Orangutan survival is threatened by habitat loss and illegal killing. Most wild populations will disappear over the next few decades unless threats are abated. Saving orangutans is ultimately in the hands of the governments and people of Indonesia and Malaysia, which need to ensure that habitats of viable orangutan populations are protected from deforestation and well managed to ensure no hunting takes place. Companies working in orangutan habitat also have to play a much bigger role in habitat management. Although the major problems and the direct actions required to solve them-reducing forest loss and hunting-have been known for decades, orangutan populations continue to decline. Orangutan populations in Sumatra and Borneo have declined by between 2,280 and 5,250 orangutans annually over the past 25 years. As the total current population for the two species is some 60,000 animals in an area of about 90,000 km(2) , there is not much time left to make conservation efforts truly effective. Our review discusses what has and has not worked in conservation to guide future conservation efforts.
Today the majority of wild great ape populations are found outside of the network of protected areas in both Africa and Asia, therefore determining if these populations are able to survive in forests that are exploited for timber or other extractive uses and how this is managed, is paramount for their conservation.
Nine isolated fossil Pongo teeth from two cave sites in Peninsular Malaysia are reported. These are the first fossil Pongo specimens recorded in Peninsular Malaysia and represent significant southward extensions of the ancient Southeast Asian continental range of fossil Pongo during two key periods of the Quaternary. These new records from Peninsular Malaysia show that ancestral Pongo successfully passed the major biogeographical divide between mainland continental Southeast Asia and the Sunda subregion before 500 ka (thousand years ago). If the presence of Pongo remains in fossil assemblages indicates prevailing forest habitat, then the persistence of Pongo at Batu Caves until 60 ka implies that during the Last Glacial Phase sufficient forest cover persisted in the west coast plain of what is now Peninsular Malaysia at least ten millennia after a presumed corridor of desiccation had extended to central and east Java. Ultimately, environmental conditions of the peninsula during the Last Glacial Maximum evidently became inhospitable for Pongo, causing local extinction. Following post-glacial climatic amelioration and reforestation, a renewed sea barrier prevented re-colonization from the rainforest refugium in Sumatra, accounting for the present day absence of Pongo in apparently hospitable lowland evergreen rainforest of Peninsular Malaysia. The new teeth provide further evidence that Pongo did not undergo a consistent trend toward dental size reduction over time.
Orangutans are believed to have close biological affinities to humans. Teeth being the hardest tissue provide useful information on primate evolution. Furthermore, knowledge of the pulp chamber and root canal morphology is important for dental treatment. A female Bornean orangutan and a Sumatran male orangutan skull were available for this study. Both of their dentitions, comprising 50 teeth, were scanned employing the cone-beam computed tomography for both metrical and nonmetrical analyses. Measurements included tooth and crown length, root length, enamel covered crown height, root canal length (posterior teeth), length of pulpal space (anterior teeth), and root canal width. Nonmetrical parameters included number of canals per root, number of foramina in each root, and root canal morphology according to Vertucci's classification. It was found that the enamel covered crown height was the longest in the upper central incisors although the canine was the longest amongst the anterior teeth. Both the upper premolars were three-rooted while the lower second premolar of the Sumatran orangutan was two-rooted, with two foramina. The mandibular lateral incisors of the Bornean orangutan were longer than the central incisors, a feature similar to humans. In addition, secondary dentine deposition was noticed, a feature consistent with aged humans.
Six extant species of non-human great apes are currently recognized: Sumatran and Bornean orangutans, eastern and western gorillas, and chimpanzees and bonobos . However, large gaps remain in our knowledge of fine-scale variation in hominoid morphology, behavior, and genetics, and aspects of great ape taxonomy remain in flux. This is particularly true for orangutans (genus: Pongo), the only Asian great apes and phylogenetically our most distant relatives among extant hominids . Designation of Bornean and Sumatran orangutans, P. pygmaeus (Linnaeus 1760) and P. abelii (Lesson 1827), as distinct species occurred in 2001 [1, 2]. Here, we show that an isolated population from Batang Toru, at the southernmost range limit of extant Sumatran orangutans south of Lake Toba, is distinct from other northern Sumatran and Bornean populations. By comparing cranio-mandibular and dental characters of an orangutan killed in a human-animal conflict to those of 33 adult male orangutans of a similar developmental stage, we found consistent differences between the Batang Toru individual and other extant Ponginae. Our analyses of 37 orangutan genomes provided a second line of evidence. Model-based approaches revealed that the deepest split in the evolutionary history of extant orangutans occurred ∼3.38 mya between the Batang Toru population and those to the north of Lake Toba, whereas both currently recognized species separated much later, about 674 kya. Our combined analyses support a new classification of orangutans into three extant species. The new species, Pongo tapanuliensis, encompasses the Batang Toru population, of which fewer than 800 individuals survive. VIDEO ABSTRACT.
Habitat loss and hunting are major threats to the long-term survival of the viable orangutan population in Batang Toru. East Batang Toru Forest Block (EBTFB) is the most threatened area due to low forest cover and high encroachment. Based on a preliminary survey in 2008, Hopong forest which is located in EBTFB, had the highest orangutan density (0.7 ind/km2). However illegal logging and hunting of protected species were occuring in this unprotected forest. Since this location has been gazetted as unprotected forest from the first survey until this study was conducted, it is important to assess orangutans population trends. This study aims to provide an updated density of orangutan in Hopong forest. The study included the location of the original survey but covered a wider overall area. The line transect method was used to record orangutan nests, ficus and trees bearing fruits. A quadrat method was used to record vegetation. The encounter rate of orangutan declined from 0.7 ind/km2 to 0.4 ind/km2 between 2008 and 2015. Forest cover has also changed in the seven years between surveys and this has influenced orangutan and orangutan nest encounter rates in Hopong. Since unprotected forest is at more risk in comparison with protected forest, allocation status of the Hopong forest is critical to reduce the threats it faces.
Behavioural observations suggest that orang-utans are semi-solitary animals with females being philopatric and males roaming more widely in search of receptive partners, leading to the prediction that females are more closely related than males at any given site. In contrast, our study presents evidence for male and female philopatry in the orang-utan. We examined patterns of relatedness and parentage in a wild orang-utan population in Borneo using noninvasively collected DNA samples from animals observed to defecate, and microsatellite markers to assess dispersal and mating strategies. Surprisingly, resident females were equally as related to other resident females (mean r(xy) = 0.303) as resident males were to other resident males (mean r(xy) = 0.305). Moreover, resident females were more related to each other and to the resident males than they were to nonresident females, and resident males were more related to each other (and resident females) than they were to nonresident males. We assigned genetic mothers to 12 individuals in the population, while sires could be identified for eight. Both flanged males and unflanged males achieved paternity, similar to findings reported for Sumatran orang-utans.
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.
The conservation of charismatic and functionally important large species is becoming increasingly difficult. Anthropogenic pressures continue to squeeze available habitat and force animals into degraded and disturbed areas. Ensuring the long-term survival of these species requires a well-developed understanding of how animals use these new landscapes to inform conservation and habitat restoration efforts. We combined 3 y of highly detailed visual observations of Bornean orangutans with high-resolution airborne remote sensing (Light Detection and Ranging) to understand orangutan movement in disturbed and fragmented forests of Malaysian Borneo. Structural attributes of the upper forest canopy were the dominant determinant of orangutan movement among all age and sex classes, with orangutans more likely to move in directions of increased canopy closure, tall trees, and uniform height, as well as avoiding canopy gaps and moving toward emergent crowns. In contrast, canopy vertical complexity (canopy layering and shape) did not affect movement. Our results suggest that although orangutans do make use of disturbed forest, they select certain canopy attributes within these forests, indicating that not all disturbed or degraded forest is of equal value for the long-term sustainability of orangutan populations. Although the value of disturbed habitats needs to be recognized in conservation plans for wide-ranging, large-bodied species, minimal ecological requirements within these habitats also need to be understood and considered if long-term population viability is to be realized.
Conservation benefits from understanding how adaptability and threat interact to determine a taxon's vulnerability. Recognizing how interactions with humans have shaped taxa such as the critically endangered orangutan (Pongo spp.) offers insights into this relationship. Orangutans are viewed as icons of wild nature, and most efforts to prevent their extinction have focused on protecting minimally disturbed habitat, with limited success. We synthesize fossil, archeological, genetic, and behavioral evidence to demonstrate that at least 70,000 years of human influence have shaped orangutan distribution, abundance, and ecology and will likely continue to do so in the future. Our findings indicate that orangutans are vulnerable to hunting but appear flexible in response to some other human activities. This highlights the need for a multifaceted, landscape-level approach to orangutan conservation that leverages sound policy and cooperation among government, private sector, and community stakeholders to prevent hunting, mitigate human-orangutan conflict, and preserve and reconnect remaining natural forests. Broad cooperation can be encouraged through incentives and strategies that focus on the common interests and concerns of different stakeholders. Orangutans provide an illustrative example of how acknowledging the long and pervasive influence of humans can improve strategies to preserve biodiversity in the Anthropocene.
The integration of Bayesian analysis into existing great ape survey methods could be used to generate precise and reliable population estimates of Bornean orang-utans. We used the Marked Nest Count (MNC) method to count new orang-utan nests at seven previously undocumented study sites in Sarawak, Malaysia. Our survey teams marked new nests on the first survey and revisited the plots on two more occasions; after about 21 and 42 days respectively. We used the N-mixture models to integrate suitability, abundance and detection models which account for zero inflation and imperfect detection for the analysis. The result was a combined estimate of 355 orang-utans with the 95% highest density interval (HDI) of 135 to 602 individuals. We visually inspected the posterior distributions of our parameters and compared precisions between study sites. We subsequently assess the strength or reliability of the generated estimates using identifiability tests. Only three out of the seven estimates had <35% overlap to indicate strong reliability. We discussed the limitations and advantages of our study design, and made recommendations to improve the sampling scheme. Over the course of this research, two of the study sites were gazetted as extensions to the Lanjak-Entimau Wildlife Sanctuary for orang-utan conservation.
A recent report, published by the Government of Indonesia with support from the Food and Agricultural Organization and Norway's International Climate and Forest Initiative, states that orangutan populations (Pongo spp.) have increased by more than 10% in Indonesia from 2015 to 2017, exceeding the government target of an annual 2% population increase . This assessment is in strong contrast with recent publications that showed that the Bornean orangutan (P. pygmaeus) lost more than 100,000 individuals in the past 16 years  and declined by at least 25% over the past 10 years . Furthermore, recent work has also demonstrated that both Sumatran orangutans (P. abelii) and the recently described Tapanuli orangutan (P. tapanuliensis) lost more than 60% of their key habitats between 1985 and 2007, and ongoing land use changes are expected to result in an 11-27% decline in their populations by 2020 [4,5]. Most scientific data indicate that the survival of these species continues to be seriously threatened by deforestation and killing [4,6,7] and thus all three are Critically Endangered under the International Union for Conservation of Nature's Red List.
We examined mitochondrial DNA control region sequences of 73 Kinabatangan orangutans to test the hypothesis that the phylogeographical structure of the Bornean orangutan is influenced by riverine barriers. The Lower Kinabatangan Wildlife Sanctuary contains one of the most northern populations of orangutans (Pongo pygmaeus) on Borneo and is bisected by the Kinabatangan River, the longest river in Sabah. Orang-utan samples on either side of the river were strongly differentiated with a high Phi(ST) value of 0.404 (P < 0.001). Results also suggest an east-west gradient of genetic diversity and evidence for population expansion along the river, possibly reflecting a postglacial colonization of the Kinabatangan floodplain. We compared our data with previously published sequences of Bornean orangutans in the context of river catchment structure on the island and evaluated the general relevance of rivers as barriers to gene flow in this long-lived, solitary arboreal ape.
The reproductive success of male primates is not always associated with dominance status. For example, even though male orangutans exhibit intra-sexual dimorphism and clear dominance relationships exist among males, previous studies have reported that both morphs are able to sire offspring. The present study aimed to compare the reproductive success of two male morphs, and to determine whether unflanged males sired offspring in a free-ranging population of Bornean orangutans, using 12 microsatellite loci to determine the paternity of eight infants. A single flanged male sired most of the offspring from parous females, and an unflanged male sired a firstborn. This is consistent with our observation that the dominant flanged male showed little interest in nulliparous females, whereas the unflanged males frequently mated with them. This suggests that the dominant flanged male monopolizes the fertilization of parous females and that unflanged males take advantage of any mating opportunities that arise in the absence of the flanged male, even though the conception probability of nulliparous females is relatively low.
The Sumatran orangutan is currently listed by the IUCN as critically endangered and the Bornean species as endangered. Unless effective conservation measures are enacted quickly, most orangutan populations without adequate protection face a dire future. Two main strategies are being pursued to conserve orangutans: (i) rehabilitation and reintroduction of ex-captive or displaced individuals; and (ii) protection of their forest habitat to abate threats like deforestation and hunting. These strategies are often mirrored in similar programs to save other valued and endangered mega-fauna. Through GIS analysis, collating data from across the literature, and combining this information within a modelling and decision analysis framework, we analysed which strategy or combination of strategies is the most cost-effective at maintaining wild orangutan populations, and under what conditions. We discovered that neither strategy was optimal under all circumstances but was dependent on the relative cost per orangutan, the timescale of management concern, and the rate of deforestation. Reintroduction, which costs twelve times as much per animal as compared to protection of forest, was only a cost-effective strategy at very short timescales. For time scales longer than 10-20 years, forest protection is the more cost-efficient strategy for maintaining wild orangutan populations. Our analyses showed that a third, rarely utilised strategy is intermediate: introducing sustainable logging practices and protection from hunting in timber production forest. Maximum long-term cost-efficiency is achieved by working in conservation forest. However, habitat protection involves addressing complex conservation issues and conflicting needs at the landscape level. We find a potential resolution in that well-managed production forests could achieve intermediate conservation outcomes. This has broad implications for sustaining biodiversity more generally within an economically productive landscape. Insights from this analysis should provide a better framework to prioritize financial investments, and facilitate improved integration between the organizations that implement these strategies.
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 analysed the reproductive parameters of free-ranging female orangutans at Sepilok Orangutan Rehabilitation Centre (SORC) on Borneo Island, Sabah, Malaysia. Fourteen adult females produced 28 offspring in total between 1967 and 2004. The average censored interbirth interval (IBI) (i.e. offspring was still alive when mother produced a next offspring) was 6 years. This was shorter than censored IBIs reported in the wild but similar to IBIs reported for those in captivity. The nonparametric survival analysis (Kaplan-Meier method) revealed a significantly shorter IBI at SORC compared with wild orangutans in Tanjung Putting. The infant (0-3 years) mortality rate at SORC of 57% was much higher than rates reported both in the wild and captivity. The birth sex-ratio was significantly biassed toward females: 24 of the 27 sex-identified infants were females. The average age at first reproduction was 11.6 years, which is younger than the age in the wild and in captivity. The high infant mortality rate might be caused by human rearing and increased transmission of disease due to frequent proximal encounters with conspecifics around the feeding platforms (FPs). This young age of first reproduction could be because of the uncertainty regarding estimated ages of the female orangutans at SORC. It may also be affected by association with other conspecifics around FPs, which increased the number of encounters of the females with males compared with the number of encounters that would take place in the wild. Provision of FPs, which improves the nutritional condition of the females, caused the shorter IBI. The female-biassed birth sex-ratio can be explained by the Trivers and Willard hypothesis. The female-biassed sex ratio could be caused by the mothers being in poor health, parasite prevalence and/or high social stress (but not food scarcity) due to the frequent encounters with conspecifics around FPs.