Biotic and abiotic drivers of dispersion dynamics in a large-bodied tropical vertebrate, the Western Bornean orangutan.

Understanding of animal responses to dynamic resource landscapes is based largely on research on temperate species with small body sizes and fast life histories. We studied a large, tropical mammal with an extremely slow life history, the Western Bornean orangutan (Pongo pygmaeus wurmbii), across a heterogeneous natural landscape encompassing seven distinct forest types. Our goals were to characterize fluctuations in abundance, test hypotheses regarding the relationship between dispersion dynamics and resource availability, and evaluate how movement patterns are influenced by abiotic conditions. We surveyed abundance in Gunung Palung National Park, West Kalimantan, Indonesia, for 99 consecutive months and simultaneously recorded weather data and assessed fruit availability. We developed a Bayesian hierarchical distance sampling model to estimate population dispersion and assess the roles of fruit availability, rainfall, and temperature in driving movement patterns across this heterogeneous landscape. Orangutan abundance varied dramatically over space and time. Each forest type was important in sustaining more than 40% of the total orangutans on site during at least one month, as animals moved to track asynchronies in fruiting phenology. We conclude that landscape-level movements buffer orangutans against fruit scarcity, peat swamps are crucial fallback habitats, and orangutans' use of high elevation forests is strongly dependent on abiotic conditions. Our results show that orangutans can periodically occupy putative-sink habitats and be virtually absent for extended periods from habitats that are vitally important in sustaining their population, highlighting the need for long-term studies and potential risks in interpreting occurrence or abundance measures as indicators of habitat importance.

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.

Flanged males have higher reproductive success in a completely wild orangutan population.

Male orangutans (Pongo spp.) exhibit bimaturism, an alternative reproductive tactic, with flanged and unflanged males displaying two distinct morphological and behavioral phenotypes. Flanged males are larger than unflanged males and display secondary sexual characteristics which unflanged males lack. The evolutionary explanation for alternative reproductive tactics in orangutans remains unclear because orangutan paternity studies to date have been from sites with ex-captive orangutans, provisioning via feeding stations and veterinary care, or that lack data on the identity of mothers. Here we demonstrate, using the first long-term paternity data from a site free of these limitations, that alternative reproductive tactics in orangutans are condition-dependent, not frequency-dependent. We found higher reproductive success by flanged males than by unflanged males, a pattern consistent with other Bornean orangutan (Pongo pygmaeus) paternity studies. Previous paternity studies disagree on the degree of male reproductive skew, but we found low reproductive skew among flanged males. We compare our findings and previous paternity studies from both Bornean and Sumatran orangutans (Pongo abelii) to understand why these differences exist, examining the possible roles of species differences, ecology, and human intervention. Additionally, we use long-term behavioral data to demonstrate that while flanged males can displace unflanged males in association with females, flanged males are unable to keep other males from associating with a female, and thus they are unable to completely mate guard females. Our results demonstrate that alternative reproductive tactics in Bornean orangutans are condition-dependent, supporting the understanding that the flanged male morph is indicative of good condition. Despite intense male-male competition and direct sexual coercion by males, female mate choice is effective in determining reproductive outcomes in this population of wild orangutans.

Possible Male Infanticide in Wild Orangutans and a Re-evaluation of Infanticide Risk.

Infanticide as a male reproductive tactic is widespread across mammals, and is particularly prevalent in catarrhine primates. While it has never been observed in wild orangutans, infanticide by non-sire males has been predicted to occur due to their extremely long inter-birth intervals, semi-solitary social structure, and the presence of female counter-tactics to infanticide. Here, we report on the disappearance of a healthy four-month-old infant, along with a serious foot injury suffered by the primiparous mother. No other cases of infant mortality have been observed at this site in 30 years of study. Using photographic measurements of the injury, and information on the behavior and bite size of potential predators, we evaluate the possible causes of this injury. The context, including the behavior of the female and the presence of a new male at the time of the injury, lead us to conclude that the most likely cause of the infant loss and maternal injury was male infanticide. We suggest that in orangutans, and other species where nulliparous females are not preferred mates, these females may be less successful at using paternity confusion as an infanticide avoidance tactic, thus increasing the likelihood of infanticide of their first-born infants.