Affiliative relationships and reciprocity among adult male bonnet macaques (Macaca radiata) at Arunachala Hill, India

In captivity, male bonnet macaques (Macaca radiata) frequently express "friendship" toward one another, including affiliative behavior such as huddling, grooming, coalitionary support, and sitting in close proximity. The purpose of this study was to determine whether wild adult male bonnet macaques also express "friendship" by investigating whether or not (1) adult male bonnet macaques have affiliative social relationships with other males, (2) the strength of social relationships varies among dyads, (3) there is time-matched reciprocity in allogrooming among dyads, and if so, whether the level of reciprocity occurs within a bout of grooming, a day, or over 2 months (the limit of this study), and (4) a correlation exists between the strength of social relationships and dominance ranks among adult males. Focal samples totaling 150 hr on all seven adult males in one study group were conducted to record both agonistic and affiliative interactions. Agonistic interactions were used to construct a dominance hierarchy, whereas affiliative interactions (sitting in proximity to within 1 m with and without grooming) were used to quantify the existence and strength of social bonds within dyads. Results show that adult male bonnet macaques had differentiated affiliative relationships with other males in their group. There was little reciprocity of grooming within a bout of grooming or within a day, but greater reciprocity over the study period of 2 months. There was no correlation between dominance ranking distance and the strength of affiliative relationship within dyads; however, within dyads lower-ranking males groomed higher-ranking males more than vice versa. This study suggests that friendships in male bonnet macaques are characterized not by immediate tit-for-tat reciprocal altruism, but by reciprocity over a longer time span, and that affiliative social relationships may be less constrained by agonistic relationships than is the case in more despotic species of macaques.     

Male–Male Relationships in Lion-tailed Macaques (<Emphasis Type="Italic">Macaca silenus</Emphasis>) and Bonnet Macaques (<Emphasis Type="Italic">Macaca radiata</Emphasis>)

Socioecology suggests that female distribution in space is determined by the distribution of food resources and the male distribution is influenced by female distribution. Though studies have traditionally focused on females, males have received increasing attention in recent years. We compared male–male relationships in lion-tailed macaques and bonnet macaques. Because bonnet macaques have a high adult male:female sex ratio and are seasonal breeders whereas lion-tailed macaques have a low adult male:female sex ratio and are largely aseasonal breeders, we predicted that bonnet macaque males would be spatially and socially more tolerant of each other and would have less linear dominance relationships than lion-tailed macaques. We recorded male–male and male–female relationships in 1 group of wild macaques of each species via scan sampling and 1–0 sampling. The results revealed that lion-tailed macaque males largely remained at a distance from each other whereas bonnet macaque males remained in close proximity to one another. Lion-tailed macaque males were more agonistic toward each other whereas bonnet macaque males showed more affiliative interactions. The dominance hierarchy among lion-tailed macaque males was more linear than among bonnet macaque males. Our data support the hypothesis that the study of spatial structuring, temporality of interactions, and linearity of social relationships may contribute to a better understanding of macaque social systems.     

Outcast males and social structure among bonnet macaques (Macaca radiata)

Outcast males are a regular feature of rhesus (Macaca mulatta) and Japanese (M. fuscata) macaque societies but are rare among bonnet macaques (M. radiata). Apparently bonnets have other solutions than ostracism for their social problems. For example, in many troops the male-female ratio ranges from 0.7?1.0+; and, as the young adult males grow to full social and physical maturity, they remain in the troop and participate actively in social relationships. A few troops of bonnet macaques, however, have a network of social relationships which prevents the young males from being integrated into the social system; in some of these cases, male isolation occurs. A comparison of the social relations between males in bonnet macaque societies of both types suggests that societies with strong male-male bonds accommodate their growing males better than those with highly individual males.     

Allogrooming in two species of macaque (Macaca nemestrina andMacaca radiata)

This paper analyzes allogrooming (social grooming) data collected from two large, fully integrated and long established social groups of macaques (one of pigtail macaques and one of bonnet macaques). The data demonstrates a species and sex difference for total allogrooming given with females of both species giving more grooming than the males and with pigtails as a species giving more grooming than bonnets. Also, pigtail females gave more allogrooming to clan members than to nonclan members, but this was not true for pigtail males, bonnet males, and bonnet females. Allogrooming given and allogrooming received by age class for both species showed development of a sexual dichotomy at three to four years. The analysis characterized some of the social structures extant in two closely related species of macaque, particularly the somewhat different use of allogrooming in pigtail females as opposed to the other three categories of animals. Social implications with relation to macaque societies are discussed.     

Of Least Concern? Range Extension by Rhesus Macaques (<Emphasis Type="BoldItalic">Macaca mulatta</Emphasis>) Threatens Long-Term Survival of Bonnet Macaques (<Emphasis Type="BoldItalic">M. radiata</Emphasis>) in Peninsular India

Rhesus and bonnet macaques are the 2 most common and widely distributed of the 8 macaque species of India. Rhesus macaques are widely distributed across southern and southeastern Asia, whereas bonnet macaques are restricted to peninsular India. We studied the current distributional limits of the 2 species, examined patterns of their coexistence in the interspecific border zones, and evaluated losses in the distributional range of bonnet macaques over the last 3 decades. Our results indicate that whereas rhesus macaques have extended their geographical range into the southern peninsula, bonnet macaques have been displaced from many areas within their former distributional range. The southern and the northern distributional limits for rhesus and bonnet macaques, respectively, currently run parallel to each other in the western part of the country, are separated by a large gap in central India, and converge on the eastern coast of the peninsula to form a distribution overlap zone. This overlap region is characterized by the presence of mixed-species troops, with pure troops of both species sometimes occurring even in close proximity to one another. The range extension of rhesus macaque—a natural process in some areas and a direct consequence of introduction by humans in other regions—poses grave implications for the endemic and declining populations of bonnet macaques in southern India.     

Vomiting in Wild Bonnet Macaques

Researchers have not studied vomiting that is not associated with pathological behavior in nonhuman primates. Scientists generally assume that vomiting provides protection by ejecting toxins from the body, but the explanation is not fully supported by the literature. We report on 163 instances of vomiting from 2 groups of free-ranging bonnet macaques in southern India. The macaques routinely reingested their vomit. The reingestion was unlike that in abnormal behavior of both humans and nonhuman primates. We suggest that the tendency to hoard food in their cheek pouches explains why they reingested the vomit.     

Locomotion and Posture During Terminal Branch Feeding

We investigated locomotor and postural behavior during terminal branch feeding in order to gain a better understanding of the motor capabilities of primates. We videotaped wild, juvenile bonnet macaques (Macaca radiata) in India as they fed on flower nectar in a simal tree (Bombax malabaricum). Kinematic analysis revealed that they select specific support surfaces and movements that, for their body design and postures, maximize lateral stability and minimize the chances of falling. These choices are made even though the distance and duration of travel to a selected target are frequently increased. Our discussion focuses on particular concepts of how primates contend with balance problems arboreally, potential reasons for changes in footfall patterns, and how the tail contributes to arboreal locomotion and posture. We concluded that balance problems due to the ratio of body size to branch size are usually avoided, at least among juvenile bonnet macaques, by placing the hands and feet on branches extending laterally from the central support branch and not on the central branch itself. The lateral branches permit a wide base of support, which increases lateral stability. Second, juvenile bonnet macaques have a striking ability to rapidly and repeatedly adapt their gait patterns to changing substrate design with minimal interruption to overall progression. Third, primate tails that are not morphologically specialized for prehension nevertheless have important prehensile and sensory functions in arboreal locomotion and posture.     

Emergence of complex social networks from spatial structure and rules of thumb: a modelling approach

Individual-based computer models show that simple heuristic governing individuals’ behavior may suffice to generate complex patterns of social behavior at the group level such as those observed in animal societies. ‘GrooFiWorld’ is an example of such kind of computer models. In this model, self-organization and simple behavioral rules generate complex patterns of social behavior like those described in tolerant and intolerant societies of macaques. Social complexity results from the socio-spatial structure of the group, the nature of which is, in turn, a side-effect of intensity of aggression. The model suggests that a similar mechanism may give rise to complex social structures in macaques. It is, however, unknown if the spatial structure of the model and that of macaques are indeed similar. Here we used social networks analysis as a proxy for spatial structure of the group. Our findings show that the social networks of the model share similar qualitative features with those of macaques. As group size increases, the density and the average individual eigenvector centrality decrease and the modularity and centralization of the network increase. In social networks emerging from simulations resembling intolerant societies the density is lower, the modularity and centralization are higher, and the individuals ranking higher in the dominance hierarchy are more central than in the social networks emerging from simulations resembling egalitarian societies. Given the qualitative similarity between the social networks of the model and that of empirical data, our results suggest that the spatial structure of macaques is similar to that of the model. It seems thus plausible that, as in the model, the spatial structure combined with simple behavioral rules plays a role in the emergence of complex social networks and complex social behavior in macaques.     

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 km². 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.     

A case of simple tool use in wild liontailed macaques (Macaca silenus)

Three members of a group of liontailed macaques (Macaca silenus) were seen to use leaves for food preparation. Other examples of prey-selection and hunting behaviour in liontailed macaques reflect individual- and group-specific skills. The absence of similar patterns in bonnet macaques (Macaca radiata) living in the same habitat might be related to differences in the social design and indicate the high significance of social aspects for the occurrence and manifestation of innovative behaviour.     

Social interactions of juvenile female bonnet monkeys,Macaca radiata

This study reports observations on juvenile female bonnet macaques (Macaca radiata) in a field cage environment. Interaction and proximity data collected on different age classes of juvenile females indicate distinct changes in social behavior from the time a female is weaned until her first pregnancy. Behavioral development includes an increase in solitary behavior, an early disinterest in younger animals, and a surprisingly low interaction rate with relatives. Important aspects of ontogeny are lost in analyses that treat all juvenile females as a single age-class. This research was supported partially by US PHS Grant No. RR00169.     

Sexual behavior during successive ejaculations in bonnet and pigtail macaques

During tests of sexual behavior which permitted multiple ejaculations, male bonnet and pigtail macaques were found to differ on a number of traditionally defined behavioral parameters. Although pigtail tests were terminated by a more stringent criterion, they were longer in duration and more ejaculations were recorded than in the bonnet tests. Pigtail males required relatively more intromissions for initial ejaculations and somewhat fewer on successive ejaculations. Bonnets, on the other hand, generally ejaculated on the first mount of a test and then exhibited increased numbers of intromissions to successive ejaculations. Although both species exhibited increases in successive postejaculatory intervals, the changes were more gradual for the pigtails.     

Dispersal,nepotism, and primate social behavior

High degrees of relatedness within primate social groups are thought to promote the evolution of altruistic behavior via kin selection. Dispersal, for whatever reason, should limit opportunities for nepotistic behaviors. Conversely, emigration is usually attributed to the avoidance of inbreeding depression. Actual dispersal patterns may result from a balance of these forces. Systematic behavioral differences are expected between taxa that differ in such patterns. In fact, comparisons of (a) colobines vs. cercopithecines, (b) bonnet, stumptailed, and Barbary macaques vs. Japanese and rhesus macaques, and (c) red vs. mantled howler monkeys yield a perplexing blend of unexplained differences and unmet theoretical expectations. Kin selection may be less important than generally believed, and/or methodological standardization more so.     

Species differences in the interest shown in infants by juvenile female macaques (Macaca radiata andM. mulatta)

Using a focal animal technique, 16 juvenile female rhesus and bonnet macaques (2–3 years old) were observed with respect to the frequency and duration of their contacts with infants (0–1 year old). Each of these subjects was a member of one of four seminatural groups of macaques housed in 0.5-acre field cages at the California Primate Research Center. Rhesus juvenile females became very interested in infants when the latter were quite young, and maintained that interest throughout the study. When the infants were approximately 3–4 months old, the rhesus juveniles spent, on the average, 3.9 min/hr with those infants. By the time the infants were yearlings, the time spent with infants was approximately 6 min/hr. Relative to the rhesus, the bonnet juveniles showed less interest (about 2.2 min/hr) in young (3- to 4- month-old) infants, although by the time the infants were yearlings, the bonnets spent about 5.6 min/hr with them. Rhesus juveniles preferred sibling over nonsibling infants, but the presence of a sibling was not a necessary requirement for infant-directed behavior. The two bonnet juveniles who had infant siblings showed no preference for the latter. These data were interpreted within the framework of Quiatt’s [(1979) Am. Anthropol. 81: 310–319 conclusions about allomaternal behavior. Specifically, it was argued that, to the extent that there are species differences in maternal behavior, there should also be species differences in allomaternal behavior. The species differences in the maternal styles of rhesus and bonnet macaques provide a basis for an examination of this hypothesis.     

Flexibility in Food Extraction Techniques in Urban Free-Ranging Bonnet Macaques,Macaca radiata

Non-human primate populations, other than responding appropriately to naturally occurring challenges, also need to cope with anthropogenic factors such as environmental pollution, resource depletion, and habitat destruction. Populations and individuals are likely to show considerable variations in food extraction abilities, with some populations and individuals more efficient than others at exploiting a set of resources. In this study, we examined among urban free-ranging bonnet macaques, Macaca radiata (a) local differences in food extraction abilities, (b) between-individual variation and within-individual consistency in problem-solving success and the underlying problem-solving characteristics, and (c) behavioral patterns associated with higher efficiency in food extraction. When presented with novel food extraction tasks, the urban macaques having more frequent exposure to novel physical objects in their surroundings, extracted food material from PET bottles and also solved another food extraction task (i.e., extracting an orange from a wire mesh box), more often than those living under more natural conditions. Adults solved the tasks more frequently than juveniles, and females more frequently than males. Both solution-technique and problem-solving characteristics varied across individuals but remained consistent within each individual across the successive presentations of PET bottles. The macaques that solved the tasks showed lesser within-individual variation in their food extraction behavior as compared to those that failed to solve the tasks. A few macaques appropriately modified their problem-solving behavior in accordance with the task requirements and solved the modified versions of the tasks without trial-and-error learning. These observations are ecologically relevant – they demonstrate considerable local differences in food extraction abilities, between-individual variation and within-individual consistency in food extraction techniques among free-ranging bonnet macaques, possibly affecting the species’ local adaptability and resilience to environmental changes.     

The relationship between maternal rank and companion choice in immature macaques (Macaca mulatta andM. radiata)

Eight bonnet and eight rhesus macaques, ranging from 5–8 months of age, were observed with respect to their choices of companions. These data were then correlated with the dominance ranks of the mothers of the companions involved. Preliminary results showed that the maternal ranks of the rhesus infants were significantly correlated with the maternal ranks of their companions; no such correlation could be demonstrated for the bonnet monkeys. Thus the partner choices of rhesus infants appeared to be more constrained than were those of the bonnets. The results were discussed in relation to the social systems of bonnet and rhesus macaques. Supported in part by faculty research grant D-552, Chancellor's Patent Fund DG #44, and by RR00169 to the California Primate Research Center.     

Hematological and serum biochemical indices in healthy bonnet macaques (Macaca radiata)

Background Blood reference values for bonnet macaques (Macaca radiata) are limited. The goal of this study was to determine reference ranges for hematological and serum biochemical indices in healthy, socially housed bonnet macaques for males and females over a range of ages. Methods Blood hematological and serum biochemical values were obtained from 50 healthy bonnet macaques of both sexes and aged 10–234 months. Results Age and sex differences were present in a number of measures. Globulins, total protein, and creatinine (CREAT) values were highest among older subjects, while alkaline phophatase, albumin, and phosphorus values were higher in juveniles. Sex differences were present in concentrations of red blood cells and CREAT, with higher values in males. Conclusion The blood parameter data reported here as age‐specific reference values for laboratory‐housed, healthy bonnet macaques may be used to inform clinical care and laboratory primate research.     

Distribution and Abundance of Primates in Rain Forests of the Western Ghats,Karnataka, India and the Conservation of Macaca silenus

We assessed the distribution and abundance of 4 species of diurnal primates viz. lion-tailed macaque, bonnet macaque, Nilgiri langur and Hanuman langur, in 2 areas—Brahmagiri-Makut and Sirsi-Honnavara—in rain forests of the Western Ghats in the state of Karnataka, India. The Nilgiri langurs in Brahmagiri-Makut and the lion-tailed macaques in Sirsi-Honnavara are the northernmost populations of the 2 species in the Western Ghats. The 2 regions represent changes in ecological zones in the Western Ghats. In Brahmagiri-Makut, Hanuman langurs and bonnet macaques occupy lower elevations whereas Nilgiri langurs live in relatively higher altitudes. Only one group of lion-tailed macaques was in Brahmagiri-Makut. In the forests of Sirsi-Honnavara, 3 species of primates viz. lion-tailed macaque, bonnet macaque and Hanuman langur were in larger numbers throughout the forest. A self-sustainable single population of 32 groups of lion-tailed macaques occurred in Sirsi-Honnavara. Few subspecies of Hanuman langurs exist in the study regions. Due to local hunting practices, the relative abundance of primates in Brahmagiri-Makut is lower than that in Sirsi-Honnavara.     

Ranking network of a captive rhesus macaque society: a sophisticated corporative kingdom

We develop a three-step computing approach to explore a hierarchical ranking network for a society of captive rhesus macaques. The computed network is sufficiently informative to address the question: Is the ranking network for a rhesus macaque society more like a kingdom or a corporation? Our computations are based on a three-step approach. These steps are devised to deal with the tremendous challenges stemming from the transitivity of dominance as a necessary constraint on the ranking relations among all individual macaques, and the very high sampling heterogeneity in the behavioral conflict data. The first step simultaneously infers the ranking potentials among all network members, which requires accommodation of heterogeneous measurement error inherent in behavioral data. Our second step estimates the social rank for all individuals by minimizing the network-wide errors in the ranking potentials. The third step provides a way to compute confidence bounds for selected empirical features in the social ranking. We apply this approach to two sets of conflict data pertaining to two captive societies of adult rhesus macaques. The resultant ranking network for each society is found to be a sophisticated mixture of both a kingdom and a corporation. Also, for validation purposes, we reanalyze conflict data from twenty longhorn sheep and demonstrate that our three-step approach is capable of correctly computing a ranking network by eliminating all ranking error.     

Blood groups of bonnet macaques (Macaca radiata), with a brief introduction to seroprimatology

The human-type A-B-O blood groups of 52 bonnet macaques (Macaca radiata) were determined. Application of method of population genetics indicated the gene frequences to be O = 0.173, A = 0.480 and B = 0.347. Cross testing of sera and red cells of the bonnet macaques revealed two blood-type-specific isoagglutinins, one of them strong enough for use as a blood typing reagent. No blood group polymorphism was revealed by testing bonnet macaque red cells with isoantisera produced in rhesus monkeys (M. mulatta) and in crab-eating macaques (M. fascicularis). The rhesus and crab-eating macaque isoantisera reacted either with all or with none of the bonnet macaque red cells tested.