Alternative reproductive tactics of unflanged and flanged male orangutans revisited.

In many slowly developing mammal species, males reach sexual maturity well before they develop secondary sexual characteristics. Sexually mature male orangutans have exceptionally long periods of developmental arrest. The two male morphs have been associated with behavioral alternative reproductive tactics, but this interpretation is based on cross-sectional analyses predominantly of Northwest Sumatran populations. Here we present the first longitudinal analyses of behavioral changes of 10 adult males that have been observed in both unflanged and flanged morph. We also analyzed long-term behavioral data on an additional 143 individually identified males from two study sites, Suaq (Sumatra, Pongo abelii) and Tuanan (Borneo, Pongo pygmaeus wurmbii), to assess male mating tactics cross-sectionally in relation to population, male morph (unflanged and flanged), and other socio-ecological factors. Both our longitudinal and cross-sectional results confirm and refine previous cross-sectional accounts of the differences in mating tactics between the unflanged and the flanged male morphs. In the unflanged morph, males exhibit higher sociability, particularly with females, and higher rates of both copulation and sexual coercion than in the flanged morph. Based on our results and those of previous studies showing that females prefer flanged males, and that flanged males have higher reproductive success, we conclude that unflanged males face a trade-off between avoiding male-male contest competition and gaining mating access to females, and thus follow a "best-of-a-bad-job" mating strategy.

Stable isotope evidence (Fe, Cu) suggests that sex, but not aging is recorded in rhesus macaque (Macaca mulatta) bone.

OBJECTIVES: Here, we examine (1) if the sex-related differences in iron (Fe) and copper (Cu) isotope ratios, represented as δ56 Fe and δ65 Cu values, respectively observed in humans exist in bulk occipital bone and incisors of male and female non-human primates, and (2) if the variation of Fe and Cu isotope ratios, known to vary in human blood as a factor of age are similar in non-human primate bone. MATERIALS AND METHODS: Isotope ratios were measured from the skeletal elements of 20 rhesus macaques (Macaca mulatta) with known life history traits. The metals were purified by column chromatography and their isotope ratios measured by MC-ICP-MS. Data were analyzed using generalized additive models (GAM). RESULTS: When accounting for age and sex independently, we found a significant relationship between δ65 Cu values and occipital bone, but not in incisors. There were no significant relationships observed between δ56 Fe values, occipital bone, or incisors. Similarly, there were no significant relationships observed between δ56 Fe values, δ65 Cu values, and age. DISCUSSION: We suggest that Cu and Fe isotope ratios have the potential to be useful supplementary tools in future research in biological anthropology, but additional studies are needed to further verify the relationship between sex, age, δ65 Cu, and δ56 Fe values in primates.

Fermented food consumption in wild nonhuman primates and its ecological drivers.

OBJECTIVES: Although fermented food use is ubiquitous in humans, the ecological and evolutionary factors contributing to its emergence are unclear. Here we investigated the ecological contexts surrounding the consumption of fruits in the late stages of fermentation by wild primates to provide insight into its adaptive function. We hypothesized that climate, socioecological traits, and habitat patch size would influence the occurrence of this behavior due to effects on the environmental prevalence of late-stage fermented foods, the ability of primates to detect them, and potential nutritional benefits. MATERIALS AND METHODS: We compiled data from field studies lasting at least 9 months to describe the contexts in which primates were observed consuming fruits in the late stages of fermentation. Using generalized linear mixed-effects models, we assessed the effects of 18 predictor variables on the occurrence of fermented food use in primates. RESULTS: Late-stage fermented foods were consumed by a wide taxonomic breadth of primates. However, they generally made up 0.01%-3% of the annual diet and were limited to a subset of fruit species, many of which are reported to have mechanical and chemical defenses against herbivores when not fermented. Additionally, late-stage fermented food consumption was best predicted by climate and habitat patch size. It was more likely to occur in larger habitat patches with lower annual mean rainfall and higher annual mean maximum temperatures. DISCUSSION: We posit that primates capitalize on the natural fermentation of some fruits as part of a nutritional strategy to maximize periods of fruit exploitation and/or access a wider range of plant species. We speculate that these factors contributed to the evolutionary emergence of the human propensity for fermented foods.

Evaluating Ketosis in Primate Field Studies: Validation of Urine Test Strips in Wild Bornean Orangutans (Pongo pygmaeus wurmbii).

The use of urine test strips (e.g., Roche Chemstrip®) has become the standard for quickly assessing the physiological condition and/or health of wild primates. These strips have been used to detect ketosis as a marker of fat catabolism in several primate taxa in their natural environments in response to changing food availability. However, the use of urine strips to determine ketosis has only been validated in human studies, and thus it remains unclear whether these strips accurately detect and quantify ketone bodies in nonhuman primates. We examined variations in ketone body concentrations in urine samples collected from wild Bornean orangutans at the Tuanan Orangutan Research Station. We assessed the accuracy of qualitative results from Chemstrip test strips in the field (i.e., negative, small, moderate, and large) using an enzyme-linked assay in the laboratory to determine the concentrations of acetoacetate of the same urine samples. Urine samples that tested positive for ketones in the field had significantly higher levels of ketones in the enzymatic assay compared to those that tested negative. There was significant variation in acetoacetate concentrations among the 4 Chemstrip values; however, post hoc tests revealed no significant differences between negative and small samples. We conclude that urinary test strips provide a useful tool for determining ketotic state in wild orangutans, but caution should be taken when interpreting results from samples showing only small levels of ketones on these strips.

Correction to: development of foraging skills in two orangutan populations: needing to learn or needing to grow?

[This corrects the article DOI: 10.1186/s12983-016-0178-5.].

Toughness of the Virunga mountain gorilla (Gorilla beringei beringei) diet across an altitudinal gradient.

The robust masticatory system of mountain gorillas is thought to have evolved for the comminution of tough vegetation, yet, compared to other primates, the toughness of the mountain gorilla diet is unremarkable. This may be a result of low plant toughness in the mountain gorilla environment or of mountain gorillas feeding selectively on low-toughness foods. The goal of this paper is to determine how the toughness of the mountain gorilla diet varies across their habitat, which spans a large altitudinal range, and whether there is a relationship between toughness and food selection by mountain gorillas. We collected data on the following variables to determine whether, and if so how, they change with altitude: leaf toughness of two plant species consumed by mountain gorillas, at every 100 m increase in altitude (2,600-3,700 m); toughness of consumed foods comprising over 85% of the gorilla diet across five vegetation zones; and toughness of unconsumed/infrequently consumed plant parts of those foods. Although leaf toughness increased with altitude, the toughness of the gorilla diet remained similar. There was a negative relationship between toughness and consumption frequency, and toughness was a better predictor of consumption frequency than plant frequency, biomass, and density. Consumed plant parts were less tough than unconsumed/infrequently consumed parts and toughness of the latter increased with altitude. Although it is unclear whether gorillas select food based on toughness or use toughness as a sensory cue to impart other plant properties (e.g., macronutrients, chemicals), our results that gorillas maintain a consistent low-toughness dietary profile across altitude, despite toughness increasing with altitude, suggest that the robust gorilla masticatory apparatus evolved for repetitive mastication of foods that are not high in toughness.

Nutritional ecology of wild Bornean orangutans (Pongo pygmaeus wurmbii) in a peat swamp habitat: Effects of age, sex, and season.

The spatial and temporal variation in food abundance has strong effects on wildlife feeding and nutrition. This variation is exemplified by the peatland forests of Central Kalimantan, which are characterized by unpredictable fruiting fluctuations, relatively low levels of fruit availability, and low fruit periods (<3% of trees fruiting) that can last nearly a year. Challenged by these environments, large, arboreal frugivores like orangutans must periodically rely on non-preferred, lower-quality foods to meet their nutritional needs. We examined variation in nutrient intake among age-sex classes and seasons over a 7-year period at the Tuanan Orangutan Research Station in Central Kalimantan. We conducted 2,316 full-day focal follows on 62 habituated orangutans (Pongo pygmaeus wurmbii). We found differences in total energy and macronutrient intake across age-sex classes, controlling for metabolic body mass. Intake of both total energy and macronutrients varied with fruit availability, and preference of dietary items increased with their nutritional quality. Foraging-related variables, such as day journey length, travel time, and feeding time, also varied among age-sex classes and with fruit availability. Our results add to the growing body of literature suggesting that great variation in foraging strategies exists among species, populations, and age-sex classes and in response to periods of resource scarcity. RESEARCH HIGHLIGHTS: The spatial and temporal variation in food abundance has strong effects on wildlife feeding and nutrition. Here we present the first long term study of the effects of variation in fruit availability and age/sex class on nutritional ecology of wild Bornean orangutans. We examined variation in nutrient intake of wild orangutans in living in a peat swamp habitat over a 7-year period at the Tuanan Orangutan Research Station in Central Kalimantan. We conducted 2,316 full-day focal follows on 62 habituated orangutans (Pongo pygmaeus wurmbii). We found differences in total energy and macronutrient intake across age-sex classes, controlling for metabolic body mass. Intake of both total energy and macronutrients varied with fruit availability, and preference of dietary items increased with their nutritional quality. Foraging-related variables, such as day journey length, travel time, and feeding time, also varied among age-sex classes and with fruit availability. Our results add to the growing body of literature suggesting that great variation in foraging strategies exists among species, populations, and age-sex classes and in response to periods of resource scarcity.

Chewing efficiency and occlusal functional morphology in modern humans.

The reduction of occlusal dimensions in early Homo is often proposed to be a functional adaptation to diet. With their smaller occlusal surfaces, species of early Homo are suggested to have reduced food-processing abilities, particularly for foods with high material properties (e.g., increased toughness). Here, we employ chewing efficiency as a measure of masticatory performance to test the relationships between masticatory function and food properties. We predicted that humans are more efficient when processing foods of lower toughness and Young's modulus values, and that subjects with larger occlusal surfaces will be less efficient when processing foods with higher toughness and Young's modulus, as the greater area spreads out the overall bite force applied to food particles. Chewing efficiency was measured in 26 adults using high-speed motion capture and surface electromyography. The dentition of each subject was cast and the occlusal surface was quantified using dental topographic analysis. Toughness and displacement-limited index were negatively correlated with chewing efficiency, but Young's modulus was not. Increased occlusal two-dimensional area and surface area were positively correlated with chewing efficiency for all foods. Thus, larger occlusal surface areas were more efficient when processing foods of greater toughness. These results suggest that the reduction in occlusal area in early Homo was associated with a reduction in chewing efficiency, particularly for foods with greater toughness. Further, the larger occlusal surfaces of earlier hominins such as Australopithecus would have likely increased chewing efficiency and increased the probability of fracture when processing tough foods.

Primate dietary ecology in the context of food mechanical properties.

Substantial variation exists in the mechanical properties of foods consumed by primate species. This variation is known to influence food selection and ingestion among non-human primates, yet no large-scale comparative study has examined the relationships between food mechanical properties and feeding strategies. Here, we present comparative data on the Young's modulus and fracture toughness of natural foods in the diets of 31 primate species. We use these data to examine the relationships between food mechanical properties and dietary quality, body mass, and feeding time. We also examine the relationship between food mechanical properties and categorical concepts of diet that are often used to infer food mechanical properties. We found that traditional dietary categories, such as folivory and frugivory, did not faithfully track food mechanical properties. Additionally, our estimate of dietary quality was not significantly correlated with either toughness or Young's modulus. We found a complex relationship among food mechanical properties, body mass, and feeding time, with a potential interaction between median toughness and body mass. The relationship between mean toughness and feeding time is straightforward: feeding time increases as toughness increases. However, when considering median toughness, the relationship with feeding time may depend upon body mass, such that smaller primates increase their feeding time in response to an increase in median dietary toughness, whereas larger primates may feed for shorter periods of time as toughness increases. Our results emphasize the need for additional studies quantifying the mechanical and chemical properties of primate diets so that they may be meaningfully compared to research on feeding behavior and jaw morphology.

Development of foraging skills in two orangutan populations: needing to learn or needing to grow?

BACKGROUND: Orangutans have one of the slowest-paced life histories of all mammals. Whereas life-history theory suggests that the time to reach adulthood is constrained by the time needed to reach adult body size, the needing-to-learn hypothesis instead suggests that it is limited by the time needed to acquire adult-level skills. To test between these two hypotheses, we compared the development of foraging skills and growth trajectories of immature wild orangutans in two populations: at Tuanan (Pongo pygmaeus wurmbii), Borneo, and Suaq Balimbing (Pongo abelii), Sumatra. We collected behavioral data on diet repertoire, feeding rates and ranging competence during focal follows, and estimated growth through non-invasive laser photogrammetry. RESULTS: We found that adult-like diet repertoires are attained around the age of weaning and that female immatures increase their repertoire size faster than their male peers. Adult-level feeding rates of easy techniques are reached just after weaning, but several years later for more difficult techniques, albeit always before adulthood (i.e. age at first reproduction). Independent immatures had faster feeding rates for easy to process items than their mothers, with male immatures achieving faster feeding rates earlier in development relative to females. Sumatran immatures reach adult-level feeding rates 2-3 years later than their Bornean peers, in line with their higher dietary complexity and later weaning. The range-use competence of independently ranging and weaned immatures is similar to that of adult females. Body size measurements showed, immatures grow until female age of first reproduction. CONCLUSIONS: In conclusion, unlike in humans, orangutan foraging skills are in place prior to reproduction. Growth trajectories suggest that energetic constraints, rather than skills, best explain the length of immaturity. However, skill competence for dietary independence is reached later where the adult niche is more complex, which is consistent with the relatively later weaning age with increasing brain size found generally in primates, and apes in particular.

Age-related variation in the mechanical properties of foods processed by Sapajus libidinosus.

OBJECTIVES: The diet of tufted capuchins (Sapajus) is characterized by annual or seasonal incorporation of mechanically protected foods. Reliance on these foods raises questions about the dietary strategies of young individuals that lack strength and experience to access these resources. Previous research has demonstrated differences between the feeding competencies of adult and juvenile tufted capuchins. Here we test the hypothesis that, compared to adults, juveniles will process foods with lower toughness and elastic moduli. MATERIALS AND METHODS: We present data on variation in the toughness and elastic modulus of food tissues processed by Sapajus libidinosus during the dry season at Fazenda Boa Vista, Brazil. Food mechanical property data were collected using a portable universal mechanical tester. RESULTS: Results show that food tissues processed by the capuchins showed significant differences in toughness and stiffness. However, we found no relationship between an individual's age and mean or maximum food toughness or elastic modulus, indicating both juvenile and adult S. libidinosus are able to process foods of comparable properties. DISCUSSION: Although it has been suggested that juveniles avoid mechanically protected foods, age-related differences in feeding competence are not solely due to variation in food toughness or stiffness. Other factors related to food type (e.g., learning complex behavioral sequences, achieving manual dexterity, obtaining physical strength to lift stone tools, or recognizing subtle cues about food state) combined with food mechanical properties better explain variation in juvenile feeding competency.

Food mechanical properties, feeding ecology, and the mandibular morphology of wild orangutans.

Bornean orangutan mandibular morphology has been functionally linked to the exploitation of hard and tough foods, based on evidence that Pongo pygmaeus wurmbii spends a greater percentage of time feeding on bark, seeds and vegetation compared with Pongo abelii (Sumatran orangutans) and the assumption that these tissues are more challenging to process than fruit pulp. We measured and compared toughness (R) and Young's modulus (E) of ripe and unripe foods exploited by P. abelii and P. p. wurmbii. Additionally, we recorded and compared the percentage of time these orangutans fed on plants/plant parts of varying degrees of R and E. Compared with P. abelii, P. p. wurmbii consumed significantly tougher and more displacement limited (R/E)(0.5) fruit parts, leaves and inner bark, and spent a significantly greater percentage of time feeding on immature leaves, unripe fruit and other vegetation. Modulus did not vary as expected between species, likely because we failed to capture the high-end range of modulus values for tissues consumed by P. p. wurmbii. Notably, P. p. wurmbii spent ∼40% of its feeding time on the toughest foods consumed (between 1000 and 4000 J m(-2)). Thus, the hypothesis that mandibular robusticity in P. p. wurmbii is functionally linked to feeding on tough foods is supported and is likely related to countering relatively larger external forces and/or repetitive loads required to process the toughest tissues. The importance of elastic modulus on morphological divergence awaits future studies capturing the full range of this material property for P. p. wurmbii. Finally, phenophase and fruit availability influence orangutan species differences in food material properties and percentage of time spent feeding on various foods, emphasizing the importance of incorporating these variables in future studies of feeding ecology and craniodental morphology in extant taxa.

Wildfire smoke linked to vocal changes in wild Bornean orangutans.

Tropical peatlands are the sites of Earth's largest fire events, with outsized contributions to greenhouse gases, toxic smoke, and haze rich with particulate matter. The human health risks from wildfire smoke are well known, but its effects on wildlife inhabiting these ecosystems are poorly understood. In 2015, peatland fires on Borneo created a thick haze of smoke that blanketed the region. We studied its effects on the long call vocalizations of four adult male Bornean orangutans (Pongo pygmaeus wurmbii) in a peat swamp forest. During the period of heavy smoke, orangutans called less often and showed reduced vocal quality-lower pitch, increased harshness and perturbations, and more nonlinear phenomena-similar to changes in human smokers. Most of these changes persisted for two months after the smoke had cleared and likely signal changes in health. Our work contributes valuable information to support non-invasive acoustic monitoring of this Critically Endangered primate.

Reproductive success of Bornean orangutan males: scattered in time but clustered in space.

Abstract: The social and mating systems of orangutans, one of our closest relatives, remain poorly understood. Orangutans (Pongo spp.) are highly sexually dimorphic and females are philopatric and maintain individual, but overlapping home ranges, whereas males disperse, are non-territorial and wide-ranging, and show bimaturism, with many years between reaching sexual maturity and attaining full secondary sexual characteristics (including cheek pads (flanges) and emitting long calls). We report on 21 assigned paternities, among 35 flanged and 15 unflanged, genotyped male Bornean orangutans (Pongo pygmaeus wurmbii), studied from 2003 to 2018 in Tuanan (Central Kalimantan, Indonesia). All 10 infants born since mid-2003 with an already identified sire were sired by flanged males. All adult males ranged well beyond the study area (c. 1000 ha), and their dominance relations fluctuated even within short periods. However, 5 of the 10 identified sires had multiple offspring within the monitored area. Several sired over a period of c. 10 years, which overlapped with siring periods of other males. The long-calling behavior of sires indicated they were not consistently dominant over other males in the area around the time of known conceptions. Instead, when they were seen in the area, the known sires spent most of their time within the home ranges of the females whose offspring they sired. Overall, successful sires were older and more often resident than others. Significance statement: It is difficult to assess reproductive success for individuals of long-lived species, especially for dispersing males, who cannot be monitored throughout their lives. Due to extremely long interbirth intervals, orangutans have highly male-skewed operational sex ratios and thus intensive male-male competition for every conception. Paternity analyses matched 21 immature Bornean orangutans with their most likely sire (only 10 of 50 genotyped males) in a natural population. Half of these identified sires had multiple offspring in the study area spread over periods of at least 10 years, despite frequently ranging outside this area. Dominance was a poor predictor of success, but, consistent with female mating tactics to reduce the risk of infanticide, known "sires" tended to have relatively high local presence, which seems to contribute to the males' siring success. The results highlight the importance of large protected areas to enable a natural pattern of dispersal and ranging. Supplementary Information: The online version contains supplementary material available at 10.1007/s00265-023-03407-6.

Bornean orangutans on the brink of protein bankruptcy.

Protein is a limiting resource that is essential to the growth, maintenance and reproduction of tropical frugivores, yet few studies have examined how wild animals maintain protein balance. During chronic periods of fruit scarcity, Bornean orangutans (Pongo pygmaeus) often catabolize their own fat reserves despite unusually low metabolic requirements. Such energy deficits suggest a marginal existence, and raise the possibility that orangutans also endure periods of negative protein balance. To test this hypothesis, we conducted the first study of protein cycling in a wild primate. Our five year analysis of urinary metabolites revealed evidence of protein recycling when fruit was scarce. During these periods, orangutans consumed more leaves and bark, proteinaceous but tough foods that yielded a mean daily intake of 1.4 g protein kg(-1) metabolic mass. Such an amount is inadequate for humans and one-tenth the intake of mountain gorillas, but sufficient to avert, perhaps narrowly, a severe protein deficit. Our findings highlight the functional and adaptive value of traits that maximize protein assimilation during periods of ecological exigency.

Slow loris (Nycticebus borneanus) consumption by a wild Bornean orangutan (Pongo pygmaeus wurmbii).

Vertebrate predation and consumption by wild Bornean orangutans (Pongo pygmaeus spp.) is rare. In contrast to recorded observations of slow loris consumption by Sumatran orangutans (Pongo abelii), no cases of this have been previously published for Bornean orangutans in the wild. In 2017, we observed the capture and consumption of a slow loris (Nycticebus borneanus) by an adult unflanged male Bornean orangutan at Tuanan Orangutan Research Station, which is located in the Kapuas region of Central Kalimantan. The unflanged male was together with an adult female and her 3.5-year-old offspring throughout the event. However, despite the mother and her offspring watching the male closely and occasionally begging while he consumed the loris, he resisted all food-taking attempts. This study reports, to the best of our knowledge, the first documented case of slow loris predation and consumption by a Bornean orangutan, and thus provides an important data point for understanding primate predation on other primate species.

The development and maintenance of sex differences in dietary breadth and complexity in Bornean orangutans.

ABSTRACT: Orangutans show a pronounced sexual dimorphism, with flanged males (i.e., males with fully grown secondary sexual characteristics) reaching twice the size of adult females. Furthermore, adult orangutans show sex-specific dispersal and activity patterns. This study investigates sex differences in adult foraging behavior and sheds light on how these differences develop in immatures. We analyzed 11 years of feeding data on ten adult female, seven flanged male, and 14 immature Bornean orangutans (Pongo pygmaeus wurmbii) at Tuanan in Central Kalimantan, Indonesia. We found that the diets of the adult females were significantly broader and required more processing steps before ingestion than the diets of flanged males. We also found evidence for a similar difference in overall diet repertoire sizes. For the immatures, we found that whereas females reached 100% of their mothers' diet spectrum size by the age of weaning, males reached only around 80%. From the age of 4 years on (i.e., years before being weaned) females had significantly broader daily diets than males. We found no difference in daily or overall diet processing intensity of immature males and females but found preliminary evidence that immature males included fewer items of their mother's diet in their own diets that were processing-intensive. Overall, our results suggest that by eating a broader variety and more complex to process food items, female orangutans go to greater lengths to achieve a balanced diet than males do. These behavioral differences are not just apparent in adult foraging behavior but also reflected in immature development from an early age on. SIGNIFICANCE STATEMENT: In many species, males and females have different nutritional needs and are thus expected to show sex-specific foraging behavior. Sex differences in several aspects of foraging behavior have been found in various species, but it remains largely unclear when and how those develop during ontogeny, which is especially relevant for long-lived altricial species that learn foraging skills over many years. In our study, we analyzed a cross-sectional and longitudinal data set containing more than 750,000 feeding events of adult and immature Bornean orangutans (Pongo pygmaeus wurmbii). We found that adult females had significantly broader and more complex diets than males. We also found that these differences started to develop during infancy, suggesting that immature orangutans prepare for their sex-specific foraging niches long before those become physiologically relevant while they are still in constant association with their mothers and before being frequently exposed to other role models. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s00265-021-03014-3.

Wild Bornean orangutans experience muscle catabolism during episodes of fruit scarcity.

Pronounced temporal and spatial variation in the availability of food resources can produce energetic deficits in organisms. Fruit-dependent Bornean orangutans face extreme variation in fruit availability and experience negative energy and protein balance during episodes of fruit scarcity. We evaluate the possibility that orangutans of different sexes and ages catabolize muscle tissue when the availability of fruit is low. We assess variation in muscle mass by examining the relationship between urinary creatinine and specific gravity and use the residuals as a non-invasive measure of estimated lean body mass (ELBM). Despite orangutans having a suite of adaptations to buffer them from fruit scarcity and associated caloric deficits, ELBM was lower during low fruit periods in all age-sex classes. As predicted, adult male orangutans had higher ELBM than adult females and immatures. Contrary to expectation, flanged and unflanged males did not differ significantly in ELBM. These findings highlight the precarity of orangutan health in the face of rapid environmental change and add to a growing body of evidence that orangutans are characterized by unique metabolic traits shaped by their unpredictable forest environment.

The cost of associating with males for Bornean and Sumatran female orangutans: a hidden form of sexual conflict?

ABSTRACT: Sexual coercion, in the form of forced copulations, is relatively frequently observed in orangutans and generally attributed to their semi-solitary lifestyle. High ecological costs of association for females may be responsible for this lifestyle and may have prevented the evolution of morphological fertility indicators (e.g., sexual swellings), which would attract (male) associates. Therefore, sexual conflict may arise not only about mating per se but also about associations, because males may benefit from associations with females to monitor their reproductive state and attempt to monopolize their sexual activities. Here, we evaluate association patterns and costs for females when associating with both males and females of two different orangutan species at two study sites: Suaq, Sumatra (Pongo abelii), and Tuanan, Borneo (Pongo pygmaeus wurmbii). Female association frequency with both males and females was higher in the Sumatran population, living in more productive habitat. Accordingly, we found that the cost of association, in terms of reduced feeding to moving ratio and increased time being active, is higher in the less sociable Bornean population. Males generally initiated and maintained such costly associations with females, and prolonged associations with males led to increased female fecal cortisol metabolite (FCM) levels at Tuanan, the Bornean population. We conclude that male-maintained associations are an expression of sexual conflict in orangutans, at least at Tuanan. For females, this cost of association may be responsible for the lack of sexual signaling, while needing to confuse paternity. SIGNIFICANCE STATEMENT: Socioecological theory predicts a trade-off between the benefits of sociality and the ecological costs of increased feeding competition. Orangutans' semi-solitary lifestyle has been attributed to the combination of high association costs and low predation risk. Previous work revealed a positive correlation between association frequencies and habitat productivity, but did not measure the costs of association. In this comparative study, we show that females likely incur costs from involuntary, male-maintained associations, especially when they last for several days and particularly in the population characterized by lower association frequencies. Association maintenance therefore qualifies as another expression of sexual conflict in orangutans, and especially prolonged, male-maintained associations may qualify as an indirect form of sexual coercion. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s00265-020-02948-4.