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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.

Current approaches to quantify phalangeal curvature assume that the long axis of the bone's diaphysis approximates the shape of a portion of a circle (included angle method) or a parabola (second‐degree polynomial method). Here we developed, tested, and employed an alternative geometric morphometrics‐based (GM) approach to quantify diaphysis shape of proximal phalanges in humans, apes and monkeys with diverse locomotor behaviors. One hundred landmarks of the central longitudinal axis were extracted from 3D surface models and analyzed using 2DGM methods, including generalized Procrustes analyses. Principal components analyses were performed and PC1 scores (>80% of variation) represented the dorsopalmar shape of the bone's central longitudinal axis and separated taxa consistently and in accord with known locomotor behavioral profiles. The most suspensory taxa, including orangutans, hylobatids and spider monkeys, had significantly lower PC1 scores reflecting the greatest amounts of phalangeal curvature. In contrast, bipedal humans and the quadrupedal cercopithecoid monkeys sampled (baboons, proboscis monkeys) exhibited significantly higher PC1 scores reflecting flatter phalanges. African ape (gorillas, chimpanzees and bonobos) phalanges fell between these two extremes and were not significantly different from each other. PC1 scores were significantly correlated with both included angle and theacoefficient of a second‐degree polynomial calculated from the same landmark dataset, but had a significantly higher correlation with included angles. Our alternative approach for quantifying diaphysis shape of proximal phalanges to investigate dorsopalmar curvature is replicable and does not assume a priori either a circle or parabola model of shape, making it an attractive alternative compared with existing methodologies.