One-male groups and intergroup interactions of mountain baboons

The behavioral ecology of mountain baboons, Papio ursinus,cuts across the traditional dichotomy between savannah and desert baboons, providing fresh opportunities to examine socioecological relationships. Mountain baboons were studied at a site where covariation in altitude and group size helps to clarify the influence of ecology on social behavior. One-male groups and lone males were regularly found in the highest-altitude zones. In encounters with multimale groups, a one-male group retreated; the group’s single male attacked his females, herding them far away from the multimale group. The male himself then approached and watched (and often loud-called at) this group. In a two-male group, the younger male herded the females, while the older sometimes defended the group. One-male groups did not show regular, stable aggregations, but two such groups seasonally coordinated their ranging “in tandem.” Ecological explanations for these behaviors and their relationship to the behavior of hamadryas baboons are evaluated, and a new theory of the origin of one-male groups in baboons is developed.     

The variable social organization of hanuman langurs(Presbytis entellus), infanticide,and the monopolization of females

Data from 24 wild populations of hanuman langurs (Presbytis entellus)in south Asia are used to test hypotheses seeking to explain variation in troop structure and the incidence of infanticide. The occurrence of infanticide is associated with a one-male troop structure and not with a high density. The density, predation, and economic-advantage hypotheses, as explanations for the occurrence of one-male and multimale troops, are not supported by the review. However, the monopolization hypothesis is not contradicted; the number of adult males per troop is significantly correlated with troop size and with the number of adult females per troop. Therefore it is suggested that a one-male troop structure will arise if a male is able to monopolize a group of females, a multimale troop if he cannot. One-male troops may predispose to infanticide because of high variance in male mating success and high intermale competition between groups rather than within troops. If female dispersion determines troop structure, it is speculated that females could manipulate males to form a multimale society if the advantages in terms of infant survival and intertroop conflict exceeded the costs in terms of not producing infanticidal “sexy sons.”     

Preliminary report on the grouping of mandrills (Mandrillus sphinx) in Cameroon

A field study of mandrill was carried out from 1979 to 1983 in Cameroon for 27 months. Group size of mandrills ranged from 15 to 95 and composition was estimated by direct and indirect observations. The ratio of group size to one adult male was 13.9, larger than other baboon species except the drill. Solitary males were seen. The mandrill may have two types of groups: one-male and multi-male. The multi-male group was observed to split into subgroups, one of which was sometimes a one-male group. The multi-male subgroup was assumed to be composed of several one-male groups. The home range area of mandrills was estimated to be at least 5 km⁵ to 28 km². Their large home range size could be related to their characteristic feeding pattern in the forest.     

Impact of Male Infanticide on the Social Structure of Mountain Gorillas

Infanticide can be a major influence upon the social structure of species in which females maintain long-term associations with males. Previous studies have suggested that female mountain gorillas benefit from residing in multimale groups because infanticide occurs when one-male groups disintegrate after the dominant male dies. Here we measure the impact of infanticide on the reproductive success of female mountain gorillas, and we examine whether their dispersal patterns reflect a strategy to avoid infanticide. Using more than 40 years of data from up to 70% of the entire population, we found that only 1.7% of the infants that were born in the study had died from infanticide during group disintegrations. The rarity of such infanticide mainly reflects a low mortality rate of dominant males in one-male groups, and it does not dispel previous observations that infanticide occurs during group disintegrations. After including infanticide from causes other than group disintegrations, infanticide victims represented up to 5.5% of the offspring born during the study, and they accounted for up to 21% of infant mortality. The overall rates of infanticide were 2–3 times higher in one-male groups than multimale groups, but those differences were not statistically significant. Infant mortality, the length of interbirth intervals, and the age of first reproduction were not significantly different between one-male versus multimale groups, so we found no significant fitness benefits for females to prefer multimale groups. In addition, we found limited evidence that female dispersal patterns reflect a preference for multimale groups. If the strength of selection is modest for females to avoid group disintegrations, than any preference for multimale groups may be slow to evolve. Alternatively, variability in male strength might give some one-male groups a lower infanticide risk than some multimale groups, which could explain why both types of groups remain common.     

Group encounters and territoriality in wild Alaotran gentle lemurs (Hapalemur griseus alaotrensis)

During a 3 month field study, 18 group encounters between four groups of Alaotran gentle lemurs (Hapalemur griseus alaotrensis) were observed in the Lake Alaotra marshland in Madagascar. Behaviors observed during group encounters are described, and quantitative data on intergroup interactions and ranging patterns are discussed in the context of territoriality. Intergroup interactions varied in their intensity, ranging from visual monitoring, scent marking, and display locomotion to penetrating the neighboring range. A quarter (27%) of the encounters were aggressive, involving chases and confrontation displays, and no affiliative interactions were observed. Both adult males and females were involved in intergroup encounters, with males playing more active roles. All encounters took place in the small overlapping areas of neighboring home ranges. The outcome of an aggressive encounter was determined by its location, with the resident group always driving out the intruding one. The Alaotran gentle lemur actively advertised and consistently defended a large area of its home range against intrusion of neighboring groups and can therefore be considered territorial. Am. J. Primatol. 46:251–258, 1998. © 1998 Wiley-Liss, Inc.     

Variation in diet and ranging of black and white colobus monkeys in Kibale National Park,Uganda

Recently, considerable intraspecific variation in the diets and ranging behavior of colobine monkeys has been described, although in most cases this has involved documenting variation between, not within, sites. Some African colobines, such as guerezas (Colobus guereza), are relatively abundant in disturbed habitats that are very heterogeneous, raising the intriguing possibility that even groups with overlapping home ranges may exhibit large behavioral differences. If such differences occur, it will be important to understand what temporal and spatial scales adequately portray a species’ or population’s diet and ranging behavior. This study documents within-site variation in the diet and ranging behavior of guerezas in the habitat types in which they are described to be most successful—forest edge and regenerating forest. We collected data on eight groups of guerezas with overlapping home ranges for 3–5 months each in Kibale National Park, Uganda. The guerezas were highly folivorous, with leaves constituting 78.5–94.0% of the groups’ diets. The percentage of mature leaves and fruit in the diet varied widely among and within groups. We show that differences among groups in the intensity with which they fed on specific tree species were not just related to phenology, but also to differences in the forest compositions of groups’ core areas. Range size estimates varied more than fivefold among groups and the minimum distance from groups’ core areas to eucalyptus forest (which all groups regularly fed in) was a better predictor of range size than was group size. These results reveal considerable variation in the diet and ranging behavior among groups with overlapping ranges and have implications for comparative studies, investigations of within- and between-group feeding competition, and the potential for populations to adapt to anthropogenic or natural environmental change.     

Female sexual swellings in the Asian colobine Simias concolor

During a preliminary study of pig-tailed langurs (Simias concolor) in the Pagai Islands, Indonesia, it was discovered that females exhibited conspicuous swelling of the urogenital triangle. The pig-tailed langur is the first Asiatic colobine found to have prominent sexual swellings and the only colobine with sexual swellings that lives in one-male groups. Because all anthropoids with conspicuous sex skin typically live in groups having female-biased adult sex ratios, it is possible that females might compete amongst themselves for the male in one-male groups, or the best males in multimale groups. Sexual swellings may therefore have resulted from sexual selection for signals attractive to males in female-female competition, as suggested earlier by Bercovitch (California Anthropologist 8:9–12, 1978). Prolonged observation of recognizable individuals will be required to test this hypothesis in the pig-tailed langur and other species.     

Shifts in Social Structure of Black Howler (Alouatta pigra) Groups Associated with Natural and Experimental Variation in Population Density

We examined variation in the group structure of black howlers (Alouatta pigra) using the adult composition of 48 social groups. We compared the structure of groups at 5 sites with different population densities and variation in group structure over time with rising population density. In addition, we examined changes in the group structure of monkeys that were translocated from an area of high population density to an area with a much lower population density. We found at low population densities, groups comprised either heterosexual pairs or a single male with two females. At high population densities groups tended to be multimale and often contained >2 adult females. We suggest the relative costs and benefits of dispersal by maturing adults varies with population density, and in Alouatta pigra results in a shift from single to multimale groups of larger size with increasing population density.     

Paternity analysis of alternative male reproductive routes among the langurs (Semnopithecus entellus) of Ramnagar

Because primate males usually invest very little in offspring, male reproductive success will mainly be determined by access to fertile females with differences in access leading to differential male reproductive success. To determine the outcome of alternative male reproductive routes, we investigated a wild population of Hanuman langurs at Ramnagar, South Nepal, where groups were either one-male or multimale. Paternity was established by DNA analysis from faeces for 42 infants in five groups. In one-male groups all infants were sired by the only resident male in the group. In multimale groups the alpha male fathered significantly more infants (57%) than all other resident males. Nonresident males sired at least 21% of all infants born into multimale groups. Because of the lower mean number of infants sired by alpha males during their whole tenure as alpha compared with males in one-male groups (2.3 versus 6.8) and the higher maximum value (8.9 versus 6.0), the alpha male route is considered the riskier option. Based on demographic data we suggest that the considerable variance in short-term reproductive success might not translate to the same degree into differences in long-term reproductive success. Copyright 2001 The Association for the Study of Animal Behaviour.     

Activity and Ranging Patterns of Guerezas in the Kakamega Forest: Intergroup Variation and Implications for Intragroup Feeding Competition

From March 1997 to February 1998, I investigated the activity patterns of 2 groups and the ranging patterns of 5 groups of eastern black-and-white colobus (Colobus guereza), aka guerezas, in the Kakamega Forest, Kenya. Guerezas at Kakamega spent more of their time resting than any other population of colobine monkeys studied to date. In addition, I recorded not one instance of intragroup aggression in 16,710 activity scan samples, providing preliminary evidence that intragroup contest competition may be rare or absent among guerezas at Kakamega. Mean daily path lengths ranged from 450 to 734 m, and home range area ranged from 12 to 20 ha, though home range area may have been underestimated for several of the study groups. Home range overlap was extensive with 49–83% of each group's range overlapped by the ranges of other groups. Despite the high level of home range overlap, the frequently entered areas (quadrats entered on 30% of a group's total study days) of any one group were not frequently entered by any other study group. Mean daily path length is not significantly correlated with levels of availability or consumption of any plant part item. Mean daily path length is also not significantly correlated with group size, though the largest group did have the longest mean daily path length. This finding suggests that intragroup scramble competition may have been rare or absent among guerezas at Kakamega except perhaps in the largest group, which was unusually large.     

Intraspecific Variation in Home Range Overlap with Habitat Quality: A Comparison among Brown Bear Populations

We developed a conceptual model of spatial organization in vertebrates based upon changes in home range overlap with habitat quality. We tested the model using estimates of annual home ranges of adult females and densities for 30 populations of brown bears (Ursus arctos) in North America. We used seasonality as a surrogate of habitat quality, measured as the coefficient of variation among monthly actual evapotranspiration values for areas in which study populations were located. We calculated home range overlap for each population as the product of the average home range size for adult females and the estimated population density of adult females. Home range size varied positively with seasonality; however, home range overlap varied with seasonality in a nonlinear manner. Areas of low and high seasonality supported brown bears with considerable home range overlap, but areas of moderate seasonality supported brown bears with low home range overlap. These results are consistent with behavioural theory predicting a nonlinear relationship between food availability and territoriality.     

Even adult sex ratios in lemurs: Potential costs and benefits of subordinate males in Verreaux's sifaka (Propithecus verreauxi) in the Kirindy Forest CFPF,Madagascar

Optimal group size and composition are determined by both the costs and benefits of group living for the group's members. Verreaux's sifakas (Propithecus verreauxi), a diurnal lemur, form multimale multifemale groups with the tendency toward even adult sex ratios despite a small average number of females per group. The unexpected presence of multiple adult males may be explained by tolerance of other group members if subordinate males provide benefits to the group that outweigh the costs associated with their presence. Results based on both demographic data collected over a 13‐year period and behavioral observations suggest that subordinate males provide no benefits in terms of infant survival and defense against group takeover by outside males. Although groups with more males are more likely to win intergroup encounters, subordinate males do not participate in these encounters more often than expected. Subordinate males are not costly to other group members in terms of direct intragroup feeding competition, but aggression rates between dominant and immigrated subordinate males increase in the mating season. Even though subordinate males provide very few benefits to the group, they are not very costly either and thus may be tolerated by resident females and dominant males. This tolerance may help to partially explain the tendency towards their unusual adult sex ratio. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Deferred agonistic behavior in a long-lived scincid lizard Eumeces laticeps

Summary A laboratory experiment with the broad-headed skink (Eumeces laticeps) involving staged agonistic encounters demonstrates that larger males have an advantage over smaller ones in agonistic bouts. Field data on head wounds produced by intraspecific fighting during the breeding season show a much higher frequency of new wounds among males over 100 mm in snout-vent-length than in smaller males. The significant difference in new-wound frequency strongly suggests avoidance of fights by the small males, which is corroborated by laboratory and field observations. Access by males to reproductively active females depends on the ability to defeat other males in aggressive contests virtually always involving head biting if the males are of nearly equal size. Because the probability of winning agonistic encounters increases with size, young males avoid fights with older males. Aggressive contests with larger males and reproductive attempts other than courtship in the absence of larger males are deferred.Aggressive behavior in E. laticeps may be employed in direct defense of females, but might also be expressed in defense of specific sites and/or territories. In the laboratory, males in their home cages were significantly more likely to win encounters with males of similar size than were males fighting in the home cages of opponents. This suggests that encounter site could be important in determining encounter outcome and that field study of possible site defense or territoriality is needed.     

Home Range Size and Use in Allocebus trichotis in Analamazaotra Special Reserve, Central Eastern Madagascar

No information is currently available on the space needs of hairy-eared dwarf lemurs (Allocebus trichotis), classified as Data Deficient. The data are crucial for their conservation and comparison with other nocturnal primates. I conducted the first radiotracking study of the species from January to December 2007 in the Analamazaotra Special Reserve of Central Eastern Madagascar. I used nocturnal focal individual follows and daytime nest locations to determine home ranges. I followed 1 full sleeping group (4 adults) for 8 mo and 1 partial sleeping group (2 females) for 3 mo. Group home ranges, calculated via 100% minimum convex polygons (MCP), were 35.5 ha and 16.0 ha, respectively. The 95% kernel method of analysis yielded group home ranges of 15.2 ha and 7.1 ha respectively. The mean home range size for individuals was 15.4 ha (MCP) and 5.4 ha (kernel). This is much larger than for other Cheirogaleidae and could be due to a more insectivorous diet or the use of patchily distributed gum-producing trees. There were small nonsignificant monthly variations in home range size. The mean home range size per individual per month was 5.2 ha (MCP) and 2.2 ha (kernel). Important individual differences in overall and monthly home range size could be due to variations in the individual reproductive cycles and survival strategies. Overlap analyses and the lack of sexual difference in home range size suggest the social unit is a family or multimale/multifemale sleeping group with monogamous or promiscuous mating. The Analamazaotra Special Reserve probably holds ca. 100 adult individuals. Additional research is urgently needed to clarify the habitat needs of this rare species.     

Spacing system of the Mysore slender loris (Loris lydekkerianus lydekkerianus)

Loris lydekkerianus lydekkerianus has been shown to have a promiscuous copulatory pattern, to maintain social networks via frequent loud calls, to interact socially throughout the night with all age classes, and to sleep socially. Though these behaviors point towards a multimale social system, no study of their spacing system has yet been provided to support this view. From October 1997-August 1998, I conducted a study of the Mysore slender loris in Ayyalur, India. During 1,400 field hours, data were collected on range use of 3 adult females, 3 adult males, 1 subadult female, and 1 subadult male. Lorises slept in groups averaging 4 individuals, composed of an adult female, her offspring, and 1-2 adult and subadult males. Sleeping sites for three groups were located within 1.9 ha in the center of the study area. The minimum convex polygon in hectares encompassing each animal's range was determined, as well as overlap among home ranges of individual lorises. Average home range sizes were: adult males, 3.6 ha +/- 0.09; subadult/smaller males, 1.17 ha +/- 0.26; and adult and subadult females, 1.59 ha +/- 0.24. Male ranges overlapped with at least 2-3 other adult males (0.72 ha +/- 0.23). Female ranges overlapped slightly with at least 2 other female ranges (0.22 ha +/- 0.25). Male ranges overlapped those of at least 3 females (0.82 ha +/- 0.51). Patterns of home range and sleeping site support previous suggestions of a multimale social system, similar to aye ayes and some galagos.     

Social dynamics of intergroup encounters in the capped langur (Presbytis pileata)

Data from a 15–month field study of the capped langur (Presbytis pileata) in moist deciduous forest in Bangladesh show that during intergroup encounters males respond more aggressively to extra-group males and to the resident males of unfamiliar one-male groups than they do toward familiar males. Lone males followed established one-male groups and attempted to interact with group females. During intergroup encounters males responded to the approach of unfamiliar males with significantly more displays than were given to familiar (frequently encountered) males. The frequency of displays given to approaching groups showed a negative correlation with the frequency of encounters with those groups. In 50% of intergroup encounters males pushed and bit their group females if the females strayed from the immediate proximity of the group male. Further observations suggest that males obtain females that have emigrated from established groups or attempt to interact with them in established onemale groups. Males had a tenure from at least 13 months to at least 26 months, and between 1986 and 1988 no male changes were seen. Capped langur males defended females from other males, but females did not defend resources or participate in intergroup encounters. The implications of this sex difference in behavior during encounters is discussed with regard to ecological models of female-bonded primate groups.     

Ranging Behavior of Proboscis Monkeys (Nasalis larvatus) in the Lower Kinabatangan, Northern Borneo

I studied the ranging behavior of proboscis monkeys (Nasalis larvatus) at two sites in the Lower Kinabatangan Region of northern Borneo. I collected data on ranging behavior via scan sampling during group follows. Groups of Nasalis larvatus had ranges overlapping those of other groups in each area. I observed no territorial behavior. Groups of Nasalis larvatus occasionally swam across the Kinabatangan River, and frequently across its tributaries. The home range size of a focal one-male group (SU1) was 220.5 ha. The group traveled farther on days when the proportion of young leaves in the diet was higher. In addition, SU1 used particular areas when they fed on flowers and fruits. Apparently, rainfall and phenology did not influence ranging patterns.     

Variance in the reproductive success of dominant male mountain gorillas

Using 30 years of demographic data from 15 groups, this study estimates how harem size, female fertility, and offspring survival may contribute to variance in the siring rates of dominant male mountain gorillas throughout the Virunga Volcano Region. As predicted for polygynous species, differences in harem size were the greatest source of variance in the siring rate, whereas differences in female fertility and offspring survival were relatively minor. Harem size was positively correlated with offspring survival, even after removing all known and suspected cases of infanticide, so the correlation does not seem to reflect differences in the ability of males to protect their offspring. Harem size was not significantly correlated with female fertility, which is consistent with the hypothesis that mountain gorillas have minimal feeding competition. Harem size, offspring survival, and siring rates were not significantly correlated with the proportion of dominant tenures that occurred in multimale groups versus one-male groups; even though infanticide is less likely when those tenures end in multimale groups than one-male groups. In contrast with the relatively small contribution of offspring survival to variance in the siring rates of this study, offspring survival is a major source of variance in the male reproductive success of western gorillas, which have greater predation risks and significantly higher rates of infanticide. If differences in offspring protection are less important among male mountain gorillas than western gorillas, then the relative importance of other factors may be greater for mountain gorillas. Thus, our study illustrates how variance in male reproductive success and its components can differ between closely related species.     

Harem structure and female territoriality in the dwarf hawkfish <Emphasis Type="Italic">Cirrhitichthys falco</Emphasis> (Cirrhitidae)

We investigated the territoriality and the spatial and mating relationships of the haremic hawkfish, Cirrhitichthys falco, on a reef off Kuchierabu-jima Island in southern Japan. Each individual maintained a territorial home range which was defended against same-sex conspecifics at the boundary of the home range. The territory of each male encompassed the territories of 2–3 females, allowing the male to completely monopolize mating opportunities with those females. Based on our observations, we classified the harem type of C. falco as a territorial female type. Large juveniles maintained independent home ranges outside the female territories. In contrast, small juveniles were allowed to cohabit within the territory of an adult female. Stomach contents analysis revealed that the smallest size class of C. falco fed primarily on copepods. In contrast, all other size classes fed primarily on decapods. Together, these results suggest that female territoriality plays an important role in defending food resources.     

Field ecology and behaviour of the rhesus macaque (Macaca mulatta): I. Group composition,home range,roosting sites,and foraging routes in the Asarori Forest

The data presented here resulted from about 1,100 observation hours in a nine-month field study of rhesus monkey from January to October 1976 in the Asarori Forest, Dheradun. Six groups were studied, the group size varied from 6–90 individuals. With the exception of one group, all were of the bisexual multimale type. The number of adult males per group varied from two to seven, and of adult females from 4 to 27. The adult sex ratio was male 1 to females 2.2–3.7 (mean 1:2.7). The exceptional group originally consisted of five juveniles only (3 ♂, 2 ♀); later on an adult male joined the group. Home ranges varied from 1.3–13.4 km² and overlapped extensively with another. The area of home range showed straight line relationship with group size. Roosting sites were not fixed but changed from night to night. The straight line distance between one night to the next varied from 75–2,500 m. Foraging routes per day ranged from 1,050–3,500 m (mean 1,803.3±160.2). There is a significant relationship between the foraging route (L) and the straight line distance between roosting sites (D). The ratio D/L varies from 0.14–0.87.