Reports of wild great ape fatalities have been very limited, and only two have described wild orangutan deaths. We found a wounded juvenile female Bornean orangutan on 7 October 2006 in the Danum Valley, Sabah, Malaysia, and observed the individual's behavior for 7 days until her death on 13 October 2006. The 5-6-year-old orangutan, which we had observed since 2004, was wounded in the left brachium, back, and right hand. The individual's behavior changed after injury; the mean nest-nest active time became significantly shorter than before injury (from 12 h 3 min to 9 h 33 min), the mean waking time became significantly later (0552-0629 hours) and the mean bedtime became significantly earlier (from 1747 to 1603 hours). In the activity budget, resting increased significantly from 28.0 to 53.3%. Traveling and feeding decreased significantly from 23.5 to 12.7% and from 45.6 to 32.8%, respectively. The rate of brachiation during traveling and nest making decreased, whereas ground activity increased from 0 to 9%. We observed one vomiting incident and four occurrences of watery diarrhea during the 7 days before the individual died. The results of an autopsy performed by a local veterinarian suggested that the cause of death was septicemia because of Pseudomonas aeruginosa infection of the severely contaminated wounds. The morphology and distribution of the wounds suggested they had been incurred during an attack by a large animal with fangs and/or claws. This juvenile female became independent of its mother at ~4-5 years of age, slightly earlier than average. This individual might have been vulnerable to predatory attack because of her small body size (~5 kg at death) and lack of the mother's protection.
We observed the diet and activity of Bornean orangutans (Pongo pygmaeus morio) in the primary lowland dipterocarp forests of Danum Valley, Sabah, Malaysia, during 2005-2007, including two mast fruitings. We collected 1,785 hr of focal data on 26 orangutans. We identified 1,466 samples of their food plants and conducted a fallen fruit census to monitor fruit availability in the study area. Their activity budget was 47.2% feeding, 34.4% resting, and 16.9% traveling. Fruits accounted for the largest part (60.9%) of feeding time, especially during mast fruiting periods (64.0-100%), although the percentages of leaves (22.2%) and bark (12.3%) were higher than those reported for P. abelii and P. pygmaeus wurmbii. Although 119 genera and 160 plant species were consumed by focal animals, only 9 genera accounted for more than 3% of feeding time (total: 67.8% for 9 genera). In particular, the focal orangutans fed intensively on Ficus and Spatholobus during most of the study period, especially in periods of fruit shortage. The percentage of fruit feeding changed markedly from 11.7 to 100% across different months of the year, and was positively correlated with the amount of fallen fruit. When fruit feeding and availability decreased, orangutans fed primarily on leaves of Spatholobus and Ficus, and the bark of Spatholobus and dipterocarp. The percentage of time devoted to feeding during mast fruitings, when the orangutans foraged almost exclusively on fruits, was lower than during seasons when the orangutan diet included leaves and bark as well as fruits. Resting increased as feeding decreased in the late stage of each fruiting season, suggesting that the orangutans adopted an energy-minimizing strategy to survive the periods of fruit shortage by using energy stored during the fruit season.
We investigated the population density of Bornean orangutans (Pongo pygmaeus morio) and fruit availability for 10 years (2005-2014), in primary lowland dipterocarp forests in the Danum Valley, Sabah, Malaysia. During the research period, two mast fruitings and three other peak fruiting events of different scales occurred in the study area. The orangutan population density, estimated every 2 months by the marked nest count method, changed between 0.3 and 4.4 ind/km(2) and the mean population density was 1.3 ind/km(2) ± SE 0.1 (n = 56). The population density increased markedly during mast and peak fruiting periods. A significant positive correlation was observed between the population density and fruit availability in the study period (Spearman, R = 0.3, P
Knowledge of niche partitioning with respect to habitat is indispensable to understand the mechanism of coexistence of multiple species. Among primates, however, data are still deficient because repeated survey for a sufficiently long time, covering seasonal changes over a large area, is the only way to clarify habitat segregation within a seasonally fluctuating environment. Southeast Asia is particularly interesting because of the supra-annual, highly unpredictable seasonality in fruiting known as mast fruiting. We conducted repeated route census, habitat monitoring, and group tracking for 25 months in two study sites (ca. 10 km apart) in the largely primary lowland dipterocarp forest of the Danum Valley Conservation Area, eastern Sabah, northern Borneo, Malaysia. The five species of diurnal primates (Bornean orangutan Pongo pygmaeus, Müeller's gibbon Hylobates muelleri, red leaf monkey Presbytis rubicunda, long-tailed macaque Macaca fascicularis, and southern pig-tailed macaque M. nemestrina) did not show horizontal spatial segregation. Red leaf monkeys showed preferences for places with short tree height, but their distribution was not confined to such places. In response to the fruiting peak observed once during the study period, orangutans increased their numbers simultaneously in the two study sites. The average tree height used by the five species was different, but their range overlapped substantially. Compared with other primate communities, the lack of horizontal spatial segregation and the suggested long-distance movement of orangutans seem to be unique characteristics in Borneo, although the use of different forest strata is a widespread phenomenon among primate communities throughout the world.
In order to obtain basic data on parasitic infections of Bornean orangutans, Pongo pygmaeus morio (Owen, 1837), in Danum Valley, Sabah, Malaysia, fecal examinations were conducted. Based on a total of 73 fecal samples from 25 individuals, cysts of Entamoeba coli, Entamoeba spp., and Chilomastix mesnili, cysts and trophozoites of Balantidium coli, and eggs of Trichuris sp. or spp., unknown strongylid(s), Strongyloides fuelleborni, and an unknown oxyurid, plus a rhabditoid larva of Strongyloides sp., were found. Mature and immature worms of Pongobius hugoti Baruš et al., 2007 and Pongobius foitovae n. sp. (Oxyuridae: Enterobiinae) were recovered from fecal debris and described. Pongobius foitovae is readily distinguished from P. hugoti by having a much longer esophageal corpus, a longer and distally hooked spicule in males, and a more posteriorly positioned vulva in female. Presence of plural species of non- Enterobius pinworms is a remarkable feature of the orangutan-pinworm relationship, which may reflect speciation process of the orangutans, host switching, and coevolution by pinworms.
Orangutans have a long period of immaturity and the longest inter-birth interval (IBI) of all mammals, which can be explained by their solitary life style, preventing the mother from rearing two offspring simultaneously (solitary life hypothesis) [corrected]. We collected data on mother-offspring dyads living in a primary lowland forest in Danum Valley, East Borneo in an effort to examine the developmental and behavioral patterns of the subspecies Pongo pygmaeus morio. We analyzed developmental changes in mother-offspring distance, contact, and activity budgets in orangutans ranging from 1 to 7 years of age. The results indicated decreased resting and playing with increasing age, whereas feeding, traveling and social play all increased significantly. Mothers' feeding and traveling time were good predictors of their offspring's feeding and traveling activities. Mother-offspring contact lasted longer in resting contexts; contact during traveling was almost non-existent after 4 years of age. Comparisons with previously published data on the Sumatran species Pongo abelli revealed no fundamental differences in these behavioral measures. However, a shorter association time with the mother after behavioral independence is documented for this East Bornean population in comparison to Sumatran populations. These results are best explained by the solitary life hypothesis, in agreement with previous studies. We suggest that environmental constraints in Bornean forests, as well as a lower population density, should be considered when interpreting the differences between Sumatran and Bornean orangutans in both the period of association with mother and the IBI.
To propose proper conservation measures and to elucidate coexistence mechanisms of sympatric carnivore species, we assessed temporal activity patterns of the sympatric carnivore species using 37,379 photos collected for more than 3 years at three study sites in Borneo. We categorized activity patterns of nine carnivore species (one bear, three civets, two felids, one skunk, one mustelid, one linsang) by calculating the photo-capturing proportions at each time period (day, night, twilight). We then evaluated temporal activity overlaps by calculating the overlap coefficients. We identified six nocturnal (three civets, one felid, one skunk, one linsang), two diurnal (one felid, one mustelid), and one cathemeral (bear) species. Temporal activity overlaps were high among the nocturnal species. The two felid species possessing morphological and ecological similarities exhibited clear temporal niche segregation, but the three civet species with similar morphology and ecology did not. Broad dietary breadth may compensate for the high temporal niche overlaps among the nocturnal species. Despite the high species richness of Bornean carnivores, almost half are threatened with extinction. By comparing individual radio-tracking and our data, we propose that a long-term study of at least 2 or 3 years is necessary to understand animals' temporal activity patterns, especially for sun bears and civets, by camera-trapping and to establish effective protection measures.
The orangutan is the world's largest arboreal mammal, and images of the red ape moving through the tropical forest canopy symbolise its typical arboreal behaviour. Records of terrestrial behaviour are scarce and often associated with habitat disturbance. We conducted a large-scale species-level analysis of ground-based camera-trapping data to evaluate the extent to which Bornean orangutans Pongo pygmaeus come down from the trees to travel terrestrially, and whether they are indeed forced to the ground primarily by anthropogenic forest disturbances. Although the degree of forest disturbance and canopy gap size influenced terrestriality, orangutans were recorded on the ground as frequently in heavily degraded habitats as in primary forests. Furthermore, all age-sex classes were recorded on the ground (flanged males more often). This suggests that terrestrial locomotion is part of the Bornean orangutan's natural behavioural repertoire to a much greater extent than previously thought, and is only modified by habitat disturbance. The capacity of orangutans to come down from the trees may increase their ability to cope with at least smaller-scale forest fragmentation, and to cross moderately open spaces in mosaic landscapes, although the extent of this versatility remains to be investigated.