Partitioning variance in a signaling trade-off under sexual selection reveals among-individual covariance in trait allocation.

Understanding the evolution of traits subject to trade-offs is challenging because phenotypes can (co)vary at both the among- and within-individual levels. Among-individual covariation indicates consistent, possibly genetic, differences in how individuals resolve the trade-off, while within-individual covariation indicates trait plasticity. There is also the potential for consistent among-individual differences in behavioral plasticity, although this has rarely been investigated. We studied the sources of (co)variance in two characteristics of an acoustic advertisement signal that trade-off with one another and are under sexual selection in the gray treefrog, Hyla chrysoscelis: call duration and call rate. We recorded males on multiple nights calling spontaneously and in response to playbacks simulating different competition levels. Call duration, call rate, and their product, call effort, were all repeatable both within and across social contexts. Call duration and call rate covaried negatively, and the largest covariance was at the among-individual level. There was extensive plasticity in calling with changes in social competition, and we found some evidence for among-individual variance in call rate plasticity. The significant negative among-individual covariance in trait values is perpendicular to the primary direction of sexual selection in this species, indicating potential limits on the response to selection.

The transition from the female-like great calls to male calls during ontogeny in southern yellow-cheeked gibbon males (Nomascus gabriellae).

It is well known that gibbons emit a pattern of vocalizations, which is specific for species and sex. A previous study showed, however, that immature southern yellow-cheeked gibbon (Nomascus gabriellae) males produce only female-like great calls from 2.3 to 5.3 years of age in co-singing interactions with their mothers. To date, nothing is known about how the vocal repertoire of a male changes from the female-like call (great call) to the male call (staccato notes and multi-modulation phrase) during vocal ontogeny. The goal of this study was to describe the transition from the female-like great call to the male call and the ontogeny of the male call. We predicted that the transition from the female-like great call to the male-specific call and the development of the male call is a normal part of the aging proces. If this is the case, the following phenomena will occur: (a) female vocalization should no longer be produced with the mature form of the multi-modulation phrase and (b) all stages of the male vocalization should occur gradually as the young male ages. Young males regularly emit both female-like great calls and male-specific calls between the ages of 5.6 to 7.1 years. Once the young males reached 7.1 years of age, they emitted male calls exclusively, and they continued to do so until the end of the observation period (at 8.11 years of age). It was confirmed that the young males emitted only female-like great calls during periods when they produced non-mature forms of a multi-modulation phrase (Fm0,1-none or one frequency modulation in second notes). Furhermore, the decrease in the number of female-like great calls was attributed to the development of the mature form of the multi-modulation phrase (Fm2-two or more frequency modulation in second notes), which developed with age. We also confirmed that the multi-modulation phrase developed gradually, while the development of the staccato notes occurred in leaps. A multi-modulation phrase developed as the initial part of the male-specific call. It was evolved from a simpler to a more complex form as the maximum frequency and age of the young males increased. Staccato notes subsequently developed in certain young males. Possible explanations for such vocal ontogeny in young males are discussed in this work.

Habitat monitoring and conservation prioritization of Western Hoolock Gibbon in upper Brahmaputra Valley, Assam, India.

The present study aimed at predicting the potential habitat of Western Hoolock Gibbon (Hoolock hoolock) in the upper Brahmaputra River Valley, Assam, India, and identifying priority conservation areas for the species, taking canopy cover into account. We used the maximum entropy algorithm for the prediction of the potential habitat of the gibbon using its current distribution with 19 environmental parameters as primary predictors. Spatio-temporal analyses of the habitat were carried out using satellite-based remote sensing and GIS techniques for two decades (1998-2018) along with Terra Modis Vegetation Continuous Field product to examine land use land cover (LULC), habitat fragmentation, Normalized Difference Vegetation Index (NDVI) and tree cover percentage of the study area. To identify the conservation priority area, we applied a cost-effective decision-making analysis using systematic conservation prioritization in R programming. The model predicted an area of 6025 km2 under high potential habitat, a major part of which was found to overlap with dense forest (80%), followed by moderately open forest (74%) and open forest (66%). The LULC change matrix showed a reduction of forest area in the predicted high potential habitat during the study period, while agricultural class showed an increasing trend. The fragmentation analysis indicated that the number of patches and patch density increased from 2008 to 2018 in the 'very dense' and 'dense' canopy regions of the gibbon habitat. Based on the conservation priority analysis, a 640 km2 area has been proposed to conserve a minimum of 10% of gibbon habitat. The current analysis revealed that in the upper Brahmaputra Valley most areas under dense forest and dense canopy have remained intact over the last two decades, at least within the high potential habitat zone of gibbons independent of the degree of area change in forest, agriculture and plantation.

Resource partitioning and niche overlap between hoolock gibbon (Hoolock hoolock) and other frugivorous vertebrates in a tropical semi-evergreen forest.

Understanding how niche differences evolve in ecologically similar species and how these differences are maintained is a fundamental question in ecology. We studied resource partitioning and niche overlap between the hoolock gibbon (Hoolock hoolock) and other frugivorous vertebrates from April 2016 to January 2018 in Satchari National Park, Sylhet, Bangladesh. We examined the differences in their diet, niche breadth, niche overlap, and characterization of patch use. We recorded feeding events of gibbons and other frugivorous vertebrates using ad libitum sampling. Gibbons consumed 76 plant species including 32 non-fig fruits and 14 figs. Twenty-one competing frugivorous vertebrate species shared 10-70% of their food species with hoolocks. Competition for fruits was intense among gibbons, macaques, and hornbills, as fruits comprised more than 50% of their diet. The niche breadth of the gibbons varied across seasons. It was lowest during the rainy season (BA = 0.39) when fruits were more readily available, and highest in winter (BA = 0.58) when gibbons were less selective due to food scarcity. The niche overlap was highest between gibbons and northern pig-tailed macaques (Ojk = 0.70), followed by gibbons and hornbills (Ojk = 0.68). Feeding heights and substrate used varied significantly between gibbons and competitors. Gibbons may minimize competition by specializing on various food resources and using different forest patches.

Sleeping trees and sleep-related behaviours of the siamang (Symphalangus syndactylus) in a tropical lowland rainforest, Sumatra, Indonesia.

Sleeping tree selection and related behaviours of a family group and a solitary female siamang (Symphalangus syndactylus) were investigated over a 5-month period in northern Sumatra, Indonesia. We performed all day follows, sleeping tree surveys and forest plot enumerations in the field. We tested whether: (1) physical characteristics of sleeping trees and the surrounding trees, together with siamang behaviours, supported selection based on predation risk and access requirements; (2) the preferences of a solitary siamang were similar to those of a family group; and (3) sleeping site locations within home ranges were indicative of home range defence, scramble competition with other groups or other species, or food requirements. Our data showed that (1) sleeping trees were tall, emergent trees with some, albeit low, connectivity to the neighbouring canopy, and that they were surrounded by other tall trees. Siamangs showed early entry into and departure from sleeping trees, and slept at the ends of branches. These results indicate that the siamangs' choice of sleeping trees and related behaviours were strongly driven by predator avoidance. The observed regular reuse of sleeping sites, however, did not support anti-predation theory. (2) The solitary female displayed selection criteria for sleeping trees that were similar to those of the family group, but she slept more frequently in smaller trees than the latter. (3) Siamangs selected sleeping trees to avoid neighbouring groups, monopolise resources (competition), and to be near their last feeding tree. Our findings indicate selectivity in the siamangs' use of sleeping trees, with only a few trees in the study site being used for this purpose. Any reduction in the availability of such trees might make otherwise suitable habitat unsuitable for these highly arboreal small apes.

A first report of separation calls in southern yellow-cheeked gibbons (Nomascus gabriellae) in captivity.

The effects of social separation, including vocalization, have been studied for a very long time in non-human primates under laboratory conditions. As part of the long-term research on the vocal behaviour of Nomascus gibbons in zoos, this study provides the first record of calls of the southern yellow-cheeked gibbon (Nomascus gabriellae) in response to involuntary separation. Our study revealed that calls were also emitted by an infant (aged 1 year 8 months), and that the acoustic structure of the infant's calls was similar to that of older individuals' calls. Separation-induced calls seem to have a shorter developmental convergence than vocalizations with a stable pattern (which are specific for species and sex). The acoustic structure of the calls reported here comprised simple syllables, and differed from the sex- and species-specific vocal patterns of this species. Our findings demonstrate a novel paradigm in this genus, and provide evidence of the ability of gibbons to express distress when socially separated.

Comparison of point transect distance and traditional acoustic point-count sampling of hoolock gibbons in Htamanthi Wildlife Sanctuary, Myanmar.

Effective conservation demands more accurate and reliable methods of survey and monitoring of populations. Surveys of gibbon populations have relied mostly on mapping of groups in "listening areas" using acoustical point-count data. Traditional methods of estimating density in have usually used counts of gibbon groups within fixed-radius areas or areas bounded by terrain barriers to sound transmission, and have not accounted for possible decline in detectability with distance. In this study we sampled the eastern hoolock gibbon (Hoolock leucogenys) population in Htamanthi Wildlife Sanctuary (WS), Myanmar, using two methods: the traditional point-count method with fixed-radius listening areas, and a newer method using point-transect Distance analysis from a sample point established in the center of each listening point array. The basic data were obtained by triangulating on singing groups from four LPs for 4 days, in 10 randomly selected sample areas within the sanctuary. The point transect method gave an average density of 3.13 groups km-2 , higher than the estimates of group density within fixed-radius areas without correction for detectability. A new method of analysis of singing probability per day (p[1]) gave an estimate of 0.547. Htamanthi WS is an important conservation area containing an estimated 7000 (95% confidence interval: 5000-10,000) hoolock groups. Surveys at Htamanthi WS and locations in the Hukaung Valley suggest that the extensive evergreen forests in northern Myanmar have the capacity to support 2-4 (average about 3) groups of hoolock gibbons per km2 , but most forests in its range have yet to be surveyed.

Monitoring occurrence, extinction, and colonization probabilities for gibbon populations.

All gibbon species (Family: Hylobatidae) are considered threatened with extinction and recognized on the International Union for Conservation of Nature Red List of Threatened Species. Because gibbons are one of the most threatened families of primates, monitoring their status is now critically important. Long-term monitoring programs applying occupancy approaches, in addition to assessing occurrence probability, improves understanding of other population parameters such as site extinction or colonization probabilities, which elucidate temporal and spatial changes and are therefore important for guiding conservation efforts. In this study, we used multiple season occupancy models to monitor occurrence, extinction, and colonization probabilities for northern yellow-cheeked crested gibbon Nomascus annamensis in three adjacent protected areas in the Central Annamites mountain range, Vietnam. We collected data at 30 listening posts in 2012, 2014, and 2016 using the auditory point count method. Occurrence probabilities were highest in 2012 (0.74, confidence interval [CI]: 0.56-0.87) but slightly lower in 2014 (0.66, CI: 0.51-0.79) and 2016 (0.67, CI: 0.49-0.81). Extinction probabilities during the 2012-2014 and 2014-2016 intervals were 0.26 (0.14-0.44) and 0.25 (0.12-0.44), respectively. Colonization probabilities during 2012-2014 were 0.44 (0.19-0.73) and between 2014 and 2016 was 0.51 (0.26-0.75). Although local site extinctions have occurred, high recolonization probability helped to replenish the unoccupied sites and kept the occurrence probability stable. Long-term monitoring programs which use occurrence probability alone might not fully reveal the true dynamics of gibbon populations. We strongly recommend including multiple season occupancy models to monitor occurrence, extinction, and colonization probabilities in long-term gibbon monitoring programs.

Strength properties of extant hominoid hallucal and pollical metapodials.

Functional comparisons of cortical bone strength properties between hominoid hallucal and pollical metapodials (Mt1 and Mc1, respectively) are lacking. Determining which of these two elements is stronger, and by how much, could be informative because the hallux and pollex are used differently both within and among extant hominoids during locomotion and manipulation (i.e., functional differentiation between autopod pairs). Here, we compare Mt1 and Mc1 midshaft cortical area, polar section modulus, and polar second moment of area, calculated from high-resolution computed tomography images in humans (n = 21), chimpanzees (n = 47), gorillas (n = 24), orangutans (n = 20), siamangs (n = 8), and gibbons (n = 21). Intraindividual comparisons between bones within species were made using paired t-tests. Log10-transformed Mt1:Mc1 ratios were created to assess relative strength asymmetry between bones, and interspecific comparisons of these proportions were made using analyses of variance. Absolute strength differences between the Mt1 and Mc1 for all variables were significantly larger in the Mt1 for all species (p < 0.05). Significant differences across species in Mt1:Mc1 proportions were also found, thereby demonstrating that strength asymmetry between bones differs among taxa (p < 0.05); asymmetry was lowest in orangutans, intermediate in gorillas, and greatest in humans, chimpanzees, siamangs, and gibbons. These findings support the hypothesis that the Mt1 is better adapted structurally than the Mc1 for bearing mechanical loads during weight support of locomotion in all extant hominoids and that pedal hallucal grasping likely engenders higher loads than manual pollical grasping in nonhuman hominoids. Thus, functional differentiation in autopod use within and among hominoids is reflected in hallucal and pollical metapodial strength properties.

Influence of food availability and climate on behavior patterns of western black crested gibbons (Nomascus concolor) at Mt. Wuliang, Yunnan, China.

Food abundance and climatic factors can significantly affect the behavior of animals and constrain their activity budgets. The population of western black crested gibbons (Nomascus concolor) in Mt. Wuliang lives in montane forest and is close to the northern extreme of the distribution for gibbons (Hylobatidae). Their habitats show remarkable seasonal variation in terms of food availability, temperature, and rainfall. To understand behavioral adaptations of western black crested gibbons to different sets of ecological conditions, we examined relationships among food availability, mean temperature, rainfall, and behavior patterns by observing two groups for 1 year each. Our results revealed that activity budget was affected by food availability and mean temperature. The gibbons spent more time eating flowers when that resource was more available and spent less time moving when fruit was more available. The gibbons spent less time feeding and more time resting, and spent less feeding time on fruit and leaves when the mean temperature was lower. These results suggest that the gibbons displayed a pronounced preference for flowers as a food resource and adopted a time minimizer strategy when high-nutrient food items (i.e., fruit) were more available. In addition, the gibbons adopted an energy-conserving strategy during periods of low temperature. The flexibility of behavioral patterns in responding to food availability and temperature may potentially improve the gibbons' prospects of surviving and reproducing in a northern montane forest.

Effects of cold weather on the sleeping behavior of Skywalker hoolock gibbons (Hoolock tianxing) in seasonal montane forest.

Considering the high energetic costs of maintaining constant body temperature, mammals must adjust their thermoregulatory behaviors in response to cold temperatures. Although primate daytime thermoregulation is relatively well studied, there is limited research in relation to nighttime strategies. To investigate how Skywalker hoolock gibbons (Hoolock tianxing) cope with the low temperatures found in montane forests, we collected sleep-related behavior data from one group (NA) and a single female (NB) at Nankang (characterized by extensive tsaoko plantations) between July 2010 and September 2011, and one group (BB) at Banchang (relatively well-managed reserve forest) between May 2013 and May 2015 in Mt. Gaoligong, Yunnan, China. The annual mean temperature was 13.3°C at Nankang (October 2010 to September 2011) and 13.0°C at Banchang (June 2013 to May 2015) with temperatures dropping below -2.0°C at both sites, making them the coldest known gibbon habitats. The lowest temperatures at both sites remained below 5.0°C from November to March, which we, therefore, defined as the "cold season". The hoolock gibbons remained in their sleeping trees for longer periods during the cold season compared to the warm season. Sleeping trees found at lower elevations and closer to potential feeding trees were favored during cold seasons at both sites. In addition, the gibbons were more likely to huddle together during cold seasons. Our results suggest that cold temperatures have a significant effect on the sleeping behavior of the Skywalker hoolock gibbon, highlighting the adaptability of this threatened species in response to cold climates.

Infant loss during and after male replacement in gibbons.

According to the sexual selection hypothesis, infanticide during resident male replacement is an adaptive strategy that has evolved because the killing of unweaned offspring sired by previous males shortens the inter-birth intervals of the mothers whose infants are targeted and thereby increases the reproductive fitness of the perpetrator. To test this hypothesis, we describe previously unreported cases of primary male replacement for two gibbon species (Hylobates lar and Nomascus nasutus), and review all other reported cases of primary male replacement in gibbons. Overall, infants were present in nearly half of all cases (16/33, 48%) and of the 18 infants present during replacement, 50% (N = 9) disappeared within 2 months of the event. In four of the five cases where there was sufficient demographic information to identify the likely sire of the subsequent offspring of females that lost infants, the new male was believed to be the sire. Infants were also less likely to die or disappear if the new male and original resident male were possible kin. However, there was no significant difference in the age of infants between those that died or disappeared following replacement and those that survived to weaning (p = .630). Our review of takeover-related infant loss in gibbons confirms that periods of male instability are risky for unweaned infants and that replacing males benefit from infant loss. Nevertheless, variability in the context of infant loss and difficulties related to data collection in the field make it difficult to test competing hypotheses concerning the mechanisms and functions of infanticide in the small apes.

Fur color change and hormonal development in captive females of northern white-cheeked (Nomascus leucogenys) and buff-cheeked (Nomascus gabriellae) gibbons.

Gibbons of the genus Nomascus exhibit strong sexual dichromatism in fur color. Change of fur color in sub-adult wild Nomascus females is associated with the onset of puberty and the time of their dispersal. The variability in fur change may be influenced by social factors. In this study, we determined whether in captive females of crested gibbons begin reproductive maturity prior to dispersing and with association to their fur color. We collected 287 fecal extracts to analyze pregnandiol -3- glucuronide and 17ß estradiol profiles of 4 sub-adult females (Nomascus leucogenys and Nomascus gabriellae) and 183 samples from their mothers, using enzyme immunoassays. The sub-adult females were monitored from 4 years of age. Their hormone profiles showed the onset of ovulatory cycling between 4.6 and 5.8 years. Based on the information about the estrogen influence to the secondary sex characteristic (fur color of female) the positive link between estrogen concentration and age of the sub-adult females was found. However, the amount of the estrogen can apparently be influenced by the presence of mother. If the mother was presented, the level of estrogen was higher than if the mother was missing. Our findings suggest that the probability of changing to beige fur color by the sub-adult females increased with increased age and if they were without mother. This initial study presents the maternal influence as a possible social factor affecting the fur color change of female offspring.

Space Use by Eastern Hoolock Gibbons (Hoolock leuconedys) in Nankang, Mt. Gaoligong, Southwest Yunnan, China.

To understand the differences in space use by eastern hoolock gibbons (Hoolock leuconedys) for meeting different behavioural needs and adapting to a disturbed environment, an eastern hoolock gibbon family in Nankang, Mt. Gaoligong, China, was observed, and data on their behaviour, spatial position and substrate use were collected through instantaneous scanning from October 2, 2010, to September 15, 2011. Our results suggested that connectivity was a primary factor influencing the movements of eastern hoolock gibbons. However, there were differences between spatial location and substrate use with different behaviours. Sitting and hanging were the main resting behaviours, but spatial position and substrate use for these two postures were different. Because tsaoko cultivation in this area has made understory plants rare, gibbons primarily used long, thin, horizontal branches in the first and second layers of the tree canopy for their movement. Although gibbons can selectively use the habitat and alter their spatial utilisation pattern accordingly, they are facing increased difficulty in movement and risk falling because of increased human disturbance and spread of tsaoko plantations. Therefore, it is important to limit the scope of tsaoko plantation to maintain a suitable habitat for Nankang gibbons.

Dispersal and female philopatry in a long-term, stable, polygynous gibbon population: Evidence from 16 years field observation and genetics.

Gibbons are generally reported to live in small socially monogamous family groups in which both sexes disperse when they reach maturity. For the first time, we documented the dispersal pattern in a population of gibbons living in stable polygynous groups (Nomascus concolor) integrating 16 years' field observation and genetic information from fecal DNA. All subadult males except for one dispersed at 9.8 ± 1.4 years of age (range: 8-12, N = 10). The last male remained in his natal group and obtained the breeding position at age 11 by evicting the original dominant male. Females reached sexual maturity (as evidenced by the change in body color from black to yellow) at 8 years (N = 4). Three of them dispersed and one obtained a position as a breeding female and bred in her natal group. We also observed one female returning to her natal group with her infant after her presumed father was taken over by a neighboring male. We identified only three mtDNA haplotypes from 22 individuals at Dazhaizi. Individuals in one group shared the same haplotype, with only one exception. Genetic results showed that the two breeding females were mother-daughter pairs in all three study groups at the end of this study, implying some degree of female philopatry. We argue that in the case of black crested gibbons, dispersal decisions appear to represent highly opportunistic events in response to reproductive opportunities in their natal and neighboring groups.

A distance sampling approach to estimate density and abundance of gibbon groups.

All 18 species of gibbons are considered threatened with extinction and listed on the IUCN Red List of Threatened Species. Because gibbons (Hylobatidae) are one of the most threatened primate families, a great need exists to determine and monitor their status effectively. To meet this need, we employed distance sampling methods to estimate the density and number of gibbon groups. We focused on southern yellow-cheeked crested gibbon in the Nam Cat Tien sector of Cat Tien National Park, Vietnam, from July to October, 2016. We used an auditory point count method at 48 listening posts to detect gibbon groups. We estimated our detection probability of calling groups of gibbons within 1,200 m of a listening post was 0.63 (95% CI: 0.54-0.74). In addition, we estimated the daily calling probability of a gibbon group to be 0.46 (95% CI: 0.33-0.59) and interpreted this as the probability that a group was available to be detected. We adjusted our group density and abundance estimates by both detection and availability probabilities. Ultimately we estimated 325 (95% CI: 232-455) gibbon groups in the Nam Cat Tien sector. Our results suggest that Cat Tien National Park contains one of the largest populations of southern yellow-cheeked crested gibbon in Vietnam. Our methods are one way of avoiding underestimation of gibbon group density and abundance by incorporating uncertainty in gibbon group availability and detection.

Look in the trees: Hylobatids as evolutionary models for extinct hominins.

Studying extant apes is of central importance to paleoanthropology. This approach is informative in inferring how hominin skeletal morphology reflects phylogeny, behavior, development, and ecological context. Traditionally, great apes have dominated the paleoanthropological literature as extant analogs for extinct hominins, to the exclusion of their phylogenetic sister group, the hylobatids. Phylogenetic proximity, large body size, and high encephalization quotients may have contributed to decisions to use great apes as models for hominins. However, if we reexamine hylobatids as extant models for extinct hominins-using modern phylogenetic, behavioral, and ecological data-this clade is uniquely poised to inform future frameworks in paleoanthropology. The following features make hylobatids strong analogs for extinct hominins: taxonomic diversity, the timing of diversification, hybridization between species, small body size, and reduced sexual dimorphism. Based on these shared features, hylobatids offer future opportunities to paleoanthropology, and provide a much richer extant analog than is currently recognized.

The gibbon's Achilles tendon revisited: consequences for the evolution of the great apes?

The well-developed Achilles tendon in humans is generally interpreted as an adaptation for mechanical energy storage and reuse during cyclic locomotion. All other extant great apes have a short tendon and long-fibred triceps surae, which is thought to be beneficial for locomotion in a complex arboreal habitat as this morphology enables a large range of motion. Surprisingly, highly arboreal gibbons show a more human-like triceps surae with a long Achilles tendon. Evidence for a spring-like function similar to humans is not conclusive. We revisit and integrate our anatomical and biomechanical data to calculate the energy that can be recovered from the recoiling Achilles tendon during ankle plantar flexion in bipedal gibbons. Only 7.5% of the required external positive work in a stride can come from tendon recoil, yet it is delivered at an instant when the whole-body energy level drops. Consequently, an additional similar amount of mechanical energy must simultaneously dissipate elsewhere in the system. Altogether, this challenges the concept of an energy-saving function in the gibbon's Achilles tendon. Cercopithecids, sister group of the apes, also have a human-like triceps surae. Therefore, a well-developed Achilles tendon, present in the last common 'Cercopithecoidea-Hominoidea' ancestor, seems plausible. If so, the gibbon's anatomy represents an evolutionary relict (no harm-no benefit), and the large Achilles tendon is not the premised key adaptation in humans (although the spring-like function may have further improved during evolution). Moreover, the triceps surae anatomy of extant non-human great apes must be a convergence, related to muscle control and range of motion. This perspective accords with the suggestions put forward in the literature that the last common hominoid ancestor was not necessarily great ape-like, but might have been more similar to the small-bodied catarrhines.

Hip extensor mechanics and the evolution of walking and climbing capabilities in humans, apes, and fossil hominins.

The evolutionary emergence of humans' remarkably economical walking gait remains a focus of research and debate, but experimentally validated approaches linking locomotor capability to postcranial anatomy are limited. In this study, we integrated 3D morphometrics of hominoid pelvic shape with experimental measurements of hip kinematics and kinetics during walking and climbing, hamstring activity, and passive range of hip extension in humans, apes, and other primates to assess arboreal-terrestrial trade-offs in ischium morphology among living taxa. We show that hamstring-powered hip extension during habitual walking and climbing in living apes and humans is strongly predicted, and likely constrained, by the relative length and orientation of the ischium. Ape pelves permit greater extensor moments at the hip, enhancing climbing capability, but limit their range of hip extension, resulting in a crouched gait. Human pelves reduce hip extensor moments but permit a greater degree of hip extension, which greatly improves walking economy (i.e., distance traveled/energy consumed). Applying these results to fossil pelves suggests that early hominins differed from both humans and extant apes in having an economical walking gait without sacrificing climbing capability. Ardipithecus was capable of nearly human-like hip extension during bipedal walking, but retained the capacity for powerful, ape-like hip extension during vertical climbing. Hip extension capability was essentially human-like in Australopithecus afarensis and Australopithecus africanus, suggesting an economical walking gait but reduced mechanical advantage for powered hip extension during climbing.

Body mass estimation in hominoids: Age and locomotor effects.

While there are a number of methods available for estimation of body mass in adult nonhuman primates, very few are available for juveniles, despite the potential utility of such estimations in both analyses of fossils and in museum collection based research. Furthermore, because of possible scaling differences, adult based body mass estimation equations may not be appropriate for non-adults. In this study, we present new body mass estimation equations for both adult and immature nonhuman hominoids based on joint and metaphyseal dimensions. Articular breadths of the proximal and distal femur, distal humerus and tibial plateau, and metaphyseal breadths of the distal femur and humerus were collected on a reference sample of 159 wild Pan, Gorilla, Pongo, Hylobates, and Symphalangus specimens of known body mass from museum and research collections. Scaling of dimensions with body weight was assessed in both the adult and the ontogenetic sample at several taxonomic levels using reduced major axis regression, followed by regression of each dimension against body mass to generate body mass estimation equations. Joint dimensions were found to be good predictors of body mass in both adult and immature hominoids, with percent prediction errors of 10-20%. However, subtle scaling differences between taxa impacted body mass estimation, suggesting that phylogeny and locomotor effects should be considered when selecting reference samples. Unlike patterns of joint growth in humans, there was little conclusive evidence for consistently larger joints relative to body mass in the non-adult sample. Metaphyseal breadths were strong predictors of body mass and, with some exceptions, gave more precise body mass estimates for non-adults than epiphyseal breadths.