Female social dynamics as viewed from grooming networks in the Central Himalayan Langur (Semnopithecus schistaceus).

Enhanced survival and reproduction are associated with an individual's direct and indirect social connections with members of a group. Yet, the role of these connections is little known in a vast range of primate species. We studied female Central Himalayan Langur (CHL) to investigate the link between four specific attributes (dominance rank, age, genetic relatedness, and the presence of females carrying infants) and a female's direct and indirect social relationships. By analyzing grooming networks, we revealed different behavioral strategies: high-ranking females form relationships with many females (high degree), whereas females with dependent infants have strong relationships (high strength and eigenvector). Subadult females are important individuals that hold the social network together (high betweenness), while an immigrant female strategy is to integrate herself into the group by forming strong bonds with females who themselves have strong bonds (high eigenvector). Our study sheds light on how behavioral strategies shape female CHL grooming networks, which may help them to secure fitness and survival advantages.

Adaptations to a cold climate promoted social evolution in Asian colobine primates.

The biological mechanisms that underpin primate social evolution remain poorly understood. Asian colobines display a range of social organizations, which makes them good models for investigating social evolution. By integrating ecological, geological, fossil, behavioral, and genomic analyses, we found that colobine primates that inhabit colder environments tend to live in larger, more complex groups. Specifically, glacial periods during the past 6 million years promoted the selection of genes involved in cold-related energy metabolism and neurohormonal regulation. More-efficient dopamine and oxytocin pathways developed in odd-nosed monkeys, which may have favored the prolongation of maternal care and lactation, increasing infant survival in cold environments. These adaptive changes appear to have strengthened interindividual affiliation, increased male-male tolerance, and facilitated the stepwise aggregation from independent one-male groups to large multilevel societies.

Apparent diet digestibility of captive colobines in relation to stomach types with special reference to fibre digestion.

Colobine monkeys are known for the anatomical complexity of their stomachs, making them distinct within the primate order. Amongst foregut fermenters, they appear peculiar because of the occurrence of two different stomach types, having either three ('tripartite') or four ('quadripartite', adding the praesaccus) chambers. The functional differences between tri and quadripartite stomachs largely remain to be explained. In this study, we aim to compare the apparent digestibility (aD) in tripartite and quadripartite colobines. Hence, we measured the aD in two colobine species, Nasalis larvatus (quadripartite) and Trachypithecus cristatus (tripartite), in two zoos. We also included existing colobine literature data on the aD and analysed whether the aD of fibre components is different between the stomach types to test the hypothesis of whether quadripartite colobines show higher aD of fibre components than tripartite colobines did. Our captive N. larvatus specimen had a more distinctively varying nutrient intake across seasons with a larger seasonal variation in aD than that of a pair of T. cristatus, which mostly consumed commercial foods with a lower proportion of browse and less seasonal variation. We observed higher aD of dry matter (DM), neutral detergent fibre (NDF) and acid detergent fibre (ADF) in the N. larvatus specimen, suggesting a higher gut capacity of N. larvatus provided by the additional praesaccus forestomach chamber. Based on the analysis of literature data for aD, we also found that quadripartite species achieved higher fibre digestibility at similar dietary fibre levels compared with tripartite species, supporting the hypothesis that the additional gut capacity offered by the praesaccus facilitates a longer retention and hence more thorough microbial fermentation of plant fibre.

Diet, food availability, and climatic factors drive ranging behavior in white-headed langurs in the limestone forests of Guangxi, southwest China.

Changes in abiotic and biotic factors can affect the efficiency of biological systems in animals, forcing them to adjust their behaviors in response to daily and seasonal variations. From September 2016 to August 2017, we collected ranging behavior data on four groups of white-headed langurs ( Trachypithecus leucocephalus) in the Guangxi Chongzuo White-Headed Langur National Nature Reserve, Guangxi, southwest China. We simultaneously analyzed how multiple ecological factors affect langur ranging behavior, which should facilitate our understanding of the potential mechanisms underlying their adaptation to limestone habitats. Results showed that langur ranging behavior was significantly affected by diet composition, food availability, and climatic factors. Specifically, moving time and daily path length increased with the increase in dietary diversity. Furthermore, moving time and daily path length were positively associated with the availability of fruit and relative humidity of the forest, and moderately associated with temperature and relative humidity of bare rock. Our study demonstrated that langurs maintain stable moving and feeding times and exhibit a short daily travel distance, likely adopting an energy-conserving behavioral strategy in response to food shortages and high temperatures in the fragmented karst forest. These results highlight the importance of food availability and temperature in shaping the ranging behavior of these karst-dwelling primates.

Allometric escape and acoustic signal features facilitate high-frequency communication in an endemic Chinese primate.

The principle of acoustic allometry-the larger the animal, the lower its calls' fundamental frequency-is generally observed across terrestrial mammals. Moreover, according to the Acoustic Adaptation Hypothesis, open habitats favor the propagation of high-frequency calls compared to habitats with complex vegetational structures. We carried out playback experiments in which the calls of the Guizhou snub-nosed monkey (Rhinopithecus brelichi) were used as stimuli in sound attenuation and degradation experiments to test the hypothesis that propagation of Guizhou snub-nosed monkey calls is favored above vs through the forest floor vegetation. We found that low-pitched Guizhou snub-nosed monkey vocalizations suffered less attenuation than its high-pitched calls. Guizhou snub-nosed monkeys were observed emitting high-pitched calls from 1.5 to 5.0 m above the ground. The use of high-pitched calls from these heights coupled with the concomitant behavior of moving about above the understory may provide a signal for receivers which maximizes potential transmission and efficacy. Our results support the Acoustic Adaptation Hypothesis and suggest that by uncoupling its vocal output from its size, this monkey can produce a high-pitched call with a broad spectral bandwidth, thereby increasing both its saliency and the frequency range over which the animal may more effectively communicate in its natural habitat.

Mixed-species associations and attempted mating suggest hybridization between purple-faced and tufted gray langurs of Sri Lanka.

Hybridization is increasingly recognized as an important mechanism of evolutionary change in the order Primates. Here, we present the first observational data supporting natural hybridization between the critically endangered purple-faced langur (Semnopithecus vetulus philbricki) and the threatened tufted gray langur (Semnopithecus priam thersites) in Kaludiyapokuna Forest Reserve in Sri Lanka. In one case study, we observed a long-term (> 1 year) mixed-species group consisting of one adult tufted gray langur male coexisting with seven adult purple-faced females. Although copulations were not observed, two infants were conceived during the male's tenure, and the coat color of one of these infants transitioned into that intermediate between those of the two langur species. The tufted gray langur male was also aggressive toward extra-group males of both species, as well as towards purple-faced juveniles within his group. However, we never witnessed the male exhibiting aggression towards the infants conceived during his tenure. In a second case study, a female purple-faced langur visited and sexually solicited a tufted gray langur male in a known study group of this species over the course of 2 days, in what resembled a sexual consortship. Taken together, the observed mixed-species association and attempted interspecific mating suggest that hybridization is very likely in these sympatric species. Genetic data are needed to confirm and determine the extent of hybridization in the dry zone of Sri Lanka where purple-faced langurs live in sympatry with tufted gray langurs.

Mitogenomic phylogeny of the Asian colobine genus Trachypithecus with special focus on Trachypithecus phayrei (Blyth, 1847) and description of a new species.

Trachypithecus, which currently contains 20 species divided into four groups, is the most speciose and geographically dispersed genus among Asian colobines. Despite several morphological and molecular studies, however, its evolutionary history and phylogeography remain poorly understood. Phayre's langur ( Trachypithecus phayrei) is one of the most widespread members of the genus, but details on its actual distribution and intraspecific taxonomy are limited and controversial. Thus, to elucidate the evolutionary history of Trachypithecus and to clarify the intraspecific taxonomy and distribution of T. phayrei, we sequenced 41 mitochondrial genomes from georeferenced fecal samples and museum specimens, including two holotypes. Phylogenetic analyses revealed a robustly supported phylogeny of Trachypithecus, suggesting that the T. pileatus group branched first, followed by the T. francoisi group, and the T. cristatus and T. obscurus groups most recently. The four species groups diverged from each other 4.5-3.1 million years ago (Ma), while speciation events within these groups occurred much more recently (1.6-0.3 Ma). Within T. phayrei, we found three clades that diverged 1.0-0.9 Ma, indicating the existence of three rather than two taxa. Following the phylogenetic species concept and based on genetic, morphological, and ecological differences, we elevate the T. phayrei subspecies to species level, describe a new species from central Myanmar, and refine the distribution of the three taxa. Overall, our study highlights the importance of museum specimens and provides new insights not only into the evolutionary history of T. phayrei but the entire Trachypithecus genus as well.

Factors influencing riverine utilization patterns in two sympatric macaques.

Many species of terrestrial animals, including primates, live in varied association with the aquatic (e.g., riverine or coastal) environment. However, the benefits that each species receive from the aquatic environment are thought to vary depending on their social and ecological characteristics, and thus, elucidating those benefits to each species is important for understanding the principles of wild animal behaviour. In the present study, to gain a more complete picture of aquatic environment use, including social and ecological factors in primates, factors affecting riverine habitat utilization of two macaque species (Macaca nemestrina and M. fascicularis) were identified and qualitative comparisons were made with sympatric proboscis monkeys (Nasalis larvatus), which have different social and ecological characteristics. Temporal variation in sighting frequency of macaques at the riverbanks was positively related to the fruit availability of a dominant riparian plant species and negatively related to the river water level which affects the extent of predation pressure. Riverine utilization of macaques was greatly influenced by distribution and abundance of food (especially fruit) resources, possibly in association with predation pressure. Additionally, qualitative ecological comparisons with sympatric proboscis monkeys suggest that the drivers of riverine utilization depend on the feeding niches of the species, and different anti-predator strategies resulting from their differing social structures.

Preliminary study on dietary selection in Shortridge's langurs ( Trachypithecus shortridgei) from China.

Understanding dietary selection and feeding strategies is important for the conservation and management of endangered primate species. Here, we conducted a preliminary study on the diet and feeding behavior of endangered Shortridge's langurs ( Trachypithecus shortridgei) within the Drung River Valley (Dulongjiang) in southwestern China. The study site lies at a high latitude (N27°47.5') and elevation (1 900 m a.s.l.) and is characterized by substantial annual rainfall (2 745.1 mm). From August 2012 to September 2013, we observed five groups of langurs and analyzed their overall food composition and dietary variation in spring and autumn. To understand their dietary adaptations to the distinctive habitat of the Drung River Valley, we also compared the diet of Shortridge's langurs to that of other Trachypithecus species inhabiting different environments. Results indicated that T. shortridgei fed on 52 plant species, 23 of which each accounted for ≥1% of their annual feeding time. Their primary dietary components included leaves (46.2%, young, mature, and petioles), fruits (28.7%, unripe 17.6%, ripe 11.1%), and mosses (10.2%). The langurs mainly consumed mature (34.2%) and young leaves (27.5%) in spring and ripe fruits (39.4%) and mature leaves (24.7%) in autumn. Two species of moss ( Macrothamnium macrocarpum and Scapania verrucosa, 21.2% of annual feeding time), which are usually found growing together on cliffs, played a relatively important role in the diet of T. shortridgei. The langurs mainly consumed ripe fruits of Saurauia napaulensis (7.1%) and Dendropanax burmanicus (7.1%), which were abundant at lower elevations. Trachypithecus species in temperate forests consumed more fruits and seeds but fewer leaves (similar mature leaves but fewer young leaves) than those species in tropical forests, which may be related to their availability and abundance. Compared to Trachypithecus species in temperate forests, the higher proportion of mosses and mature leaves but fewer young leaves in the annual diet of T. shortridgei are likely a response to the distinctive Drung River Valley habitat. Therefore, conservation of the main food plants of this threatened species could be vital for its survival and conservation management.

Influence of habitat conditions on group size, social organization, and birth pattern of golden langur (Trachypithecus geei).

We studied endangered golden langurs in fragmented and altered habitats to understand the consequences of habitat conditions on group size, social organization, and birth seasonality. We selected 12 groups inhabiting forest edge and forest core of Chakrashila Wildlife Sanctuary (henceforth Chakrashila WLS) and adjoining the Abhaya rubber plantation. Each group was monitored every month from May 2013 to September 2016 and recorded the age-sex of individuals in the group. The births were recorded with the individual identity of females in five focal groups. The overall group size of golden langur was 11.3 ± 3.5SD, and ranged between 5 and 18. The mean group size in forest core, forest edge, and rubber plantation differed significantly. We recorded a total of 46 births in 12 groups across the three different habitats. The number of infants correlates positively with adult females and group size across all the 12 groups for all the years. The number of births that occurred in all the months varied significantly across the months. Births occurred in all the months but peaked between May and September (82.6%). The mean number of births positively correlated with mean monthly rainfall. Mean inter-birth interval was 24.5 ± 1.6SD months that did not vary between the females. It therefore appears that group size is sensitive to forest type, and births are positively related to social and environmental factors. The behavioral parameters may influence life-history traits if continuous habitat alteration persists.

Odd-nosed monkey scapular morphology converges on that of arm-swinging apes.

Odd-nosed monkeys 'arm-swing' more frequently than other colobines. They are therefore somewhat behaviorally analogous to atelines and apes. Scapular morphology regularly reflects locomotor mode, with both arm-swinging and climbing anthropoids showing similar characteristics, especially a mediolaterally narrow blade and cranially angled spine and glenoid. However, these traits are not expressed uniformly among anthropoids. Therefore, behavioral convergences in the odd-nosed taxa of Nasalis, Pygathrix, and Rhinopithecus with hominoids may not have resulted in similar structural convergences. We therefore used a broad sample of anthropoids to test how closely odd-nosed monkey scapulae resemble those of other arm-swinging primates. We used principal component analyses on size-corrected linear metrics and angles that reflect scapular size and shape in a broad sample of anthropoids. As in previous studies, our first component separated terrestrial and above-branch quadrupeds from clambering and arm-swinging taxa. On this axis, odd-nosed monkeys were closer than other colobines to modern apes and Ateles. All three odd-nosed genera retain glenoid orientations that are more typical of other colobines, but Pygathrix and Rhinopithecus are closer to hominoids than to other Asian colobines in mediolateral blade breadth, spine angle, and glenoid position. This suggests that scapular morphology of Pygathrix may reflect a significant reliance on arm-swinging and that the morphology of Rhinopithecus may reflect more reliance on general climbing. As 'arm-swinging' features are also found in taxa that only rarely arm-swing, we hypothesize that these features are also adaptive for scrambling and bridging in larger bodied anthropoids that use the fine-branch component of their arboreal niches.

Cafeteria-style feeding trials provide new insights into the diet and nutritional strategies of the black snub-nosed monkey (Rhinopithecus strykeri): Implications for conservation.

Anthropogenic changes and fragmentation of natural habitats often exert a negative effect on resource availability and distribution, and the nutritional ecology and feeding behavior of nonhuman primates. The goals of this study are to examine food choice and to identify the nutritional profile of foods consumed by the Critically Endangered black snub-nosed monkey (Rhinopithecus strykeri). To accomplish our study goals, we presented cafeteria-style feeding trials of fresh food items collected in the home range of wild black snub-nosed monkeys to the only two captive R. strykeri, and compared the nutritional profiles of the leafy foods (buds, young, and mature leaves, 100 i23tems from 70 plant species) selected with those avoided (54 items from 48 plant species). Overall, the results indicate that captive R. strykeri selected foods that were higher in moisture (Mo; 77.7%), crude protein (CP; 21.2%), total nonstructural carbohydrates (TNC; 34.9%), and phosphorus (P; 0.37%) while tending to avoid foods with a neutral detergent fiber (NDF) content of greater than 46.8%. Leaves collected in autumn and selected by the monkeys were characterized by a slightly higher amount of metabolizable energy (ME) than those rejected (1,350 kJ/100 g vs. 1,268 kJ/100 g). In contrast, the protein content of foods collected and consumed during the spring was greater (22.9%) than in autumn (16.4%). Random Forests modeling, an ensemble learning method, indicated that the proportion of Mo, NDF, ME, CP, P, and TNC were among the most important factors in predicting which items were consumed by the captive R. strykeri during spring and autumn. On the basis of the nutritional profile of foods consumed across the two seasons, we identified 18 nutrient-rich native plant species that we recommend for use in ex- and in-situ conservation management and reforestation programs to provide long-term access to a nutritionally adequate diet.

Responses to Spatial and Temporal Variation in Food Availability on the Feeding Ecology of Proboscis Monkeys (Nasalis larvatus) in West Kalimantan, Indonesia.

Folivorous primates have long been assumed to experience food competition less acutely than frugivores due to their ability to eat leaves, an abundant resource in most forest systems. Consequently, the behavioural responses of leaf-eating primates to variation in food availability are less well characterised than those of frugivores. Recent empirical studies have demonstrated that many colobine species are more affected by food availability and distribution than previously thought; they employ multiple strategies to survive during periods of food scarcity. We studied a population of proboscis monkeys (Nasalis larvatus) over 16 months in three forest types in West Kalimantan, Indonesia, to examine their responses to temporal fluctuation and spatial variation in food availability. We examined how feeding behaviour was influenced by the availability of plants in botanical plots to identify important and preferred foods of proboscis monkeys across months and in different forest types. Proboscis monkeys consumed foods from 68 genera, comprising 35% young leaves, 27% unripe fruit, 12% flowers and 6% mature leaves. Consumption of plant parts and genera by proboscis monkeys varied in response to monthly changes in food availability but did not vary among forest types despite substantial differences in phenology and floristics among them. The monkeys preferred unripe fruits and flowers and used young and mature leaves as fallback foods in mangrove forests. Documentation of proboscis monkey responses to variation in food availability contributes to our understanding of how monkeys respond to changes in their environments due to climate change and habitat degradation.

Reproductive strategy of bachelors in a snub-nosed monkey (Rhinopithecus bieti) all-male unit.

Snub-nosed monkeys exhibit a rare multilevel social system composed of several one-male units (OMU) and at least one all-male unit (AMU). The AMU comprises males who are blocked from access to females by resident males in the OMUs, and how these satellite males achieve reproductive success is still unclear. To investigate their reproductive strategies, we focused on the AMU in a band of provisioned black-and-white snub-nosed monkeys (Rhinopithecus bieti) in Yunnan, China. Behaviors that AMU males use to gain access to females (i.e. immigration, male takeover, and sexual interaction with females) were recorded and compared with resident OMU males to explore how AMU bachelors achieve reproductive success when they are denied stable access to females. We found that in response to solicitation from females, adult and sub-adult members of the AMU responded more actively than resident males, and the bachelors actively initiated mating with females when the latter's resident male was temporarily absent. These mating opportunities mostly coincided with the peak mating season in OMUs, and probably allowed bachelors to sire some offspring. We also found that for some AMU adults, taking over an OMU is the main strategy used to gain stable access to females, and these males repeatedly migrate between bands. AMU members therefore show multiple strategies that allow them to gain some degree of reproductive success.

Proximate causes of dispersal for female Yunnan snub-nosed monkeys.

Individual dispersal trends, unquestionably important for species ecology and evolution, are affected by multiple factors. Understanding the factors that influence female dispersal strategies offers important insight into primate dispersal mechanisms and female choice. To investigate the proximate causes of dispersal in female Yunnan snub-nosed monkeys (Rhinopithecus bieti), we observed and analyzed nine years of detailed dispersal and demographic data from a population of R. bieti in Xiangguqing, Baimaxueshan Nature Reserve, Yunnan Province, China. Results showed that females who lived long-term in a one-male unit (OMU), without giving birth and with few or no relatives, were more likely to leave that OMU. In addition, an OMU led by an outgroup male and containing more female relatives was significantly more likely to be chosen for immigration. Conversely, greater male age, longer male tenure, and more potentially fertile females discouraged immigration into an OMU. These results suggest that reproduction, male quality, and kin cooperation play the largest roles in female Yunnan snub-nosed monkey dispersal.

Suitable habitat prediction of Sichuan snub-nosed monkeys (Rhinopithecus roxellana) and its implications for conservation in Baihe Nature Reserve, Sichuan, China.

As an endemic primate species with one of the highest priorities in wildlife conservation in China, Sichuan snub-nosed monkeys (Rhinopithecus roxellana) have undergone a sharp decline and range reduction in recent centuries. Here, we used maximum entropy modelling (MaxEnt) integrated with four types of environmental variables, including three biological climate variables (Bio17, precipitation of the driest quarter; Bio6, min. temperature of the coldest month; and Bio2, mean diurnal range), three topographic variables (altitude, slope, and aspect), two anthropogenic variables (Human Footprint Index and human disturbance), and three vegetation-related variables (enhanced vegetation index, normalized difference vegetation index, and Wet Index) to identify the spatial distribution of suitable habitats for Sichuan snub-nosed monkeys in Baihe Nature Reserve (BNR), which is located in the Minshan Mountains. The average training AUC of our model performance is 0.929 ± 0.003. The model predicted 9.6 km2 of high suitability habitats and 14.1 km2 of moderate suitability habitats for Sichuan snub-nosed monkeys, adding up to only 11.7% of the total area of concern for the study in the BNR. The top four variables ranked in the model (altitude, Human Footprint Index, human disturbance, and Bio17) accounted for relative gain contributions of 23.3%, 19.3%, 14.2%, and 13.4%, respectively. The predicted suitable habitats were confined to an altitude range of 1971-3198 m, Human Footprint Index of mainly 3-5 values, low human disturbance (mainly livestock), and precipitation of the driest (or coldest) quarter of 9-22 mm. Additionally, the suitable habitats were mainly distributed in the core zone (36.1%), buffer zone (26.8%), and experimental zone (29.5%). The remaining habitats (7.6%) were distributed in the 0.5-km buffer zone of the reserve border. The predicted suitable habitats indicated limited suitable habitat space for the Sichuan snub-nosed monkeys, with most of the suitable habitat distributed outside the core zone in the BNR. Our findings highlighted that human activities in all three functional zones could be the most negative factor on suitable habitat distribution of Sichuan snub-nosed monkeys in the BNR.

Diet of the Indochinese silvered langur (Trachypithecus germaini) in Kien Luong Karst area, Kien Giang Province.

The Indochinese silvered langur (Trachypithecus germaini) is distributed to the west of Mekong River in Cambodia, Lao PDR, Thailand and Vietnam. During a two-year study, from May 2014 to May 2016, we collected 320.44 hr of behavior, with 17,040 feeding bouts recorded (142 hr) for T. germaini on Chua Hang Karst Mountain, Kien Luong District, Kien Giang Province, Vietnam. Feeding accounted for 45% of the Indochinese silvered langurs' activity budget. The plant diet of the Indochinese silvered langurs was principally composed of young leaves (58%), followed by mature leaves (9.5%), fruits (22.7%), flowers (4.7%), buds (3.3%), petioles (1.2%), and other (0.5%). A total of 58 plant species were fed on by the silvered langurs, and leaves of eight species (Phyllathus reticulatus, Ficus rumphii, Ficus tinctoria, Ficus microcarpa, Cayratia trifolia, Streblus ilicifolia, Combretum latifolium, and Streblus asper) were fed on throughout the year. P. reticulatus was most frequently eaten (13.9% feeding time, n = 1,733). Food selection differed significantly between months and seasons. The Indochinese silvered langurs ate 27 plant species in the wet season compared with 23 plant species in the dry season. Leaf chemical composition of two food categories, 16 eaten species (with 10 most frequently consumed species and six least consumed species), and four noneaten species, were analyzed. Feeding samples from eaten species in the Indochinese silvered langurs's diet contained lower amounts of condensed tannin, lignin, protein, ash, and lipids, but a higher amount of total sugar compared with samples from noneaten species. Furthermore, the most frequently consumed species contained lower amounts of lignin compared with the less frequently consumed species. Using a generalized linear model with five variables, including neutral detergent fiber (NDF), total sugar, lignin, lipid, and calcium (Ca) indicated that NDF positively correlated and lignin content negatively correlated with feeding records in the diet of these langur.

Population abundance and distribution of the endangered golden langur (Trachypithecus geei, Khajuria 1956) in Bhutan.

Reliable population estimates are lacking for many South Asian primate species, including the golden langur (Trachypithecus geei), which is endangered and restricted to Bhutan and northeast India. Although well studied in India, few studies exist on this species in Bhutan. In November 2017, we undertook a nationwide survey of golden langurs in Bhutan using double observers along trail-based transects in 17 blocks within its habitat, and modeled its distribution using MaxEnt. A total of 2439 golden langurs in 222 groups were collectively encountered by 17 teams of double observers, from which, an overall population of 2516 ± SE 363 individuals and 236 ± SE 9 groups were estimated. Group sizes varied from 2 to 35 individuals with a mean of 11 ± SD 0.38 individuals. A total of 468 adult males (19%), 924 adult females (38%), 649 juveniles (27%), and 398 infants (16%) were counted. Adult male-to-female sex ratio was 1:1.97 and adult female-to-infant ratio was 1:0.43. We determined 2848 km2 of suitable area for golden langurs in Bhutan and estimated a density of 0.88 individuals/km2. Our population estimate of golden langurs in Bhutan is much lower than the current IUCN estimate of 4000 individuals for Bhutan, necessitating a reassessment of its current conservation status due to threats from road kills, electrocution, and development activities like road construction, hydropower, and electrical transmission lines. We further recommend our refined double-observer survey method to reliably estimate primate populations in rugged terrain.

Seed dispersal by proboscis monkeys: the case of Nauclea spp.

Frugivorous vertebrates such as primates are important dispersal agents in tropical forests, although the role of folivorous colobines is generally not considered. However, recent studies reported seed dispersal by endo- and epizoochory in colobine primates, including the proboscis monkey (Nasalis larvatus), suggesting that the role colobines play in seed dispersal might have been underestimated. In the Lower Kinabatangan Floodplain in Sabah, Malaysian Borneo, we investigated whether seeds were still able to germinate after being ingested by proboscis monkeys. Faecal samples (n = 201) from proboscis monkeys were collected between 2015 and 2017. Intact seeds belonging to eight plant species were found in 77% of the faecal samples. Nauclea spp. were the most abundant plant species, accounting for 98% of all intact seeds. This study is the first to conduct germination trials on seeds defecated by proboscis monkeys. Higher germination success was recorded in ingested Nauclea spp. seeds than in control seeds, from both ripe and unripe Nauclea orientalis fruits (P < 0.001). Therefore, we suggest that proboscis monkeys play a role in seed dispersal by enhancing the germination success of defecated seeds for at least some plant species. Similar to other colobines, although proboscis monkeys may provide a lower contribution to seed dispersal (low seed diversity over short distances) than other sympatric frugivores, this study emphasises that proboscis monkeys do contribute to the dispersal of intact seeds, such as Nauclea spp., in potentially suitable riverine habitats.