Proboscis monkey (Nasalis larvatus) social organization: Intergroup patterns of association

Proboscis monkey (Nasalis larvatus) social organization was studied at the Natai Lengkuas station, Tanjung Puting National Park in Kalimantan Tengah, Indonesia. Data were collected on ten one-male groups and two all-male groups using scan sampling and event sampling during follows, and through evening census surveys. Proboscis monkeys were found to form stable one-male groups. Specific groups associated at their sleeping sites, in a manner similar to the fission-fusion pattern described for hamadryas and gelada baboons. Analyses performed on these associations indicated that the groups formed two separate bands. These separate bands appear to be a secondary level of organization, distinct from the primary level of organization, the one-male and all-male groups. This is the first evidence for such a two-tiered system of organization in an arboreal colobine species. Proboscis monkeys closely resemble gelada baboons in their organizational pattern. In both species, adult females direct the majority of their affiliative behavior towards their offspring and other adult females, males do not engage in active herding, all-male groups are present, and one-male groups do spend time apart from other groups. Home ranges of both groups and of bands of proboscis monkeys overlapped extensively; temporal avoidance between bands may have been mediated by early morning vocalizations and branch shaking displays.     

Proboscis monkey (Nasalis larvatus) social organization: Group structure

Proboscis monkey (Nasalis larvatus) social organization was studied at the Tanjung Puting National Park in Kalimantan Tengah, Indonesia, for 12 months. Data were collected on one-male groups and all-male groups by using scan sampling and event sampling while following groups. Evening census surveys were also conducted. Proboscis monkeys were found to follow the typical Asian colobine pattern of one-male social groups, with extra-group males forming all-male groups. These groups appear stable, with few changes in membership. One-male groups appear to be “female bonded”; adult females direct affiliative behaviors towards their offspring and other adult females, not towards the male. Nearest-neighbor data indicate some peripheralization of juvenile animals.     

Feeding ecology of the proboscis monkey (Nasalis larvatus)

Proboscis monkey (Nasalis larvatus) feeding behavior and ecology were studied at the Natai Lengkuas Station, Tanjung Puting National Park, Kalimantan Tengah, Indonesia. Data on feeding behavior were collected using scan sampling during group follows. Three vegetational plots containing 1,732 trees were established and monitored monthly for changes in fruit, flower, and young leaf production. Basal area and canopy cover were calculated and used in estimating food abundance. Proboscis monkeys were found to be folivore/frugivores, specializing in seed consumption. At least 55 different plant species were used as food sources, with a marked preference for Eugenia sp. 3/4,Ganua motleyana and Lophopetalum javanicum. These tree species were among the most frequent and most dominant. However, proboscis monkeys were selective feeders; use of tree species as food sources was not based simply on relative density. During times of low food abundance and/or availability proboscis monkeys switched dietary strategies and increased dietary diversity. The average total home range was estimated to be 130.3 ha, with an average group density of 5.2 groups per km². The average biomass per km² was estimated to be 499.5 kg. Given their high biomass and predilection for consuming seeds of dominant species, proboscis monkeys may help to maintain and increase vegetational diversity.     

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.     

Regurgitation and remastication in the foregut-fermenting proboscis monkey (Nasalis larvatus)

Although foregut fermentation is often equated with rumination in the literature, functional ruminants (ruminants, camelids) differ fundamentally from non-ruminant foregut fermenters (e.g. macropods, hippos, peccaries). They combine foregut fermentation with a sorting mechanism that allows them to remasticate large particles and clear their foregut quickly of digested particles; thus, they do not only achieve high degrees of particle size reduction but also comparatively high food intakes. Regurgitation and remastication of stomach contents have been described sporadically in several non-ruminant, non-primate herbivores. However, this so-called 'merycism' apparently does not occur as consistently as in ruminants. Here, to our knowledge we report, for the first time, regurgitation and remastication in 23 free-ranging individuals of a primate species, the foregut-fermenting proboscis monkey (Nasalis larvatus). In one male that was observed continuously during 169 days, the behaviour was observed on 11 different days occurring mostly in the morning, and was associated with significantly higher proportions of daily feeding time than on days when it was not observed. This observation is consistent with the concept that intensified mastication allows higher food intake without compromising digestive efficiency, and represents an expansion of the known physiological primate repertoire that converges with a strategy usually associated with ruminants only.     

Feed intake,digestion and passage of the proboscis monkey (Nasalis larvatus) in captivity

Feeding trials were conducted with a troop of six proboscis monkeys (Nasalis larvatus) at the New York Zoological Park to quantify aspects of digestive physiology. Diets consumed comprised, on an as-fed basis, 55% browse and green produce, 11% fruit, 9% commercial primate diets, 9% root vegetables, and 16% miscellaneous items. Total feed intake averaged 12% of body mass; dry matter intake was 3%. Dry matter and plant cell wall disappearance exceeded 80%. Passage marker studies (three 6-day trials) revealed: transit time=14 hr; mean passage time=49 hr; and 5–80% retention time=52 hr. Digestion coefficients and retention times were greater than expected based on body size. These results might reflect the low fiber content of the diets fed, or alternatively, provide evidence that proboscis monkeys are efficient digesters of cell wall constituents.     

Ecology and behavior of the wild proboscis monkey,Nasalis larvatus (Wurmb), in Sabah,Malaysia

Proboscis monkeys,Nasalis larvatus (Wurmb), were investigated in the Padas Bay and Kinabatangan River areas, Sabah, from July 1968 to March 1969. The proboscis monkey is a common, widely distributed species of coloboids in Sabah, living in the various vegetations that extend from the coastal swamp to the inland plain. It appears to have a terrestrial tendency, compared with other coloboids, moving around on the ground in the forest and extending its activity, sometimes, to the open land along the river beach. The number of monkey in a troop, we observed, ranged from 11 to 32, and almost all troops were composed of several adult males, many more adult females, and immature animals. The adult males appear to play the role of leaders in the troop through their characteristic behavior, attacking, threatening, warning, watching, etc., other troop members or invaders. Each troop always emitted various boisterous vocalizations which were divided into 14 types.     

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.     

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.     

Population structure and social organization of Southern Elephant Seals, Mirounga leonina (L.)

The population structure and social organization of the Southern Elephant Seal, Mirounga leonina , were studied at South Georgia principally by extensive field census work and determination of age and reproductive history from sections of teeth taken from samples of bulls and cows.
The adult males of the South Georgia population were exploited from 1910 to 1964, mainly at the maximum sustainable yield for this population.
The present data are compared with similar information obtained from studies at South Georgia in 1951 during the exploitation phase and at Macquarie Island in the 1950's where sealing ended in 1919 and the population had stabilized.
Changes have been noted in the time of bull haul out, number of bulls ashore, cow: bull ratio, harem size and the age of harem bulls. These changes can all be attributed to the ending of exploitation. In contrast, the structure of the cow herd has not changed appreciably in the same period.
In addition, differences in growth, body size and population structure still persist between the South Georgia and Macquarie Island populations and it is likely that most of them may reflect differences in food availability at the two locations.     

Habitat use,ranging behaviour,and food habits of the proboscis monkey,Nasalis larvatus (van Wurmb), in Sarawak

Studies on the ecology of the proboscis monkey were conducted in seven areas of Sarawak during 1980–1981. Groups were found in a variety of riparian and coastal habitats but where high forests occurred they appeared to be preferred over other adjacent types. Undisturbed tidal forests were widely used but were preferred only where high forest was lacking or limited in extent. Clear-felled tidal forests and areas around human settlements were completely avoided, although some use was made of selectively felled tidal forests, remnant tidal forests adjacent to agricultural land, and selectively logged high forest. Monkeys slept primarily and perhaps exclusively along river edges where these were available, moving inland up to 750m during the day and returning to the riverside in late afternoon. Groups often split up into distinct subunits during daily movements. Feeding occurred throughout the day and in all habitats occupied. Proboscis monkeys were observed feeding on at least 90 plant species, including leaves or shoots of 75 species, fruits of 15 species, seeds of 10 species, and flowers of 4 species. Certain tidal forest species (Sonneratia alba, Avicennia alba, Bruguiera gymnorrhiza, Rhizophora spp.) appeared to be the single most important food plants, although where high forests were available more feeding time was spent in these types than in tidal forests or other habitats.     

Habitat preference reflects social organization of humpback whales (Megaptera novaeangliae) on a wintering ground

Habitat preference is driven by a complex interaction among behavioural patterns, biological requirements, and environmental conditions. These variables are difficult to determine for any species but are further complicated for migratory marine mammals, such as humpback whales Megaptera novaeangliae . Patterns of habitat use in relation to social organization potentially exist for this species on their wintering grounds. Using an integrated GIS approach, we examined the degree to which spatial patterns of habitat stratification are correlated within different humpback whale group types from 6 years of sighting data (1996–2001) collected on the Antongil Bay, Madagascar, wintering ground. Stratification of humpback whale sightings by behavioural classification showed significant variation in depth and distance from shore. Distribution by depth could not be described as a function of group size but could be described as a function of social organization, with mother–calf pairs showing a strong preference for shallower water compared to all other group types. Group size and social organization seem to be factors in distribution by distance from shore. Significant diurnal patterns in distribution by depth and distance from shore also exist, where mother–calf groups maintain a relatively stable distribution and pairs and competitive groups are the most variable. Patterns of habitat preference on this wintering ground appear to be guided by social organization, where distribution by depth and distance from shore highlight areas critical to conservation.     

A new interpretation of the social organization and mating system of the Callitrichidae

Tamarins and marmosets are usually described as monogamous, or as living in extended family groups. Field research on a variety of callitrichid species shows, however, that immigration and emigration by adults are common and that groups are likely to be composed largely of unrelated adults of both sexes. Despite the number of adult females present in a group, only a single female is usually reproductively active. This female may mate with more than one male. Care for the young is provided by all group members but principally by adult males. This generally is referred to as a “communal breeding system. ” However, a vertebrate social system in which more than two reproductively active adults participate in infant care is rare and has otherwise been reported only in a small number of avian and canid species. Wild callitrichid groups virtually always include helpers, but the number of adults of each sex is highly variable. We propose that the Callitrichidae tend to live in small multimalemultifemale groups, communally rear the young of one female, and exhibit a mating pattern characterized by female promiscuity but a breeding system that is functionally polyandrous. There is a significant correlation between the number of males and a group and the total number of surviving young. The possible adaptive advantages of this social organization are discussed. An erratum to this article is available at .     

Observations on the fetus and placenta of a proboscis monkey (Nasalis Iarvatus)

The villous bidiscoid placenta in utero of a proboscis monkey with a fetus of 9.0 cm crown rump length was examined. The immature male fetus weighed 56 g and had a 5.4 cm head circumference. Of particular interest was the stomach, consisting of three parts, large fundus, a narrow tubular portion, and a small pyloric region. The fetal adrenal glands were remarkable in that they were larger than the kidneys. Thus, the adrenal/kidney weight ratio approximated 1.18. The marginally inserted umbilical cord measured 15 cm. Owing to the superficial implantation in the proboscis monkey, penetration of syncytiotrophoblast and a fibrin layer were not observed in the basal plate of the placenta; cytotrophoblastic cell columns attached to the basal plate, consisting of decidua basalis without invasion of trophoblastic giant cells. Intraarterial trophoblastic migration was found in the uterus.     

Genetic diversity and distinctiveness of the proboscis monkeys (Nasalis larvatus) of the Klias Peninsula, Sabah, Malaysia

In this study, we sequenced a partial segment of the mitochondrial control region from 21 proboscis monkeys of the Klias peninsula, the last large population remaining on the west coast of Sabah, Malaysia. Our results showed that this population retains substantial genetic variation, and subpopulations from different river systems in the central and southern portions of the Klias share multiple haplotypes. We also compared our data with previously generated sequences from 2 eastern populations of proboscis monkeys in Sabah and found little evidence of regional genetic structure. Based on these results, we argue that conservation efforts should focus on restoring connectivity between central and southern Klias peninsula proboscis monkeys and discuss future analyses needed to better understand the mitochondrial structure of proboscis monkeys in Sabah.     

Spatial organization in a dimorphic ant: caste specificity of clustering patterns and area marking

Living in groups constitutes the root of social organizationin animals. Likewise, the spatial aggregation between membersof insects societies plays a crucial role in social cohesionand division of labor, namely, in polymorphic ant species. Inthe present paper, we show caste-specific aggregation patternsin the strictly dimorphic Pheidole pallidula ant species. Weinvestigate the influence on the clustering of ants exertedby direct contacts between nest mates as well as by indirectcues through chemical marking. In a homogeneous environmentdeprived of chemical cues, majors show a higher aggregationlevel than minors and a centripetal behavior. By contrast, minorsare more scattered in the experimental arena and display a centrifugalbehavior. In addition, area marking laid by minors enhancestheir own aggregative behavior while contributing to the localizationof the spontaneously aggregating majors. Such differences inaggregative patterns as well as their adaptive value have tobe coupled with the mobility level and the task performanceefficiency of each worker caste. Contrary to majors that arelikely to aggregate, highly mobile minors, scattered insideand outside the nest colony, can detect colony needs and cancarry out most of the daily tasks for which they are more efficientthan majors.     

Female transfer between one-male groups of proboscis monkey (<Emphasis Type="Italic">Nasalis larvatus</Emphasis>)

Successful or unsuccessful female transfers were observed seven times during a 32-month field study of proboscis monkeys (Nasalis larvatus) inhabiting a riverine forest along a tributary of the Kinabatangan River, Sabah, Malaysia. In all cases, the females voluntarily left their own groups and immediately joined with another one. When adult females tried to shift to other groups, adult males called them back to their own groups, but appeared to be indifferent to subadult females. When the adult females returned, the males never attacked the females physically, but instead often emitted herding sounds to them. One subadult female was repelled by a resident adult female. When one adult female transferred into a new one-male group, she left her behind son in an all-male group. The number of females often fluctuated in most study groups, with this fluctuation being more prominent among subadult females than adult females. It is likely that female transfer in proboscis monkeys is not a rare occurrence and that it is especially common among sub-adult females.