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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.

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.

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.

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.

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.

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.

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 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.

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.

Life history variables such as the age at first reproduction and the interval between consecutive births are measures of investment in growth and reproduction in a particular population or species. As such they allow for meaningful comparisons of the speed of growth and reproduction between species and between larger taxa. Especially in primates such life history research has far reaching implications and has led for instance to the "grandmother hypothesis". Other links have been proposed with respect to dietary adaptations: Because protein is essential for growth and one of the primary sources of protein, leaves, occurs much less seasonally than fruits, it has been predicted that folivorous primates should grow faster compared to frugivorous ones. However, when comparing folivorous Asian colobines with frugivorous Asian macaques we recently documented a longer, instead of a shorter gestation length in folivores while age at first reproduction and interbirth interval did not differ. This supports earlier findings for Malagasy lemurs in which all life history variables tested were significantly longer in folivores compared to frugivores. Wondering why these trends were not apparent sooner, we tried to reconstruct our results for Asian primates with data from four popular life history compilations. However, this attempt failed; even the basic, allometric relationship with adult female body mass that is typical for life history variables could not be recovered. This negative result hints at severe problems with data quality. Here we show that data quality can be improved significantly by standardizing the variables and by controlling for factors such as nutritional conditions or infant mortality. Ideally, in the future, revised primate life history data should be collated in a central database accessible to everybody. In the long run such an initiative should be expanded to include all mammalian species.

Recent advances in DNA and isotope analyses have allowed tentative reconstructions of dispersal strategies of Plio-Pleistocene hominins.(1,2) Comparing their findings to dispersal patterns of some extant apes and humans suggested groups of related males and unrelated females in Neandertals indicating patrilocality(2) and Pan-like male philopatry in australopiths.(1) Here we review the demographic, ethnographic, and genetic evidence of dispersal patterns in extant apes and humans and compare the results to the suggestions for Plio-Pleistocene hominins. We find that alternative dispersal patterns, for example among gorillas or gibbons, could explain the findings of related or natal males in a confined geographic area. Based on sexual size dimorphism, we speculate that gorillas might currently be the best model for reconstructing dispersal in robust australopiths. Given that the sexual size dimorphism in other australopiths is still hotly debated, the question of which hominoid model best matches their dispersal pattern must remain unanswered. Neandertal dispersal patterns have been compared to patrilocality of modern humans. However, the latter is related to the advent of food production. Consequently, hunter-gatherers exhibiting primarily multilocality appear to be the better comparison for Neandertals. Overall, human-like patrilocality and Pan-like male philopatry appear to be poor models for the reconstruction of dispersal patterns in Plio-Pleistocene hominins.

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.

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.

In Asian colobines, small one-male groups (OMG) seem to predominate alongside all-male groups (AMG), while larger multimale groups (MMG) are rare, but are reported for Hanuman langurs and red-shanked douc langurs. Recently, however, it has been speculated that the genus Pygathrix could have multilevel societies based on (1) a theoretical extension of the multilevel societies found in Rhinopithecus to all odd-nosed colobines and (2) first data for black-shanked douc langurs. This assumes bands composed of small OMG with a skewed adult sex ratio. Band size may vary with seasonal food availability resulting in smaller bands when feeding competition is increased. To investigate the social organization of red-shanked douc langurs and potential seasonal influences, we observed 2 unhabituated groups at Hin Namno National Protected Area, Lao PDR from March 2007 to August 2008 for 803 hr. We recorded births and performed group counts and scan sampling of feeding behavior. Most births (79% of N = 15) occurred from June-September, indicating a 4-month peak conception season from November to February. Group size averaged 24.5 individuals (range 17-45) with 2.45 adult males (range 1-4). Although the smaller group remained at a stable size (about 18 individuals), the larger group reduced from about 45 to 25 individuals during the 7-months long lean season, when less than 50% of the feeding time was spent on fruits. This suggests feeding competition as a potential cause of seasonal variation in group size. With 1.9 females per male the skew in adult sex ratio was much lower compared with Rhinopithecus, indicating MMG rather than multilevel societies. However, data on the spacing and interaction patterns between recognized individuals need to be collected and analyzed before the social organization can be determined. Detailed ecological data are furthermore required to investigate the basis for the seasonal changes in group size found.

The ecological model of Carel van Schaik provides clear predictions for female-female relationships in relation to scramble or contest within-group competition and contest between-group competition. These predictions were applied to data from a 12-year field study on Hanuman langurs (Presbytis entellus) that ranged freely around Jodhpur (India). It appears that hierarchical relationships between females (unstable, inconsistent, individualistic, with low rates of agonistic coalitions) reflect scramble within-group competition. Such competition, however, results in individual evolutionary advantages (differential feeding time; differential reproductive success) so that dominance effects mirror contest within-group competition. Between-group contest competition at Jodhpur is strong (low predator pressure, high population density, good food defensibility, essential role of females during intergroup encounters, loud vocalization of males). The results are discussed in the light of langur feeding habits.

Selection and use patterns of sleeping sites in nonhuman primates are suggested to have multiple functions, such as predation avoidance, but they might be further affected by range defense as well as foraging constraints or other factors. Here, we investigate sleeping tree selection by the male and female members of one group of pileated gibbons (Hylobates pileatus) at Khao Ang Rue Nai Wildlife Sanctuary, Thailand. Data were collected on 113 nights, between September 2006 and January 2009, yielding data on 201 sleeping tree choices (107 by the female and 94 by the male) and on the characteristics of 71 individual sleeping trees. Each sleeping tree and all trees ≥40 cm diameter at breast height (DBH) in the home range were assessed (height, DBH, canopy structure, liana load) and mapped using a GPS. The gibbons preferentially selected tall (mean=38.5 m), emergent trees without lianas. The majority of the sleeping trees (53.5%) were used only once and consecutive reuse was rare (9.5%). Sleeping trees were closer to the last feeding tree of the evening than to the first feeding tree in the morning, and sleeping trees were located in the overlap areas with neighbors less often than expected based on time spent in these areas. These results suggest avoidance of predators as the main factor influencing sleeping tree selection in pileated gibbons. However, other non-mutually exclusive factors may be involved as well. Am. J. Primatol. 72:617–625, 2010. © 2010 Wiley-Liss, Inc.

Human females, unlike most mammals, are sexually active outside of fertile periods. This decoupling of sexual behavior from its conceptive function has had an enormous impact on human social relationships, and yet we know little about why there was selection for nonconceptive mating. Here we examine one form of nonconceptive mating, the mating that occurs during pregnancy or post-conceptive (PC) mating, in wild western gorillas (Gorilla gorilla). Using a near complete mating record for five females during gestation, we show that pregnant females varied in the timing and frequency of mating, and used PC mating conditionally, synchronizing copulations to occur on days when other females mated, and refraining from mating for lengthy periods when no other females mated. As pregnant females mated exclusively with the same male before and after conception, and mated in response to group female (and not male) behavior, we conclude that western gorillas used PC mating as a form of female competition, and not to confuse paternity or to obtain immediate benefits from the male, as suggested earlier. The male initiated copulations preferentially with females of high rank, rather than distinguishing between pregnant and cycling females. Therefore, PC mating appears to be a strategy by which high-ranking pregnant females attempt to minimize male interest in other females, while reinforcing their own status and potentially delaying conception in others. These findings indicate that female-mating competition is more important than considered earlier, and may be a factor in the evolution of nonconceptive mating in humans.

Within phylogenetic limits reproductive characteristics of a given species may vary between populations in response to ecological and social factors. For instance, in environments where high quality nutrition is readily available, the onset and speed of reproduction are often accelerated. Other influencing factors might be maternal experience or the sex of the infant. Here we present data on reproductive characteristics for the silvered leaf monkey (Trachypithecus cristatus), a medium-sized Asian colobine housed at the Wildlife Conservation Society's Bronx Zoo as a one-male group. To place the species into an appropriate phylogenetic context, we limited our comparison to other colobine species. Demographic data span 21.4 years (October 1985 to March 2007) and derive from 30 adult females (128.0 female years). Detailed behavioral data stem from a 2.2 years study (November 2002 to January 2005; 734 days, 4,225 hr). As in other Asian colobines, receptive periods were short (mean=4.3 days, n=68). This is expected for one-male groups where receptivity likely indicates, rather than conceals, ovulation. Gestation length was estimated based on a change in the pattern of sexual behavior (mean=194.6 days, n=7). It fell within the range reported for the taxon. Births occurred year round, at an early age (mean=2.9 years, n=8), at short intervals (mean=14.9 months, n=59) in combination with a short lactation (mean 12.1 months, n=9) likely due to the nearly unlimited availability of nutrition in this zoo setting. Primiparous females tended to have a longer first interbirth interval but infant survival rates were similar to multipara possibly due to the absence of predators. Maternal investment was independent of the infant's sex and birth sex ratio was even. Our results emphasize that when interpreted with caution, zoo populations yield realistic reproductive characteristics that can help fill the gap in our knowledge about colobine life history. Am. J. Primatol. 71:852–859, 2009. © 2009 Wiley-Liss, Inc.

Female-female mounting was studied for 3233 hr in a bisexual one-male troop of free-ranging Hanuman langurs in northwestern India over a period of 6 years. The population breeds throughout the year and female-female mounts (n = 524) occurred during all months. All 15 adult females mounted and were mounted, although only 20% of the individuals exhibited more than half of all mounter activity and 33% more than half of all mountee activity. Various similarities with heterosexual mounting suggest that sexual arousal is a stimulus of the activity. Cycling, pregnant, and lactating females acted as mounters as well as mountees. However, ovulating individuals were greatly overrepresented concerning their likelihood of mounting and being mounted. Females of any given rank in the dominance hierarchy showed mounter as well as mountee activity, although mounters were higher ranking than mountees in 84% of cases. Young mounters were overrepresented and middle- to old-aged mounters underrepresented, whereas no such differences existed for mountees. Mounting relations among four young adult paternal half-sisters who occupied top positions in the dominance hierarchy were significantly overrepresented. Although the ultimate function of homosexual female mounting is not fully understood, it is suspected to be connected with intrasexual competition: If "pseudocopulations" induce mountees to reduce the number of solicitations addressed to the male, the probability of insemination decreases and mounters thus reduce the number of future competitors.

Data from four Asian colobine populations, wild (Ramnagar) and provisioned (Jodhpur) Hanuman langurs, wild Phayre's leaf monkeys (Phu Khieo Wildlife Sanctuary), and captive silvered langurs (Wildlife Conservation Society's Bronx Zoo) were analyzed with respect to reproductive and behavioral trends of aging. All but one female (from a total of seven) became menopausal 1.8 to 5+ years prior to death with a lag time between last parturition and death ranging from 3.0 to 9.0 years (n=8). Hormonal analysis revealed constantly low and acyclic levels of immunoreactive pregnanediol-3-glucuronide (iPdG) in one old, wild female supporting the notion of true menopause. Nonsexual behavior was analyzed based on quantitative data collected for 6 old and 26 younger, adult females simultaneously (1,378.5 h). No support was found for social disengagement, but old females were more inactive and spent less time feeding. Altogether the data support earlier claims of menopause and extended postreproductive life spans in colobine monkeys. Future behavioral studies should aim at longitudinal assessments of behavioral changes and incorporate energetic data as well as testing of social continuity and selectivity.

Socioecological theory suggests a link between the strength of competition for food/safety, rates of agonism, structure of dominance hierarchies, and dispersal among group-living females. This study presents preliminary data on agonistic behavior and dominance relationships for female Phayre's leaf monkeys (Trachypithecus phayrei), a species in which females routinely disperse. Behavioral observations were conducted on two groups (four adult females, and five adult females plus two juvenile females, respectively) at Phu Khieo Wildlife Sanctuary, northeast Thailand. Rates of agonistic behavior were analyzed from focal continuous recordings, while dominance hierarchies were constructed from all agonistic behaviors (focal and ad libitum sampling). Overall, female-female agonistic behaviors (aggression, submission, and displacements) occurred at a rate of < 0.25 interactions per hour. Agonistic interactions involving food occurred more frequently than expected based on feeding time. Females in both groups exhibited linear dominance hierarchies with some reversals, and possibly an age-inversed hierarchical structure in the larger group. The results fit well with previous results for colobine monkeys regarding frequency of interactions, displacements predominating agonistic behavior, and the possibility of an age-inversed hierarchy. The results contradict the suggested link between linearity of hierarchies and female philopatry. Future studies should consider the notion that female dispersal may coexist with linear dominance hierarchies.