Monandrous social organization of pigtailed langurs ( Simias concolor ) in the Pagai Islands, Indonesia

Between June 1986 and October 1992, in 3 periods totaling 12 months, we studied social organization of pigtailed langurs (Simias concolor)in the Pagai Islands, Indonesia. With one possible exception, all of 20 family groups contained only 1 adult male each. Nine of them contained only 1 adult female,and 11 contained ≥2 adult females. Mean group size is 4.1 individuals. Home ranges varied from 7 to 20 ha, and population density averages 21 animals/km ² . Simias concoloris sexually dimorphic⤖n a museum sample males average 29% heavier than females, and their canine teeth are on average nearly twice the length of female canines. Combined data from several studies indicate that 60% of S. concolorgroups contain an adult pair plus young,and 40% are multifemale groupings. We sggest that any 1-male mating system that comprises a mixture of 1-female and multifemale groups in the same population should be termed monandry.     

Intergroup calls of male pig-tailed langurs (Simias concolor)

Pig-tailed langurs (Simias concolor) were observed in the Pagai Islands and in Siberut. They lived in 1-male groups containing from one to five or more females plus their young. Adult males produced loud vocalizations of the sortGautier andGautier (1977) called “type 1 loud calls.” The pig-tailed langur's type 1 loud call (TOLC) was a series of 2 to 25 nasal barks, the loudest of which could be heard for 500 m or more in the rain forest. Exchange of TOLC's between males was the only frequent intergroup interaction. Fifty-one percent of TOLC's were emitted spontaneously, 33% occurred in response to TOLC's of other males, 13% to falling trees, 4% in response to thunder, and 1% during a fight between males of different groups. Although TOLC's occurred throughout the day, they exhibited a bimodal distribution, with a peak of calling early in the morning, and another in late afternoon. Tape-recorded TOLC's played back in the field stimulated males hearing them to emit TOLC's of their own. Barks in experimentally-elicited TOLC's were delivered at a faster rate than barks in other TOLC's were. While TOLC's mediate intergroup communication among males, field playback experiments are needed to truly elucidate their functions.     

The social organization of forest hanuman langurs(Presbytis entellus)

The social organization of hanuman langurs (Presbytis entellus;Colobinae) was studied in Kanha Tiger Reserve, Central Indian Highlands, for 2300 hr (1980–1985), in a mosaic of moist deciduous forest and anthropogenic meadow. The langur population density was 46.15/km ² and the mean troop and band sizes were 21.7 and 14.0, respectively. Of 14 troops, 13 were one-male and 1 was trimale. The population adult sex ratio was 1:2.5. The majority of female sexual solicitations was directed toward the harem male. The birth season was December to May, with an estimated gestation of 171–224 days. A review of langur reproductive seasonality suggests that breeding throughout the year is confined to those populations able to exploit human food sources. Mortality during the first year of life was 40%, including infanticide. A significant positive correlation was found between the age of an infant at death or disappearance and the mother’s subsequent interbirth interval. Five cases of social change are described, including female transfer, one-male to multimale change, troop formation, and gradual and rapid replacement of troop males. Takeover-associated infant killing by band males, in an undisturbed moderate-density population, supported the sexual-selection/infanticide hypothesis but not the social-pathology hypothesis. However, it could not be directly confirmed that an invading infanticidal male gains a reproductive advantage. The male tenure of harems was estimated to be 45 months.     

Demography of Simakobu (Simias concolor) and the impact of human disturbance

Asian colobines typically live in small one-male groups (OMGs) averaging five adult females, but Simias concolor (simakobu or pig-tailed langur) is considered an exception because mostly adult male-female pairs have been reported. However, based on their phylogenetic position and marked sexual dimorphism, simakobu are also expected to form OMGs with multiple females. The preponderance of small groups could be the result of human disturbance (hunting or habitat disturbance) reducing group size in the recent past. To investigate this possibility, we documented the demography of ten wild simakobu groups from January 2007 until December 2008 at an undisturbed site, the Peleonan Forest, Siberut Island, Indonesia. We assessed the population-specific size and composition of groups and documented demographic changes due to births, disappearances, and dispersals throughout our 2-year study. We found OMGs with 3.0 adult females on average in addition to all-male groups, but no adult male-female pairs. The ratio of 0.5 infants per adult female (and 0.64 births per female-year in focal groups) suggested that birth rates were similar to those of other Asian colobines. In 5.1 group-years, we observed six dispersal events and six temporary presences (i.e., less than 3 months' residency). Both males and females dispersed, and juveniles seemed to disperse more frequently than adults. To assess the impact of human disturbance on simakobu demography, we compiled data for seven additional populations from the literature and compared them using multiple regressions. Adult sex ratio and the number of immatures per group were influenced negatively by hunting and positively by habitat disturbance while reproductive rates were not significantly affected by either variable. These findings suggest that adult male-female pairs may result from hunting pressure reducing group size, and that conservation action to reduce hunting in the Mentawai Islands is needed to ensure the survival of this critically endangered species.     

Annual Variation in Ecology and Reproduction of Wild Simakobu (Simias concolor)

Seasonal breeding in primates is related to the degree of environmental seasonality, particularly the availability and predictability of food. Southeast Asian species in general show moderate birth seasonality due to either low environmental seasonality or unpredictable fluctuations of mast-fruiting food resources. One Southeast Asian primate, the simakobu (Simias concolor), however, has been reported to be a strict seasonal breeder with births occurring in June and July only. It is unclear whether these observations are characteristic of the species or result from a sampling bias. To address this question, we documented the annual distribution of 11 births in eight groups of simakobu over two consecutive years at Pungut, an undisturbed site on Siberut Island, Indonesia. We assessed annual variation in ecology and reproduction via rainfall, temperature, food availability, feeding time, physical condition, conceptions, and births. Mean monthly temperature was nearly constant (26.3–27.1?°C), and monthly precipitation always high (219–432?mm). Although simakobu foods were abundant year-round, there were two fruit-feeding peaks in June and September. In contrast to previous reports, we documented births in 7?mo. Most births occurred in October (45?%), the wettest month of the year, and most conceptions in March and April, following a peak in unripe fruit availability. Although sample sizes are very small, females seemed to conceive when their physical condition was best, suggesting that simakobu time conceptions flexibly to the recovery of energy reserves. Across study sites, births occurred in 10 calendar months, indicating that simakobu reproduction is not strictly seasonal.     

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.     

Habitat Use and Trophic Niche Overlap of Two Sympatric Colobines, Presbytis potenziani and Simias concolor, on Siberut Island, Indonesia

According to ecological theory, the concept of niche differentiation is essential to our understanding of how sympatric species can limit competition over resources. We here examine ecological niche differentiation in 2 sympatric Asian langurs, Presbytis potenziani and Simias concolor, both endemic to the Mentawai Islands off the west coast of Sumatra. We collected data (home range size, canopy use, activity budgets, diet, and niche breadth and overlap) using GPS and scan/focal animal sampling methods on 2 groups of Presbytis potenziani and 3 groups of Simias concolor living in a mixed primary rain forest in northern Siberut. Results show that home ranges of the 2 species overlapped completely and that the home range size of Presbytis potenziani was ca. 4 times larger than that of Simias concolor. Lower canopy levels (<20 m) were used more often by Simias concolor, whereas Presbytis potenziani preferred the canopy >20 m. Apart from foraging and other activities, there was little difference in overall activity budgets of the 2 species. Regarding diet, although 60% of all food species examined were used by both langur species, they shared only 3 of the 10 most commonly eaten species. Presbytis potenziani fed more selectively on fruits, whereas Simias concolor fed predominantly on leaves. Levin’s niche breadth indices revealed that the diet of Simias concolor is more general (0.34) than that of Presbytis potenziani (0.22). Based on a Pianka index of 0.32, we conclude that there is a relatively small food niche overlap between the 2 colobine species and that diet represents an important mechanism enabling their coexistence.     

Patterns of lateralized hand use in an arboreal primate, Simias concolor

Studies of hand use in nonhuman primates suggest that several species exhibit hand preferences for a variety of tasks. The majority of studies, however, focus on the lateralized hand use of captive nonhuman primate populations. Although captive settings offer a more controlled environment for assessing hand preferences, studies of wild populations provide important insights into how handedness is affected by natural environmental conditions and thus potential insights into the evolution of handedness. To investigate handedness in a population of wild nonhuman primates, we studied patterns of lateralized hand use during feeding in four simakobu monkeys (Simias concolor), an arboreal species inhabiting the Mentawai Islands, Indonesia. Our data show that individual variation in hand preferences for feeding existed among our study animals. In addition, each simakobu expressed a significant hand preference for supporting itself on a branch during feeding, an uncoordinated bimanual task. This bias was most prevalent when the branch used for support was a main branch rather than a terminal branch. When both hands were employed in a coordinated bimanual feeding activity (bimanual manipulation), only two subjects showed a significant bias for feeding. Our data suggest that these individuals are more likely to express significant hand preferences when feeding from stable, rather than precarious, positions within the canopy.     

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.”     

Photosymbiotic ascidians from Pari Island (Thousand Islands,Indonesia)

Photosymbiotic ascidian fauna were surveyed in the subtidal zone off Pari Island in the Thousand Islands (Java Sea, Indonesia). Nine species were recorded: Didemnum molle, Trididemnum miniatum, Lissoclinum patella, L. punctatum, L. timorense, Diplosoma gumavirens, D. simile, D. simileguwa, and D. virens. All of these species have been previously recorded in the Ryukyu Archipelago, Japan. Diplosoma gumavirens and D. simileguwa were originally described from the Ryukyu Archipelago in 2009 and 2005, respectively, and all of the observed species are potentially widely distributed in Indo–West Pacific coral reefs.     

The organization of song in the Hainan black gibbon (Hylobates concolor hainanus)

There have never been any documented studies on the vocal or non-vocal behaviour of wild or captive Hainan black gibbons, which are highly endangered in the wild (Hainan Island, China), and very scarce in captivity. The singing behaviour of the only pair of captive Hainan black gibbons outside Southeast Asia was observed, recorded and analyzed in detail. This adult pair produced duets only, with the organizational features of their duets found here to be similar to that in two other concolor gibbon sub-species; however, the acoustical features were very different from any other concolor gibbon pair studied previously. Some functional and evolutionary implications of the singing behaviour of the Hainan black gibbon are presented and discussed.     

Interspecific spacing between gibbons (Hylobates klossii) and langurs (Presbytis potenziani) on Siberut Island,Indonesia

Spatial arrangement and social interactions of two sympatric and ecologically similar primate species, Hylobates klossii and Presbytis potenzianai, are described from field observations made between July 1972 and October 1974 on Siberut Island, Indonesia. Gibbon territories and langur home ranges overlap extensively. Because gibbons have the ability to supplant langurs at shared food sources, langurs are at a competitive disadvantage. To avoid or decrease the frequency of hostile interactions with gibbons, langurs locate their core areas on boundaries between adjacent gibbon territories, which permits langurs to retreat across these barriers in response to gibbon movements. Langurs further enhance segregration by leaving their sleeping trees earlier than gibbons, gaining additional feeding time on contested food sources. This form of interspecific spatial organization between gibbons and langurs resembles certain predator-prey spacing systems, where territorial boundaries between adjacent predators serve as sanctuaries for prey populations.     

Social organization of white‐headed langurs (Trachypithecus leucocephalus) in the Nongguan Karst Hills,Guangxi, China

The number of males per group is the most variable aspect of primate social organization and is often related to the monopolizability of females, which is mainly determined by the number of females per group and their reproductive synchrony. Colobines show both inter‐specific and intra‐specific variations in the number of males per group. Compared with other colobine species, little is known about the social organization of white‐headed langur (Trachypithecus leucocephalus), despite its endangered status and unusual limestone habitat. As a part of a long‐term study of the white‐headed langurs in the Nongguan Karst Hills, Guangxi, China, we quantitatively investigated their social organization by analyzing census data from 1998 to 2003. The population censuses revealed that the predominant social organization of bisexual groups was the one‐male group, similar to a previous report on this species and many other Asian colobines. In such groups, one adult male associated with 5.1 adult females, 0.1 sub‐adult males, 2.6 juveniles and 2.9 infants on average, with a mean group size of 11.7 individuals. In addition, three multi‐male groups were recorded, consisting of 2–3 adult males, 1–5 adult females, 0–2 sub‐adult males, 0–7 juveniles and 0–2 infants. They did not contain more adult females than the one‐male groups and were unstable in group membership. The langurs outside bisexual groups were organized into small nonreproductive groups or lived as solitaries. The nonreproductive groups averaged 1.3 adult males, 1.3 sub‐adult males and 2.6 juveniles. Juvenile females were present in such groups on 52.4% of all occasions. As predicted by the monopolization model, the prevalence of the one‐male pattern in this species may mainly be attributed to the small number of females in the group. The possible reasons for the occurrence of multi‐male groups and the presence of juvenile females in nonreproductive groups are also discussed. Am. J. Primatol. 71:206–213, 2009. © 2008 Wiley‐Liss, Inc.     

云南无量山印支灰叶猴社会结构初步研究

乌叶猴属物种主要以一雄多雌的小群活动,其群体大小一般不超过20只。泰国东北部的印支灰叶猴生活在一雄多雌或多雄多雌群的小群中,而云南无量山的印支灰叶猴却生活在多雄多雌的大群中,有些群大小超过100只,但是对无量山印支灰叶猴的社会结构缺乏研究。2012年9月至2015年7月,在云南无量山国家级自然保护区大寨子片区开展一群习惯化印支灰叶猴的社会结构研究。采用10min间隔的焦点动物瞬时记录法,记录焦点成年雄性5 m范围内不同性别年龄个体的数量。观察期间共有594次准确识别焦点雄性5 m范围内其它个体年龄性别的有效数据 (至少有1只其他个体)。其中,焦点雄性周围出现其它成年雄性的几率为44.2%,数量为(0.62 ± 0.85)只;同时出现成年雄性和成年雌性的几率是32.5%。取食时焦点雄性周围出现其它雄性的比例 (60.0%) 和数量 (平均0.93只) 都大于休息和移动时,数量最多为5只成年雄性。结果表明该群灰叶猴的社会结构不是由多个OMU和AMU组成的重层社会,而是多雄多雌的社会结构。无量山印支灰叶猴群内成年雄性间相互容忍度较高,个体的空间分布在取食时尤其密集,说明无量山印支灰叶猴的食物丰富 (至少在某些季节),这也部分解释了为什么无量山印支灰叶猴能够生活在大群中。本文为进一步研究亚洲疣猴社会结构的多样性和进化提供了重要的基础资料。     

Weaving a tight social net: Allogrooming in free-ranging female langurs (Presbytis entellus)

We studied grooming among adults of a one-male multifemale troop of free-ranging Hanuman langurs (Presbytis entellus)living near Jodhpur, India, for 9 years. The 11–13 females devoted about 6% of their day to allogrooming. Adult males, whose tenures averaged 2.2 years, were transient figures in the troop's history, as reflected by their rather peripheral role in the grooming network. Females groomed males 4–40 times more frequently (1006 episodes) than vice versa- (176 episodes). Adult females received 97% of all grooming from other adult females (6655 episodes). Although females exhibited an age- inversed dominance hierarchy, they did not compete for grooming access to particular troop mates. Dyads of all possible rank differences occurred as frequently as expected: 51% of grooming was directed up the hierarchy and 49% down it. Young, high- ranking individuals gave and received significantly more grooming than the oldest, low- ranking females did. The pattern seemed to be influenced by kin selection because of the presumably high degree of female relatedness. They invested most in troopmates with the highest reproductive value, i.e., the youngest individuals. This trend was coupled with a preference of closest kin (mothers and daughters). Reciprocity was the outstanding feature since all adult females groomed and were groomed by all others. Such a tight social net might establish the necessary cohesion during frequent territorial disputes with neighboring troops.     

Ecological correlates of lion social organization (Panthers, leo)

Data on lion home range size, pride size, density, group size, cub survival, litter size and adult sex ratio were extracted from studies carried out in 10 habitats. The data were compared with five measures of food supply: mean prey biomass, prey biomass during the season of greatest abundance, prey biomass during the season of least abundance, biomass of middle-sized prey species (i.e. those with mean adult live-weights between 50 and 250 kg), and the mean size of carcasses fed on by lions. The results indicated that range size is inversely correlated with the abundance of prey during the period of least abundance, but not with overall prey abundance. Range size showed no consistent variation with pride size or with minimum metabolic requirements. Pride size (measured in terms of average number of animals per pride and average number of adult females) and cub survival strongly correlated with lean season food abundance. No relationship was found between group size or litter size and any of the measures of food supply Adult sex ratio did not vary consistently with food supply or lion density, although the data did suggest that prides inhabiting small, circumscribed reserves may experience less inter-male competition and this, in turn, may affect the adult sex ratio.