Female-biased birth sex ratio in a female dispersal primate suggests local resource competition.

Group living may entail local resource competition (LRC) which can be reduced if the birth sex ratio (BSR) is biased towards members of the dispersing sex who leave the group and no longer compete locally with kin. In primates, the predicted relationship between dispersal and BSR is generally supported although data for female dispersal species are rare and primarily available from captivity. Here, we present BSR data for Phayre's leaf monkeys (Trachypithecus phayrei crepusculus) at the Phu Khieo Wildlife Sanctuary, Thailand (N = 104). In this population, nearly all natal females dispersed, while natal males stayed or formed new groups nearby. The slower reproductive rate in larger groups suggests that food can be a limiting resource. In accordance with LRC, significantly more females than males were born (BSR 0.404 males/all births) thus reducing future competition with kin. This bias was similar in 2-year-olds (no sex-differential mortality). It became stronger in adults, supporting our impression of particularly fierce competition among males. To better evaluate the importance of BSR, more studies should report sex ratios throughout the life span, and more data for female dispersal primates need to be collected, ideally for multiple groups of different sizes and for several years.

The effect of data provenance on estimates of gestation length in African and Asian colobines.

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.

The function of geophagy in Nepal gray langurs: Sodium acquisition rather than detoxification or prevention of acidosis.

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.

Offspring loss after male change in wild siamangs: the importance of abrupt weaning and male care.

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.

Characteristics of alpha males in Nepal gray langurs.

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.

Transparency, usability, and reproducibility: Guiding principles for improving comparative databases using primates as examples.

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.

Dominance relationships in male Nepal gray langurs (Semnopithecus schistaceus).

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 < 0.091) 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 (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 160:208-219, 2016. © 2016 Wiley Periodicals, Inc.

The meaning of weaning in wild Phayre's leaf monkeys: last nipple contact, survival, and independence.

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.

Intra- and interspecific variation in macaque molar enamel thickness.

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.

Reproductive characteristics of wild Sanje mangabeys (Cercocebus sanjei).

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.

Timing of conceptions in Phayre's leaf monkeys: Energy and phytochemical intake.

OBJECTIVES: Raising offspring imposes energetic costs, especially for female mammals. Consequently, seasons favoring high energy intake and sustained positive energy balance often result in a conception peak. Factors that may weaken this coordinated effect include premature offspring loss and adolescent subfertility. Furthermore, seasonal ingestion of phytochemicals may facilitate conception peaks. We examined these factors and potential benefits of a conception peak (infant survival and interbirth interval) in Phayre's leaf monkeys (Trachypithecus phayrei crepusculus). MATERIALS AND METHODS: Data were collected at Phu Khieo Wildlife Sanctuary, Thailand (78 conceptions). We estimated periods of high energy intake based on fruit and young leaf feeding and via monthly energy intake rates. Phytochemical intake was based on fecal progestin. We examined seasonality (circular statistics and cox proportional hazard models) and compared consequences of timing (infant survival and interbirth intervals, t-test, and Fisher exact test). RESULTS: Conceptions occurred in all months but peaked from May to August. This peak coincided with high fecal progestin rather than presumed positive energy balance. Primipara conceived significantly later than multipara. Neither infant survival nor interbirth intervals were related to the timing of conception. DISCUSSION: Periods of high energy intake may not exist and would not explain the conception peak in this population. However, the presumed high intake of phytochemicals was tightly linked to the conception peak. Timing conceptions to the peak season did not provide benefits, suggesting that the clustering of conceptions may be a mere by-product of phytochemical intake. To confirm this conclusion, seasonal changes in phytochemical intake and hormone levels need to be studied more directly.

Costs and benefits of allomaternal care to mothers and others in wild Phayre's leaf monkeys.

OBJECTIVES: Allomaternal care (AMC) is suggested to be energetically beneficial to mothers and costly to allomothers. However, among primates, AMC is a heterogeneous phenomenon and its implications are less clear especially in female dispersal species. Here, we investigated infant care in a female dispersal species, Phayre's leaf monkeys (Trachypithecus phayrei crepusculus), to evaluate whether mothers were constrained by infant care and benefitted energetically from AMC, whether AMC was energetically costly for allomothers and how maternal experience was associated with AMC. MATERIALS AND METHODS: Data were collected via instantaneous focal animal sampling between 2004 and 2005 for juvenile and adult females (N = 18) from two groups at the Phu Khieo Wildlife Sanctuary, Thailand (440 h). We used generalized linear mixed models to determine how infant care during the first month after birth affected the time mothers and allomothers spent feeding, socializing, resting, and locomoting and how AMC varied. RESULTS: In the first month, infants spent 26% of their time with an allomother. We found no differences in mothers' overall activity before versus after birth, although mothers fed significantly more and rested less when without their infant. Allomothers fed and rested less when with an infant. AMC varied between 0.0% and 20.5%, with immature females being most active. DISCUSSION: Mothers appear to benefit energetically from AMC such that their overall activity after birth remained unchanged. Costs and benefits for allomothers seem to be variable. Some very active immature females may be benefitting from learning-to-mother. The overall low cost of AMC may facilitate a reciprocal social network among unrelated females.

Hominoid dispersal patterns and human evolution.

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.

Juggling priorities: female mating tactics in Phayre's leaf monkeys.

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.

Demography of Simakobu (Simias concolor) and the impact of human disturbance.

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.

Phytochemicals and reproductive function in wild female Phayre's leaf monkeys (Trachypithecus phayrei crepusculus).

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.

Primate life histories and dietary adaptations: a comparison of Asian colobines and macaques.

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.

Seasonality of group size, feeding, and breeding in wild red-shanked douc langurs (Lao PDR).

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.

Reproductive characteristics of wild female Phayre's leaf monkeys.

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.

Selection of sleeping trees in pileated gibbons (Hylobates pileatus).

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 > or =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.