Phayre's Leaf Monkeys Research Articles

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

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. 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. 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. 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.

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.

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

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). 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). 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. 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.

Age and reproductive status influence dominance in wild female Phayre's leaf monkeys

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

Aggressive Behavior of Phayre's Leaf Monkeys Towards Domestic Dogs in Cachar District of Assam, India

Phayre’s leaf-monkeys (Trachipithecus phayrei) or Phayre’s langurs are old world monkeys that inhabit South-East Asian tropical forests. The species is under a severe threat due to large scale habitat destruction and disturbances by people living near the habitat of langurs. The present study recorded the aggressive behavior of male langurs towards domestic dogs in the Cachar district of Assam. Response of each member in the troop was observed. The sophisticated behavior of males in safeguarding the weaker members was observed. In conclusion, the harassment by domestic dogs may result in the expulsion of Phayre’s langurs from their native habitat.

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.

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.

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.

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 20months 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.

Agonistic behavior and dominance relationships in female phayre's leaf monkeys – preliminary results

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.

Temporary Absence and Dispersal in Phayre's Leaf Monkeys (Trachypithecus phayrei)

Temporary Absence and Dispersal in Phayre's Leaf Monkeys (Trachypithecus phayrei)