Male-female affiliative relationships in naturally occurring ringtailed lemurs (Lemur catta) at the Beza-Mahafaly Reserve,Madagascar

Affiliative behavior between adult male and female ringtailed lemurs was examined as part of a project concerning male affiliation with conspecifics of all age/sex classes. Males in three social groups were studied over a 12 month period. Male-female preferred partnerships existed, and were variable according to reproductive season. Dominance rank, age, or tenure of the male did not appear to affect either the number of partnerships or frequency of affiliative behaviors that males had with females. However, males residing in groups with fewer males exhibited both higher frequencies of affiliative interactions with females and were nearest neighbors to females more often than males living in a group containing more males. Females were found to be responsible for proximity maintenance of male-female dyads in the majority of cases. Neither reproductive season nor seasonal availability of food resources strongly affected the frequency of affiliative interactions between males and females, it is proposed that an important aspect of successful group membership for male ringtailed lemurs relates to the development of social relationships with adult females. Males can benefit from such relationships in terms of greater centrality to the spatial core of the group, which can result in enhanced predator protection, greater opportunities for social contact, and potentially greater access to estrous females. © 1996 Wiley-Liss, Inc.     

Social organization in the dusky-footed woodrat (Neotoma fuscipes): A field and laboratory study

The dusky-footed woodrat has been characterized as solitary and asocial. In this research I sought to determine some of the parameters of woodrat social organization. In the field, I live trapped, marked and released members of a distinct group of woodrats at bi-weekly intervals from February to June. Twenty-two residents were identified and 21 out of 34 houses were found to be occupied. Visiting occurred regularly, most often at non-occupied houses. Males were unlikely to be visited at their house and were more likely to visit occupied houses than females. Females were visited at home and visited each other. In June the population was removed to the laboratory where paired encounters with familiar and unfamiliar woodrats were used to examine the effect of sex and familiarity on social interactions. Both females and males interacted little with familiar same sex conspecifics, with the behaviour of one member of such pairs being very inhibited. Females with strange females were more interactive and spent more time in contact. Neither sex clearly differentiated between familiar and strange conspecifics of the opposite sex. Males were agonistic or affiliative in encounters with females. The type of response was consistent for a given male, and females responded differently to the two types. It is concluded that woodrats differentiate between same sex conspecifics and show sufficient individual variability to make individual recognition possible and adaptive.     

Social lives of adult Mysore slender lorises (Loris lydekkerianus lydekkerianus)

Despite the persistent use of the word "solitary" to describe nocturnal primate social behavior, increasing numbers of studies are revealing sophisticated levels of social interactions among nocturnal primates. This study explores the relationships among 11 adult Mysore slender lorises (Loris lydekkerianus lydekkerianus) studied over 10(1/2) months in Tamil Nadu, India. When all observations regarding dependent offspring are excluded, the animals spent on average 38% of their activity in various forms of neutral, affiliative, and agonistic behaviors. Affiliative behaviors were the most common type of social interaction, and males in general were more social than females. Low values for Cole's index (CI) of association emphasize that females rarely interacted with same-sex conspecifics, but commonly interacted with males. In turn, males also formed strong affiliative relationships with other adult males. This index also indicates that levels of affiliation are strongest among animals that share sleeping sites. The Hinde index (HI) suggests that males control proximity to females more than the reverse. A female's tolerance of multiple males in her home range and at a sleeping site may be related to high spatial variability of food resources. Such resources may constrain females with costly reproductive strategies (up to two sets of twins per annum) to a small home range. With their larger home ranges, males may be able to monopolize females by initiating social interactions, and also provide a benefit to females by contributing to parental care.     

Territorial intrusions by pikas (Ochotona princeps) as a function of occupant activity

Pikas (Ochotona princeps) defend individual territories on talus. We tested the short-term stability of pika territories and found that although animals may intrude on the territories of conspecifics at any time, they were much more likely to do so if the occupant was inactive. Pikas usually waited until occupants were unavailable to defend their territory before intruding, apparently to avoid detection and repulsion by the occupant. Intruder avoidance of active occupants was most pronounced among same-sex dyads and non-nearest neighbours. Instances of territory intrusion were most frequent among nearest neighbour heterosexual dyads. Most cases of intrusion appeared to be related to eventual relocation of territories, deterring settlement of unfamiliar conspecifics on nearby vacant territories, and/or increasing familiarity with nearest neighbours of the opposite sex.     

Exclusive core areas and intrasexual territoriality in Eurasian red squirrels (<Emphasis Type="Italic">Sciurus vulgaris</Emphasis>) revealed by incremental cluster polygon analysis

When animal home ranges overlap extensively in species lacking overt territorial behaviours, identifying exclusive core areas within individual ranges can be difficult. By analysing the size and overlap of successively smaller core areas among individual Eurasian red squirrels (Sciurus vulgaris), we determined exclusive areas within the home ranges of resident males and females. Possible effects of habitat composition and food supplies were explored by monitoring squirrels in different conifer forests and during years with low and high tree seed production. Using outlier-exclusive cores (OEC) revealed that the total ranges consisted of large sally zones (on average, 35% of the total minimum convex polygon [MCP] range) around home ranges with multi-nucleate cores. The mean OEC home range size did not differ between the sexes but was larger with poor food availability. Home ranges (99% incremental cluster polygons [ICP]) overlapped extensively between sexes (average overlap high food–low food: males by females 21–40%, females by males 43–45%) and among males (males by males 26–44%), while intrasexual overlap among females was low (9–10%). The overlap of inner cores among females rapidly approached zero, suggesting the intrasexual territoriality of 75% core areas. This was not the case among male squirrels, for which intrasexual overlap averaged only 4% at 50% but 18% at 75% core areas. Even the smallest inner cores had some degree of intersexual overlap, indicating that complete territoriality did not occur in this species. Female home ranges were more strongly affected by annual fluctuations in food supplies than male ranges. Females reduced the size of their food-based intrasexual territories when food availability increases. Males probably benefit from using larger home ranges and core areas, which overlap with the ranges of several females, by increasing their probability of successful mating.     

Social organization of the Milne-Edward's potto

Nocturnal prosimian primates are often seen alone during their nocturnal activities, and are therefore categorized as "solitary". Recent research has shown that these animals actually possess social networks that differ among species. Here we present new information on one of the lesser studied prosimian primates, the potto, derived from fieldwork in Cameroon that employed radiotelemetry and behavioral observations. An analysis of association patterns and home-range overlaps between animals revealed that pottos associated more frequently with conspecifics than expected for a supposedly solitary primate. Certain males and females that were seen together regularly throughout the study, but were not observed to have contact with other pottos of the opposite sex, were labeled as "pairs". These pairs were found to have higher levels of association with each other than with other conspecifics. The only affiliative and sexual behaviors observed between members of the opposite sex occurred within the pairs of pottos. Although given the secretive nature of these primates, the possibility of extrapair copulations cannot be ruled out, their relatively small testis size indicates that sperm competition is not so important for this species.     

Dynamics in space use of American mink (Neovison vison) in a fishpond area in Northern Germany

Telemetry of American mink Neovison vison revealed seasonal and sexual variations in home-range size, intersexual- and partially intrasexual range overlaps and spatial dynamic of territorial system. From 2003 to 2006, 14 mink (nine males and five females) were radio-tracked in a fishpond area in northern Germany. We observed comparatively large home ranges. Both sexes reduced their large summer home ranges to smaller ones in the winter period. In both seasons, male mink used significantly larger areas than female mink. During the mating season, males considerably enlarged their home ranges and roamed nearly through the entire study area. These variations and diversity of range size were discussed in connection with thermoregulatory behaviour, prey availability and distribution, population density and stability of the social system. Analysing intraspecific static interactions, we observed high intersexual overlapping rates in both the summer and winter periods. But there were no or rather very low levels of intrasexual overlapping. However, in the summer period, large home ranges of neighbouring males also overlapped to some extent, but temporal avoidance of adjacent territory holders was observed. Consequently, our results reveal intrasexual territoriality, with males having larger territories, overlapping those of females. The shifting of stable temporary home ranges observed within one season or between the same seasons of consecutive years demonstrates the highly dynamic nature of spatial behaviour among American mink. All these recorded characteristic features of spatial behaviour should be considered when planning mink control projects and measures for the management of this invasive carnivore.     

Male chimpanzee behavior in relation to female ano-genital swelling

This study examined the relationships between male agonistic, affiliative, and sexual behaviors and female estrus condition in captive adolescent and young-adult chimpanzees (Pan troglodytes). Data on agonistic, affiliative, and sexual behaviors of 11 males living in three social groups were collected during daily 45 minute observations over a 5 month period. Female estrus condition was assessed daily using the relative size of the female's ano-genital swelling. It was hypothesized that the presence of maximally tumescent females would generate conflicts between males, so an increase in inter-male agonism was predicted. Males exhibited higher rates of agonism toward other males when at least one female in the group was maximally tumescent. Male affiliative behavior directed toward other males and social play with males were affected by the presence and number of maximally tumescent females. Male sexual behavior increased when maximally tumescent females were present.     

Noninvasive genetic sampling reveals intrasex territoriality in wolverines

Due to its conspicuous manifestations and its capacity to shape the configuration and dynamics of wild populations, territorial behavior has long intrigued ecologists. Territoriality and other animal interactions in situ have traditionally been studied via direct observations and telemetry. Here, we explore whether noninvasive genetic sampling, which is increasingly supplementing traditional field methods in ecological research, can reveal territorial behavior in an elusive carnivore, the wolverine (Gulo gulo). Using the locations of genotyped wolverine scat samples collected annually over a period of 12 years in central Norway, we test three predictions: (1) male home ranges constructed from noninvasive genetic sampling data are larger than those of females, (2) individuals avoid areas used by other conspecifics of the same sex (intrasexual territoriality), and (3) avoidance of same‐sex territories diminishes or disappears after the territory owner's death. Each of these predictions is substantiated by our results: sex‐specific differences in home range size and intrasexual territoriality in wolverine are patently reflected in the spatial and temporal configuration of noninvasively collected genetic samples. Our study confirms that wildlife monitoring programs can utilize the spatial information in noninvasive genetic sampling data to detect and quantify home ranges and social organization.     

Home range and ranging pattern in the slender loris(Loris tardigradus lydekkerianus)

The home range and ranging pattern of the slender loris (Loris tardigradus lydekkerianus) was studied for 21 months in a scrub jungle in Dindigul, Tamil Nadu, south India. Sixteen individuals were observed for a total of 2261 hours. Home ranges were measured for eight adult individuals and eight juvenile and subadult individuals. Males had significantly larger home ranges than the females, and home range size increased post-weaning. The ranging patterns involved minimal female intrasexual overlap, large male intrasexual overlap and large intersexual range overlap.     

Effects of Pair-Bond and Social Context on Male–Female Interactions in Captive Titi Monkeys (Callicebus moloch, Primates: Cebidae)

In monogamous species, an abiding relationship between a specific adult male and a specific adult female is a defining feature of the social system. The interactions between these individuals are influenced by many factors, including not only the history of their relationship (for example, development of a mutual bond), but also the immediate effects of the prevailing social context (for example, presence and sex of extra‐pair conspecifics). In this study we examined the effects of an existing bond and of social context on interactions between adult heterosexual pairs of the monogamous titi monkeys (Callicebus moloch). Twelve adult males and 12 adult females were tested with their cagemates and with an unfamiliar partner of the opposite sex in five social contexts: (1) mated male–female pair; (2) unfamiliar pair; (3) single female; (4) single male; and (5) empty stimulus cage. Results show that mated pairs were more affiliative than unfamiliar pairs and differentiated social contexts more sharply. Males were more responsive to context than females. Distance between mates was less and physical contact was more frequent in the presence of male–female pairs or a single male, than in the presence of a single female or an empty cage. These findings suggest that the presence and sex of strangers have a stronger influence on male–female interactions when the pair has an existing relationship.     

Interindividual interactions of Molina’s hog-nosed skunksConepatus chinga in the Pampas grassland of Argentina

During one year, we radiotracked two female and two male Molina’s hog-nosed skunksConepatus chinga (Molina, 1782), a little studied mephitid, in the Pampas of central Argentina, to analyze the static and dynamic interactions between individuals. Mean home range overlap was large (44.5%), but males shared a smaller proportion of their home ranges than females. The average percentage of localizations in the overlap area (39.3%) indicates that these areas were not marginal sections of individual home ranges, but this value was greater for individuals of different genders than intrasexual dyads. The Coefficient of Spatial Association confirmed that the distances between individuals of different sexes were smaller than between animals of the same sex. During simultaneous locations, females were closer than males, but reciprocal distances between individuals varied over the time. These patterns are congruent with those described as typical for mustelids and mephitids. Our results confirm that an analysis of home range overlap should not be considered complete without the study of dynamic interactions among individuals and their temporal variations, which are necessary to overcome the limitations of spatial overlap analysis.     

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.     

Bridging behavior and other affiliative interactions among male tibetan macaques (Macaca thibetana)

I describe bridging behavior and social relationships between adult males and infants in a free- ranging group of Tibetan macaques (Macaca thibetana)at Mt. Huangshan, China. The subjects performed bridging in which two adult males simultaneously lifted up an infant, sucked or touched its genitalia, and then groomed each other in nonagonistic contexts. Males also expressed social behaviors with other males, such as mounting, penis-sucking, and embracing while touching each other’s penes. Males also employ bridging while exploiting an infant as a social tool, not only to reduce the probability of an aggressive response from dominant males (agonistic buffering), but also to develop and to maintain affiliative social relationships with other males. Use of male infants in bridging contributed to frequent male-infant interactions such as holding,grooming, and penis-sucking. Although these interactions might not have a positive influence on infant survival, they may facilitate the maintenance of affiliative relationships with adult males until they reach maturity. The development of bridging might have a close relation to the high socionomic sex ratio (adult male/adult female) and frequent affiliative interactions between males, especially among the adolescents and adults.     

Experimental field studies of Asian ape social systems

The Asian apes, orangutans and gibbons, possess unusual social systems among anthropoid primates. Social groups of gibbons consist of mated adult pairs and their offspring; mature orangutans are primarily solitary. Recent experimental field research has begun to yield insights into the behavioral mechanisms employed by these animals to maintain their characteristic patterns of social dispersion. While spatial separation between female orangutans appears to be maintained passively, aggression, which is manifest during direct encounters and long-distance vocal interactions, mediates male asociality. Male-male aggression is the result of intense intrasexual competition occurring between animals for mating access to females. To reduce intrasexual competition, male orangutans have adopted alternative mating tactics. In contrast to female orangutans, female gibbons show marked agonistic tendencies toward conspecifics. Female territoriality contributes to preventing males from becoming polygynous. Male gibbons, restricted to monogamous relationships, attempt to ensure their paternity through intrasexual aggression. These observations suggest that the spatial dispersion of females constrains male mating options in both species. However, variations between orangutan and gibbon social systems can be understood as consequences of the temporal dispersion of sexually receptive females. The temporal clumping of females, due to relatively high operational sex ratios, limits the ability of male gibbons to acquire multiple mates. Conversely, an extremely low operational sex ratio in orangutans creates a strong selection pressure for intrasexual competition and polygamous mating. These considerations provide a novel framework for interpreting the social systems of the African apes.     

Spider monkey home ranges: A comparison of radio telemetry and direct observation

The ranging patterns of two male and five female spider monkeys (Ateles geoffroyi) were studied with the use of radio telemetry in Santa Rosa National Park, Costa Rica. The average size of a spider monkey home range was 62.4 hectares; however, range size varied with sex, and, for females, with the presence of a dependent infant. The probability of encountering a radio-collared spider monkey in a three-hour search using radio telemetry (0.91) was much greater than using a visual search (0.20), and telemetric data resulted in a larger estimate of mean home range size than did observational data, when all subjects were compared. However, the difference appeared to be owing to the presence of male ranges in the telemetric, but not the observational, data. When the size of home ranges derived from radio-tracking data for adult females was compared to size of ranges for adult females derived from observations, the results were not significantly different. Adult males had larger home ranges than adult females, thus lending support to the hypothesis that males have adapted to the dispersion of females by occupying a large home range that overlaps the ranges of several adult females. The smallest home ranges were occupied by low-weight females with dependent infants, perhaps reflecting social and energetic constraints.     

Male–Female Association Patterns Among Free-ranging Chimpanzees (Pan troglodytes schweinfurthii)

Although male–female relationships can offer a number of advantages such as protection or social support, they are poorly studied among primates compared to same-sex relationships. We used 12 yr of data from the Kanyawara chimpanzee community to compare three independent measures of association (party association, 5m association, and grooming) among all adult dyads. Party association exhibited by male–female dyads was of intermediate strength between strong male–male and weak female–female association. Male–female dyads were less likely to be within 5m of one another and to groom as male–male dyads, but equally likely to be within 5m and more likely to groom as female–female dyads. Variation in male–female association strength was not related to male rank but was affected by female ranging patterns and female reproductive states. Females with core areas in the center of the home range were more likely to be in parties with males but did not show higher spatial proximity or grooming indices compared to females ranging in the periphery. Party association and 5m indices were higher for dyads of males and estrous females compared to those with anestrous females. These results indicate that male–female dyads are likely to associate with one another more often than female–female dyads because of overlapping ranging patterns and short-term changes in female reproductive state. We conclude that male and female chimpanzees do not exhibit proximity and grooming patterns indicative of strong affiliative bonds. This study also highlights the importance of using multiple independent measures of bond strength in studies of primate social dynamics.     

Space use of common genets<Emphasis Type="Italic">Genetta genetta</Emphasis> in a Mediterranean habitat of northeastern Spain: differences between sexes and seasons

The seasonal home range size and spatial relationships of 16 adult genetsGenetta genetta Linnaeus, 1758 (6 males and 10 females) were estimated in a Mediterranean habitat of northeastern Spain. Genets minimum density was estimated as 0.98/km². Mean annual home range was 113.1 ha in males and of 72.0 ha in females. Males had larger home ranges than females in all seasons, but differences were only significant in winter. Home range size changed seasonally and showed a similar pattern in both sexes, with lower values in summer (males — 41.2 ha, females — 29.0 ha) and maximum ones in spring (males — 78.8 ha, females — 56.1 ha). Animals displayed spatial fidelity throughout the year. Core areas (MCP50) represented 27% and 19% of total home range size for males and females, respectively. Resting home ranges (based on locations of inactive animals) were 9 times lower than overall home range size. Individuals of the same sex overlapped less than individuals of different sexes, especially with regard to core areas, which showed almost no overlap. The results obtained suggest that (1) different factors are likely to affect the space use of genets, such as body mass, food abundance and reproductive cycle; (2) genets use space in a heterogeneous way, with areas of greater activity than others within their home range; (3) there was intrasexual segregation with regard to space use.     

Polygamous mating system and protogynous sex change in the gobiid fish <Emphasis Type="Italic">Fusigobius neophytus</Emphasis>

Whereas mating behaviors and social structure have been studied extensively in monogamous hermaphroditic gobiid species, such studies are relatively limited for polygamous gobiid species. To investigate the reproductive strategy of polygamous gobies, mating groups of the common fusegoby Fusigobius neophytus were observed on reefs of Kuchierabu-jima Island, southern Japan. Males established mating nests on flat-rock surfaces within their territorial home ranges on sandy rubble flats. Females maintained independent home ranges outside the male home ranges during nonreproductive periods, but they shifted their home ranges to overlap with male ranges and actively visited male mating nests during their reproductive periods (1–3 days at ca. 7-day intervals). Females often changed mates during their serial mating. The mating system used by the common fusegoby fits with the definition of male-territory-visiting polygamy. The sex ratio within the study population was female-biased. Nest-holding males were significantly larger than females and were polygynous (mating with up to eight females). These characteristics fit well with the prediction of protogyny by the size-advantage model. Some of the females were observed to undergo functional sex changes to nest-holding males. In addition, small floating males demonstrated sneaking behavior. None of the floating males were derived from females that had changed sex, suggesting a diandric life-history pathway for F. neophytus.     

Use of space by Necromys lasiurus (Rodentia,Sigmodontinae) in a grassland among Atlantic Forest fragments

The spatial patterns presented by the rodent Necromys lasiurus were studied through capture-mark-recapture in a grassland among Atlantic Coastal Forest fragments in Southeastern Brazil. Males moved farther than females. Males moved similar distances in the dry and wet seasons, whereas females moved significantly longer distances in the dry season. Home ranges sizes were estimated for 32 individuals, and varied from 0.02 to 0.52 ha. Males had larger home ranges than females when all data are pooled and in the wet season; in the dry season home ranges of both sexes were of similar sizes. Home range sizes did not vary significantly between seasons for either sex. Home ranges of males overlapped extensively among themselves and with females, whereas females showed mutually exclusive home ranges. The spatial patterns described here are consistent with the results found for this species in different habitats and they suggest that N. lasiurus has a promiscuous mating system.