A tough nut to crack? Proficiency in coconut processing of the Nicobar long-tailed macaque Macaca fascicularis umbrosus.

The ability to manipulate objects enables macaques to utilize resources well. Coconut (Cocos nucifera) is one such food that has high energy and nutrient value but requires complex motor skills for extraction. In this study, we examined the proficiency of a single group of Macaca fascicularis umbrosus from Campbell Bay, India, in feeding on different types of coconuts. The group comprised 16 individuals with 3 adult males, 6 adult females, 3 sub-adult males, and 4 infants. We observed individuals feeding on tender, mature, dry, and dry-dehusked coconuts in 90 feeding bouts between August 2013 and December 2015. Their feeding behaviour was divided into four types of behavioural acts: pluck, process, drink, and feed. We prepared an ethogram consisting of 32 behavioural acts associated with feeding on coconuts, and recorded the occurrence of these acts during focal animal sampling. We used NMDS to examine differences among age/sex classes in the occurrence of behavioural acts associated with feeding on coconuts. Although females and males showed no significant separation in these behaviours, adults and young individuals formed distinct groups. However, only 3 out of 32 acts were exclusive to age classes. Using Discrete Time Markov Chain analysis, we found that adults tended to transition back to processing behaviours after drinking and feeding behaviours more often than young individuals. An almost similar repertoire of behaviours among age and sex classes indicates that there is no single model behaviour. Further, this study indicates that among different age/sex classes, adults tend to maximise the nutritional gains during single feeding bouts than younger individuals.

Community Reserves: Their significance for the conservation of mammals in a mosaic of community-managed lands in Meghalaya, Northeast India.

Community Reserves (CRs) have been advocated for increasing the protected area coverage in northeast India where the land is primarily owned and managed by local indigenous institutions. To understand the significance of these reserves for the conservation of mammals, we investigated the diversity and abundance of mammals in five CRs in the Khasi Hills of Meghalaya as well as interviewed 75 local villagers to assess the hunting practices and perceptions of the Indigenous Khasis on mammals. We employed 60 camera traps in the CRs and undertook a recce survey (day-time and night-time) for capturing the diversity in the CRs. We used photo-capture rate and encounter rate as indices of relative abundance in the CRs. We used an exact multinomial test to test differences of opinion among the respondents of the five CRs. We found a relatively low abundance of mammals in the CRs, yet they persist. A total of 28 species were detected through camera trapping and recce survey and an additional 12 species were reported by respondents to also occur in the CRs. Among the respondents, it was believed that the decline in mammal populations was largely driven by habitat loss and degradation (82.67%) while only a few believed it was also driven by hunting (5.33%). Respondents also believed that the presence of the reserves and awareness programs taken under them had also led to a reduction in hunting (20%) in their area. Although, some attributed it to a general decline in wildlife populations and forest cover (21.33%). Thus, despite these CRs being small (<2 km2) and isolated, they still harbour mammal species and are important for retaining remnant forest patches in a landscape that is highly fragmented.

Determinants of habitat occupancy and spatial segregation of primates in the central Western Ghats, India.

Primates are among the globally imperiled fauna requiring urgent conservation interventions to protect their habitat. Information on species distribution and factors influencing it are vital to species management and habitat protection. In this study, we assessed habitat occupancy of the lion-tailed macaque (Macaca silenus), bonnet macaque (M. radiata), and black-footed gray langur (Semnopithecus hypoleucos) that occur in the Kudremukh Wildlife Division, a large protected area network in the central Western Ghats. We examined the influence of habitat variables on the occupancy probability of these primates. We carried out four temporally replicated detection/non-detection surveys to assess detection probability and site occupancy of the primate species. We surveyed 244 sites of 5 Km2, with each site surveyed for 4 days, to assess detection probability and site occupancy. Among the three species, the langur had the highest habitat occupancy estimate (0.66 ± 0.05SE) and the lion-tailed macaque had the lowest estimate (0.28 ± 0.08SE). The habitat occupancy estimate for the bonnet macaque was (0.56 ± 0.05SE). Wet and semi-evergreen forest cover and mean elevation positively influenced the lion-tailed macaque's occurrence. Covariates influencing bonnet macaque's occurrence were plantations, semi-evergreen and moist deciduous forests, and non-forest areas. Mean elevation negatively affected its occurrence. Wet evergreen forests and plantations positively influenced the occurrence of the langur. We examined spatial segregation between the species based on their site occupancy. Pairwise comparisons revealed a significant negative association between the bonnet macaque and the other two primates. However, we found a significant positive association between the lion-tailed macaque and the langur. We discuss these results and their implications for conservation of primates in the region. Given the cost-effectiveness of carrying out surveys at large spatial scales, we recommend occupancy surveys for future surveys of forest primates.

Responses of birds and mammals to long-established wind farms in India.

Wind turbines have been recognised as an alternative and clean-energy source with a low environmental impact. The selection of sites for wind-farm often creates serious conservation concerns on biodiversity. Wind turbines have become a serious threat to migratory birds as they collide with the turbine blades in some regions across the globe, while the impact on terrestrial mammals is relatively less explored. In this context, we assessed the responses of birds and mammals to the wind turbines in central Karnataka, India from January 2016 to May 2018 using carcass searches to quantify animal collisions (i.e., birds and bats), fixed radius point count for bird population parameters, and an occupancy framework for assessing the factor that determines the spatial occurrence of terrestrial mammals. The mean annual animal fatality rate per wind turbine was 0.26/year. Species richness, abundance, and unique species of birds were relatively higher in control sites over wind turbine sites. Species and functional compositions of birds in control sites were different from wind turbine sites, explaining the varied patterns of bird assemblages of different feeding guilds. Blackbuck, Chinkara, Golden Jackal, and Jungle Cat were less likely to occupy sites with a high number of wind turbines. The study indicates that certain bird and mammal species avoided wind turbine-dominated sites, affecting their distribution pattern. This is of concern to the management of the forested areas with wind turbines. We raised conservation issues and mitigating measures to overcome the negative effects of wind turbines on animals.

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.

Does rank rule? Rank-related grooming patterns in Nicobar long-tailed macaques Macaca fascicularis umbrosus.

Primates maintain social bonds with specific individuals in the group by directing grooming toward them. Social grooming is often targeted toward individuals with whom the most benefits can be exchanged, which are usually the high-ranking individuals. We used the Seyfarth model to investigate whether dominance rank alters the distribution of grooming in a group of Macaca fascicularis umbrosus at Great Nicobar Island. We investigated whether dominance rank predicted grooming rate in both males and females. We used social network analysis to examine whether high-ranking animals maintain a central position in the grooming network. We also investigated whether adult individuals exchange grooming for social tolerance, as indicated by physical proximity. We found that more social grooming was directed toward high-ranking females, but not high-ranking males. Social network analysis suggested that high-ranking animals are not central individuals in the distribution of grooming; rather, middle-ranking animals were major contributors to the distribution of grooming. There was no relation between the grooming rate and proximity in both males and females. Overall, our findings suggest that females prefer to direct grooming toward high-ranking partners, although the dominance hierarchy is less steep than expected for Macaca fascicularis. Thus rank-related grooming patterns may be a relatively conserved trait in this isolated island population.

Do males bond? A study of male-male relationships in Nicobar long-tailed macaques Macaca fascicularis umbrosus.

In primates, males compete for a mate, which is a non-sharable resource. This makes the conditions less conducive for males to have stable relationships. One such special kind of relationship is a bond where the interactions are reciprocated, equitable and differentiated. Bonds in macaque societies are based on the degree of within-group contest competition for mates which is dependent on the synchronization of female fertile phase and reliability of fertility signals. Species of the Fascicularis group, including Nicobar subspecies, show intermediate reliability in the signals with mild peaks, and studies have shown reciprocity but no differentiation. We conducted a study on a group of wild Nicobar long-tailed macaques Macaca fascicularis umbrosus to understand the existing patterns of male-male relationships. We examined whether there is reciprocity in affiliation among the individuals and whether the rate of affiliation is balanced. We also measured the dominance linearity and steepness in the group to understand the monopolizability of females. We used social network analysis to understand whether the relations are differentiated based on hierarchical position and whether the high-ranking individuals are the most central individuals in the distribution of grooming in the group. We found that there is reciprocity among the males although that is not equitable. There was no rankrelated differentiation of affiliation among the males of the group. Instead, the identities of individuals influenced affiliation patterns. Our results correspond to the existent strong relationships but lack of social bond otherwise found in the Fascicularis group of macaques.

Multiple infestations of gastrointestinal parasites - Probable cause for high mortality of Spot-billed Pelican (Pelecanus philippensis) at Kokrebellur Community Reserve, India.

We witnessed mortalities of Spot-billed Pelicans Pelecanus philippensis between December 2017 and May 2018 in Mandya and Mysuru districts of Karnataka, especially at Kokrebellur Community Reserve in Mandya district. The region has experienced severe drought in recent years with negligible water in all the water tanks. A total of 67 Spot-billed Pelicans died in five locations, of which 55 adult birds died at Kokrebellur. We collected four dead pelicans along with 97 fecal samples of live birds at Kokrebellur, water samples from nine water tanks around Kokrebellur, and six fish samples. We isolated the endoparasite eggs by following sedimentation and flotation technique, and counted the eggs from the water and fecal samples, and identified at the genus level using light microscope. We approximately counted the endoparasites by dissecting the fish and conducting a necropsy on dead pelicans. Endoparasite eggs were detected in seven of the nine water tanks. Each fish sample had at least 50-100 L3 stage worms of Contracaecum sp., and 880.0 ±â€¯459.3SD of Contracaecum sp., worms in the digestive tracts and 60.0 ±â€¯36.5SD worms of Echinostoma sp. in the intestine of the four dead pelicans. The endoparasite prevalence was 84.5% (N = 83) with a mean abundance of 368.2 ±â€¯561.5SD eggs/g in the fecal samples of live pelicans. Contracaecum sp., Echinostoma sp. and Opisthorchis viverrini were recorded in 51, 67 and nine fecal samples respectively. The high load of endoparasite eggs in the water tanks, an infestation of Contracaecum sp. in fishes and a heavy load of fully-grown worms of Contracaecum sp. and Echinostoma sp. in the adult pelicans are indicative of their high mortality in Kokrebellur Community Reserve. The coordinated program was initiated with the support of all stakeholders to control the endoparasites in water, fish, and pelicans.

Deceased-infant carrying in nonhuman anthropoids: Insights from systematic analysis and case studies of bonnet macaques (Macaca radiata) and lion-tailed macaques (Macaca silenus).

Existing models of attachment do not explain how death of offspring affects maternal behavior. Previous descriptions of maternal responsiveness to dead offspring in nonhuman anthropoids have not expounded the wide variation of deceased-infant carrying (DIC) behavior. Through the current study, we attempt to (a) identify determinants of DIC through a systematic survey across anthropoids, (b) quantitatively assess behavioral changes of mother during DIC, and (c) infer death perception of conspecifics. Firstly, we performed phylogenetic regression using duration of DIC as the dependent variable. Secondly, we undertook case studies of DIC in the bonnet monkey and the lion-tailed monkey through behavioral sampling. Results of phylogenetic Generalized Linear Mixed Model (Nspecies = 18; Ncases = 48) revealed a strong homology (H2 = 0.86). We also obtained a high intraspecific variation in DIC and found DIC to be affected by mother's age, context of death, habitat condition, and degree of arboreality. We found bonnet mothers to carry their deceased offspring for 3.56 ± 4.03 SD days (N = 7) with diminished feeding, enhanced passivity, and social isolation during DIC and progressive decline in protection/attentiveness of corpse and attachment. Following Anderson (2016)'s framework of death perception, we interpreted repeated sensory investigation of corpses by mothers as comprehending causality of death, inanimate handling of corpse and its defense as comprehension of non-functionality, and a progressive disinterest of mothers in them as perceiving irreversibility of death. Lastly, we integrated DIC with mother-infant attachment theories and proposed a conceptual model characterizing DIC with causal determinants. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Prevalence of gastrointestinal parasites in bonnet macaque and possible consequences of their unmanaged relocations.

Relocation is one of the mitigating measures taken by either local people or related officers to reduce the human-bonnet macaque Macaca radiata conflict in India. The review on relocations of primates in India indicates that monkeys are unscreened for diseases or gastrointestinal parasites (henceforth endoparasites) before relocation. We collected 161 spatial samples from 20 groups of bonnet macaque across their distribution range in south India and 205 temporal samples from a group in Chiksuli in the central Western Ghats. The isolation of endoparasite eggs/cysts from the fecal samples was by the centrifugation flotation and sedimentation method. All the sampled groups, except one, had an infection of at least one endoparasite taxa, and a total of 21 endoparasite taxon were recorded. The number of helminth taxon (16) were more than protozoan (5), further, among helminths, nematodes (11) were more common than cestodes (5). Although the prevalence of Ascaris sp. (26.0%), Strongyloides sp. (13.0%), and Coccidia sp. (13.0%) were greater, the load of Entamoeba coli, Giardia sp., Dipylidium caninum and Diphyllobothrium sp. were very high. Distant groups had more similarity in composition of endoparasites taxon than closely located groups. Among all the variables, the degree of provisioning was the topmost determinant factor for the endoparasite taxon richness and their load. Temporal sampling indicates that the endoparasite infection remains continuous throughout the year. Monthly rainfall and average maximum temperature in the month did not influence the endoparasite richness. A total of 17 taxon of helminths and four-taxon of protozoan were recorded. The prevalence of Oesophagostomum sp., and Strongyloides sp., and mean egg load of Spirurids and Trichuris sp. was higher than other endoparasite taxon. The overall endoparasite load and helminth load was higher in immatures than adults, where, adult females had the highest protozoan load in the monsoon. The findings indicate that relocation of commensal bonnet macaque to wild habitat can possible to lead transmission of novel endoparasites that can affect their population. Thus, we suggest avoidance of such relocations, however, if inevitable the captured animals need to be screened and treated for diseases and endoparasites before relocations.

Extractive foraging and tool-aided behaviors in the wild Nicobar long-tailed macaque (Macaca fascicularis umbrosus).

Macaques possess a repertoire of extractive foraging techniques that range from complex manipulation to tool-aided behaviors, to access food items that increase their foraging efficiency substantially. However, the complexity and composition of such techniques vary considerably between species and even between populations. In the present study, we report seven such complex manipulative behaviors that include six extractive foraging behaviors, and teeth flossing, in a population of Nicobar long-tailed macaques. The apparent purpose of these behaviors was an extraction of encased food, processing food, foraging hidden invertebrates, and dental flossing. Among these behaviors, three behaviors viz. wrapping, wiping, and teeth-flossing were tool-aided behaviors, where macaques used both natural and synthetic materials as tools. Occasionally macaques also modified those tools prior to their use. The substrate use patterns of leaf rubbing and teeth flossing were similar to that observed in other macaques. The spontaneous tool modification to perform wrapping was a first time observation. These observations suggest that Nicobar long-tailed macaques have a high level of sensorimotor intelligence which helps to evolve such innovative foraging solutions.

Losing its ground: A case study of fast declining populations of a 'least-concern' species, the bonnet macaque (Macaca radiata).

The populations of many species that are widespread and commensal with humans have been drastically declining during the past few decades, but little attention has been paid to their conservation. Here, we report the status of the bonnet macaque, a species that is considered 'least-concern' for conservation. We show that the widely ranging rhesus macaque is expanding its range into the distributional range of the bonnet macaque, a species endemic only to southern India. Bonnet macaques have very low abundance in forests of all types indicating that it is not a typically forest dwelling species. The traditionally preferred habitats of bonnet macaques have been Hindu temples/ tourist spots but our data reveal that nearly 50% population of bonnet macaques has disappeared from such previously occupied spots. Another preferred habitat of bonnet macaques has been roadsides with abundant Ficus trees adjoining croplands. We found that between 2003 and 2015, the roadsides have drastically changed where vegetation has been replaced with barren lands and urbanization. Consequently, the populations of bonnet macaques have declined by more than 65% over the past 25 years, and by more than 50% between 2003 and 2015 alone. We, therefore, conclude that this 'least-concern' species is actually facing serious conservation challenges. We also identify a few places such as small hillocks with natural vegetation and a few temples/tourist spots which are likely to remain stable and thus can serve as 'bonnet macaque conservation reserves'. Since the bonnet macaque shares many traits with several other commensal and 'low-risk' species, it can serve as a model for the development of long-term conservation strategies for most such species.

Distribution of the Grey Slender Loris (Loris lyddekerianus Cabrera, 1908) in Tamil Nadu, Southern India.

The grey slender loris Loris lydekkerianus, one of only two nocturnal primates of India, is found in the southern part of the country. Our understanding of its geographical distribution is largely based on historical records and short surveys, and little is known of its occurrence in southern India today. We sought to establish the relative abundance of this species in 26 districts in the state of Tamil Nadu and the union territory of Pondicherry in southern India. We sighted lorises in 19 districts, and their relative abundance ranged from 0.01 to 2.21/km. The south-central districts of Tamil Nadu showed the highest densities of lorises, while the western districts showed the lowest. Based on these results, we recommend increased protection measures for the forest patches of the Eastern Ghats mountains in order to ensure the long-term survival of the grey slender loris.

Demography and reproductive output in langurs of the Western Ghats, India.

Life history traits evolve such that the reproductive output of an organism is maximized. Demographic characteristics, a consequence of life history traits, indicate the reproductive output per individual in group-living species. Both phylogenetic and ecological factors influence demographic traits. In the forests of the Western Ghats, India, we studied the demography of three langur species: Semnopithecus johnii, a wet forest-dwelling species; Semnopithecus hypoleucos, largely a wet forest-dwelling species; and Semnopithecus priam, a primarily dry forest-dwelling species. S. hypoleucos and S. priam are genetically closer to each other than to S. johnii. We sampled a total of 193 groups of the three species of langurs. The group size was smaller in the two wet forest-dwelling species, S. johnii (median = 10) and S. hypoleucos (nine), than in the dry forest-dwelling species, S. priam (18). The number of adult females per group was higher in S. priam (seven) and S. johnii (six) than in S. hypoleucos (four). On the other hand, the adult female:immature ratio, indicating reproductive output and life history, was highest in S. johnii (1:0.33) followed by S. hypoleucos (1:1) and S. priam (1:1.09). Our results suggest that reproductive output is lowest in the arboreal wet forest species and increases as the species become somewhat dry deciduous forest dwellers, or almost facultative dry forest dwellers, and relatively more terrestrial. Some traits, such as group size, appear to be more sensitive to ecological factors, and some other traits such as age-sex ratios and reproductive output appear to be more conservative.

Population Recovery of Nicobar Long-Tailed Macaque Macaca fascicularis umbrosus following a Tsunami in the Nicobar Islands, India.

Natural disasters pose a threat to isolated populations of species with restricted distributions, especially those inhabiting islands. The Nicobar long tailed macaque.Macaca fascicularis umbrosus, is one such species found in the three southernmost islands (viz. Great Nicobar, Little Nicobar and Katchal) of the Andaman and Nicobar archipelago, India. These islands were hit by a massive tsunami (Indian Ocean tsunami, 26 December 2004) after a 9.2 magnitude earthquake. Earlier studies [Umapathy et al. 2003; Sivakumar, 2004] reported a sharp decline in the population of M. f. umbrosus after thetsunami. We studied the distribution and population status of M. f. umbrosus on thethree Nicobar Islands and compared our results with those of the previous studies. We carried out trail surveys on existing paths and trails on three islands to get encounter rate as measure of abundance. We also checked the degree of inundation due to tsunami by using Normalized Difference Water Index (NDWI) on landsat imageries of the study area before and after tsunami. Theencounter rate of groups per kilometre of M. f. umbrosus in Great Nicobar, Little Nicobar and Katchal was 0.30, 0.35 and 0.48 respectively with the mean group size of 39 in Great Nicobar and 43 in Katchal following the tsunami. This was higher than that reported in the two earlier studies conducted before and after the tsunami. Post tsunami, there was a significant change in the proportion of adult males, adult females and immatures, but mean group size did not differ as compared to pre tsunami. The results show that population has recovered from a drastic decline caused by tsunami, but it cannot be ascertained whether it has reached stability because of the altered group structure. This study demonstrates the effect of natural disasters on island occurring species.

Pre-Historic and Recent Vicariance Events Shape Genetic Structure and Diversity in Endangered Lion-Tailed Macaque in the Western Ghats: Implications for Conservation.

Genetic isolation of populations is a potent force that helps shape the course of evolution. However, small populations in isolation, especially in fragmented landscapes, are known to lose genetic variability, suffer from inbreeding depression and become genetically differentiated among themselves. In this study, we assessed the genetic diversity of lion-tailed macaques (Macaca silenus) inhabiting the fragmented landscape of Anamalai hills and examined the genetic structure of the species across its distributional range in the Western Ghats. We sequenced around 900 bases of DNA covering two mitochondrial regions-hypervariable region-I and partial mitochondrial cytochrome b-from individuals sampled both from wild and captivity, constructed and dated phylogenetic trees. We found that the lion-tailed macaque troops in the isolated forest patches in Anamalai hills have depleted mitochondrial DNA diversity compared to troops in larger and continuous forests. Our results also revealed an ancient divergence in the lion-tailed macaque into two distinct populations across the Palghat gap, dating to 2.11 million years ago. In light of our findings, we make a few suggestions on the management of wild and captive populations.

Of how much concern are the 'least concern' species? Distribution and conservation status of bonnet macaques, rhesus macaques and Hanuman langurs in Karnataka, India.

We assessed the distribution and conservation status of bonnet macaques (Macaca radiata), rhesus macaques (Macaca mulatta) and Hanuman langurs (Semnopithecus entellus) in the state of Karnataka, India. Karnataka is situated in southwest India with an area of 191,791 km(2). A total of 9697 km of vehicular survey was made from November 2001 to July 2004. We also visited 107 temples/tourist spots to determine the presence of primates. Bonnet macaques and Hanuman langurs were widely distributed, whereas rhesus macaques were not found in the state. However, bonnet macaques were absent in a few districts in the northern plains and Hanuman langurs were absent in some districts of the southern plains. A total of 205 groups of bonnet macaques and 139 groups of Hanuman langurs were sighted. The relative encounter rate of both species differed across biogeographic zones. Bonnet macaques were largely encountered in the Western Ghats and the Southern Plateau whereas Hanuman langurs were abundant in the Western Ghats and Northern Plains. We found that bonnet macaques have been eliminated from about 48% temples/tourist spots where they occurred in the recent past. The Hanuman langur population of Dharwar-Haliyal Road was assessed during April 2003, and we found that the present population size was about 38% of a previous survey in 1961. Habitat change, hunting/trapping and translocation were the major factors causing a decline in the langur population.