despite its traditional categorization as a propulsor of limited functional importance, the salmoniform pectoral fin exhibits a diverse locomotor repertoire comparable to that of higher teleostean fishes.

New Sivapithecus postcranial specimens from the Siwaliks of Pakistan.

Several new postcranial elements of Sivapithecus from the Siwaliks of Pakistan are described. These include a distal femur from the U-level of the Dhok Pathan Formation, a navicular from the Chinji Formation, and seven manual and pedal phalanges from the Nagri Formation. The functional morphology of these elements adds new detail to the reconstruction of Sivapithecus positional behavior. Femoral cross-sectional geometry indicates that the shaft was adapted to support mediolaterally directed loading. Femoral condylar asymmetry and a broad but shallow trochlea are distinctly ape-like, revealing capabilities for both rotation and withstanding eccentric loading in the knee. The navicular is characterized by features relating to a broad mid-tarsus and broad distal articulations for the cuneiforms. It also lacks a navicular tubercle as in Pongo. These features suggest that the foot was capable of a powerful grip on large supports, with an inversion/supination capability that would permit foot placement in a variety of positions. The morphology of the new phalanges, including evidence for a relatively large pollex, similarly suggests powerful grasping, consistent with prior evidence from the hallux and tarsus. The functional features of the new specimens permit refinement of previous interpretations of Sivapithecus positional capabilities. They suggest a locomotor repertoire dominated by pronograde activities and also such antipronograde activities as vertical climbing and clambering, but not by antipronograde suspensory activities as practiced by extant apes.

Wake dynamics and fluid forces of turning maneuvers in sunfish.

While experimental analyses of steady rectilinear locomotion in fishes are common, unsteady movement involving time-dependent variation in heading, speed and acceleration probably accounts for the greatest portion of the locomotor time budget. Turning maneuvers, in particular, are key elements of the unsteady locomotor repertoire of fishes and, by many species, are accomplished by generating asymmetrical forces with the pectoral fins. The development of such left-right asymmetries in force production is a critical and as yet unstudied aspect of aquatic locomotor dynamics. In this paper, we measure the fluid forces exerted by the left and right pectoral fins of bluegill sunfish (Lepomis macrochirus) during turning using digital particle image velocimetry (DPIV). DPIV allowed quantification of water velocity fields, and hence momentum, in the wake of the pectoral fins as sunfish executed turns; forces exerted during turning were compared with those generated by the immediately preceding fin beats during steady swimming. Sunfish generate the forces required for turning by modulating two variables: wake momentum and pectoral fin stroke timing. Fins on opposite sides of the fish play functionally distinct roles during turning maneuvers. The fin nearer the stimulus inducing the turn (i.e. the strong side fin) generates a laterally oriented vortex ring with a strong central jet whose associated lateral force is four times greater than that produced during steady swimming. Little posterior (thrust) force is generated by the strong-side fin, and this fin therefore acts to rotate the body away from the source of the stimulus. The contralateral (weak-side) fin generates a posteriorly oriented vortex ring with a thrust force nine times that produced by the fin during steady swimming. Minimal lateral force is exerted by the weak-side fin, and this fin therefore acts primarily to translate the body linearly away from the stimulus. Turning with the paired fins is not simply steady swimming performed unilaterally. Instead, turning involves asymmetrical fin movements and fluid forces that are distinct in both direction and magnitude from those used to swim forward at constant speed. These data reflect the plasticity of the teleost pectoral fin in performing a wide range of steady and unsteady locomotor tasks.

Locomotor repertoire of the larval zebrafish: swimming, turning and prey capture.

Larval zebrafish (Brachydanio rerio) are a popular model system because of their genetic attributes, transparency and relative simplicity. They have approximately 200 neurons that project from the brainstem into the spinal cord. Many of these neurons can be individually identified and laser-ablated in intact larvae. This should facilitate cellular-level characterization of the descending control of larval behavior patterns. Towards this end, we attempt to describe the range of locomotor behavior patterns exhibited by zebrafish larvae. Using high-speed digital imaging, a variety of swimming and turning behaviors were analyzed in 6- to 9-day-old larval fish. Swimming episodes appeared to fall into two categories, with the point of maximal bending of the larva's body occurring either near the mid-body (burst swims) or closer to the tail (slow swims). Burst swims also involved larger-amplitude bending, faster speeds and greater yaw than slow swims. Turning behaviors clearly fell into two distinct categories: fast, large-angle escape turns characteristic of escape responses, and much slower routine turns lacking the large counterbend that often accompanies escape turns. Prey-capture behaviors were also recorded. They were made up of simpler locomotor components that appeared to be similar to routine turns and slow swims. The different behaviors observed were analyzed with regard to possible underlying neural control systems. Our analysis suggests the existence of discrete sets of controlling neurons and helps to explain the need for the roughly 200 spinal-projecting nerve cells in the brainstem of the larval zebrafish.

New skeletal remains of Omomys(Primates, Omomyidae): functional morphology of the hindlimb and locomotor behavior of a Middle Eocene primate

New associated craniodental and postcranial remains of Omomys carteri from Bridger C beds in Uinta County, Wyoming represent the largest and most nearly complete single taxon sample of omomyid postcranial fossils presently known. They allow, for the first time, a description and detailed analysis of the functional morphology of the hindlimb of this middle Eocene omomyine primate. Comparisons of pelvic, femoral, tibial, and pedal morphology and metrics with a large sample of prosimian primates of known locomotor behavior suggest that Omomys possessed a highly distinctive mosaic of functional adaptations related to active quadrupedalism and leaping. Traits suggestive of quadrupedalism include the lengths of the ischium and ilium, position of the femoral trochanters, and lengths and features of the tarsal bones. Morphological traits that suggest leaping include a semi-cylindrical femoral head with moderate posterior expansion of the articular surface, greater trochanter projecting anterior to the femoral shaft, deeper than wide femoral condyles, narrow and deep patellar groove with prominent lateral border, elongated calcaneus, and close aposition of distal tibia and fibula. While Omomys most closely resembles active quadrupedal cheirogaleids like Cheirogaleus and Mirza, leaping must have been an important component of its locomotor repertoire. In this respect, Omomys closely resembles other North American omomyines (notablyHemiacodon ), and is significantly more generalized postcranially than European microchoerines (e.g.,Microchoerus , Nannopithex, and Necrolemur).

A partial skeleton of Proteopithecus sylviae (Primates, Anthropoidea): first associated dental and postcranial remains of an Eocene anthropoidean

Recent excavation in the Late Eocene quarry L-41 (Fayum Depression, Egypt) revealed two tibiae and a femur in direct association with a mandible of Proteopithecus sylviae, arguably the most generalized African anthropoidean known from cranial remains. This discovery represents the first association of dental and postcranial material belonging to an Eocene anthropoidean, and provides new insights into the functional anatomy and phylogenetic position of Proteopithecus. The hindlimb morphology of Proteopithecus is most similar to small-bodied platyrrhines among living and extinct primates and is consistent with a locomotor repertoire that included a considerable amount of running and pronograde leaping. In certain dental and postcranial features, Proteopithecus differs from the other Fayum anthropoideansand shows a greater resemblance to living and extinct platyrrhines, but it is unclear whether these features are of particular phylogenetic significance.

Ontogeny and phylogeny of femoro-tibial characters in humans and hominid fossils: functional influence and genetic determinism.

Three different human femoro-tibial characters are selected as functionally relevant and derived hominid characters: femoral bicondylar angle, shape of the femoral distal epiphysis, and the tibial insertion of the lateral meniscus. The timing and mode of formation of these characters are investigated during human ontogeny and are shown to differ considerably. The available hominid fossils (Australopithecus afarensis and early Homo) are interpreted in the light of this ontogenetic analysis with the conclusion that, during hominid evolution, different modes of selection of these features must have occurred. In modern humans, the femoral bicondylar angle proves to be an epigenetic functional feature, which develops during early childhood growth. It is present in all australopithecines and we suggest that it developed following a change in their locomotor behavior and not upon a genomic change: the early practice of bipedal walking, with adducted knee joints, in the locomotor repertoire of infant australopithecines, was sufficient to promote this angle. Later in hominid evolution, the knee joint evolved from having a single insertion of the lateral meniscus on the tibia to a double one. While Australopithecus afarensis exhibits a single insertion, early Homo clearly exhibits a double insertion of the lateral meniscus on the tibia. The double insertion restricts the mobility of the meniscus on the tibial plateau, indicating a habitual practice of full extension movements of the knee joint. Among modern humans, the posterior insertion of the lateral meniscus appears early in fetal life. Consequently in early Homo, this new selected feature developed directly as a result of a genomic change. The derived shape of human distal femoral epiphysis includes a prominence of the lateral lip of the femoral trochlea, an elliptical profile of the lateral condyle, and an anteroposterior lengthening of the epiphysis. Analysis of human fetal and neonatal distal epiphyses shows that the prominence of the lateral lip of the trochlea arises before any use, and thus appears to be genetically determined. However, the postnatal development of this joint shows that this feature is also modified epigenetically by use. It is argued that the hominid femoro-patellar joint would have been reshaped following the process of genetic assimilation (Waddington [1942] Nature 3811:563-565). The prominence of the lateral lip of the femoral trochlea was probably selected following a two-staged process-first epigenetic, then genetic. Far from being a Lamarckian explanation, this concept applies precisely to adaptive characters that are induced by an external stimulus during a single lifetime and are replaced through natural selection by genetically based equivalent characters. The nature of the structures involved in the studied features is shown to be an important parameter determining their mode of development and selection.

Morphological and taxonomic affinities of the Olduvai ulna (OH 36).

The OH 36 ulna derives from Upper Bed II in the Olduvai Gorge, and is dated to circa 1.1-1.2 Myr. Multivariate analyses incorporating data from samples of modern humans, common and pygmy chimpanzees, gorillas, orangutans, and two other early hominin ulnae, Omo L40-19 and KNM-BK 66, suggest that OH 36 belonged to an individual with powerful forearms consistent with a locomotor repertoire that included arboreal locomotion. However, there is no compelling evidence that it made regular use of its forelimbs as supports when travelling on the ground. When compared with levels of intra- and intertaxon size and shape variation in the comparative sample (humans, chimpanzees, gorillas), the differences between OH 36, KNM-BK 66, and Omo L40-19 are compatible with OH 36 differing from the other two fossil hominin ulnae to the extent that modern humans differ from modern great apes. KNM-BK 66 and Omo L40-19 differ from each other in overall size and shape only to the degree that would be expected within any of the individual modern comparative samples. Based on these analyses, there is no evidence to support the hypothesis that OH 36 and Omo L40-19 belong to the same species of fossil hominin, or to two species that shared a similar forelimb locomotor repertoire. We suggest that OH 36 has the greater claim to be assigned to Paranthropus boisei, and we recommend that for the time being the latter be referred to the tribe Hominini gen. et sp. indet. The surprising result of these analyses is the overall size and shape similarity between Omo L40-19 and KNM-BK 66, two fossils that are separated in time by more than 1.5 million years, and which have traditionally been assumed to represent hominin species with quite different locomotor patterns.

A newly discovered Kenyapithecus skeleton and its implications for the evolution of positional behavior in Miocene East African hominoids.

The middle Miocene stem kenyapithecine Nacholapithecus kerioi (16–15 Ma; Nachola, Kenya) is represented by a large number of isolated fossil remains and one of the most complete skeletons in the hominoid fossil record (KNM-BG 35250). Multiple fieldwork seasons performed by Japanese–Kenyan teams during the last part of the 20th century resulted in the discovery of a large sample of Nacholapithecus fossils. Here, we describe the new femoral remains of Nacholapithecus. In well-preserved specimens, we evaluate sex differences and within-species variation using both qualitative and quantitative traits. We use these data to determine whether these specimens are morphologically similar to the species holotype KNM-BG 35250 (which shows some plastic deformation) and to compare Nacholapithecus with other Miocene hominoids and extant anthropoids to evaluate the distinctiveness of its femur. The new fossil evidence reaffirms previously reported descriptions of some distal femoral traits, namely the morphology of the patellar groove. However, results also show that relative femoral head size in Nacholapithecus is smaller, relative neck length is longer, and neck–shaft angle is lower than previously reported for KNM-BG 35250. These traits have a strong functional signal related to the hip joint kinematics, suggesting that the morphology of the proximal femur in Nacholapithecus might be functionally related to quadrupedal-like behaviors instead of more derived antipronograde locomotor modes. Results further demonstrate that other African Miocene apes (with the exception of Turkanapithecus kalakolensis) generally fall within the Nacholapithecus range of variation, whose overall femoral shape resembles that of Ekembo spp. and Equatorius africanus. Our results accord with the previously inferred locomotor repertoire of Nacholapithecus, indicating a combination of generalized arboreal quadrupedalism combined with other antipronograde behaviors (e.g., vertical climbing).

What's your angle? Size correction and bar-glenoid orientation in "Lucy" (A.L. 288-1)

There has been much debate as to the locomotor repertoire of Lucy (A.L. 288-1) and other specimens of Australopithecus afarensis, ranging from fully committed bipeds to species that spent a significant time in the trees as well as on the ground. We examined the bar–glenoid angle, a character purported to indicate arboreal propensities, and its implications for this specific debate and the more general challenge of extracting behavioral information from fossils. We examined the bar–glenoid angle in ontogenetic samples of Pan paniscus, Pan troglodytes, Gorilla gorilla gorilla, Gorilla gorilla beringei, Pongo pygmaeus, Homo sapiens, and A.L. 288-1 (“Lucy”). We found that there is no allometry in the bar–glenoid angle for the great apes, but a weak correlation for humans. Moreover, the data scatters for the African apes and humans converge at the smaller size ranges, and Lucy's value for bar–glenoid angle falls precisely in this area of overlap. Therefore, we conclude that the bar–glenoid angle is not tightly correlated with function and, as such, cannot be used as a morphological signal of arboreal behavior, especially in the smaller size ranges, at which arboreal and nonarboreal species overlap. Our work does not resolve issues concerning Lucy's precise locomotor repertoire but adds new information to consider. The total morphological pattern, plus an appreciation of the underlying variance in morphological and behavioral characters in extant species, is key for making functional inferences from the morphology of fossils.