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.

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.

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.

Department of Anthropology, Stony Brook University, SUNY, Stony Brook, NY, United States, 2 Interdepartmental Doctoral Program in Anthropological Science, Stony Brook University, SUNY, Stony Brook, NY, United States, Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, CT, United States, Department of Botany and Plant Pathology, Oregon State University, Corvallis, OR, United States, Department of Anthropology, University of Georgia, Athens, GA, United States, Graduate Program in Ecology and Evolution, Stony Brook University, SUNY, Stony Brook, NY, United States

Male-female agonism varies throughout the primate order with males often dominating females, especially in sexually dimorphic species. While intersexual agonism has been attributed to sexually coercive contexts, it can also occur for other reasons and intersexual dominance may be influenced by the adult sex ratio. If the proportion of males is high, certain males will regularly lose against other males. Loser-effects may then pave the way for some females to dominate these males, an effect that has been described in a few primate species. Here we investigated the frequency, general style, and context of agonism among gray langurs ( Semnopithecus spp.). Data were collected at two study sites, at Jodhpur, India (one group), and at Ramnagar, Nepal (two groups). The adult sex ratio varied between 0.077 and 1.000 males to females. At both sites, data on agonistic interactions (aggression and submission) were collected in continuous focal animal and ad libitum sampling techniques during 1,945 contact hours (including 1,220 focal animal hours, total). Although aggression intensity was low, high directional consistency and the rare occurrence of counteraggression suggested a despotic dominance style, a likely prerequisite for dominance effects based on adult sex ratio. Aggression by females against males was very rare and mainly occurred in the defense of offspring. We found little evidence for partial female dominance regardless of adult sex ratio. In a few cases in which a female had a higher dominance index than a male, she did not dominate this male in dyadic encounters. Agonism by males directed at females occurred mainly in a feeding context while male policing and a sexual context were both rare. The latter was mostly restricted to females after they had harassed a sexual interaction or after they had behaved proceptively toward a male. Our study suggests that across species the effect of adult sex ratio on female dominance might be more variable than previously suggested. The fact that most agonism between males and females occurred over food identifies intersexual feeding competition as a new research avenue with potentially important consequences for existing ideas on the costs and benefits of group life and composition.

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.

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.

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.

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.

Citation for published item: Borries, C. and Sandel, A. and Koenig, A. and Fernandez-Duque, E. and Kamilar, J. and Amoroso, C. and Barton, R. and Bray, J. and Di Fiore, A. and Gilby, I. and Gordon, A. and Mundry, R. and Port, M. and Powell, L. and Pusey, A. and Spriggs, A. and Nunn, C. (2016) 'Transparency, usability, and reproducibility : guiding principles for improving comparative databases using primates as examples.', Evolutionary anthropology., 25 (5). pp. 232-238.

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.