Phylogenetic differences in the morphology and shape of the central sulcus in great apes and humans: implications for the evolution of motor functions.

The central sulcus divides the primary motor and somatosensory cortices in many anthropoid primate brains. Differences exist in the surface area and depth of the central sulcus along the dorso-ventral plane in great apes and humans compared to other primate species. Within hominid species, there are variations in the depth and aspect of their hand motor area, or knob, within the precentral gyrus. In this study, we used post-image analyses on magnetic resonance images to characterize the central sulcus shape of humans, chimpanzees (Pan troglodytes), gorillas (Gorilla gorilla), and orangutans (Pongo pygmaeus and Pongo abelii). Using these data, we examined the morphological variability of central sulcus in hominids, focusing on the hand region, a significant change in human evolution. We show that the central sulcus shape differs between great ape species, but all show similar variations in the location of their hand knob. However, the prevalence of the knob location along the dorso-ventral plane and lateralization differs between species and the presence of a second ventral motor knob seems to be unique to humans. Humans and orangutans exhibit the most similar and complex central sulcus shapes. However, their similarities may reflect divergent evolutionary processes related to selection for different positional and habitual locomotor functions.

The role of great ape behavioral ecology in One Health: Implications for captive welfare and re-habilitation success.

Behavior is the interface through which animals interact with their environments, and therefore has potentially cascading impacts on the health of individuals, populations, their habitats, and the humans that share them. Evolution has shaped the interaction between species and their environments. Thus, alterations to the species-typical "wild-type" behavioral repertoire (and the ability of the individual to adapt flexibly which elements of the repertoire it employs) may disrupt the relationship between the organism and its environment, creating cascading One Health effects. A good example is rehabilitant orangutans where, for example, seemingly minor differences from wild conspecifics in the time spent traveling on the ground rather than in the forest canopy can affect an individual's musculoskeletal and nutritional health, as well as social integration. It can also increase two-way transmission of infectious diseases and/or pathogens with local human populations, or potentially with neighboring wild populations if there are no geographical barriers and rehabilitants travel far enough to leave their release area. Primates are well known ecosystem engineers, reshaping plant communities and maintaining biodiversity through seed dispersal, consuming plants, and creating canopy gaps and trails. From the habitat perspective, a rehabilitant orangutan which does not behave like a wild orangutan is unlikely to fulfill these same ecosystem services. Despite the importance of the diversity of an ape's behavioral repertoire, how it compares to that of wild conspecifics and how it alters in response to habitat variation, behavior is an often under-appreciated aspect of One Health. In this review, focusing on orangutans as an example of the kinds of problems faced by all captive great apes, we examine the ways in which understanding and facilitating the expression of wild-type behavior can improve their health, their ability to thrive, and the robustness of local One Health systems.

Low relationship quality predicts scratch contagion during tense situations in orangutans (Pongo pygmaeus).

Primates show various forms of behavioral contagion that are stronger between kin and friends. As a result, behavioral contagion is thought to promote group coordination, social cohesion, and possibly state matching. Aside from contagious yawning, little is known about the contagious effect of other behaviors. Scratching is commonly observed during arousal and as such may play a role within group dynamics. While the Bornean orangutan (Pongo pygmaeus) is commonly considered the least social great ape, orangutans do engage in social interactions. Therefore, their social organization makes them a suitable case for studying the social function of behavioral contagion. Through behavioral observations of captive orangutans, we recorded all yawn and scratch events together with the corresponding behavior of all bystander group-members. As yawning was rarely observed, no conclusions could be drawn regarding this behavior. Scratching was contagious and occurred within 90 s after the triggering scratch. Specifically, orangutans showed increased scratch contagion when they had seen a weakly bonded individual scratch during tense contexts. When the orangutan had not seen the triggering scratch, the contagiousness of scratching was not affected by context or relationship quality. Our results indicate that behavioral contagion is not simply higher between individuals with stronger social relationships, but that the contagiousness of behaviors may vary based on the context and on social factors. We discuss these findings in light of an adaptive function that may reduce aggression.

Sociosexual behavioral patterns involving nulliparous female orangutans (Pongo sp.) reflect unique challenges during the adolescent period.

The primate adolescent period is characterized by a series of changes in physiology, behavior, and social relationships. Orangutans have the slowest life history and the longest period of dependency of all primates. As members of a semisolitary species with high levels of sexual coercion, adolescent female orangutans face a unique combination of challenges when achieving independence from their mother. This study examined the mating behavior of adolescent female orangutans and compared it with that of adult females to assess whether mating behavior reflects distinct strategies at these different points in the life cycle. Data were collected in Gunung Palung National Park on the island of Borneo over 20 years. Mating events from adolescent (n = 19) and adult females (n = 26) were scored and compared. Adolescent female mating events had significantly higher mating scores (indicating more proceptivity) than those of adult females (ß = 1.948, p = .001). Adolescent females also engaged in elaborate sociosexual interactions with different flanged males, behaviors that were never observed during mating events of adult females. These interactions involved characteristic behavior on the part of both the adolescent females and the flanged males. Given these findings and the documentation of similar accounts of adolescent female-flanged male mating from the island of Sumatra, we propose that adolescent female orangutans display distinctive behavioral repertoires throughout the genus Pongo which serves to overcome male ambivalence toward nulliparous females, establish familiarity, and evaluate coercive tendencies in flanged males. We suggest that these behavioral patterns are an integral part of female social development in a female philopatric, but highly dispersed species where consistent social support is absent after ranging independence is achieved.

Wild Bornean orangutan (Pongo pygmaeus wurmbii) feeding rates and the Marginal Value Theorem.

The Marginal Value Theorem (MVT) is an integral supplement to Optimal Foraging Theory (OFT) as it seeks to explain an animal's decision of when to leave a patch when food is still available. MVT predicts that a forager capable of depleting a patch, in a habitat where food is patchily distributed, will leave the patch when the intake rate within it decreases to the average intake rate for the habitat. MVT relies on the critical assumption that the feeding rate in the patch will decrease over time. We tested this assumption using feeding data from a population of wild Bornean orangutans (Pongo pygmaeus wurmbii) from Gunung Palung National Park. We hypothesized that the feeding rate within orangutan food patches would decrease over time. Data included feeding bouts from continuous focal follows between 2014 and 2016. We recorded the average feeding rate over each tertile of the bout, as well as the first, midpoint, and last feeding rates collected. We did not find evidence of a decrease between first and last feeding rates (Linear Mixed Effects Model, n = 63), between a mid-point and last rate (Linear Mixed Effects Model, n = 63), between the tertiles (Linear Mixed Effects Model, n = 63), nor a decrease in feeding rate overall (Linear Mixed Effects Model, n = 146). These findings, thus, do not support the MVT assumption of decreased patch feeding rates over time in this large generalist frugivore.

The slow ape: High infant survival and long interbirth intervals in wild orangutans.

Orangutans (Pongo spp.) are reported to have extremely slow life histories, including the longest average interbirth intervals of all mammals. Such slow life history can be viable only when unavoidable mortality is kept low. Thus, orangutans' survivorship under natural conditions is expected to be extremely high. Previous estimates of orangutan life history were based on captive individuals living under very different circumstances or on small samples from wild populations. Here, we combine birth data from seven field sites, each with demographic data collection for at least 10 years (range 12-43 years) on wild orangutans to better document their life history. Using strict criteria for data inclusion, we calculated infant survival, interbirth intervals and female age at first reproduction, across species, subspecies and islands. We found an average closed interbirth interval of 7.6 years, as well as consistently very high pre-weaning survival for males and females. Female survival of 94% until age at first birth (at around age 15 years) was higher than reported for any other mammal species under natural conditions. Similarly, annual survival among parous females is very high, but longevity remains to be estimated. Current data suggest no major life history differences between Sumatran and Bornean orangutans. The high offspring survival is remarkable, noting that modern human populations seem to have reached the same level of survival only in the 20th century. The orangutans' slow life history illustrates what can be achieved if a hominoid bauplan is exposed to low unavoidable mortality. Their high survival is likely due to their arboreal and non-gregarious lifestyle, and has allowed them to maintain viable populations, despite living in low-productivity habitats. However, their slow life history also implies that orangutans are highly vulnerable to a catastrophic population crash in the face of drastic habitat change.

Stillbirth rates across three ape species in accredited American zoos.

Stillbirths, or births of infants that died in the womb, represent a failure of the materno-feto-placental unit to maintain a suitable fetal environment. Typical studies of nonhuman primate (NHP) stillbirth patterns are primarily descriptive and focus on macaques (genus Macaca). Thus, less is known about other NHP species and rarer still are studies that examine possible biological factors that influence stillbirth rates across taxa. To examine possible contributors to stillbirths in great apes, we analyzed 36 years (1980-2016) of historical data documenting births of zoo-housed chimpanzees (Pan troglodytes, N = 391), western lowland gorillas (Gorilla gorilla gorilla, N = 491), and orangutans (Pongo spp, N = 307) in accredited zoological parks in the United States. The average number of births for each of the 446 mothers was 2.7, resulting in a total of 1,189 births with 143 stillbirths (12%). Stillbirths represented 12% of chimpanzee births, 13% of gorilla births, and 10% of orangutan births. We used generalized linear mixed-effects models to assess possible relationships between stillbirth likelihood and mother origin (wild- versus captive-born), age, and genus. Across taxa, older mothers were more likely to have a stillbirth (p = 0.004). While these results are likely influenced by both biological and management-related factors (e.g., selective captive breeding), they may be useful to population managers in evaluating pregnancy risks for great apes. Captive settings and archival studbook data such as these may provide a unique opportunity to further explore this topic.

Recent origin of low trabecular bone density in modern humans.

Humans are unique, compared with our closest living relatives (chimpanzees) and early fossil hominins, in having an enlarged body size and lower limb joint surfaces in combination with a relatively gracile skeleton (i.e., lower bone mass for our body size). Some analyses have observed that in at least a few anatomical regions modern humans today appear to have relatively low trabecular density, but little is known about how that density varies throughout the human skeleton and across species or how and when the present trabecular patterns emerged over the course of human evolution. Here, we test the hypotheses that (i) recent modern humans have low trabecular density throughout the upper and lower limbs compared with other primate taxa and (ii) the reduction in trabecular density first occurred in early Homo erectus, consistent with the shift toward a modern human locomotor anatomy, or more recently in concert with diaphyseal gracilization in Holocene humans. We used peripheral quantitative CT and microtomography to measure trabecular bone of limb epiphyses (long bone articular ends) in modern humans and chimpanzees and in fossil hominins attributed to Australopithecus africanus, Paranthropus robustus/early Homo from Swartkrans, Homo neanderthalensis, and early Homo sapiens. Results show that only recent modern humans have low trabecular density throughout the limb joints. Extinct hominins, including pre-Holocene Homo sapiens, retain the high levels seen in nonhuman primates. Thus, the low trabecular density of the recent modern human skeleton evolved late in our evolutionary history, potentially resulting from increased sedentism and reliance on technological and cultural innovations.

Urinary sex steroid hormone and placental leucine aminopeptidase concentration differences between live births and stillbirth of Bornean orangutans (Pongo pygmaeus).

BACKGROUND: Under the environment of pregnancy, the placenta assumes an important steroidogenic role in the maintenance of pregnancy. METHODS: Urinary placental leucine aminopeptidase (PLAP), estrone-3-glucuronide (E1 G), and pregnanediol-3-glucuronide (PdG) concentrations were compared among five pregnancies (four live births and one stillbirth) in four orangutans. RESULTS: The gestation period of the stillbirth (223 days) was shorter than that of the live births (239-254 days). In females who gave a live birth, average PLAP and E1 G concentrations increased until the delivery. Conversely, in the female who gave a stillbirth, PLAP concentration failed to increase, and E1 G concentration was significantly low in late pregnancy period. Regarding PdG concentrations, there was no significant difference among all pregnancies. CONCLUSIONS: This is the first study reporting a change in urinary PLAP, E1 G, and PdG concentrations during orangutan stillbirth and live birth pregnancies. The findings will assist in developing pregnancy screening tests.

Bornean orangutan geophagy: analysis of ingested and control soils.

Geophagy among orangutans is the most poorly documented in contrast to the knowledge of soil-eating practices of other great ape species. Observations of soil consumption by orangutans in the Sungai Wain Forest Preserve (Wanariset) of Borneo are presented, along with physico-mineral-chemical analyses of the ingested soil in an effort to understand what might stimulate the activity. The consumed soils are: light colored, not excessively weathered by normal standards, higher in the clay size fraction relative to controls, and are comprised of a mix of clay minerals without any specificity of 1:1, 2:1 and/or 2:1:1 (Si:Al) species. The geophagic soils contain chlorides below detection limits, effectively eliminating salt as a stimulus. Soil chemical and geochemical analyses confirm that orangutans prefer soils with pH levels near or above 4.0, while controls are consistently lower (pH = 3.5-4.0), a considerable difference in acidity for at least four out of six soils consumed. Geochemical analysis shows Al, Fe and K are high in the consumed vs control samples; higher Al follows from higher clay percentages in the consumed earth. Iron and K may play physiological roles, but Fe is mostly in the ferrous form (Fe(+2)) and may not be readily taken up by the animals. The preferential choice of consumed samples, with pH above 4.0 and higher clay contents, may promote a more beneficial intestinal environment.

Acoustic Characteristics of Long Calls Produced by Male Orang-Utans (Pongo pygmaeus wurmbii): Advertising Individual Identity, Context, and Travel Direction.

Acoustic characteristics and context of the long-distance call of male orang-utans (Pongo pygmaeus wurmbii) were examined in a population of orang-utans from Central Kalimantan, Borneo. Male orang-utans produced long calls under different circumstances, including calls made spontaneously, in response to conspecifics, when accompanied by a snag crash and when travelling with a female. It was shown by acoustic analyses that there was individual discrimination between the male's calls, discrimination between the calls made under different contexts, and between calls from one individual from different years, which coincided with a change in his dominance status. We also confirm that flanged male orang-utans advertise their intended travel route, by long calling in the direction of their travel. If other orang-utans (males and females) within ear shot of the caller can identify the caller from their long call, and even obtain information about the context and status of the individual, they can then therefore choose whether to approach or avoid them. Thus, males seem to be using their long call to announce their presence, allowing them to orient themselves spatially to other orang-utans, and, potentially, to co-ordinate a network of loose associations between both males and females in the area.

Acoustic models of orangutan hand-assisted alarm calls.

Orangutans produce alarm calls called kiss-squeaks, which they sometimes modify by putting a hand in front of their mouth. Through theoretical models and observational evidence, we show that using the hand when making a kiss-squeak alters the acoustics of the production in such a way that more formants per kilohertz are produced. Our theoretical models suggest that cylindrical wave propagation is created with the use of the hand and face as they act as a cylindrical extension of the lips. The use of cylindrical wave propagation in animal calls appears to be extremely rare, but is an effective way to lengthen the acoustic system; it causes the number of resonances per kilohertz to increase. This increase is associated with larger animals, and thus using the hand in kiss-squeak production may be effective in exaggerating the size of the producer. Using the hand appears to be a culturally learned behavior, and therefore orangutans may be able to associate the acoustic effect of using the hand with potentially more effective deterrence of predators.

Body mass of wild Bornean orangutans living in human-dominated landscapes: Implications for understanding their ecology and conservation.

Body mass is a key determinant of a species' ecology, including locomotion, foraging strategies, and energetics. Accurate information on the body mass of wild primates allows us to develop explanatory models for relationships among body size, ecology, and behavior and is crucial for reconstructing the ecology and behavior of fossil primates and hominins. Information on body mass can also provide indirect information on health and can be an important tool for conservation in the context of increasingly widespread habitat disturbance. This study reports body mass data recorded for wild Northeast Bornean orangutans (Pongo pygmaeus morio) during relocation efforts in forestry and oil palm plantations in East Kalimantan, Indonesia. The average mass of flanged adult males (n = 12, 74 ± 9.78 kg) and adult females (n = 7, 35.29 ± 7.32 kg) from this study were 13.6% and 9% lower, respectively, than the only other published wild Bornean orangutan body mass measurements, but the range of weights for both males and females was larger for this study. This pattern could be due to sampling error, data collection differences, or the influence of habitat disturbance, specifically a lack of access to resources, on individual health. When necessary relocations present the opportunity, we encourage researchers to prioritize the collection of body size data for the purposes of understanding ecology but also as an indirect means of monitoring population viability. As primate habitat becomes increasingly fragmented and altered by humans such data will become critical to our ability to make informed conservation decisions.

Why do orangutans leave the trees? Terrestrial behavior among wild Bornean orangutans (Pongo pygmaeus wurmbii) at Tuanan, Central Kalimantan.

Orangutans (genus Pongo) are the largest arboreal mammals, but Bornean orangutans (P. pygmaeus spp.) also spend time on the ground. Here, we investigate ground use among orangutans using 32,000 hr of direct focal animal observations from a well-habituated wild population of Bornean orangutans (P. p. wurmbii) living in a closed-canopy swamp forest at Tuanan, Central Kalimantan, Indonesia. Ground use did not change with increasing observation time of well-habituated individuals, suggesting it was not an artifact of observer presence. Flanged males spent the most time on the ground (ca. 5% of active time), weaned immatures the least (around 1%). Females and immatures descended mainly to feed, especially on termites, whereas flanged males traveled more while on the ground. Flanged males may travel more inconspicuously, and perhaps also faster, when moving on the ground. In addition, orangutans engaged in ground-specific behavior, including drinking from and bathing in swamp pools. Supplementary records from 20 ground-level camera traps, totaling 3986 trap days, confirmed the observed age-sex biases in ground use at Tuanan. We conclude that ground use is a natural part of the Bornean orangutan behavioral repertoire, however it remains unclear to what extent food scarcity and canopy structure explain population differences in ground use.

Validation of an acoustic location system to monitor Bornean orangutan (Pongo pygmaeus wurmbii) long calls.

The long call is an important vocal communication signal in the widely dispersed, semi-solitary orangutan. Long calls affect individuals' ranging behavior and mediate social relationships and regulate encounters between dispersed individuals in a dense rainforest. The aim of this study was to test the utility of an Acoustic Location System (ALS) for recording and triangulating the loud calls of free-living primates. We developed and validated a data extraction protocol for an ALS used to record wild orangutan males' long calls at the Tuanan field site (Central Kalimantan). We installed an ALS in a grid of 300 ha, containing 20 SM2+ recorders placed in a regular lattice at 500 m intervals, to monitor the distribution of calling males in the area. The validated system had the following main features: (i) a user-trained software algorithm (Song Scope) that reliably recognized orangutan long calls from sound files at distances up to 700 m from the nearest recorder, resulting in a total area of approximately 900 ha that could be monitored continuously; (ii) acoustic location of calling males up to 200 m outside the microphone grid, which meant that within an area of approximately 450 ha, call locations could be calculated through triangulation. The mean accuracy was 58 m, an error that is modest relative to orangutan mobility and average inter-individual distances. We conclude that an ALS is a highly effective method for detecting long-distance calls of wild primates and triangulating their position. In combination with conventional individual focal follow data, an ALS can greatly improve our knowledge of orangutans' social organization, and is readily adaptable for studying other highly vocal animals.

Orangutan behavior in Kutai National Park after drought and fire damage: Adjustments to short- and long-term natural forest regeneration.

This study aimed to develop a long-term picture of orangutan (Pongo pygmaeus morio) behavioral adjustments to damaged masting forest around Mentoko, Kutai National Park, Indonesia. Mentoko is regenerating from two severe burnings and is one of few areas where orangutans were well-studied before and early after damage. We studied orangutans' feeding ecology, diet, and activity budgets 12-15 years after the second burning then compared our findings with earlier pre- and post-damage ones to assess the changes and factors involved. By our study, we predicted (1) improved feeding ecology compared to early in regeneration, (2) behavior diverging from the normal foraging strategy and (3) behavior shifting toward pre-damage patterns with improving feeding ecology. Data were behavioral observations on 42 orangutans (422 full day follows, 3,522 hr) and tree plot measures of feeding ecology. Findings were consistent with the first and third predictions but not the second: (1) feeding ecology had improved (plant food abundance was near per-damage levels, but species composition had changed); (2) foraging strategies showed no divergence from normal (fallback-preferred food switches in diet and activity budget adjustments were both normal, notably travel did not reduce), (3) diet and activity budgets had reverted to near pre-damage values by our study. Differently than post-damage studies on other orangutans but consistent with those on other primates, our comparisons showed behavioral adjustments were flexible, multiple vs. single, and influenced by multiple factors. Factors likely involved at Mentoko include type and spatial configuration of damage, duration of regeneration, and P. p. morio's recognized resilience. Findings have value for orangutan and nature conservation in showing that recovery to near-normal levels from severe habitat damage is possible within ca 15 years and in adding to understanding of the factors and processes that contribute to recovery.

Endocrinological correlates of male bimaturism in wild Bornean orangutans.

Among primates, orangutans are unique in having pronounced male bimaturism leading to two fully adult morphs that differ in both physical appearance and behavior. While unflanged males have a female-like appearance, flanged males have the full suite of secondary sexual characteristics, including cheek flanges and a large throat sac. So far, hormonal correlates of arrested development in unflanged males and the expression of secondary sexual characteristics in flanged males have only been studied in zoo-housed individuals. In this study, we investigated fecal androgen and glucocorticoid metabolites as hormonal correlates of male bimaturism in 17 wild adult Bornean orangutans (Pongo pygmaeus) in Central Kalimantan, Indonesia. We predicted and found higher androgen levels in flanged males compared to unflanged males, probably due to ongoing strong competition among flanged males who meet too infrequently to establish a clear linear dominance hierarchy. Furthermore, we found no difference in fecal glucocorticoid metabolite concentrations between flanged and unflanged males, indicating that social stress is unlikely to explain arrested development in unflanged wild orangutans. The only actively developing male in our study showed significantly higher androgen levels during the period of development than later as a fully flanged male. This supports earlier findings from zoo studies that elevated androgen levels are associated with the development of secondary sexual characteristics.

Hair as a long-term retrospective cortisol calendar in orang-utans (Pongo spp.): new perspectives for stress monitoring in captive management and conservation.

This study examined whether the method of hair cortisol analysis is applicable to orang-utans (Pongo spp.) and can help to advance the objective monitoring of stress in non-human primates. Specifically, we examined whether fundamental prerequisites for hair cortisol analysis are given in orang-utans and, subsequently, whether segmental hair analysis may provide a retrospective calendar of long-term cortisol levels. For this, hair samples were examined from 71 zoo-living orang-utans (38 males, mean age=22.5years; 33 females, mean age=24years) for which detailed records of past living conditions were available. Hair samples were cut from defined body regions and were analyzed either in full length or in segments. Results showed that hair cortisol concentrations (HCC) were unrelated to age or sex of the individual animal. HCC were found to be higher in orang-utans, with perceived long-term stressful periods (mean HCC=43.6±26.5pg/mg, n=13) compared to animals without perceived stressful periods (19.3±5.5pg/mg, n=55, P<0.001). In non-stressed animals, segmental hair analyses revealed that HCC was stable along the hair shaft even when hair reached >40cm. The possibility of obtaining a retrospective calendar of stress-related cortisol changes through hair analysis was further supported by data of three case studies showing close correspondence between the segmental HCC results and keeper reports of stress exposure during the respective time periods. Finally, low within-animal variation in HCC from different body regions (CV%: 14.3) suggested that this method may also be applicable to naturally shed hair, e.g., as found in nests of wild orang-utans and other great apes. Therefore, using HCC may provide an ideal non-invasive tool for both captive management as well as conservation in orang-utans and potentially other great apes.

Mapping ecosystem services for land use planning, the case of Central Kalimantan.

Indonesia is subject to rapid land use change. One of the main causes for the conversion of land is the rapid expansion of the oil palm sector. Land use change involves a progressive loss of forest cover, with major impacts on biodiversity and global CO2 emissions. Ecosystem services have been proposed as a concept that would facilitate the identification of sustainable land management options, however, the scale of land conversion and its spatial diversity pose particular challenges in Indonesia. The objective of this paper is to analyze how ecosystem services can be mapped at the provincial scale, focusing on Central Kalimantan, and to examine how ecosystem services maps can be used for a land use planning. Central Kalimantan is subject to rapid deforestation including the loss of peatland forests and the provincial still lacks a comprehensive land use plan. We examine how seven key ecosystem services can be mapped and modeled at the provincial scale, using a variety of models, and how large scale ecosystem services maps can support the identification of options for sustainable expansion of palm oil production.

Fine-scale habitat use by orang-utans in a disturbed peat swamp forest, central Kalimantan, and implications for conservation management.

This study was conducted to see how orang-utans (Pongo pygmaeus wurmbii) were coping with fine-scale habitat disturbance in a selectively logged peat swamp forest in Central Kalimantan, Borneo. Seven habitat classes were defined, and orang-utans were found to use all of these, but were selective in their preference for certain classes over others. Overall, the tall forest classes (≥20 m) were preferred. They were preferred for feeding, irrespective of canopy connectivity, whereas classes with a connected canopy (canopy cover ≥75%), irrespective of canopy height, were preferred for resting and nesting, suggesting that tall trees are preferred for feeding and connected canopy for security and protection. The smaller forest classes (≤10 m high) were least preferred and were used mainly for travelling from patch to patch. Thus, selective logging is demonstrated here to be compatible with orang-utan survival as long as large food trees and patches of primary forest remain. Logged forest, therefore, should not automatically be designated as 'degraded'. These findings have important implications for forest management, forest classification and the designation of protected areas for orang-utan conservation.