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OBJECTIVES It seems to be commonly accepted that gestation length within the subfamily Colobinae lasts several weeks longer in the African tribe (Colobini) than in the Asian tribe (Presbytini) even though closely related taxa of similar body mass should have similar life histories. Suspecting problems with data provenance to cause the difference, we revisited the published records expecting similar gestation lengths in both tribes if based on vetted, accurate data. MATERIALS AND METHODS We compiled published gestation length data for Colobini and Presbytini, labeling them as "unspecified" (n = 16) if the primary reference could not be located, methods were not described, and/or conceptions, the beginning of gestation, were determined based on sporadic observations of mating. If conceptions were determined based on changing hormone levels or patterns of daily mating records, we labeled the data as "accurate" (n = 12). We analyzed the ln transformed data in a phylogenetic framework in relation to adult female body mass. RESULTS In the unspecified dataset, gestation length in the two tribes overlapped extensively and did not differ significantly. However, in the accurate dataset, gestation length was significantly shorter in Colobini (not longer, as previously assumed). DISCUSSION Data provenance had a strong impact on the comparison, reversing the relationship in gestation length in the two sister tribes. It remains to be determined why gestation lengths differ, whether, relative to the other primates, Colobini have a shortened gestation or Presbytini a lengthened gestation, and whether similar differences exist in other closely related taxa. Addressing these questions will require additional, broader, comparative analyses.

Variation in the relationship between gestation length and body mass can arise because different types of tissue require varying amounts of energy to build, and not all species build such tissues in the same proportions. Given that a pregnant female has a finite amount of energy, trade-offs between investment in different tissues may occur. Here we examine if dental precocity accounts for variation in primate gestation length. If true, this could explain why folivorous species with precocial dentition have longer gestation lengths than predicted by neonatal brain and body mass. We compiled data on gestation length, neonate and adult female body and brain mass from the literature. We used published postcanine eruption schedules at 4 months of age and measured the total occlusal area as dental endowment to approximate dental precocity at birth. Species with embryonic delay in growth or altricial neonates were not considered because they represent grade shifts regarding gestation length. Consequently, our data were biased toward Simiiformes and Old World monkeys, specifically. We performed a phylogenetic generalized least squares regression (pGLS) of neonate brain mass in relation to neonate body mass, and a second pGLS with dental endowment as an additional predictor variable. Including dental endowment in the pGLS did not improve the model. Dental endowment did not systematically impact primate gestation length. Concordant with results from previous studies, this indicates that the energetically expensive period of tooth mineralization may occur postnatally. More data are required to examine if the results are typical across primates.

Aggression rarely escalates to physical conflict because doing so puts individuals at risk of injury. Escalation only pays off when the potential benefits outweigh the potential costs, that is, when resources critical to fitness are at stake. Here, we investigated the occurrence of injury in 2 Asian colobine species: Nepal gray langurs (Semnopithecus schistaceus) and Phayre’s leaf monkeys (Trachypithecus phayrei crepusculus). In both species, younger individuals are higher-ranking and might have greater incentive to fight. However, Nepal gray langurs have a strict breeding season, which may magnify male mating competition, and Phayre’s leaf monkeys, unlike Nepal gray langurs, have female-biased dispersal, which may increase female injury risk during subadulthood. Using long-term data on observed injuries (Nepal gray langurs: n = 208; Phayre’s leaf monkeys: n = 225), we modeled the monthly occurrence of injury (Y/N) and found that males received more injuries than females in both species. Also, subadults generally experienced frequent injury, as young individuals likely face challenges when competing for group membership and/or establishing rank. In Nepal gray langurs, males received 3 times more injuries during the mating season, suggesting strong competition for mates during this period, and females experienced more injuries before conception, suggesting competition to meet the nutritional requirements for reproduction. Unexpectedly, females in smaller groups received more injuries in Nepal gray langurs. Overall, these results indicate that injuries are most likely when fighting may aid in establishing group membership, achieving high rank, and reproducing. Future research should investigate the influence of injuries on fitness outcomes.

Studies in multiple host species have shown that gut microbial diversity and composition change during pregnancy and lactation. However, the specific mechanisms underlying these shifts are not well understood. Here, we use longitudinal data from wild Phayre’s leaf monkeys to test the hypothesis that fluctuations in reproductive hormone concentrations contribute to gut microbial shifts during pregnancy. We described the microbial taxonomic composition of 91 fecal samples from 15 females (n = 16 cycling, n = 36 pregnant, n = 39 lactating) using 16S rRNA gene amplicon sequencing and assessed whether the resulting data were better explained by overall reproductive stage or by fecal estrogen (fE) and progesterone (fP) concentrations. Our results indicate that while overall reproductive stage affected gut microbiome composition, the observed patterns were driven by reproductive hormones. Females had lower gut microbial diversity during pregnancy and fP concentrations were negatively correlated with diversity. Additionally, fP concentrations predicted both unweighted and weighted UniFrac distances, while reproductive state only predicted unweighted UniFrac distances. Seasonality (rainfall and periods of phytoprogestin consumption) additionally influenced gut microbial diversity and composition. Our results indicate that reproductive hormones, specifically progestagens, contribute to the shifts in the gut microbiome during pregnancy and lactation.

OBJECTIVES Geophagy has several hypothesized functions, among them (1) detoxification of plant secondary compounds, for example, tannins, (2) buffering stomach pH to alleviate acidosis caused by high sugar intake, and (3) supplementing the diet with mineral nutrients. We tested these hypotheses in Nepal gray langurs (Semnopithecus schistaceus), a foregut fermenter for which fruits and leaves dominate the diet at different times of the year. MATERIALS AND METHODS We collected data on feeding time per food item for 21 unprovisioned adult langurs at Ramnagar, Nepal, for 1 year using instantaneous sampling (773 observation hours). We combined these data with relative sugar and tannin content of food items to estimate diet content. We collected rainfall data to distinguish the wet season (May-September) from the dry season (October-April). We collected soil samples from consumption and control sites to test for pH and sodium, potassium, magnesium, and calcium concentrations. RESULTS Langurs consumed soil from two sources: termite structures (in almost all months) and river banks and beds (mainly in the dry season). Soil feeding was not positively correlated with dietary tannin or sugar content (GLMM). Soil pH was not high enough to buffer stomach contents. Only sodium was significantly higher in consumed soil. DISCUSSION The most likely function of geophagy was the acquisition of sodium. This conclusion is consistent with reports for other animals. Buffering stomach pH, an often-suggested function for animals with ruminant-like digestion, was not supported. Detoxification, often proposed for animals with a diet high in secondary compounds, was also not supported.

Females of several catarrhine primate species exhibit exaggerated sexual swellings that change in size and coloration during the menstrual cycle and, in some species, gestation. Although their function remains under debate, studies indicate that swellings may contain information males could use to discern ovulation and the probability that a cycle will be conceptive. Here we combine visual ratings of swellings with hormonal data for a group of Sanje mangabeys (18 adult, 3 adolescent females) to determine if their swellings provide reliable information on female fertility. In all cases where ovulation was detected (N = 7), it occurred during maximum tumescence, and in 83.3% during the first two days of the “shiny phase,” a period during maximum tumescence when the swelling was brightest. There were no significant differences in maximum tumescence and shiny phase duration among cycles of different probability of conception, although there was a trend toward conceptive cycles exhibiting shorter shiny phases than nonconceptive ones. Only 25% (N = 4) of postconceptive swellings developed the shiny phase, and adolescents displayed the longest maximum tumescence and shiny phases. The conspicuous nature of the shiny phase and the frequent overlap between its onset and ovulation suggest that its presence serves as a general signal of ovulation and that the cycle has a high probability of being conceptive. It also suggests that swellings in some Sanje mangabeys are more accurate signals of fertility than in other primates. Electronic supplementary material The online version of this article (doi:10.1007/s10764-017-9961-1) contains supplementary material, which is available to authorized users.

In most group-living mammals, reproductive success declines with increasing age and increases with increasing rank. Such effects have mainly been studied in matrilineal and in "age positive" hierarchies, which are stable and in which high ranking females often outperform low ranking ones. These relationships are less well-understood in age-inversed dominance hierarchies, in which a female's rank changes over time. We analyzed demographic data of 2 wild, unprovisioned groups of gray langurs (Semnopithecus schistaceus) near Ramnagar, Nepal covering periods of 5 years each. Female rank was unstable and age-inversed. We measured reproductive success via birth rates (57 births), infant survival (proportion of infants surviving to 2 years) and number of offspring surviving to 2 years of age (successful births) for 3 age and 3 rank classes. We found that old females performed significantly worse than expected (birth rate P = 0.04; successful births P = 0.03). The same was true for low ranking females (P = 0.04, and P < 0.01, respectively). Infant survival was highest for young and middle-aged as well as for high and middle ranking females. Overall, the results for these unstable hierarchies were rather similar to those for stable hierarchies of other mammals, particularly several nonhuman primates. Compared to a provisioned population of a closely related species, the wild and unprovisioned population examined (i) showed stronger age effects, while (ii) female reproductive success was equally affected by rank. Future comparative studies are needed to examine whether captive or provisioned populations deviate predictably from wild populations.

In group-living mammals, social dominance can be achieved and maintained via kin support, winner–loser effects and individual differences in resource-holding potential (RHP). When dominance is based on RHP, the likelihood of winning is also influenced by the incentive to compete. For females, this equation can be quite complex, as both age and reproductive state can alter the relative value of resources and the risks associated with fighting. For example, females that are young, pregnant or lactating may have high nutritional demands, while lactating females also face risks associated with offspring survival. To date, however, there have been few studies investigating how such factors influence female agonistic relationships. Here, we use 2 years of data to evaluate how age and reproductive state influence the probability of winning, focusing on two groups (5–9 and 10–12 females) of wild Phayre's leaf monkeys, Trachypithecus phayrei crepusculus, at Phu Khieo Wildlife Sanctuary, Thailand. Consistent with previous studies, we found that young females and pregnant females were more likely to win dyadic interactions in the smaller group, PA. By contrast, the likelihood of winning was unrelated to age in the larger group, PB. Moreover, differences between reproductive states were less pronounced in PB, with females that were pregnant or in early lactation being more likely to win compared to females that were cycling. The deviating results for PB were likely associated with a greater number of competitors and small age differences intensifying intrasexual competition. These results highlight the importance of individual as well as demographic factors in female dominance relationships. In this population, females with high nutritional demands are more likely to win agonistic encounters, suggesting that female–female competition occurs primarily over access to resources.

Objectives Reproductive skew is proposed to link to despotism in dominance hierarchies. While studies illustrating male skew are plentiful, demonstrating the link to despotism is rare. Likewise, it is often unknown which factors (e.g., resource holding potential, age, physical condition) affect a male's dominance rank. Here we investigated correlates of male rank and hierarchy characteristics in Nepal gray langurs (Semnopithecus schistaceus), a population with high male reproductive skew, and compared the results to other multi-male groups of nonhuman primates. Methods We collected dyadic displacement interactions from two groups (mean 3.0 and 4.1 adult males) for five years each. We assessed dominance ranks for demographically stable phases (n = 11, n = 28) and analyzed the effects of age and physical condition through linear mixed models (LMM). We analyzed hierarchy characteristics via the program MatMan. We used data from 27 primate groups (cercopithecines, colobines, hominoids) as a comparative sample. Results The highest ranks were attained by adults in one group (LMM, P 0.93) and linearity (mean >0.81) and were relatively steep (mean >0.66) when compared with other species. Discussion Dominance rank followed a pattern predicted by resource holding potential, but other individual attributes and group composition also seemed important. As predicted, hierarchy characteristics indicated a despotic system in line with the strong reproductive skew. Across primates, however, the degree of despotism did not appear to match the degree of reproductive skew. Am J Phys Anthropol, 2016. © 2016 Wiley Periodicals, Inc.

Enamel thickness has played an important role in studies of primate taxonomy, phylogeny, and functional morphology, although its variation among hominins is poorly understood. Macaques parallel hominins in their widespread geographic distribution, relative range of body sizes, and radiation during the last five million years. To explore enamel thickness variation, we quantified average and relative enamel thickness (AET and RET) in Macaca arctoides, Macaca fascicularis, Macaca fuscata, Macaca mulatta, Macaca nemestrina, and Macaca sylvanus. Enamel area, dentine area, and enamel-dentine junction length were measured from mesial sections of 386 molars scanned with micro-computed tomography, yielding AET and RET indices. Intraspecific sex differences were not found in AET or RET. Macaca fuscata had the highest AET and RET, M. fascicularis showed the lowest AET, and M. arctoides had the lowest RET. The latitudinal distribution of macaque species was associated with AET for these six species. Temperate macaques had thicker molar enamel than did tropical macaques, suggesting that thick enamel may be adaptive in seasonal environments. Additional research is needed to determine if thick enamel in temperate macaques is a response to intensified hard-object feeding, increased abrasion, and/or a broader diet with a greater range of food material properties. The extreme ecological flexibility of macaques may prohibit identification of consistent trends between specific diets and enamel thickness conditions. Such complications of interpretation of ecological variability, dietary diversity, and enamel thickness may similarly apply for fossil Homo species.

An accurate description of reproductive characteristics and ovarian endocrinology is necessary to address questions about the reproductive strategies and life history of a species and for meaningful, cross species analyses. Here we used analysis of fecal estradiol (fE) and behavioral observations to determine for the first time the reproductive characteristics and endocrinology of a wild group (N = 18 adult and 3 adolescent females) of Sanje mangabeys (Cercocebus sanjei). The study was conducted in the Udzungwa Mountains National Park, Tanzania, from October 2008 through September 2010. Average cycle length (±SD) was 29.3 ± 3.2 days in adults and 51.4 ± 5.5 days in adolescents. Menses appeared within 5.1 ± 2.1 days in adults and 4.8 ± 0.3 days in adolescents after the end of maximum tumescence, and lasted 6.7 ± 3.1 and 10.3 ± 5.0 days, respectively. Infant death tended to reduce the number of cycles to conception (4.3 ± 1.5 cycles after a surviving infant vs. 2.6 ± 1.0 cycles after infant death). Adolescents cycled for at least 16 months without conceiving. Implantation bleeding began 17.5 ± 0.7 days from the onset of detumescence, and lasted 10.0 ± 1.4 days. Gestation length averaged 171.8 ± 3.4 days. Postpartum amenorrhea lasted 6.7 ± 2.3 months while females whose infants had died resumed cycling within 14.3 ± 5.9 days. The interbirth interval after a surviving infant averaged 20.0 ± 4.3 months. These reproductive characteristics of the Sanje mangabey resembled those of other mangabeys and related cercopithecines, with the exception of an earlier onset and longer duration of menstruation and implantation bleeding. Further information on the physiology of the Sanje mangabey is needed to clarify what factors may cause the unusual characteristics of both, their menses and implantation bleeding. Am. J. Primatol. 76:1163–1174, 2014. © 2014 Wiley Periodicals, Inc.

In primates and other mammals, weaning is an equivocal concept, as is reflected in the numerous ways it is measured: a) first intake of solid food, b) conflict over access to the nipple, c) ability to survive without mother, d) maternal resumption of cycling, or e) the cessation of nipple contact. The lack of a consistent definition means that weaning age, although it falls between gestation (fetal growth) and age at first reproduction (most energy diverted from growth), is currently not a reliable life history variable capturing offspring independence. Using data for wild Phayre's leaf monkeys (Trachypithecus phayrei crepusculus) at Phu Khieo Wildlife Sanctuary, Thailand (51 offspring, four groups), we asked whether the end of nipple contact indicates offspring independence as measured by survival to 3 years. To establish a baseline for the onset of independence, we assessed the youngest age at which individuals were orphaned (15-17 months) but then survived to 3 years. Next we determined that offspring age at last nipple contact (19.0 months) was comparable to two other independently calculated measures: offspring age at mother's first postpartum ovulation (11.5 months), and age at mother's re-conception (15.6 months). Using these separate "starting points," we arrived at similar ages for nipple contact cessation (18.4 and 19.2 months, respectively). Overall, in wild (but not in provisioned) Asian colobines, age at last nipple contact was allometrically related to adult female body mass, supporting its designation as a life history variable. Future comparisons need to show if this holds for other taxa.

We examined historical and ecological factors affecting current primate biodiversity in Southeast Asia. In Africa, Madagascar and South America, but not Southeast Asia, primate species richness is positively associated with average rainfall and distance from the equator (latitude). We predicted that Southeast Asia's non-conformance may be due to the effect of dispersed Pleistocene refuges (locations of constricted tropical forests during glacial maxima which today are at least 305 m in altitude). Based on 45 forested sites (13 on large islands; 32 on the mainland) of at least 100 km(2) to minimize recent human impact, we determined correlations between extant primate species richness and rainfall, latitude and supplementary ecological variables, while controlling for refuges and islands. We found that refuge sites had significantly higher primate species richness than non-refuges (t = -2.76, P < 0.05), and distance from the nearest Pleistocene refuge was negatively correlated with species richness for non-refuge sites (r = -0.51, P < 0.05). There was no difference in species richness between sites on large islands and the mainland (t = -1.4, P = 0.16). The expected positive relationship between rainfall and species richness was not found (r = 0.17, P = 0.28). As predicted, primate species richness was negatively correlated with latitude (r = -0.39, P < 0.05) and positively correlated with mean temperature (r = 0.45, P < 0.05). General linear models indicated that a site's latitude (F1,38 = 6.18, P < 0.05) and Pleistocene refuge classification (F1,42 = 5.96, P < 0.05) were the best predictors of species richness. Both ecological and historical factors contribute to present day primate species richness in Southeast Asia, making its biodiversity less of an outlier than previously believed.

Female dominance hierarchies form as a result of individual differences in resource holding potential, social processes such as winner-loser effects or coalitions, and ecological conditions that favor contest competition. Contest competition is assumed to result in despotic, nepotistic, and stable hierarchies. However, female Hanuman langurs are exceptions to this pattern, with data from provisioned populations indicating despotic, yet individualistic (age-inversed) and unstable hierarchies despite strong within-group contest. We present data on hierarchical linearity, stability, and the determinants of female rank and rank change in a population of unprovisioned, wild Hanuman langurs (Ramnagar, Nepal). Based on 12 490 dyadic displacement interactions collected over 5 years from a medium-sized group (P group, mean = 6.9 adult females) and a large group (O group, mean = 13.6 adult females), stable periods (P group, N = 14 ; O group, N = 31 ) were identified and dominance hierarchies constructed with the program MatMan. In both groups, dominance hierarchies were linear ( p < 0 . 05 ), with high directional consistency within dyads. Rank was negatively related with age, while the presence of maternal kin had no effect. Reproductive state affected dominance rank in the larger group, with females ascending the hierarchy prior to conception, and dropping in rank after birth. Ranks were unstable, with group size and the number of juvenile females driving the effect (GLMM, p < 0 . 001 ). These results match earlier findings for provisioned populations. In female Hanuman langurs, competition seems most intense around conception and during gestation, creating rank instability, which is further exacerbated by the number of adult as well as maturing females.

Recent decades have seen rapid development of new analytical methods to investigate patterns of interspecific variation. Yet these cutting-edge statistical analyses often rely on data of questionable origin, varying accuracy, and weak comparability, which seem to have reduced the reproducibility of studies. It is time to improve the transparency of comparative data while also making these improved data more widely available. We, the authors, met to discuss how transparency, usability, and reproducibility of comparative data can best be achieved. We propose four guiding principles: 1) data identification with explicit operational definitions and complete descriptions of methods; 2) inclusion of metadata that capture key characteristics of the data, such as sample size, geographic coordinates, and nutrient availability (for example, captive versus wild animals); 3) documentation of the original reference for each datum; and 4) facilitation of effective interactions with the data via user friendly and transparent interfaces. We urge reviewers, editors, publishers, database developers and users, funding agencies, researchers publishing their primary data, and those performing comparative analyses to embrace these standards to increase the transparency, usability, and reproducibility of comparative studies.

Asian colobines typically live in small one-male groups (OMGs) averaging five adult females, but Simias concolor (simakobu or pig-tailed langur) is considered an exception because mostly adult male–female pairs have been reported. However, based on their phylogenetic position and marked sexual dimorphism, simakobu are also expected to form OMGs with multiple females. The preponderance of small groups could be the result of human disturbance (hunting or habitat disturbance) reducing group size in the recent past. To investigate this possibility, we documented the demography of ten wild simakobu groups from January 2007 until December 2008 at an undisturbed site, the Peleonan Forest, Siberut Island, Indonesia. We assessed the population-specific size and composition of groups and documented demographic changes due to births, disappearances, and dispersals throughout our 2-year study. We found OMGs with 3.0 adult females on average in addition to all-male groups, but no adult male–female pairs. The ratio of 0.5 infants per adult female (and 0.64 births per female-year in focal groups) suggested that birth rates were similar to those of other Asian colobines. In 5.1 group-years, we observed six dispersal events and six temporary presences (i.e., less than 3 months’ residency). Both males and females dispersed, and juveniles seemed to disperse more frequently than adults. To assess the impact of human disturbance on simakobu demography, we compiled data for seven additional populations from the literature and compared them using multiple regressions. Adult sex ratio and the number of immatures per group were influenced negatively by hunting and positively by habitat disturbance while reproductive rates were not significantly affected by either variable. These findings suggest that adult male–female pairs may result from hunting pressure reducing group size, and that conservation action to reduce hunting in the Mentawai Islands is needed to ensure the survival of this critically endangered species. Am. J. Primatol. 74:580–590, 2012. © 2012 Wiley Periodicals, Inc.

Extended sexual receptivity in primates is thought to facilitate paternity confusion, thus decreasing the risk of infanticide. However, females might also provide some indication of ovulation to attract preferred males during fertile periods. We examined female mate preferences across defined receptive periods (N = 59) in a group of wild Phayre's leaf monkeys (Trachypithecus phayrei crepusculus) at Phu Khieo Wildlife Sanctuary (February–September 2006; 2,603 contact hours). The group contained seven cycling adult females and three reproductively active males (one adult and two adolescents). We predicted that females would prefer the adult male during periovulatory (POP) receptive periods, but the adolescent males during nonperiovulatory (NPOP) and postconceptive (PC) periods. We collected focal and ad libitum data on sexual and agonistic behaviors to determine female preferences and male awareness of female fertility. We also determined the degree of mating overlap to assess if males were capable of monopolizing females. Our results indicate that females were more frequently proceptive and receptive toward the adult male during POP. By contrast, females were more proceptive and receptive toward one of the adolescent males during PC periods, but rarely interacted with the other adolescent. Patterns of attractivity and agonism across receptive periods suggested that the adult male could detect fertility, while the preferred adolescent could not. Finally, we found a high degree of overlap in total receptive period days, but a low degree of overlap in POP receptive days, suggesting that the adult male might have monopolized females, especially since he seemed to be aware of female fertility. Although these results suggest that females provide some information on ovulation, they also suggest that females attempt to confuse paternity, perhaps capitalizing on male differences in the ability to detect fertility. Am. J. Primatol. 74:471-481, 2012. © 2012 Wiley Periodicals, Inc.

Infanticide by males is common in mammalian species such as primates in which lactation lasts much longer than gestation. It frequently occurs in one-male groups following male takeovers and is likely a male reproductive strategy. Reported female countertactics include abrupt weaning of infants, dispersal, or paternity confusion. Here, we estimated costs of female countertactics in terms of weaning ages and interbirth intervals. We observed a population of white-headed leaf monkeys (Trachypithecus leucocephalus) in Nongguan Nature Reserve, China (1995–2006) mainly composed of one-male groups. Takeovers (N = 11) coincided with the peak conception period. Detailed data are presented for five takeovers (34 females, 29 infants, and 47 group-years) leading to six infant disappearances (42.9% of infant mortality). All presumed infanticides were in accordance with the sexual selection hypothesis. Following a takeover, females without infants or with old infants stayed with the new males, incurring no or low costs (via abrupt weaning). Females with young infants dispersing with the old males also experienced low costs. High costs (due to infant loss) were incurred by pregnant females and those with young infants who stayed with the new males indicating that paternity was not confused. Costs in terms of long interbirth intervals were also high for females leaving with the old males to later join the new males, despite infant survival. Female countertactics reflected female philopatry mediated by infant age. Presumably due to the seasonal timing of takeovers, most countertactics seemed successful given that 32.3% of females apparently incurred no costs and 41.2% incurred only low costs.

Although it is known that forest loss and degradation negatively impact most forest-dwelling primates, such relationships are difficult to quantify because many primates are difficult to survey over large areas. Furthermore, recovery times are also difficult to assess due to a lack of long-term data. Here, we determined how forest characteristics and habitat disturbance correlate with the abundance of pileated gibbons, Hylobates pileatus. We studied a population in Khao Ang Rue Nai Wildlife Sanctuary in southeastern Thailand, assessed its density using an auditory method combined with distance sampling at 24 randomly placed sample sites. In addition, we determined how simple forest structural characteristics and habitat disturbance correlate with the gibbon abundance. Average gibbon density per site was 1.02 ± 0.16 (SE) groups/km 2 (range 0—2.74). Bivariate analyses indicated that densities depended on food tree biomass, level of disturbance, evergreen forest cover, time since protection, and distance to the sanctuary boundary. Multiple regression analysis suggested evergreen forest cover and distance to boundary were the most influential factors. Because evergreen forest cover, time since protection, and habitat disturbance are correlated, these results suggest a direct dependence of gibbon densities on mature, undisturbed evergreen forest. While gibbons can persist in disturbed areas if the forest is protected, it appears that recovery to previous densities may take decades. We suggest that this is due to the slow pace of forest regeneration and/or poor recovery potential of gibbons.

Female reproduction is known to be influenced by food availability and its impact on energetic status. However, emerging evidence suggests that the phytochemical content of food may also be an important factor. Here, we investigated this hypothesis, presenting 20 months of data on fecal progestin (fP) patterns in wild female Phayre's leaf monkeys (Trachypithecus phayrei crepusculus). We examined whether (a) the availability of Vitex (a plant known to contain phytochemicals) might be linked to seasonal fP levels, (b) fP levels were associated with female reproductive performance, and (c) reproductive performance might also be linked with energetic status (as measured by physical condition). We collected fecal samples (N = 2077) from 10 adult females to analyze estrogen (fE) and progestin (fP) metabolites, behavioral data from 7 cycling females to determine receptivity, and monthly data on Vitex availability and female physical condition. Seasonally elevated fP levels were found in all females, with higher levels when Vitex leaves and fruits were abundant. During the period of high progestins, females had longer cycle lengths and follicular phases, while receptive periods did not change. Nevertheless, when ovulations occurred, females were more likely to conceive. On the other hand, conceptions were also more likely when physical condition was improving, suggesting that the effects of phytochemicals and energetic status on reproduction may be difficult to separate. Although our results support the predicted effects of Vitex on endocrine and reproductive function, future studies with detailed feeding data and chemical analyses of plants are needed to confirm this finding.

The evolution of social monogamy in larger mammals is difficult to explain because males usually do not invest much in direct offspring care and might achieve greater fitness by deserting a pregnant female to reproduce with additional females elsewhere. It has been hypothesized that socially monogamous males remain with the female year-round to protect their offspring from infanticide by new immigrant males. We investigated this idea by analyzing all cases of infant loss in a wild population of white-handed gibbons (Hylobates lar; Primates), in which most groups were socially monogamous and some polyandrous (137.5 group years). We examined the influence of (a) male intruder pressure on male immigration rates and (b) the presence of a new male in the group on infant loss. We found no relation between intruder pressure and male immigration rates. Infant loss was lowest (4.5%) for stable monogamy (probable father stayed from conception through infancy) and intermediate (25.0%; p = 0.166) for stable polyandry. If a new male immigrated after conception, however, the infant was lost in all cases (p

Understanding female reproductive characteristics is important for assessing fertility, interpreting female behavior, and designing appropriate conservation and captive management plans. In primate species lacking morphological signs of receptivity, such as most colobines, determination of reproductive parameters depends on the analysis of reproductive hormones. Here, we use fecal hormone analysis to characterize cycle patterns (N=6 females) and gestation length (N=7 females) in a group of wild Phayre's leaf monkeys (Trachypithecus phayrei crepusculus) in Phu Khieo Wildlife Sanctuary, Thailand. We found that both fecal estrogen (fE) and progestin (fP) levels showed clear biological patterns indicative of ovulation and conception. However, because fP patterns were inadequate in determining the end of the luteal phase, we used fE rather than fP patterns to delineate menstrual cycle parameters. We found a mean cycle length of 28.4 days (N=10), with follicular and luteal phases of 15.4 (N=10) and 12.5 days (N=14), respectively. On average, females underwent 3.57 (N=7) cycles until conception. Average gestation length was 205.3 days (N=7), with fE levels increasing over the course of pregnancy. Overall, the reproductive characteristics found for Phayre's leaf monkeys were consistent with results for other colobine species, suggesting that fecal hormone monitoring, particularly for fE metabolites, can provide useful reproductive information for this species. Am. J. Primatol. 72:1073–1081, 2010. © 2010 Wiley-Liss, Inc.

Primate life histories are strongly influenced by both body and brain mass and are mediated by food availability and perhaps dietary adaptations. It has been suggested that folivorous primates mature and reproduce more slowly than frugivores due to lower basal metabolic rates as well as to greater degrees of arboreality, which can lower mortality and thus fecundity. However, the opposite has also been proposed: faster life histories in folivores due to a diet of abundant, protein-rich leaves. We compared two primate taxa often found in sympatry: Asian colobines (folivores, 11 species) and Asian macaques (frugivores, 12 species). We first described new data for a little-known colobine (Phayre's leaf monkeys, Trachypithecus phayrei crepusculus) from Phu Khieo Wildlife Sanctuary, Thailand. We then compared gestation periods, ages at first birth, and interbirth intervals in colobines and macaques. We predicted that heavier species would have slower life histories, provisioned populations would have faster life histories, and folivores would have slower life histories than frugivores. We calculated general regression models using log body mass, nutritional regime, and taxon as predictor variables. Body mass and nutritional regime had the predicted effects for all three traits. We found taxonomic differences only for gestation, which was significantly longer in colobines, supporting the idea of slower fetal growth (lower maternal energy) compared to macaques and/or advanced dental or gut development. Ages at first birth and interbirth intervals were similar between taxa, perhaps due to additional factors (e.g., allomothering, dispersal). Our results emphasize the need for additional data from wild populations and for establishing whether growth data for provisioned animals (folivores in particular) are representative of wild ones.

“‘What is the matter?’ she [Alice] said, as soon as there was a chance of making herself heard. ‘Have you pricked your finger?’ ‘I haven't pricked it yet,’ the [White] Queen said, ‘but I soon shall – oh, oh, oh!’” (emphasis as in1:82)

Dominance hierarchies based on resource holding potential (RHP) or age are central to the social structure of many group-living animals. Nonhuman primate females and some other mammals are unusual because ranks can depend on kin support or follow an inverse age-graded pattern independent of kin. Whereas the former hierarchies are mediated by frequent coalitions leading to long-term stability, the latter are unstable due to individualistic contests and occasional revolutionary coalitions. We use mathematical modeling to investigate this variation, based on the interaction of triads, subgroups of the group consisting of 3 individuals. This produces 4 key situations, namely that relationships can be either stable or unstable, and involving coalitionary support or not. Varying the parameter values of our model and applying it to real-world examples, we conclude that the emergence of different dominance hierarchies depends on 6 major factors. In addition to RHP, reproductive skew and demography often suggested previously, we stress the importance of 1) the variance in relatedness; 2) the costs and likelihood of interventions; and 3) the value of resources and rank in different life stages, which might be tightly linked to the life history of a species. Key words: ageinversed hierarchies, ESS, evolutionary games, matrilineal hierarchies, triads. [Behav Ecol]

Primate population assessments provide the basis for comparative studies and are necessary prerequisites in determining conservation status. The most widely used assessment method is line transect sampling, which generates systematic data quickly and comparatively inexpensively. In contrast, the presumably most reliable method is long-term monitoring of known groups, which is both slow and costly. To assess the reliability of various analytical methods, we compared group and population densities for white-handed gibbons (Hylobates lar carpenteri) and Phayre’s leaf monkeys (Trachypithecus phayrei crepusculus) derived from transect walks with those from long-term group follows at Phu Khieo Wildlife Sanctuary, Thailand. Our assistants and we regularly walked a 4-km transect over 30 mo (480 km total), resulting in 155 gibbon sightings and 125 leaf monkey sightings. We then estimated densities via 1) DISTANCE and 2) the Kelker method based on perpendicular distances (PD) or animal-to-observer distances (AOD). We compared the 3 estimates to values based on known home ranges (95% kernels), accounting for home range overlap, combined with group size data. Analyses of line transect data consistently overestimated group densities for both species, while underestimating group size for leaf monkeys. Quality of results varied according to the group size and spread of each species. However, we found, in accordance with previous studies, that values derived via AOD (or its derivations) matched most closely with population estimates based on home range data.

Patterns of agonistic relationships among group-living mammalian females are suggested to reflect differences in feeding competition. Socioecological models predict that within-group contest (WGC) competition leads to despotic relationships that are linear, consistent, and stable, in which subordinate individuals use formal submission to acknowledge rank. Between-group contest (BGC) competition and risk of infanticide are predicted to add tolerance, as evidenced by frequent reconciliation, low-intensity aggression and counteraggression. We examined the predicted effects for formal submission and two criteria of tolerance for a group of Hanuman langurs ( Semnopithecus entellus ) in a population known to experience WGC and BGC as well as high rates of infanticide. Results were based on focal animal sampling of eight of 13 adult females (419 h) and ad libitum sampling of all 13 females (711 contact hours) in the group Kailana I (Jodhpur, India). The behaviour ‘presenting’ ( N = 281) qualified as formal submission, adhering to most proposed criteria (e.g. linear, consistent, predictive of other agonistic relationships and hierarchies, 40.9% spontaneous). Thus, regarding formal submission, the predicted effects of WGC hold, and support the explanatory value of this aspect of socioecological models. In contrast, agonistic relationships were nontolerant, with frequent high-intensity aggression (48.9% of N = 231) and almost no counteraggression (

Feeding competition is suggested as a major factor constraining group size in social foragers. It has, however, been challenging to demonstrate consequences of reduced energy gain in terms of fitness, possibly because social foragers may compensate negative effects of scramble competition via adjustments in time budgets. Herbivorous animals are particularly interesting in this context because their fibrous diet and slow digestion process may make it difficult to adjust time budgets. Here we investigate infant development and reproductive rates in Phayre’s leaf monkeys (Trachypithecus phayrei) at Phu Khieo Wildlife Sanctuary, Thailand. The diet of the species consists of 39.0% leaves (maximum 81.2% per month). Our analysis is based on data for 3 groups (185 group months) of different sizes (mean 11.4, 18.3, and 25.8 individuals, respectively). Infant development was significantly slower in the large group, in which infants changed to the adult coat later than in the medium-sized group (20.3 vs. 26.3 weeks) and were older when weaned (18.3, 19.7, and 21.4 months, respectively). The interbirth interval after a surviving infant significantly increased with group size (21.3, 22.8, and 24.5 months) while rearing success did not differ (77.8%, 76.5%, and 82.4%, survival to 2 years). Thus, infants in the large group developed more slowly were weaned later and females reproduced more slowly. With similar infant survival rates, these different reproductive rates indicate fitness differences across groups. As in other herbivores, these group-specific differences may reflect scramble competition for food or differences in habitat quality. Key words: infant development, infant mortality, interbirth interval, reproductive costs, scramble competition, Trachypithecus phayrei. [Behav Ecol]

The occurrence of genital swellings was examined in adult female silvered langurs (Trachypithecus cristatus). In contrast to the exaggerated swellings found in cercopithecines and apes, genital swellings in silvered langurs are confined to the vulva and the surrounding perineum, but they may nevertheless convey information similar to that of exaggerated swellings (i.e., correlate with the receptive period and fertility). If so, genital swellings would be expected to occur most frequently in cycling females, and sexual behavior and male interest should most frequently involve females with swellings. Swellings during gestation, if they occur at all, should be most pronounced at the beginning. Swelling sizes (in three size categories) in nine adult females were examined throughout different reproductive states (cycling, pregnant, and lactating), and in relation to proceptivity, receptivity, and attractivity. Data were collected from November 2002 through March 2004 (on 500 of the 502 calendar days) at the Wildlife Conservation Society's Bronx Zoo. Female sexual behavior (proceptivity and receptivity) and male inspection (attractivity) were recorded each day for 6 hr by video camera (2,948 hr total) and analyzed as present or absent for each female day. Swellings were assessed directly (not from videotapes). In contrast to the predictions, swellings occurred significantly less frequently in cycling females (compared to pregnant females) and no regular, cyclic pattern could be detected. Some females conceived without a swelling. Female attractivity was independent of swellings but coincided with proceptive behavior. Swellings occurred most frequently in pregnant females, especially toward the end of the gestation period. Therefore, genital swellings in silvered langurs are not similar to exaggerated swellings or the smaller genital swellings that have been described for some other primates. It is currently not clear what they signal to conspecifics. Male behavior needs to be studied in more detail, and the hormonal basis for these swellings should be explored.

Objectives Infanticide by males is assumed to promote permanent male–female associations, although its importance for social monogamy is still debated. We examined the consequences of male membership change in the largest socially monogamous primate, the siamang (Symphalangus syndactylus), a species that also forms polyandrous groups and where males may provide offspring care. We examine (a) the potential risk of infanticide by documenting changes in female–offspring relationships following male change, expecting abrupt weaning; and (b) the potential importance of male care and polyandry for offspring survival. Methods We witnessed four male changes at Way Canguk Field Station (Indonesia) in groups where the youngest offspring was estimated to be about 2 years of age. We quantified aspects of mother–offspring relationships (interindividual distance, body contact, and nipple contact) and estimated the proportion of time being carried during travel (August 2007–April 2009). Results In the two groups into which new males immigrated, offspring were weaned abruptly, suggesting infanticide by males as a potential risk. Even though no attacks on infants by males were witnessed, both infants disappeared shortly after the male change. Another infant disappeared after its main carrier (a male) emigrated. Conversely, in a group where the resident subordinate male replaced the dominant male, the offspring experienced no change in care and survived the transition. Conclusions These findings suggest that male immigration in hylobatids increases mortality risk for infants, emphasize the importance of offspring care, and cast doubts on prior classifications of 2-year-old siamangs as independently traveling individuals.

Reproductive skew is proposed to link to despotism in dominance hierarchies. While studies illustrating male skew are plentiful, demonstrating the link to despotism is rare. Likewise, it is often unknown which factors (e.g., resource holding potential, age, physical condition) affect a male's dominance rank. Here we investigated correlates of male rank and hierarchy characteristics in Nepal gray langurs (Semnopithecus schistaceus), a population with high male reproductive skew, and compared the results to other multi-male groups of nonhuman primates.We collected dyadic displacement interactions from two groups (mean 3.0 and 4.1 adult males) for five years each. We assessed dominance ranks for demographically stable phases (n = 11, n = 28) and analyzed the effects of age and physical condition through linear mixed models (LMM). We analyzed hierarchy characteristics via the program MatMan. We used data from 27 primate groups (cercopithecines, colobines, hominoids) as a comparative sample.The highest ranks were attained by adults in one group (LMM, P 0.091) and by young adults in the other group (LMM, P 0.001). With some exceptions, rank was highest for males with higher physical condition scores (LMM, P 0.05). Hierarchies had high directional consistency (mean 0.93) and linearity (mean0.81) and were relatively steep (mean0.66) when compared with other species.Dominance rank followed a pattern predicted by resource holding potential, but other individual attributes and group composition also seemed important. As predicted, hierarchy characteristics indicated a despotic system in line with the strong reproductive skew. Across primates, however, the degree of despotism did not appear to match the degree of reproductive skew. Am J Phys Anthropol 160:208-219, 2016. © 2016 Wiley Periodicals, Inc.

In species with a high risk of infanticide, a conflict of interest exists between the sexes over the amount of paternity information that is available to males. While females are expected to keep males unaware of their reproductive status in order to confuse paternity, selection should favor those male traits that enhance the males' assessment of female status and consequently of paternity probability. In Hanuman langurs (Semnopithecus entellus), a species that is extremely vulnerable to infanticide, females have been shown to successfully conceal the exact timing of ovulation from males--perhaps because they exhibit no sexual swelling and mate during all reproductive phases, including gestation. Nevertheless, it remains unclear whether males have hitherto unrecognized information about females' reproductive condition on a broader level that could still enhance male reproductive success. We investigated male assessment of female reproductive states in a population of wild Hanuman langurs as indicated by changes in male behavior, such as rates of copulations, anogenital inspections, and consortships, in relation to different female receptive periods (pregnant, fertile-nonconceptional, and conceptional). Our data indicate that males were able to discern qualitatively distinct reproductive states. Males were more interested in fertile than pregnant females, as indicated by higher copulation rates. Based on consortships, males distinguished fertile from nonfertile phases, as well as fertile, nonconceptional receptive periods from conceptional ones. Hanuman langur males are thus not as unaware of female reproductive condition as previously thought, supporting the idea of an ongoing battle of the sexes over paternity information. However, granting some knowledge while at the same time concealing the exact day of ovulation may also reflect a pure female strategy of balancing paternity concentration with paternity confusion, which is the most likely strategy in this system with high infanticide risk and male defense of infants.

In polygynous, sexual dimorphic species with higher variance in male reproductive success compared with females, females are expected to invest more heavily in sons than daughters within the constraints imposed by their physical condition (Science 1973; 179:90). Mothers in good condition, usually those of high rank, should produce more sons than females in poor condition or of low rank. We investigated sex allocation and sex-biased maternal investment in a population of wild Hanuman langurs using rank and group size as approximations of female physical condition. Our results show that reproductive costs of sons were higher with both significantly longer interbirth intervals following male births and longer lactational periods for sons. Not in all groups did analyses of rank-dependent sex allocation reveal the expected pattern of high-ranking mothers producing more sons. However, sex ratio was significantly influenced by group size, with females from larger groups, i.e., in worse physical condition, producing a daughter-biased sex ratio. In fact, only females of population-wide superior physical condition can be expected to produce sons, because in Hanuman langurs males disperse and compete population-wide. Thus, our results support the Trivers–Willard model and may explain the mixed evidence accruing from studies of single groups. We present a graphical model of how group size and dominance-related differences in energy gain may influence sex allocation under different competitive regimes relative to overall resource availability. Tests of adaptive sex allocation models should consider whether reproductive competition of the preferred sex takes place primarily within a group or within the population.

Infanticide by primate males was considered rare if groups contain more than one adult male because, owing to lower paternity certainty, a male should be less likely to benefit from infanticide. Guided by recent evidence for strong variation of infanticide in primate multi-male groups, we modelled the conditions for when infanticide should occur for a group with a resident and an immigrant male. Setting the parameters (e.g. infant mortality, reduction of interbirth interval, life-time reproductive success, genetic representation) to fit the conditions most commonly found in nature, we develop a game-theoretic model to explore the influence of age and dominance on the occurrence of infanticide and infant defence. Male age strongly impacts the likelihood of an attack which is modified by the father's defence. If the new male is dominant he is likely to attack under most circumstances whereas a subordinate male will only attack if the father does not defend. These model scenarios fit the conditions under which infanticide is known to occur in primate multi-male groups and offer an explanation why infanticide is common in some multi-male groups and rare in others. Overall, the benefits for infanticidal males are strongly governed by a reduced interbirth interval while advantages via improved genetic representation in the gene pool contribute but a minor fraction.

The time spent between sleeping periods, which is called the active period, has to accommodate all essential activities, including feeding, resting, social behavior, and reproduction. To minimize costs in terms of, e.g., predation risk, suboptimal foraging, or sleep deficiency, the active period of diurnal animals should be less than or equal to the daylight period. Thus, the active period of an animal should be shaped by local environmental conditions as well as by metabolic and reproductive demands. Chimpanzees, which exhibit reduced predator pressure and a flexible fission-fusion society, were chosen as a model to explore these links. We investigated the influence of sex, female reproductive status, dominance rank, and season on the duration of the active period of adult chimpanzees at Gombe National Park, Tanzania (1975-1992). Sexually nonreceptive females had shorter active periods compared to males, while receptive females had even longer active periods than males. Dominance rank did not influence the duration of the active period of nonreceptive females, but high- and middle-ranking males had shorter active periods compared to low-ranking males. Nonreceptive females exhibited longer active periods during the dry season than in the wet season. No seasonal effect was discovered for males, perhaps because they already had long active periods in the wet season. Nonreceptive females seem to be able to accommodate all essential activities in the daylight period available, probably because they live less socially than males. Thus, the active period does not reflect differences in female competitive abilities, but does reflect such differences in males. The duration of the active period appears to be a simple, reliable tool for exploring basic responses and constraints in animal societies.

In species with strong male-male competition, access to females in multimale-multifemale groups is usually regulated via a dominance hierarchy. The highest ranking (alpha) male often has priority of access and sires most offspring. The alpha male can change in three basic ways: (i) a recent immigrant or a resident challenges and becomes the new alpha; (ii) formation of a new group; (iii) succession-becoming alpha after higher ranking males have left. When, in a given primate population, the alpha male changes in different ways, two questions arise: (a) which is the most successful tactic and (b) do male attributes, such as age, aggressiveness or propensity to commit infanticide, affect the outcome? We examined these questions in the seasonally breeding Nepal gray langurs (Semnopithecus schistaceus) at Ramnagar, where new alpha males were either recent immigrants or residents. Success was measured as alpha tenure, residency duration, and the number of offspring sired (paternity exclusion based on DNA analysis, 28 infants). We documented 12 alpha-male tenures across two multimale-multifemale groups between 1991 and 1997. The predominant mode of change was the immigrant tactic. Age had no effect perhaps because alpha males were among the youngest adult males in their group. As expected, infanticidal males performed similarly to non-infanticidal ones. Alpha tenure was highly variable and longer for immigrant alphas and hyper-aggressive ones. However, none of the tactics or attributes examined resulted in significantly longer residencies or more offspring, likely because of the timing of immigrations and stochastic effects (i.e., the number of conceptions occurring per alpha tenure). The influence of female mate choice on male reproductive success requires further investigation. Furthermore, it remains to be examined why resident alpha males-with their presumed better knowledge of their opponents -performed so poorly. Am. J. Primatol. 79:e22437, 2017. © 2015 Wiley Periodicals, Inc.

To determine nutritional and social influences on life history traits and mating behavior in female langur monkeys, new long-term data for a feral population living in multi-male groups under poor nutritional conditions are presented and compared with published long-term data for the same species living in one-male groups under excellent nutritional conditions. Under poor nutritional conditions, conceptions were confined to 5 months per year, coinciding with the period of the highest mating activity. Age at first parturition, gestation periods, lactation periods, and interbirth intervals were all significantly increased resulting in a more than 50% reduced reproductive rate under poor conditions. The pattern of sexual behavior during gestation was influenced by neither nutritional nor social conditions but the frequency was higher in multi-male groups. Furthermore, estrus length was significantly increased in multi-male groups where females tended to mate with as many resident males as available. Both phenomena, together with a high percentage of sexual behavior occurring outside the mid-cycle estrus, are interpreted in the light of paternity confusion in multi-male groups in a species where infanticide by males is regularly observed. The extreme variation in life history traits and mating patterns disclosed here in one species suggests that future cross-species comparisons should strictly select datasets based on local ecological and social conditions.

Hypotheses about the evolution of gregariousness and social organisation in primates are based on ecological explanations as well as on social factors such as conspecific threat (especially infanticide by males). The social explanation fits well with the conditions found in strepsirrhine primates and furthermore explains why infanticide in anthropoid primates living in one-male groups mainly occurs when the resident male (protector) is replaced. However, whether it likewise fits to the conditions in multimale groups will depend on the role of resident males as infant protectors, which has rarely been examined. We investigated long-term data of wild Hanuman langurs (Presbytis entellus) inhabiting a forest near the village of Ramnagar (southern Nepal). Twenty-two eye-witnessed attacks on infants by males were analysed in connection with male residency, paternity (DNA analyses) and sexual behaviour. Adult males played a major role in infant defence (65%). Only the genetic father or males who had been residents when the infant was conceived were observed to protect infants. Males who immigrated after a female had conceived may later attack her infant and were never observed to defend it. lt seems that the males took only copulations with potentially fertile females but not with pregnant females as clues for paternity. In the light of these results it seems likely that the risk of infanticide is an important determinant in female-male associations even in anthropoid primate multimale groups.

Ecological conditions, feeding and ranging behaviour as well as physical condition of female Hanuman langurs (Presbytis entellus) were studied in order to analyse the major influences on the timing of reproductive events. The langurs inhabited a semi-evergreen forest dominated by Sal trees (Skorea robusta). The climate was dominated by the summer monsoon lasting from May to September. Availability of plant phytophases was markedly seasonal, with fruiting as well as flushing of the six main food plants during the monsoon. Feeding and ranging behaviour of the females varied with food availability and its distribution. High dietary quality was only obtained during a short period from May to July. Physical condition of the females varied in accordance with dietary quality and energy expenditure, with a good physical condition during the second half of the year. Births occurred from January until June, with most births around the energetically worst period of the year. The results indicate that, in contrast to several other non-human primates, most females of this population gave birth too early to meet the optimum lactation requirements. Complete weaning mainly took place during an energetically bad time of the year (October to December). However, it can be inferred that the reduction of suckling bouts prior to the next conception took place when high quality food was available. Conceptions were confined to the months of July to November, coinciding with the time of best physical condition. Moreover, females who conceived were in a better physical condition than those who failed to conceive, suggesting a primary influence of nutrition on the probability of conception. Seasonality and timing of births may be the result of a regular seasonal shortfall of resources influencing the ability of the females to ovulate and to conceive.

In the seasonally breeding langur (Presbytis entellus) population of Ramnagar, South Nepal, where multimale groups prevail, 25 attacks on 11 infants (including one actual killing) by seven adult males were witnessed in five groups by six observers between 1990 and 1996. Circumstantial evidence also indicates three additional attempts at infanticide and in seven additional cases infanticide was presumed or likely. Infanticide presumably accounted for 30.8–62.5% of infant mortality in the first 2 years of life. Most attackers (91.4%) were residents of the infants' group and had immigrated after the infants had been born (75.0%) or conceived (25.0%). Thus, they were not related to the victims. The interbirth interval was shortened if an infant died either prior to September of its 1st year of life (mean = 1.2 years), or its 2nd year (mean = 2.0) and even its 3rd year (mean = 2.4). All attackers remained in the group at least until the next mating season; high-ranking males maintained their dominance rank and lower-ranking males rose in rank. Since rank and mating success were correlated and rank and reproductive success might be correlated, all attackers had a good chance of siring the next infant of the victims' mothers and could thus have benefited by their action. Infanticide seems to be a male reproductive strategy at Ramnagar. Infanticide has never before been reported among seasonally breeding langurs living at such low densities. This is also the first detailed report of infanticide as a male reproductive strategy in a seasonally breeding primate population.

We investigated sex differences in the social behavior of immature Hanuman langurs (Presbytis entellus) in the light of sex-specifically different life-courses and Hanuman langur characteristics, such as the individualistic dominance hierarchy and the rarity of intragroup coalitions among adult females. We observed four immature female and four immature male langurs—all members of the same free-ranging multimale multifemale group in Ramnagar, South Nepal—from November 1992 to February 1993 for 288 hr via focal-animal and instantaneous sampling techniques. Immature females spent significantly more time in proximity to other group members than immature males did. They had more physical contact and groomed more. Other immature females were their preferred social partners. Immature males also preferred like-aged females. They restricted their relationships with other immature males to proximity and occasional grooming. Monitoring was directed especially toward adult males. Female behavior can be interpreted as oriented toward integration into the female social network and their age-inverted dominance hierarchy. Males seem to prepare for leaving their natal group and for future strong intrasexual competition.

In primates and other mammals, weaning is an equivocal concept, as is reflected in the numerous ways it is measured: a) first intake of solid food, b) conflict over access to the nipple, c) ability to survive without mother, d) maternal resumption of cycling, or e) the cessation of nipple contact. The lack of a consistent definition means that weaning age, although it falls between gestation (fetal growth) and age at first reproduction (most energy diverted from growth), is currently not a reliable life history variable capturing offspring independence. Using data for wild Phayre's leaf monkeys (Trachypithecus phayrei crepusculus) at Phu Khieo Wildlife Sanctuary, Thailand (51 offspring, four groups), we asked whether the end of nipple contact indicates offspring independence as measured by survival to 3 years. To establish a baseline for the onset of independence, we assessed the youngest age at which individuals were orphaned (15-17 months) but then survived to 3 years. Next we determined that offspring age at last nipple contact (19.0 months) was comparable to two other independently calculated measures: offspring age at mother's first postpartum ovulation (11.5 months), and age at mother's re-conception (15.6 months). Using these separate "starting points," we arrived at similar ages for nipple contact cessation (18.4 and 19.2 months, respectively). Overall, in wild (but not in provisioned) Asian colobines, age at last nipple contact was allometrically related to adult female body mass, supporting its designation as a life history variable. Future comparisons need to show if this holds for other taxa.

Enamel thickness has played an important role in studies of primate taxonomy, phylogeny, and functional morphology, although its variation among hominins is poorly understood. Macaques parallel hominins in their widespread geographic distribution, relative range of body sizes, and radiation during the last five million years. To explore enamel thickness variation, we quantified average and relative enamel thickness (AET and RET) in Macaca arctoides, Macaca fascicularis, Macaca fuscata, Macaca mulatta, Macaca nemestrina, and Macaca sylvanus. Enamel area, dentine area, and enamel-dentine junction length were measured from mesial sections of 386 molars scanned with micro-computed tomography, yielding AET and RET indices. Intraspecific sex differences were not found in AET or RET. Macaca fuscata had the highest AET and RET, M. fascicularis showed the lowest AET, and M. arctoides had the lowest RET. The latitudinal distribution of macaque species was associated with AET for these six species. Temperate macaques had thicker molar enamel than did tropical macaques, suggesting that thick enamel may be adaptive in seasonal environments. Additional research is needed to determine if thick enamel in temperate macaques is a response to intensified hard-object feeding, increased abrasion, and/or a broader diet with a greater range of food material properties. The extreme ecological flexibility of macaques may prohibit identification of consistent trends between specific diets and enamel thickness conditions. Such complications of interpretation of ecological variability, dietary diversity, and enamel thickness may similarly apply for fossil Homo species.

An accurate description of reproductive characteristics and ovarian endocrinology is necessary to address questions about the reproductive strategies and life history of a species and for meaningful, cross species analyses. Here we used analysis of fecal estradiol (fE) and behavioral observations to determine for the first time the reproductive characteristics and endocrinology of a wild group (N = 18 adult and 3 adolescent females) of Sanje mangabeys (Cercocebus sanjei). The study was conducted in the Udzungwa Mountains National Park, Tanzania, from October 2008 through September 2010. Average cycle length (±SD) was 29.3 ± 3.2 days in adults and 51.4 ± 5.5 days in adolescents. Menses appeared within 5.1 ± 2.1 days in adults and 4.8 ± 0.3 days in adolescents after the end of maximum tumescence, and lasted 6.7 ± 3.1 and 10.3 ± 5.0 days, respectively. Infant death tended to reduce the number of cycles to conception (4.3 ± 1.5 cycles after a surviving infant vs. 2.6 ± 1.0 cycles after infant death). Adolescents cycled for at least 16 months without conceiving. Implantation bleeding began 17.5 ± 0.7 days from the onset of detumescence, and lasted 10.0 ± 1.4 days. Gestation length averaged 171.8 ± 3.4 days. Postpartum amenorrhea lasted 6.7 ± 2.3 months while females whose infants had died resumed cycling within 14.3 ± 5.9 days. The interbirth interval after a surviving infant averaged 20.0 ± 4.3 months. These reproductive characteristics of the Sanje mangabey resembled those of other mangabeys and related cercopithecines, with the exception of an earlier onset and longer duration of menstruation and implantation bleeding. Further information on the physiology of the Sanje mangabey is needed to clarify what factors may cause the unusual characteristics of both, their menses and implantation bleeding.