Among extant great apes, orangutans are considered the most sexually dimorphic in body size. However, the expression of sexual dimorphism in orangutans is more complex than simply males being larger than females. At sexual maturity, some male orangutans develop cheek pads (flanges), while other males remain unflanged even after becoming reproductively capable. Sometimes flange development is delayed in otherwise sexually mature males for a few years. In other cases, flange development is delayed for many years or decades, with some males even spending their entire lifespan as unflanged adults. Thus, unflanged males of various chronological ages can be mistakenly identified as “subadults.” Unflanged adult males are typically described as “female-sized,” but this may simply reflect the fact that unflanged male body size has only ever been measured in peri-pubescent individuals. In this study, we measured the skeletons of 111 wild adult orangutans (Pongo spp.), including 20 unflanged males, 45 flanged males, and 46 females, resulting in the largest skeletal sample of unflanged males yet studied. We assessed long bone lengths (as a proxy for stature) for all 111 individuals and recorded weights-at-death, femoral head diameters, bi-iliac breadths, and long bone cross-sectional areas (CSA) (as proxies for mass) for 27 of these individuals, including seven flanged males, three adult confirmed-unflanged males, and three young adult likely-unflanged males. ANOVA and Kruskal–Wallis tests with Tukey and Dunn post-hoc pairwise comparisons, respectively, showed that body sizes for young adult unflanged males are similar to those of the adult females in the sample (all P ≥ 0.09 except bi-iliac breadth), whereas body sizes for adult unflanged males ranged between those of adult flanged males and adult females for several measurements (all P < 0.001). Thus, sexually mature male orangutans exhibit body sizes that range from the female end of the spectrum to the flanged male end of the spectrum. These results exemplify that the term “sexual dimorphism” fails to capture the full range of variation in adult orangutan body size. By including adult unflanged males in analyses of body size and other aspects of morphology, not as aberrations but as an expected part of orangutan variation, we may begin to shift the way that we think about features typically considered dichotomous according to biological sex.
- Award ID(s):
- 1638823
- NSF-PAR ID:
- 10320158
- Date Published:
- Journal Name:
- Northeastern Evolutionary Primatologists
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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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.
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null (Ed.)Due to extreme incongruence in parental investment, the potential for sexual conflict in orangutan reproductive strategies is high. Female orangutans prefer flanged males, but with intense male-male competition and sexual coercion, it is unclear how females are able to exert choice. We hypothesized that female orangutans use initiation and maintenance of associations with preferred males as a mechanism of female choice. We used encounter rates and behavioral measures of proximity maintenance to distinguish between the role of female choice and male coercion in male-female associations in Gunung Palung National Park, West Kalimantan, Indonesia. From May 2018-April 2019, we collected data on initiation, termination, and proximity maintenance during associations between males and cycling females (females without dependent offspring or with offspring over 6 years old). Encounters between cycling females and flanged males were more likely to be female-initiated (83.3%), while encounters with unflanged males were more likely to be male-initiated (80%) (N=16, p=0.035, Fisher’s exact test). Additionally, the Brown Index indicates significantly more female proximity maintenance when cycling females associated with flanged males than unflanged males (N=33, W = 198.5, p= 0.012). However, using long-term data on male-female associations, we found that dyads of flanged males and females were less likely to encounter other males compared to dyads of unflanged males and females (N=542, χ2= 3.3051, df=1, p=0.035). This indicates that flanged males may also use associations to mate guard females. These data indicate that there are behavioral manifestations of both female choice and male coercion in orangutan associations. Funders: NSF (DGE-1247312, BCS-1638823, BCS-0936199), Boston University, Leakey Foundation, Disney Conservation Fund, and US Fish and Wildlife Service (F18AP00898)more » « less
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null (Ed.)Due to extreme incongruence in parental investment, the potential for sexual conflict in orangutan reproductive strategies is high. Female orangutans prefer flanged males, but with intense male-male competition and sexual coercion, it is unclear how females are able to exert choice. We hypothesized that female orangutans use initiation and maintenance of associations with preferred males as a mechanism of female choice. We used encounter rates and behavioral measures of proximity maintenance to distinguish between the role of female choice and male coercion in male-female associations in Gunung Palung National Park, West Kalimantan, Indonesia. From May 2018-April 2019, we collected data on initiation, termination, and proximity maintenance during associations between males and cycling females (females without dependent offspring or with offspring over 6 years old). Encounters between cycling females and flanged males were more likely to be female-initiated (83.3%), while encounters with unflanged males were more likely to be male-initiated (80%) (N=16, p=0.035, Fisher’s exact test). Additionally, the Brown Index indicates significantly more female proximity maintenance when cycling females associated with flanged males than unflanged males (N=33, W = 198.5, p= 0.012). However, using long-term data on male-female associations, we found that dyads of flanged males and females were less likely to encounter other males compared to dyads of unflanged males and females (N=542, χ2= 3.3051, df=1, p=0.035). This indicates that flanged males may also use associations to mate guard females. These data indicate that there are behavioral manifestations of both female choice and male coercion in orangutan associations. Funders: NSF (DGE-1247312, BCS-1638823, BCS-0936199), Boston University, Leakey Foundation, Disney Conservation Fund, and US Fish and Wildlife Service (F18AP00898)more » « less
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Male orangutans exhibit bimaturism—two mature morphs—flanged and unflanged males. Flanged males are larger, have cheek pads (flanges) and large throat sacs, and produce long calls. Previous orangutan paternity studies found variation between the reproductive success of each morph and in the degrees of reproductive skew. However, these studies were limited by a lack of behavioral maternity data, the inclusion of ex-captive orangutans, and/or the presence of feeding stations. Here we present the first paternity data from completely wild orangutans with known mothers. We hypothesized that (1) flanged males would have higher reproductive success than unflanged males due to flanged male dominance and female preference and (2) a single male would not monopolize paternity due to the temporal and spatial distribution of fecund females. We used fecal samples collected in Gunung Palung National Park from 2008-2019 to genotype orangutans (13 offspring born 2002-2015, their 10 mothers, and 19 candidate sires) using 12 microsatellites. MICROCHECKER 2.2.3 and CERVUS 3.0 were used to confirm the suitability of the microsatellite panel, fidelity of individual identities, and genetic maternity. Paternity analysis was performed with both CERVUS 3.0 and COLONY 2.0.6.7. We were able to identify paternity for six offspring. Four flanged males sired five offspring, and one sire’s morph was unknown at the time of conception. We found that flanged males have higher reproductive success and that females are not monopolizable in this completely wild setting. We discuss the implications of all published orangutan paternity results for understanding bimaturism in orangutans.more » « less
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