The demise of the giant ape Gigantopithecus blacki.

The largest ever primate and one of the largest of the southeast Asian megafauna, Gigantopithecus blacki1, persisted in China from about 2.0 million years until the late middle Pleistocene when it became extinct2-4. Its demise is enigmatic considering that it was one of the few Asian great apes to go extinct in the last 2.6 million years, whereas others, including orangutan, survived until the present5. The cause of the disappearance of G. blacki remains unresolved but could shed light on primate resilience and the fate of megafauna in this region6. Here we applied three multidisciplinary analyses-timing, past environments and behaviour-to 22 caves in southern China. We used 157 radiometric ages from six dating techniques to establish a timeline for the demise of G. blacki. We show that from 2.3 million years ago the environment was a mosaic of forests and grasses, providing ideal conditions for thriving G. blacki populations. However, just before and during the extinction window between 295,000 and 215,000 years ago there was enhanced environmental variability from increased seasonality, which caused changes in plant communities and an increase in open forest environments. Although its close relative Pongo weidenreichi managed to adapt its dietary preferences and behaviour to this variability, G. blacki showed signs of chronic stress and dwindling populations. Ultimately its struggle to adapt led to the extinction of the greatest primate to ever inhabit the Earth.

The variation and evolution of complete human centromeres.

Human centromeres have been traditionally very difficult to sequence and assemble owing to their repetitive nature and large size1. As a result, patterns of human centromeric variation and models for their evolution and function remain incomplete, despite centromeres being among the most rapidly mutating regions2,3. Here, using long-read sequencing, we completely sequenced and assembled all centromeres from a second human genome and compared it to the finished reference genome4,5. We find that the two sets of centromeres show at least a 4.1-fold increase in single-nucleotide variation when compared with their unique flanks and vary up to 3-fold in size. Moreover, we find that 45.8% of centromeric sequence cannot be reliably aligned using standard methods owing to the emergence of new α-satellite higher-order repeats (HORs). DNA methylation and CENP-A chromatin immunoprecipitation experiments show that 26% of the centromeres differ in their kinetochore position by >500 kb. To understand evolutionary change, we selected six chromosomes and sequenced and assembled 31 orthologous centromeres from the common chimpanzee, orangutan and macaque genomes. Comparative analyses reveal a nearly complete turnover of α-satellite HORs, with characteristic idiosyncratic changes in α-satellite HORs for each species. Phylogenetic reconstruction of human haplotypes supports limited to no recombination between the short (p) and long (q) arms across centromeres and reveals that novel α-satellite HORs share a monophyletic origin, providing a strategy to estimate the rate of saltatory amplification and mutation of human centromeric DNA.

A high-quality bonobo genome refines the analysis of hominid evolution.

The divergence of chimpanzee and bonobo provides one of the few examples of recent hominid speciation1,2. Here we describe a fully annotated, high-quality bonobo genome assembly, which was constructed without guidance from reference genomes by applying a multiplatform genomics approach. We generate a bonobo genome assembly in which more than 98% of genes are completely annotated and 99% of the gaps are closed, including the resolution of about half of the segmental duplications and almost all of the full-length mobile elements. We compare the bonobo genome to those of other great apes1,3-5 and identify more than 5,569 fixed structural variants that specifically distinguish the bonobo and chimpanzee lineages. We focus on genes that have been lost, changed in structure or expanded in the last few million years of bonobo evolution. We produce a high-resolution map of incomplete lineage sorting and estimate that around 5.1% of the human genome is genetically closer to chimpanzee or bonobo and that more than 36.5% of the genome shows incomplete lineage sorting if we consider a deeper phylogeny including gorilla and orangutan. We also show that 26% of the segments of incomplete lineage sorting between human and chimpanzee or human and bonobo are non-randomly distributed and that genes within these clustered segments show significant excess of amino acid replacement compared to the rest of the genome.

Elemental signatures of Australopithecus africanus teeth reveal seasonal dietary stress.

Reconstructing the detailed dietary behaviour of extinct hominins is challenging1-particularly for a species such as Australopithecus africanus, which has a highly variable dental morphology that suggests a broad diet2,3. The dietary responses of extinct hominins to seasonal fluctuations in food availability are poorly understood, and nursing behaviours even less so; most of the direct information currently available has been obtained from high-resolution trace-element geochemical analysis of Homo sapiens (both modern and fossil), Homo neanderthalensis4 and living apes5. Here we apply high-resolution trace-element analysis to two A. africanus specimens from Sterkfontein Member 4 (South Africa), dated to 2.6-2.1 million years ago. Elemental signals indicate that A. africanus infants predominantly consumed breast milk for the first year after birth. A cyclical elemental pattern observed following the nursing sequence-comparable to the seasonal dietary signal that is seen in contemporary wild primates and other mammals-indicates irregular food availability. These results are supported by isotopic evidence for a geographical range that was dominated by nutritionally depauperate areas. Cyclical accumulation of lithium in A. africanus teeth also corroborates the idea that their range was characterized by fluctuating resources, and that they possessed physiological adaptations to this instability. This study provides insights into the dietary cycles and ecological behaviours of A. africanus in response to food availability, including the potential cyclical resurgence of milk intake during times of nutritional challenge (as observed in modern wild orangutans5). The geochemical findings for these teeth reinforce the unique place of A. africanus in the fossil record, and indicate dietary stress in specimens that date to shortly before the extinction of Australopithecus in South Africa about two million years ago.

Evaluating the muscle attachment hypothesis for sagittal cresting in Gorilla and Pongo.

Primate mandibular morphology is often associated with jaw functionality of the masticatory complex in the context of variation in diets. Recent research into the disparities between the diet and jaw functionality in male and female hominoids is inconclusive and suggests that sexual dimorphism in the mandible may be influenced by external factors such as temporalis and masseter muscle morphology, which in turn may be influenced by sexual selection. As the muscles associated with mastication (i.e., the type of chewing exhibited by primates and other mammals) encompass the mandible as well as the neurocranium, including the sagittal crest among some individuals, this study investigates sex-specific associations between regions of the mandibular ramus and neurocranium associated with mastication in a dentally mature sample of Gorilla and Pongo. A total of four cranial and mandibular variables were measured in two Gorilla taxa (Gorilla gorilla gorilla and Gorilla beringei graueri) and one Pongo taxon (Pongo pygmaeus pygmaeus) (n = 220). For all three taxa, we investigate (a) whether the degree of sexual dimorphism in cranial regions associated with sagittal cresting (sagittal crest size (SCS) and temporalis muscle attachment area (TMAA)) is proportional to the degree of mandibular ramus area (MRA) and coronoid process height (CPH) sexual dimorphism, (b) whether there are sex differences in scaling relationships between TMAA and MRA, and (c) whether there are sex differences in the strength of association between TMAA and CPH. We show that for G. g. gorilla, variables associated with sagittal cresting show higher sexual dimorphism values than our two mandibular ramus variables, which is not the case for G. b. graueri or for P. p. pygmaeus. All three taxa show similar sex-specific scaling relationships between TMAA and MRA, where for males this relationship does not diverge from isometry, and for females there is a negative allometric relationship. Our findings also show intraspecific sex differences in allometric slopes between MRA and TMAA for all three taxa. Only G. g. gorilla shows a significant association between TMAA and CPH, which is observed in both sexes. Although there are some statistical associations between the cranial and mandibular regions associated with mastication, our results show that among male gorillas and orangutans, patterns of variation in the sagittal crest, TMAA, mandibular ramus and the coronoid process cannot be explained by the muscle attachment hypothesis alone. These findings have implications surrounding the associations between social behaviour and the morphology of the craniofacial complex.

Evidence for the smallest fossil Pongo in southern China.

The rarity of Pongo fossils with precise absolute dating from the Middle Pleistocene hampers our understanding of the taxonomy and spatiotemporal distribution of Quaternary orangutans in southern China. Here, we report a newly discovered sample of 113 isolated teeth of fossil Pongo from Zhongshan Cave in the Bubing Basin, Guangxi, southern China. We describe the Pongo specimens from Zhongshan Cave and compare them metrically to other samples of fossil Pongo species (i.e., Pongo weidenreichi, Pongo devosi, Pongo duboisi, Pongo palaeosumatrensis, Pongo javensis, and Pongo sp.) and to extant orangutans (i.e., Pongo pygmaeus and Pongo abelii). The Zhongshan Pongo assemblage is dated using U-series and coupled electron spin resonance/U-series methods. Our results reasonably constrain the Zhongshan Pongo assemblage to 184 ± 16 ka, which is consistent with the biostratigraphic evidence. The Zhongshan Pongo teeth are only 6.5% larger on average than those of extant Pongo. The Zhongshan teeth are smaller overall than those of Pongo from all other cave sites in southern China, and they currently represent the smallest fossil orangutans in southern China. Based on their dental size, and the presence of a well-developed lingual pillar and lingual cingulum on the upper and lower incisors, an intermediate frequency of lingual cingulum remnants on the upper molars, and a higher frequency of moderate to heavy wrinkling on the upper and lower molars, we provisionally assign the Zhongshan fossils to P. devosi. Our results confirm earlier claims that P. weidenreichi is replaced by a smaller species in southern China, P. devosi, by the late Middle Pleistocene. The occurrence of P. devosi in Zhongshan Cave further extends its spatial and temporal distribution. The Pongo specimens from Zhongshan provide important new evidence to demonstrate that the dental morphological features of Pongo in southern China changed substantially during the late Middle Pleistocene.

Immature wild orangutans acquire relevant ecological knowledge through sex-specific attentional biases during social learning.

As a part of growing up, immature orangutans must acquire vast repertoires of skills and knowledge, a process that takes several years of observational social learning and subsequent practice. Adult female and male orangutans show behavioral differences including sex-specific foraging patterns and male-biased dispersal. We investigated how these differing life trajectories affect social interest and emerging ecological knowledge in immatures. We analyzed 15 years of detailed observational data on social learning, associations, and diet repertoires of 50 immatures (16 females and 34 males), from 2 orangutan populations. Specific to the feeding context, we found sex differences in the development of social interest: Throughout the dependency period, immature females direct most of their social attention at their mothers, whereas immature males show an increasing attentional preference for individuals other than their mothers. When attending to non-mother individuals, males show a significant bias toward immigrant individuals and a trend for a bias toward adult males. In contrast, females preferentially attend to neighboring residents. Accordingly, by the end of the dependency period, immature females show a larger dietary overlap with their mothers than do immature males. These results suggest that immature orangutans show attentional biases through which they learn from individuals with the most relevant ecological knowledge. Diversifying their skills and knowledge likely helps males when they move to a new area. In sum, our findings underline the importance of fine-grained social inputs for the acquisition of ecological knowledge and skills in orangutans and likely in other apes as well.

Dental findings in wild great apes from macerated skull analysis.

Oral health is a crucial aspect of overall well-being in both humans and nonhuman primates. Understanding the oral pathologies and dental conditions in apes can provide valuable insights into their evolutionary history, dietary habits, and overall health. The present study evaluates dental findings in wild great apes from museum specimens to gain insights into the influence of natural nutrition on dental health. Complete macerated skulls of wild, adult great apes from the collection of the Museum of Natural History, Berlin, Germany, were examined. We analyzed skulls of 53 gorillas (Gorilla gorilla), 63 chimpanzees (Pan troglodytes), and 41 orangutans (Pongo spp.). For each skull, we recorded wear of dental hard tissues (Lussi and Ganss index), carious lesions, and periodontal bone loss. Incisal and occlusal dental hard tissue defects were found in all skulls, as well as considerable external staining. In all species, incisors and canines showed the greatest loss of tissue, followed by molars. The wear of molars decreased from the first to the third molars, premolars showed the least pronounced defects. Some individuals had apical osteolytic defects along with severe dental hard tissue loss with pulp involvement or after dental trauma, respectively (n = 5). Our study did not observe any carious lesions among the examined great ape skulls. However, we did find evidence for localized or generalized periodontal bone loss in a subset of the specimens (n = 3 chimpanzees, n = 7 orangutans). The natural diet and foraging behavior of great apes induces abrasion and attrition of dental hard tissue but does not yield carious lesions. The occurrence of periodontitis in individual apes indicates that the natural circumstances can induce periodontal bone loss even in the wild, despite physiological nutrition.

Novel Cascade Alpha Satellite HORs in Orangutan Chromosome 13 Assembly: Discovery of the 59mer HOR-The largest Unit in Primates-And the Missing Triplet 45/27/18 HOR in Human T2T-CHM13v2.0 Assembly.

From the recent genome assembly NHGRI_mPonAbe1-v2.0_NCBI (GCF_028885655.2) of orangutan chromosome 13, we computed the precise alpha satellite higher-order repeat (HOR) structure using the novel high-precision GRM2023 algorithm with Global Repeat Map (GRM) and Monomer Distance (MD) diagrams. This study rigorously identified alpha satellite HORs in the centromere of orangutan chromosome 13, discovering a novel 59mer HOR-the longest HOR unit identified in any primate to date. Additionally, it revealed the first intertwined sequence of three HORs, 18mer/27mer/45mer HORs, with a common aligned "backbone" across all HOR copies. The major 7mer HOR exhibits a Willard's-type canonical copy, although some segments of the array display significant irregularities. In contrast, the 14mer HOR forms a regular Willard's-type HOR array. Surprisingly, the GRM2023 high-precision analysis of chromosome 13 of human genome assembly T2T-CHM13v2.0 reveals the presence of only a 7mer HOR, despite both the orangutan and human genome assemblies being derived from whole genome shotgun sequences.

USE OF HYSTEROSCOPIC MORCELLATION WITH A MYOSURE® DEVICE FOR REMOVAL OF UTERINE FOREIGN MATERIAL IN AN ORANGUTAN (PONGO SP.).

A 36-yr-old female intact hybrid orangutan (Pongo sp.) presented with evidence of chronic bacterial cystitis with no response to treatment with multiple systemic antibiotics. Abnormalities were identified within the reproductive tract on CT scan, and hysteroscopy was pursued based on the recommendation of an obstetrician-gynecologist. Hysteroscopic examination revealed a large amount of intrauterine foreign material consistent with wood wool from the orangutan's bedding. A hysteroscopic morcellator (MyoSure®) was used to remove the foreign material. Histologic examination of endometrial biopsies showed severe suppurative to pyogranulomatous endometritis with intralesional foreign (wood) fibers and bacteria. The orangutan was treated with antibiotics, but evidence of bacterial cystitis persisted. After 15 wk, additional wood wool was identified within the uterus by hysteroscopic examination and was removed similarly. Five months later, there was no recurrence of foreign material on hysteroscopy or CT. This report describes the first documentation of intrauterine foreign material in a nonhuman primate. Hysteroscopic morcellation is a useful technique for noninvasive removal of intrauterine foreign material in great apes.

Cortical bone distribution of the proximal phalanges in great apes: implications for reconstructing manual behaviours.

Primate fingers are typically in direct contact with the environment during both locomotion and manipulation, and aspects of external phalangeal morphology are known to reflect differences in hand use. Since bone is a living tissue that can adapt in response to loading through life, the internal bone architecture of the manual phalanges should also reflect differences in manual behaviours. Here, we use the R package Morphomap to analyse high-resolution microCT scans of hominid proximal phalanges of digits 2-5 to determine whether cortical bone structure reflects variation in manual behaviours between bipedal (Homo), knuckle-walking (Gorilla, Pan) and suspensory (Pongo) taxa. We test the hypothesis that relative cortical bone distribution patterns and cross-sectional geometric properties will differ both among extant great apes and across the four digits due to locomotor and postural differences. Results indicate that cortical bone structure reflects the varied hand postures employed by each taxon. The phalangeal cortices of Pongo are significantly thinner and have weaker cross-sectional properties relative to the African apes, yet thick cortical bone under their flexor sheath ridges corresponds with predicted loading during flexed finger grips. Knuckle-walking African apes have even thicker cortical bone under the flexor sheath ridges, as well as in the region proximal to the trochlea, but Pan also has thicker diaphyseal cortices than Gorilla. Humans display a distinct pattern of distodorsal thickening, as well as relatively thin cortices, which may reflect the lack of phalangeal curvature combined with frequent use of flexed fingered hand grips during manipulation. Within each taxon, digits 2-5 have a similar cortical distribution in Pongo, Gorilla and, unexpectedly, Homo, which suggest similar loading of all fingers during habitual locomotion or hand use. In Pan, however, cortical thickness differs between the fingers, potentially reflecting differential loading during knuckle-walking. Inter- and intra-generic variation in phalangeal cortical bone structure reflects differences in manual behaviours, offering a comparative framework for reconstructing hand use in fossil hominins.

Scapular morphology of great apes and humans: A three-dimensional computed tomography-based comparative study.

The primate scapula has been studied widely since its shape has been shown to correlate with how the forelimb is used in daily activities. In this study, we expand on the existing literature and use an image-based methodology that was originally developed for orthopaedic practice to quantify and compare the three-dimensional (3D) morphology of the scapula across humans and great apes. We expect that this image-based approach will allow us to identify differences between great apes and humans that can be related to differences in mobility and loading regime of the shoulder. We hypothesize that gorillas and chimpanzees will have a similar scapular morphology, geared towards stability and weight-bearing in knuckle-walking, whilst the scapular morphology of orangutans is expected to be more similar to that of humans given their high glenohumeral mobility associated with their suspensory lifestyle. We made 3D reconstructions of computed tomography scans of 69 scapulae from four hominid genera (Pongo, Gorilla, Pan and Homo). On these 3D bone meshes, the inferior glenoid plane was determined, and subsequently, a set of bony landmarks on the scapular body, coracoid, and acromion were defined. These landmarks allowed us to measure a set of functionally relevant angles which represent acromial overhang, subacromial space and coracoacromial space. The angles that were measured are: the delto-fulcral triangle (DFT), comprising the alpha, beta, and delta angle, the acromion-glenoid angle (AGA), the coracoid-glenoid centre-posterior acromial angle (CGA), the anterior tilt (TA CGA) and the posterior tilt of the CGA (PT CGA). Three observers placed the landmarks on the 3D bone meshes, allowing us to calculate the inter-observer error. The main differences in the DFT were found between humans and the great apes, with small differences between the great apes. The DFT of humans was significantly lower compared to that of the great apes, with the smallest alpha (32.7°), smallest delta (45.7°) and highest beta angle (101.6°) of all genera. The DFT of chimpanzees was significantly higher compared to that of humans (p < 0.01), with a larger alpha (37.6°) and delta angle (54.5°) and smaller beta angle (87.9°). The mean AGA of humans (59.1°) was significantly smaller (p < 0.001) than that of gorillas (68.8°). The mean CGA of humans (110.1°) was significantly higher (p < 0.001) than in orangutans (92.9°). Humans and gorillas showed mainly a posterior tilt of their coracoacromial complex whilst chimpanzees showed mainly an anterior tilt. The coracoacromial complex of the orangutans was not tilted anteriorly or posteriorly. With our image-based method, we were able to identify morphological features of the scapula that differed significantly between hominid genera. However, we did not find an overall dichotomy in scapular morphology geared towards high stability (Pan/Gorilla) or high mobility (Homo/Pongo). Further research is needed to investigate the functional implications of these differences in scapular morphology.

Middle Pleistocene Pongo from Ganxian Cave in southern China with implications for understanding dental size evolution in orangutans.

The Pongo fossil record of China extends from the Early Pleistocene to the Late Pleistocene, but to date, no late Middle Pleistocene samples of Pongo with precise absolute dating have been identified in southern China. Here, we report the recovery of 106 fossil teeth of Pongo from Ganxian Cave in the Bubing Basin, Guangxi, southern China. We dated the speleothems using Uranium-series and dated the two rhinoceros teeth using coupled electron spin resonance/Uranium-series dating methods to between 168.9 ± 2.4 ka and 362 ± 78 ka, respectively. These dates are consistent with the biostratigraphic and magnetostratigraphic age estimates. We further describe the fossil teeth from Ganxian Cave and compare them metrically to samples of fossil Pongo (i.e., Pongo weidenreichi, Pongo duboisi, Pongo palaeosumatrensis, Pongo javensis, and Pongo sp.) from the Early, Middle, and Late Pleistocene and to extant Pongo (i.e., Pongo pygmaeus and Pongo abelii) from Southeast Asia. Based on overall dental size, a high frequency of lingual cingulum remnants on the upper molars, and a low frequency of moderate to heavy wrinkling on the molars, we attribute the Ganxian fossils to P. weidenreichi. Compared with Pongo fossils from other mainland Southeast Asia sites, those from Ganxian confirm that dental size reduction of Pongo occurred principally during the Early and Middle Pleistocene. From the Middle to Late Pleistocene, all teeth except the P3 show little change in occlusal area, indicating that the size of these teeth remained relatively stable over time. The evolutionary trajectory of the Pongo dentition through time may be more complex than previously thought. More orangutan fossils with precise dating constraints are the keys to solving this issue.

Parental influence on human germline de novo mutations in 1,548 trios from Iceland.

The characterization of mutational processes that generate sequence diversity in the human genome is of paramount importance both to medical genetics and to evolutionary studies. To understand how the age and sex of transmitting parents affect de novo mutations, here we sequence 1,548 Icelanders, their parents, and, for a subset of 225, at least one child, to 35× genome-wide coverage. We find 108,778 de novo mutations, both single nucleotide polymorphisms and indels, and determine the parent of origin of 42,961. The number of de novo mutations from mothers increases by 0.37 per year of age (95% CI 0.32-0.43), a quarter of the 1.51 per year from fathers (95% CI 1.45-1.57). The number of clustered mutations increases faster with the mother's age than with the father's, and the genomic span of maternal de novo mutation clusters is greater than that of paternal ones. The types of de novo mutation from mothers change substantially with age, with a 0.26% (95% CI 0.19-0.33%) decrease in cytosine-phosphate-guanine to thymine-phosphate-guanine (CpG>TpG) de novo mutations and a 0.33% (95% CI 0.28-0.38%) increase in C>G de novo mutations per year, respectively. Remarkably, these age-related changes are not distributed uniformly across the genome. A striking example is a 20 megabase region on chromosome 8p, with a maternal C>G mutation rate that is up to 50-fold greater than the rest of the genome. The age-related accumulation of maternal non-crossover gene conversions also mostly occurs within these regions. Increased sequence diversity and linkage disequilibrium of C>G variants within regions affected by excess maternal mutations indicate that the underlying mutational process has persisted in humans for thousands of years. Moreover, the regional excess of C>G variation in humans is largely shared by chimpanzees, less by gorillas, and is almost absent from orangutans. This demonstrates that sequence diversity in humans results from evolving interactions between age, sex, mutation type, and genomic location.

Orangutans and the evolution of sharp stone tools.

Orangutans use stone tools in a variety of modes, including cutting. This behavior appears to be learned from trusted social partners.

Mother-offspring proximity maintenance as an infanticide avoidance strategy in bornean orangutans (Pongo pygmaeus wurmbii).

Sexually-selected infanticide by males is widespread across primates. Maternal protection is one of many infanticide avoidance strategies employed by female primates. Bornean orangutan (Pongo pygmaeus wurmbii) mothers with younger offspring are less social with males than mothers with older offspring. Additionally, the distance between a mother and offspring decreases in the presence of male conspecifics, but not female conspecifics. We hypothesized that mothers are responsible for the change in mother-offspring proximity when males are present. Using a year of behavioral data from orangutans in Gunung Palung National Park, we tested whether the Hinde Index, a ratio of the number of approaches and leaves between two individuals, was indicative of mother or offspring proximity maintenance across different social groupings. The semi-solitary social organization of orangutans allows us to observe different social groupings. We found that the mother-offspring Hinde Index was typically indicative of offspring maintenance of proximity. However, the presence of male conspecifics was associated with an increase in the Hinde Index which indicates that mothers are responsible for the decrease in mother-offspring distance when males are present. The decrease in mother-offspring distances and increase in Hinde Index when males are present indicates that mothers react to the presence of males in a protective manner. We suggest this may be an infanticide avoidance behavior by mother orangutans.

A composite menstrual cycle of captive orangutans, with associated hormonal and behavioral variability.

Knowledge of species-typical reproductive endocrinology profiles is crucial for testing hypotheses pertaining to the evolutionary history, reproductive parameters, and life history of a species, and for managing the well-being of individual animals in human care. Large-scale empirical measurements of ovarian hormones, however, are rare for most primate species, including orangutans. In this study, we used enzyme immunoassays (EIA) to quantify estrogen (estrone conjugates; E1 C) and progesterone (pregnanediol-3-glucuronide; PdG) levels for 98 cycles in 7 cycling zoo-housed female orangutans (10-43 years old). We use a subset of these cycles (N = 44) to create the first composite menstrual cycle for orangutans, which serves as a valuable baseline for future comparative analyses and veterinary considerations. Similar to previous studies, we determined the mean ovarian cycle length of orangutans to be 29.7 days (N = 98 cycles), although we illustrate evidence of both intra- and interindividual variation in ovarian steroid production. Given that this study took place in captivity, we consider how energetic and psychosocial aspects of the zoo environment, such as greater food availability and potential stress, may affect the reproductive physiology and sexual behavior of these females. Furthermore, we discuss the role that age and genetic background may play in producing variability. Finally, we test whether ovarian hormone levels correlate with the reproductive behaviors of these female orangutans using associated behavioral data. Our results suggest that matings are more common during the periovulatory period than outside of it, but do not support a consistent link between hormonal indices of fecundability and mating behaviors in these individuals.