Relative leg-to-arm skeletal strength proportions in orangutans by species and sex.

Among extant great apes, orangutans climb most frequently. However, Bornean orangutans (Pongo pygmaeus) exhibit higher frequencies of terrestrial locomotion than do Sumatran orangutans (Pongo abelii). Variation in long bone cross-sectional geometry is known to reflect differential loading of the limbs. Thus, Bornean orangutans should show greater relative leg-to-arm strength than their Sumatran counterparts. Using skeletal specimens from museum collections, we measured two cross-sectional geometric measures of bone strength: the polar section modulus (Zpol) and the ratio of maximum to minimum area moments of inertia (Imax/Imin), at the midshaft of long bones in Bornean (n = 19) and Sumatran adult orangutans (n = 12) using medical CT and peripheral quantitative CT scans, and compared results to published data of other great apes. Relative leg-to-arm strength was quantified using ratios of femur and tibia over humerus, radius, and ulna, respectively. Differences between orangutan species and between sexes in median ratios were assessed using Wilcoxon rank sum tests. The tibia of Bornean orangutans was stronger relative to the humerus and the ulna than in Sumatran orangutans (p = 0.008 and 0.025, respectively), consistent with behavioral studies that indicate higher frequencies of terrestrial locomotion in the former. In three Zpol ratios, adult female orangutans showed greater leg-to-arm bone strength compared to flanged males, which may relate to females using their legs more during arboreal locomotion than in adult flanged males. A greater amount of habitat discontinuity on Borneo compared to Sumatra has been posited as a possible explanation for observed interspecific differences in locomotor behaviors, but recent camera trap studies has called this into question. Alternatively, greater frequencies of terrestriality in Pongo pygmaeus may be due to the absence of tigers on Borneo. The results of this study are consistent with the latter explanation given that habitat continuity was greater a century ago when our study sample was collected.

Identification of potentially zoonotic parasites in captive orangutans and semi-captive mandrills: Phylogeny and morphological comparison.

Cysts and trophozoites of vestibuliferid ciliates and larvae of Strongyloides were found in fecal samples from captive orangutans Pongo pygmaeus and P. abelii from Czech and Slovak zoological gardens. As comparative material, ciliates from semi-captive mandrills Mandrillus sphinx from Gabon were included in the study. Phylogenetic analysis of the detected vestibuliferid ciliates using ITS1-5.8s-rRNA-ITS2 and partial 18S ribosomal deoxyribonucleic acid (rDNA) revealed that the ciliates from orangutans are conspecific with Balantioides coli lineage A, while the ciliates from mandrills clustered with Buxtonella-like ciliates from other primates. Morphological examination of the cysts and trophozoites using light microscopy did not reveal differences robust enough to identify the genera of the ciliates. Phylogenetic analysis of detected L1 larvae of Strongyloides using partial cox1 revealed Strongyloides stercoralis clustering within the cox1 lineage A infecting dogs, humans, and other primates. The sequences of 18S rDNA support these results. As both B. coli and S. stercoralis are zoonotic parasites and the conditions in captive and semi-captive settings may facilitate transmission to humans, prophylactic measures should reflect the findings.

Densities of Bornean orang-utans (Pongo pygmaeus morio) in heavily degraded forest and oil palm plantations in Sabah, Borneo.

The conversion of forest to agriculture continues to contribute to the loss and fragmentation of remaining orang-utan habitat. There are still few published estimates of orang-utan densities in these heavily modified agricultural areas to inform range-wide population assessments and conservation strategies. In addition, little is known about what landscape features promote orang-utan habitat use. Using indirect nest count methods, we implemented surveys and estimated population densities of the Northeast Bornean orang-utan (Pongo pygmaeus morio) across the continuous logged forest and forest remnants in a recently salvage-logged area and oil palm plantations in Sabah, Malaysian Borneo. We then assessed the influence of landscape features and forest structural metrics obtained from LiDAR data on estimates of orang-utan density. Recent salvage logging appeared to have a little short-term effect on orang-utan density (2.35 ind/km 2 ), which remained similar to recovering logged forest nearby (2.32 ind/km 2 ). Orang-utans were also present in remnant forest patches in oil palm plantations, but at significantly lower numbers (0.82 ind/km 2 ) than nearby logged forest and salvage-logged areas. Densities were strongly influenced by variation in canopy height but were not associated with other potential covariates. Our findings suggest that orang-utans currently exist, at least in the short-term, within human-modified landscapes, providing that remnant forest patches remain. We urge greater recognition of the role that these degraded habitats can have in supporting orang-utan populations, and that future range-wide analyses and conservation strategies better incorporate data from human-modified landscapes.

CONGENITAL HYPOTHYROIDISM IN A BORNEAN ORANGUTAN (PONGO PYGMAEUS) AND A SUMATRAN ORANGUTAN (PONGO ABELII).

Congenital hypothyroidism (CH) in humans is most commonly caused by disruption of thyroid gland development (dysgenesis) or an inherited defect in thyroid hormone biosynthesis (dyshormonogenesis). CH has not been previously documented in great apes. This report describes the clinical presentation, diagnosis, and treatment of CH in a 9-mo-old male Bornean orangutan (Pongo pygmaeus) and a 6-wk-old female Sumatran orangutan (Pongo abelii). Primary CH due to thyroid dysgenesis was confirmed in the Bornean orangutan using sonography and radioisotope scintigraphy. Although commercial thyroid immunoassays are not validated for use in orangutans, in comparison to age-matched controls, thyroid-stimulating hormone level was markedly elevated, and serum thyroxine (T4) and free T4 levels were markedly decreased in both cases. Oral supplementation with levothyroxine sodium resulted in noticeable clinical improvement in both orangutans within 30 days of initiating treatment.

SURGICAL MANAGEMENT OF UTERINE LESIONS IN TWO CAPTIVE ORANGUTANS ( PONGO SPP.).

Uterine lesions in two orangutans were effectively managed with surgical intervention. A 26-year-old hybrid orangutan ( Pongo spp.) was diagnosed with uterine adenomyosis based on advanced imaging. Histologic evaluation identified multifocal myometrial endometriosis, a variant of adenomyosis. A 27-year-old Bornean orangutan ( Pongo pygmaeus) was diagnosed with a focal uterine fibroid based on histologic examination. The animals were housed at separate institutions and initially presented with dysmenorrhea and menorrhagia. Both animals were treated intermittently for episodes of dysmenorrhea, with recurrence of clinical signs after each treatment. Due to the lack of consistent response to medical management, an ovariohysterectomy in the hybrid orangutan and a myomectomy in the Bornean orangutan were performed and resulted in complete resolution of clinical signs. Surgical management of adenomyosis and neoplasia has previously been reported in nonhuman primates. These cases are the first known documentation of surgical management of multifocal myometrial endometriosis and a fibroid in orangutans.

Orangutans, enamel defects, and developmental health: A comparison of Borneo and Sumatra.

Orangutans (Pongo sp.) show among the highest occurrence of three types of developmental enamel defect. Two are attributed to nutritional factors that reduce bone growth in the infant's face early in development. Their timing and prevalence indicate that Sumatra provides a better habitat than does Borneo. The third type, repetitive linear enamel hypoplasia (rLEH) is very common but its etiology is not understood. Our objective is to draw attention to this enigmatic, episodic stressor in the lives of orangutans. We are concerned that neglect of this possible marker of ill health may be contributing, through inaction, to their alarming decline in numbers. Width and depth of an LEH are considered proxies for duration and intensity of stress. The hypothesis that Bornean orangutans would exhibit relatively wider and deeper LEH was tested on 163 independent episodes of LEH from 9 Sumatran and 26 Bornean orangutans measured with a NanoFocus AG "µsurf Mobile Plus" scanner. Non-normally distributed data (depths) were converted to natural logs. No difference was found in width of LEH among the two island taxa; nor are their differences in width or depth between the sexes. After controlling for significant differences in LEH depths between incisors and canines, defects are, contrary to prediction, significantly deeper in Sumatran than Bornean animals (median = 28, 18 µm, respectively). It is concluded that repetitive LEH records an unknown but significant stressor present in both Sumatra and Borneo, with an average periodicity of 6 months (or multiples thereof) that lasts about 6-8 weeks. It is worse in Sumatra. Given this patterning, shared with apes from a wide range of ecological and temporal sources, rLEH is more likely attributable to disease than to malnutrition.

MAMMARY GLAND ADENOCARCINOMA IN A MALE BORNEAN ORANGUTAN (PONGO PYGMAEUS).

An adult male Bornean orangutan ( Pongo pygmaeus ) was diagnosed with invasive, poorly differentiated grade 9/9 mammary gland adenocarcinoma from a subcutaneous mass that was surgically removed during a routine preventative health examination. The tumor was tested for estrogen and progesterone receptors, human epidermal growth factor receptor 2 (HER2), and HER2 fluorescence in situ hybridization (HER2 FISH). Whole blood was tested for breast cancer 1 (BRCA1) and breast cancer 2 (BRCA2) genes. The orangutan was treated orally with two common human breast cancer drugs; tamoxifen and anastrozole. The orangutan lived for 4.5 yr postdetection, dying from an unrelated cause. This is the first reported case of mammary gland adenocarcinoma in a male great ape.

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.

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.

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.

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.

EVALUATION OF DIAGNOSTIC ACCURACY OF THE COMPARATIVE TUBERCULIN SKIN TEST IN REHABILITANT BORNEAN ORANGUTANS (PONGO PYGMAEUS).

The tuberculin skin test (TST) has been the mainstay of tuberculosis (TB) testing in primates for decades, but its interpretation in orangutans (Pongo spp.) is challenging, because many animals react strongly, without evidence of infection with Mycobacterium tuberculosis complex. One explanation is cross-reactivity with environmental nontuberculous mycobacteria (NTM). The use of a comparative TST (CTST), comparing reactivity to avian (representing NTM) and bovine (representing tuberculous mycobacteria) tuberculins aids in distinguishing cross-reactivity due to sensitization by NTM from shared antigens. The specificity of the TST can be increased with the use of CTST. We considered three interpretations of the TST in rehabilitant Bornean orangutans ( Pongo pygmaeus ) using avian purified protein derivative (APPD; 25,000 IU/ml) and two concentrations of bovine purified protein derivative (BPPD; 100,000 and 32,500 IU/ml). The tests were evaluated for their ability to identify accurately seven orangutans previously diagnosed with and treated for TB from a group of presumed negative individuals (n = 288 and n = 161 for the two respective BPPD concentrations). BPPD at 32,500 IU/ml had poor diagnostic capacity, whereas BPPD at 100,000 IU/ml performed better. The BPPD-only interpretation had moderate sensitivity (57%) and poor specificity (40%) and accuracy (41%). The comparative interpretation at 72 hr had similar sensitivity (57%) but improved specificity (95%) and accuracy (94%). However, best results were obtained by a comparative interpretation incorporating the 48- and 72-hr scores, which had good sensitivity (86%), specificity (95%) and accuracy (95%). These data reinforce recommendations that a CTST be used in orangutans and support the use of APPD at 25,000 IU/ml and BPPD at 100,000 IU/ml. The highest score at each site from the 48- and 72-hr checks should be considered the result for that tuberculin. If the bovine result is greater than the avian result, the animal should be considered a TB suspect.

Fatal metacestode infection in Bornean orangutan caused by unknown Versteria species.

A captive juvenile Bornean orangutan (Pongo pygmaeus) died from an unknown disseminated parasitic infection. Deep sequencing of DNA from infected tissues, followed by gene-specific PCR and sequencing, revealed a divergent species within the newly proposed genus Versteria (Cestoda: Taeniidae). Versteria may represent a previously unrecognized risk to primate health.

Oxygen isotopes in orangutan teeth reveal recent and ancient climate variation.

Studies of climate variation commonly rely on chemical and isotopic changes recorded in sequentially produced growth layers, such as in corals, shells, and tree rings, as well as in accretionary deposits-ice and sediment cores, and speleothems. Oxygen isotopic compositions (δ18O) of tooth enamel are a direct method of reconstructing environmental variation experienced by an individual animal. Here, we utilize long-forming orangutan dentitions (Pongo spp.) to probe recent and ancient rainfall trends on a weekly basis over ~3-11 years per individual. We first demonstrate the lack of any consistent isotopic enrichment effect during exclusive nursing, supporting the use of primate first molar teeth as environmental proxies. Comparisons of δ18O values (n=2016) in twelve molars from six modern Bornean and Sumatran orangutans reveal a high degree of overlap, with more consistent annual and bimodal rainfall patterns in the Sumatran individuals. Comparisons with fossil orangutan δ18O values (n=955 measurements from six molars) reveal similarities between modern and late Pleistocene fossil Sumatran individuals, but differences between modern and late Pleistocene/early Holocene Bornean orangutans. These suggest drier and more open environments with reduced monsoon intensity during this earlier period in northern Borneo, consistent with other Niah Caves studies and long-term speleothem δ18O records in the broader region. This approach can be extended to test hypotheses about the paleoenvironments that early humans encountered in southeast Asia.

When an animal drinks water, two naturally occurring variants of oxygen ­ known as oxygen-18 and oxygen-16 ­ are incorporated into its growing teeth. The ratio of these variants in water changes with temperature, rainfall and other environmental conditions and therefore can provide a record of the climate during an animal's life. Teeth tend to be well preserved as fossils, which makes it possible to gain insights into this climate record even millions of years after an animal's death. Orangutans are highly endangered great apes that today live in rainforests on the islands of Borneo and Sumatra. During a period of time known as the Pleistocene (around 2.6 million years to 12,000 years ago), these apes were more widely spread across Southeast Asia. Climate records from this area in the time before human-induced climate change are somewhat limited. Therefore, fossilized orangutan teeth offer a possible way to investigate past seasonal rainfall patterns and gain insight into the kind of environments early humans would have encountered. To address this question, Smith et al. measured oxygen-18 and oxygen-16 variants in thin slices of modern-day orangutan teeth using a specialized analytical system. This established that the teeth showed seasonal patterns consistent with recent rainfall trends, and that the ratio of these oxygen variants did not appear to be impacted by milk intake in young orangutans. These findings indicated that the oxygen variants could be a useful proxy for predicting prehistoric weather patterns from orangutan teeth. Further measurements of teeth from fossilized Sumatran orangutans showed broadly similar rainfall patterns to those of teeth from modern-day orangutans. On the other hand, fossilized teeth from Borneo suggested that the environment used to be drier, with less intense wet seasons. The approach developed by Smith et al. provides an opportunity for scientists to leverage new fossil discoveries as well as existing collections to investigate past environments. This could allow future research into how climate variation may have influenced the spread of early humans through the region, as well as the evolution of orangutans and other endangered animals.

Wild and captive immature orangutans differ in their non-vocal communication with others, but not with their mothers.

Abstract: In many group-living species, individuals are required to flexibly modify their communicative behaviour in response to current social challenges. To unravel whether sociality and communication systems co-evolve, research efforts have often targeted the links between social organisation and communicative repertoires. However, it is still unclear which social or interactional factors directly predict communicative complexity. To address this issue, we studied wild and zoo-housed immature orangutans of two species to assess the impact of the socio-ecological setting on the production of non-vocal signal repertoires. Specifically, we compared repertoire size, dyadic repertoire similarity, and number of social goals (i.e. observer's estimate of the signaller's intended interaction outcome) for communicative interactions with mothers versus other conspecifics, controlling for critical individual and environmental factors. In this small sample of immature orangutans, wild-captive contrasts were statistically significant only for other-directed repertoires, but not for mother-directed repertoires, and not for the number of social goals that immatures communicated towards. While the repertoires of individuals living in the same research setting were more similar than those living in contrasting settings, this difference was most pronounced for other-directed repertoires of the less socially tolerant orangutan species. These results suggest that the boosted interactional opportunities in captivity rather than mere differences in environmental affordances or communicative needs drive the wild-captive contrast in orangutan communicative repertoires. Overall, this fine-grained analysis of repertoires further underscores that not only a species' social organisation but also the targeted audience may have a profound impact on communicative behaviour. Significance statement: Navigating a dynamic social environment often requires flexible signal use. While it has repeatedly been shown that the social organisation and structure of species predict the complexity of their communication systems, the mechanisms underlying these relationships are largely unknown. Because targeted studies to assess this issue in great apes are difficult, we take an alternative approach here: we compare the same species living in the wild and in artificial habitats in captivity. This contrast allows a direct test of how repertoires respond to the relevant difference in socio-ecological conditions. Our results show that the diversity of interaction partners (i.e. social opportunities), but not the diversity of social goals (i.e. possible interaction outcomes) or the broader physical opportunities (i.e. safe ground use), predict the size and consistency of wild and captive signalling repertoires. Supplementary Information: The online version contains supplementary material available at 10.1007/s00265-023-03426-3.

Terrestriality in the Bornean orangutan (Pongo pygmaeus morio) and implications for their ecology and conservation.

Aside from anecdotal evidence, terrestriality in orangutans (Pongo spp.) has not been quantified or subject to careful study and important questions remain about the extent and contexts of terrestrial behavior. Understanding the factors that influence orangutan terrestriality also has significant implications for their conservation. Here we report on a camera trapping study of terrestrial behavior in the northeastern Bornean orangutan, Pongo pygmaeus morio, in Wehea Forest, East Kalimantan, Indonesia. We used 78 non-baited camera traps set in 43 stations along roads, trails, and at mineral licks (sepans) to document the frequency of orangutan terrestriality. Habitat assessments were used to determine how terrestrial behavior was influenced by canopy connectivity. We compared camera trapping results for P. p. morio to those for a known terrestrial primate (Macaca nemestrina), and another largely arboreal species (Presbytis rubicunda) to assess the relative frequency of terrestrial behavior by P. p. morio. A combined sampling effort of 14,446 trap days resulted in photographs of at least 15 individual orangutans, with females being the most frequently recorded age sex class (N=32) followed by flanged males (N=26 records). P. p. morio represented the second most recorded primate (N=110 total records) of seven primate species recorded. Capture scores for M. nemestrina (0.270) and P. p. morio (0.237) were similar and almost seven times higher than for the next most recorded primate, P. rubicunda (0.035). In addition, our results indicate that for orangutans, there was no clear relationship between canopy connectivity and terrestriality. Overall, our data suggest that terrestriality is relatively common for the orangutans in Wehea Forest and represents a regular strategy employed by individuals of all age-sex classes. As Borneo and Sumatra increasingly become characterized by mixed-use habitats, understanding the ecological requirements and resilience in orangutans is necessary for designing optimal conservation strategies.

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.

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.

Social tolerance and interactional opportunities as drivers of gestural redoings in orang-utans.

Communicative repair is a fundamental and universal element of interactive language use. It has been suggested that the persistence and elaboration after communicative breakdown in nonhuman primates constitute two evolutionary building blocks of this capacity, but the conditions favouring it are poorly understood. Because zoo-housed individuals of some species are more social and more terrestrial than in the wild, they should be more likely to show gestural redoings (i.e. both repetition and elaboration) after communicative failure in the coordination of their joint activities. Using a large comparative sample of wild and zoo-housed orang-utans of two different species, we could confirm this prediction for elaboration, the more flexible form of redoings. Specifically, results showed that gestural redoings in general were best predicted by the specific social action context (i.e. social play) and interaction dyad (i.e. beyond mother-offspring), although they were least frequent in captive Bornean orang-utans. For gestural elaboration, we found the expected differences between captive and wild research settings in Borneans, but not in Sumatrans (the more socially tolerant species). Moreover, we found that the effectiveness of elaboration in eliciting responses was higher in Sumatrans, especially the captive ones, whereas effectiveness of mere repetition was influenced by neither species nor setting. We conclude that the socio-ecological environment plays a central role in the emergence of communicative repair strategies in great apes. This article is part of the theme issue 'Revisiting the human 'interaction engine': comparative approaches to social action coordination'.

Preliminary findings of age and male sexual characteristics andand potential effect to semen characteristics and cryopreservation of the critically endangered Bornean orangutan in Malaysia.

Bornean orangutan is a critically endangered non-human primate; however, the threat of extinction is not merely from poaching and habitat loss. Orangutan survival is also threatened by the genetic loss and genetic bottleneck due to the low effective population, prompting the dire need for an immediate genetic preservation program through systematic biobanking and assisted reproductive technology (ART). This study aims to provide integral data to the semen characteristics, extension, and cryopreservation of the Bornean orangutan and the potential relationship to male traits. Five captive orangutans from Sepilok Orangutan Rehabilitation Centre (SORC) with a mean body weight of 52.81 ± 7.00 kg were used for this study. Semen collection was performed using electroejaculation (EE) under complete general anesthesia. Semen was subjected to macroscopic and microscopic evaluation while testicular measurement was obtained using digital calipers. The semen characteristics of the orangutans are volume (778 ± 250.21 µl), pH (7.80 ± 0.25), concentration (32.38 ± 17.40 × 106 sperm/ml), total motility (61.00 ± 12.88%), adjusted motility index (48.76 ± 11.32%), live spermatozoa (77.75 ± 6.94%) and normal spermatozoa (11.48 ± 11.34%). Analysis of variance statistical analysis test was used to compare the significant difference between means, at (p < 0.05). Spermatozoa concentration was the only significant different parameter between individuals. Testes biometry parameters are statistically significant between the flanged and unflanged individuals. Live spermatozoa are different in adult and sub-adult individual while teratospermia was found to be consistently high in all individuals. Chilled and post-thaw quality after cryopreservation suggests promising survivability of spermatozoa. Semen collection with EE yields a consistent and acceptable quality of spermatozoa for cryopreservation, biobanking purposes, and potential application of ART.