Female reproductive aging in seven primate species: Patterns and consequences.

Age-related changes in fertility have increasingly been documented in wild animal populations: In many species the youngest and oldest reproducers are disadvantaged relative to prime adults. How do these effects evolve, and what explains their diversity across species? Tackling this question requires detailed data on patterns of age-related reproductive performance in multiple animal species. Here, we compare patterns and consequences of age-related changes in female reproductive performance in seven primate populations that have been subjects of long-term continuous study for 29 to 57 y. We document evidence of age effects on fertility and on offspring performance in most, but not all, of these primate species. Specifically, females of six species showed longer interbirth intervals in the oldest age classes, youngest age classes, or both, and the oldest females also showed relatively fewer completed interbirth intervals. In addition, five species showed markedly lower survival among offspring born to the oldest mothers, and two species showed reduced survival for offspring born to both the youngest and the oldest mothers. In contrast, we found mixed evidence that maternal age affects the age at which daughters first reproduce: Only in muriquis and to some extent in chimpanzees, the only two species with female-biased dispersal, did relatively young mothers produce daughters that tended to have earlier first reproduction. Our findings demonstrate shared patterns as well as contrasts in age-related changes in female fertility across species of nonhuman primates and highlight species-specific behavior and life-history patterns as possible explanations for species-level differences.

Examining the dual hormone hypothesis in wild male mountain gorillas (Gorilla beringei beringei).

The Challenge Hypothesis is an influential framework for understanding how androgens are involved in the promotion of competitive behavior during mating-related challenges and has been tested extensively in studies across scientific disciplines. Mixed support in psychological research led scholars to develop the Dual Hormone Hypothesis as a potential path forward, which argues that glucocorticoids moderate the relationship between androgens and status-striving. In the current study, we examine the Challenge Hypothesis and the Dual Hormone Hypothesis in wild male mountain gorillas, representing the first time the latter hypothesis has been tested in a non-human primate. In a sample of 30 adult males comprising over 600 days of observation, we find some limited support for the Challenge Hypothesis. Greater daily rates of targeted aggression toward other adult males corresponded to higher fecal androgen metabolites 1-2 days following observations, though this pattern did not fully generalize to dominance rank or other competitive behaviors examined. However, we find no support for the Dual Hormone Hypothesis: neither dominance rank nor any category of competitive behavior was predicted by the interaction between androgens and glucocorticoids. We close by discussing how this initial investigation might be leveraged toward the development of an expanded Dual Hormone Hypothesis that draws on the large evidence base in primate behavioral ecology.

Maternal death and offspring fitness in multiple wild primates.

Primate offspring often depend on their mothers well beyond the age of weaning, and offspring that experience maternal death in early life can suffer substantial reductions in fitness across the life span. Here, we leverage data from eight wild primate populations (seven species) to examine two underappreciated pathways linking early maternal death and offspring fitness that are distinct from direct effects of orphaning on offspring survival. First, we show that, for five of the seven species, offspring face reduced survival during the years immediately preceding maternal death, while the mother is still alive. Second, we identify an intergenerational effect of early maternal loss in three species (muriquis, baboons, and blue monkeys), such that early maternal death experienced in one generation leads to reduced offspring survival in the next. Our results have important implications for the evolution of slow life histories in primates, as they suggest that maternal condition and survival are more important for offspring fitness than previously realized.

Multiple mechanisms for inbreeding avoidance used simultaneously in a wild ape.

Mating with close kin can have considerable negative fitness consequences, which are expected to result in selective pressure for inbreeding avoidance mechanisms, such as dispersal, mate choice and post-copulatory biases. Captive studies have suggested that inbreeding avoidance through mate choice is far less widespread than expected and may be absent where other mechanisms already limit inbreeding. However, few studies have examined multiple mechanisms of inbreeding avoidance simultaneously, particularly in the wild. We use 13 years of detailed dispersal, copulation and paternity data from mountain gorillas to examine inbreeding avoidance. We find that partial dispersal of both sexes results in high kinship in multimale groups, but that copulations between close kin occur 40% less than expected. We find strong kin discrimination in mate choice, with significant avoidance of maternal kin but more limited avoidance of paternal kin. We find no evidence for post-copulatory inbreeding avoidance. Our analyses support familiarity-based mechanisms of kin identification and age-based avoidance that limits mating between fathers and daughters in their natal group. Our findings demonstrate that multiple complementary mechanisms for inbreeding avoidance can evolve in a single species and suggest that inbreeding avoidance through mate choice may enable more flexible dispersal systems to evolve.

An empirical comparison of several commercial enzyme immunoassays for the non-invasive assessment of adrenocortical and gonadal function in mountain gorillas.

Wildlife researchers seeking to non-invasively examine endocrine function in their study species are presented with a dense and technical 'garden of forking paths' to navigate between collecting a biological sample and obtaining a final measurement. In particular, the choice of which enzyme immunoassay (EIA) to use with collected fecal samples, out of the many options offered by different manufacturers and research laboratories, may be one of the most consequential for final results. However, guidance for making this decision is still emerging. With this gap in mind, we performed a head-to-head comparison of results obtained from four different EIAs for fecal glucocorticoid metabolites (FGCMs), and three different EIAs for fecal androgen metabolites (FAMs), applied to the same set of fecal samples collected from the mountain gorillas (Gorilla beringei beringei) monitored by the Dian Fossey Gorilla Fund in Volcanoes National Park, Rwanda. We provide a) an analytical validation of the different EIAs via tests of parallelism and linearity; b) an estimate of inter-assay correlation between EIA kits designed for the same metabolites; and c) a test of the kits' ecological validity, in which we examine how well each captures endocrine changes following events that theory predicts should result in elevated FGCM and/or FAM concentrations. Our results show that kits differ to some degree in their performance; at the same time, nearly all assays exhibited at least moderate evidence of validity and covariance with others for the same analyte. Our findings, which differ somewhat from similar comparisons performed in other species, demonstrate the need to directly assess assay performance in a species- and context-specific manner as part of efforts to develop the burgeoning discipline of wildlife endocrinology.

Information seeking in western lowland gorillas (Gorilla gorilla gorilla).

Many animals will seek information when they do not know the answer to a problem, suggesting that they monitor their knowledge state. In the classic "tubes task," subjects are presented with a set of opaque tubes and either see (visible trials) or do not see (hidden trials) which tube holds a food reward on a given trial. Chimpanzees, bonobos, orangutans, and human children show information-seeking behavior on this task, looking into the tubes to find the reward more on hidden than on visible trials. However, evidence for information-seeking behavior in gorillas is limited. In Experiment 1, gorillas that were presented with a classic information-seeking tubes task showed performance patterns consistent with metacognitive behavior; they looked down tubes more on hidden than on visible trials, their accuracy on hidden trials on which they looked was higher than on hidden trials on which they did not look, and they primarily employed an appropriate search strategy when looking down the tubes. In Experiment 2, we decreased or increased the amount of effort required to look down the tubes by increasing or decreasing the height of the tubes, respectively. Gorillas were less likely to look in tubes on trials that required high effort, but continued to look more on hidden than on visible trials, indicating that their tendency to look was affected by both knowledge state and effort. Together these results provide strong evidence for logical, controlled information-seeking behavior by gorillas.

Facial asymmetry tracks genetic diversity among Gorilla subspecies.

Mountain gorillas are particularly inbred compared to other gorillas and even the most inbred human populations. As mountain gorilla skeletal material accumulated during the 1970s, researchers noted their pronounced facial asymmetry and hypothesized that it reflects a population-wide chewing side preference. However, asymmetry has also been linked to environmental and genetic stress in experimental models. Here, we examine facial asymmetry in 114 crania from three Gorilla subspecies using 3D geometric morphometrics. We measure fluctuating asymmetry (FA), defined as random deviations from perfect symmetry, and population-specific patterns of directional asymmetry (DA). Mountain gorillas, with a current population size of about 1000 individuals, have the highest degree of facial FA (explaining 17% of total facial shape variation), followed by Grauer gorillas (9%) and western lowland gorillas (6%), despite the latter experiencing the greatest ecological and dietary variability. DA, while significant in all three taxa, explains relatively less shape variation than FA does. Facial asymmetry correlates neither with tooth wear asymmetry nor increases with age in a mountain gorilla subsample, undermining the hypothesis that facial asymmetry is driven by chewing side preference. An examination of temporal trends shows that stress-induced developmental instability has increased over the last 100 years in these endangered apes.

Group structure, but not dominance rank, predicts fecal androgen metabolite concentrations of wild male mountain gorillas (Gorilla beringei beringei).

Androgens are important mediators of male-male competition in many primate species. Male gorillas' morphology is consistent with a reproductive strategy that relies heavily on androgen-dependent traits (e.g., extreme size and muscle mass). Despite possessing characteristics typical of species with an exclusively single-male group structure, multimale groups with strong dominance hierarchies are common in mountain gorillas. Theory predicts that androgens should mediate their dominance hierarchies, and potentially vary with the type of group males live in. We validated the use of a testosterone enzyme immunoassay (T-EIA R156/7, CJ Munro, UC-Davis) for use with mountain gorilla fecal material by (1) examining individual-level androgen responses to competitive events, and (2) isolating assay-specific hormone metabolites via high-performance liquid chromatography. Males had large (2.6- and 6.5-fold), temporary increases in fecal androgen metabolite (FAM) after competitive events, and most captured metabolites were testosterone or 5α-dihydrotestosterone-like androgens. We then examined the relationship between males' dominance ranks, group type, and FAM concentrations. Males in single-male groups had higher FAM concentrations than males in multimale groups, and a small pool of samples from solitary males suggested they may have lower FAM than group-living peers. However, data from two different time periods (n = 1610 samples) indicated there was no clear relationship between rank and FAM concentrations, confirming results from the larger of two prior studies that measured urinary androgens. These findings highlight the need for additional research to clarify the surprising lack of a dominance hierarchy/androgen relationship in mountain gorillas.

Western lowland gorillas (Gorilla gorilla gorilla) do not show an aversion to inequity in a token exchange task.

Although individuals in some species refuse foods they normally accept if their partner receives a more preferred one, this is not true across all species. The cooperation hypothesis proposes that this species-level variability evolved because inequity aversion is a mechanism to identify situations in which cooperation is not paying off, and that species regularly observed cooperating should be more likely to be averse to inequity. To rule out other potential explanations of inequity aversion, we need to test the converse as well: species rarely observed cooperating, especially those phylogenetically close to more cooperative species, should be less likely to be inequity averse. To this end, we tested eight zoo-housed Western lowland gorillas (Gorilla gorilla gorilla) on a token exchange task in which subjects received either the same food reward or a less-preferred reward for the same or more effort than their partner, recording both refusals to participate in the exchange and refusals to accept the reward. Supporting the cooperation hypothesis, even with procedural differences across sessions, gorillas were significantly more likely to refuse in all conditions in which they received a low-value food reward after completing an exchange, regardless of what their partner received, suggesting that gorillas were not inequity averse, but instead would not work for a low-value reward. Additionally, gorillas were more likely to refuse later in the session; while the pattern of refusals remained unchanged after accounting for this, this suggests that species should be tested on as many trials as is practical.

A partnership to build scientific capacity of Rwanda's future conservationists: The Memoirs Program.

The future of primates depends on conservationists in primate range countries having the education and opportunities needed to facilitate conservation efforts. However, most primates are found in countries where conservation is underfunded. Rwanda is home to 14 primate species, with three being listed as endangered on the International Union for Conservation of Nature (IUCN) Red List. Their remaining native habitats are scarce, highly fragmented, and surrounded by high human densities. Lacking education opportunities and the loss of qualified individuals following decades of political instability created a need for well-trained conservationists to tackle conservation challenges in Rwanda. The Memoirs Program, a science capacity-building partnership between Dian Fossey Gorilla Fund (DFGF), University of Rwanda (UR), the Rwanda Development Board, and Cleveland Metroparks Zoo, aims to train the next generation of Rwandan conservationists. Annually, the Memoirs Program provides 10-12 top students from the UR Conservation Biology undergraduate program the opportunity to conduct senior research projects (Memoirs) at DFGF addressing conservation issues in and around Rwanda's protected areas. Students receive one-on-one supervision by DFGF scientists and affiliates to supplement UR mentorship, along with targeted teaching modules that reinforce and complement UR curricula. In continuation of the Memoirs Program, participants are offered professional development, employment opportunities, and support to publish their research in peer-reviewed journals. Since inception of the Program in 2003, 123 students have participated with 83% being employed or pursuing higher education degrees in conservation-related fields in Rwanda and the region. Pre- and post-training comparison of oral presentation skills, one of the teaching modules, involving 22 students demonstrated increased post-training performance. The Memoirs Program represents an effective partnership between the government, an international conservation non-governmental organization, and a public higher-learning institution in a developing country to support high-performing students to become tomorrow's conservation leaders. This model could be replicated within Rwanda and in other regions of the world.

Daily Travel Distances of Unhabituated Grauer's Gorillas (Gorilla beringei graueri) in a Low Elevation Forest.

To accurately determine the space use of animals, we need to follow animal movements over prolonged periods, which is especially challenging for the critically endangered Grauer's gorillas (Gorilla beringei graueri) in eastern Democratic Republic of the Congo (DRC). As a consequence, we know little about Grauer's gorillas, particularly from the lower elevational parts of their range. Between 2016 and 2018, we tracked unhabituated Grauer's gorillas in lowland forests (500-1,000 m a.s.l.), at the community-managed Nkuba Conservation Area in Nord Kivu (DRC) to provide estimates of daily travel distances (DTD), daily displacement distances (DDD), and the linearity of recorded paths expressed as the Linearity Index (LI): DDD/DTD. We found an average DTD of ∼1.3 km (range 0.05-5.0 km), with temporal variation among monthly averages; specifically, an increase in travel distance over the June-August dry season resulting in peak travel distances at the beginning of the September-December wet season. Daily displacements showed similar temporal variation, which resulted in a lack of obvious temporal patterns in LI. We conclude that the movement patterns of Grauer's gorillas in lowland forests, which are characterized by larger DTD than those of Grauer's gorillas that inhabit highland habitats, show similarity to travel distances of other predominantly frugivorous gorillas. Moreover, the observed temporal patterns in space use may be tentatively linked to temporal changes in fruit availability or consumption. These observations have consequences for our understanding of the ecological role that Grauer's gorillas play and provide baseline data to estimate current and future distributions, abundances, and carrying capacities of this highly threatened animal.

Diet of Grauer's Gorillas (Gorilla beringei graueri) in a Low-Elevation Forest.

Although the vast majority of critically endangered Grauer's gorillas (Gorilla beringei graueri) inhabit low-elevation rain forests, current insights into this ape's life history and ecology stem predominantly from 2 small populations ranging in highland habitats. Here, we provide an initial and non-exhaustive overview of food items of Grauer's gorillas in the Nkuba Conservation Area (NCA), a lower-elevation (500-1,500 m) forest located between Kahuzi-Biega National Park and Maiko National Park in the Democratic Republic of the Congo. Community-based conservation efforts at the NCA aim to protect a population of unhabituated Grauer's gorillas, which we have studied since 2014. Between 2014 and 2020, we simultaneously tracked 1-3 gorilla groups and recorded a total of 10,514 feeding signs on at least 100 plant species, ants, termites, and fungi. Vegetative plant parts (plant stems, leaves, pith, bark, and roots), especially of Marantaceae and Fabaceae, made up close to 90% of recorded feeding signs, with fruit accounting for most of the remainder and a small (<1%) number of feeding signs on invertebrates and fungi. We found that the most frequently recorded food items were consumed year-round, though fruit intake seems to peak in the September-December wet season, possibly reflecting patterns in fruit phenology. The diet of Grauer's gorillas in the NCA differed from that of Grauer's gorillas in highland habitat and instead showed similarities with Grauer's gorillas at the lowland forest of Itebero and with western lowland gorillas (G. gorilla), which live under ecologically comparable conditions.

Comparing measures of social complexity: larger mountain gorilla groups do not have a greater diversity of relationships.

Social complexity reflects the intricate patterns of social interactions in societies. Understanding social complexity is fundamental for studying the evolution of diverse social systems and the cognitive innovations used to cope with the demands of social life. Social complexity has been predominantly quantified by social unit size, but newer measures of social complexity reflect the diversity of relationships. However, the association between these two sets of measures remains unclear. We used 12 years of data on 13 gorilla groups to investigate how measures of social complexity relate to each other. We found that group size was a poor proxy for relationship diversity and that the social complexity individuals experienced within the same group varied greatly. Our findings demonstrate two fundamental takeaways: first, that the number of relationships and the diversity of those relationships represent separate components of social complexity, both of which should be accounted for; and second, that social complexity measured at the group level may not represent the social complexity experienced by individuals in those groups. These findings suggest that comprehensive studies of social complexity, particularly those relating to the social demands faced by individuals, may require fine-scale social data to allow accurate comparisons across populations and species.

Inter-group relationships influence territorial defence in mountain gorillas.

Many species show territoriality, in which territory owners have exclusive or priority use of a region. In humans, tolerance of others within our space also depends greatly on our social relationships with them. This has been hypothesized as one potential driver of the evolution of long-term, inter-group relationships, through enabling shared access of resources and easing disputes over space. However, extremely little is known about the importance of social relationships between neighbouring groups in non-humans for how space is used and shared. Using 16 years of data on the simultaneous movement and interaction patterns of 17 mountain gorilla groups, we investigated how the occurrence of aggressive and affiliative behaviour during inter-group encounters was influenced by both their social and spatial context. We found evidence of territorial defence, with rates of aggression increasing towards the centre of home ranges. Groups which had previously split from each other showed higher levels of affiliation during encounters with each other and experienced lower levels of aggression when within the other's peripheral home range. However, encounters within core areas of the home range consistently elicited higher aggression, regardless of the groups' history. Our findings indicate that not only are the social relationships between groups retained after they split from one another but also that these relationships enable groups to access certain areas with a reduced risk of aggression. This suggests that reduced aggression when accessing areas within neighbours' home ranges may be an advantage for the maintenance of inter-group relationships and a potential driver in the evolution of long-term, post-dispersal relationships and complex multi-level societies.

Evaluating changes in salivary oxytocin and cortisol following positive reinforcement training in two adult male western lowland gorillas (Gorilla gorilla gorilla).

Positive reinforcement training (PRT) is associated with increases in species-typical behavior and decreases in stereotypic and abnormal behavior in participating animals. Physiological changes following PRT, for example, increases in oxytocin (OXT) and/or decreases in cortisol (CORT), may facilitate these behavioral changes. This study evaluated salivary OXT and salivary CORT concentrations in two adult male western lowland gorillas (Gorilla gorilla gorilla) following PRT with their primary animal care staff. Following PRT, no change in OXT was observed. CORT decreased in one subject following PRT. Changes in endogenous OXT are related to affiliative interactions and interact with strongly bonded conspecifics. PRT may not activate the oxytocinergic system because PRT is not a species-specific affiliative interaction and/or animal care staff are not viewed as conspecifics. Regardless, PRT may still be viewed as a positive interaction resulting in stress reduction via a decrease in CORT. Relationships are unique, thus these results only apply to these two gorillas and one animal caregiver. Larger population-level studies are needed to understand overall trends in human-animal interactions, and ultimately human-animal relationships. Further evaluation of physiological changes following human-animal interactions should be informative for understanding the human-animal relationship in zoos.

Faster growth corresponds with shallower linear hypoplastic defects in great ape canines.

Deeper or more 'severe' linear enamel hypoplasia (LEH) defects are hypothesized to reflect more severe stress during development, but it is not yet clear how depth is influenced by intrinsic enamel growth patterns. Recent work documented inter- and intraspecific differences in LEH defect depth in extant great apes, with mountain gorillas having shallower defects than other taxa, and females having deeper defects than males. Here, we assess the correspondence of inter- and intraspecific defect depth and intrinsic aspects of enamel growth: enamel extension rates, outer enamel striae of Retzius angles, and linear enamel thickness. Thin sections of great ape canines (n = 40) from Gorilla beringei beringei, Gorilla gorilla gorilla, Pan troglodytes, and Pongo spp. were analyzed. Enamel extension rates were calculated within deciles of enamel-dentine junction length. Linear enamel thickness and the angle of intersection between striae of Retzius and the outer enamel surface were measured in the imbricational enamel. Mountain gorillas have faster enamel extension rates and shallower striae angles than the other taxa examined. Mountain gorillas have thinner imbricational enamel than western lowland gorillas and orangutans, but not chimpanzees. In the combined-taxon sample, females exhibit larger striae angles and thicker imbricational enamel than males. Enamel extension rates are highly negatively correlated with striae angles and LEH defect depth. Enamel growth variation corresponds with documented inter- and intraspecific differences in LEH defect depth in great ape canines. Mountain gorillas have shallower striae angles and faster extension rates than other taxa, which might explain their shallow LEH defect morphology and the underestimation of their LEH prevalence in previous studies. These results suggest that stressors of similar magnitude and timing might produce defects of different depths in one species or sex vs. another, which has implications for interpretations of stress histories in hominins with variable enamel growth patterns.

The emergence of longevous populations.

The human lifespan has traversed a long evolutionary and historical path, from short-lived primate ancestors to contemporary Japan, Sweden, and other longevity frontrunners. Analyzing this trajectory is crucial for understanding biological and sociocultural processes that determine the span of life. Here we reveal a fundamental regularity. Two straight lines describe the joint rise of life expectancy and lifespan equality: one for primates and the second one over the full range of human experience from average lifespans as low as 2 y during mortality crises to more than 87 y for Japanese women today. Across the primate order and across human populations, the lives of females tend to be longer and less variable than the lives of males, suggesting deep evolutionary roots to the male disadvantage. Our findings cast fresh light on primate evolution and human history, opening directions for research on inequality, sociality, and aging.

Incisor tooth wear and age determination in mountain gorillas from Volcanoes National Park, Rwanda.

OBJECTIVES: Ecological factors, but also tooth-to-tooth contact over time, have a dramatic effect on tooth wear in primates. The aim of this study is to test whether incisor tooth wear changes predictably with age and can thus be used as an age estimation method in a wild population of mountain gorillas (Gorilla beringei beringei) from Volcanoes National Park, Rwanda. MATERIALS AND METHODS: In mountain gorillas of confidently known chronological age (N = 24), we measured the crown height of all permanent maxillary and mandibular incisors (I1 , I1 , I2 , I2 ) as a proxy for incisal macrowear. Linear and quadratic regressions for each incisor were used to test whether age can be predicted by crown height. Using these models, we then predicted age at death of two individual mountain gorillas of probable identifications, based on their incisor crown height. RESULTS: Age decreased significantly with incisor height for all teeth, but the upper first incisors (I1 ) provided the best results, with the lowest Akaike's Information Criterion corrected for small sample size (AICc) and lowest Standard Error of the Estimate (SEE). When the best age equations for each sex were applied to gorillas with probable identifications, the predicted ages differed 1.58 and 3.33 years from the probable ages of these individuals. CONCLUSIONS: Our findings corroborate that incisor crown height, a proxy for incisal wear, varies predictably with age. This relationship can be used to estimate age at death of unknown gorillas in the skeletal collection, and in some cases, to corroborate the identity of individual gorillas recovered from the forest postmortem at an advanced state of decomposition. Such identifications help fill gaps in the demographic database and support research that requires individual-level data.

Unexpected terrestrial hand posture diversity in wild mountain gorillas.

OBJECTIVES: Gorillas, along with chimpanzees and bonobos, are ubiquitously described as 'knuckle-walkers.' Consequently, knuckle-walking (KW) has been featured pre-eminently in hypotheses of the pre-bipedal locomotor behavior of hominins and in the evolution of locomotor behavior in apes. However, anecdotal and behavioral accounts suggest that mountain gorillas may utilize a more complex repertoire of hand postures, which could alter current interpretations of African ape locomotion and its role in the emergence of human bipedalism. Here we documented hand postures during terrestrial locomotion in wild mountain gorillas to investigate the frequency with which KW and other hand postures are utilized in the wild. MATERIALS AND METHODS: Multiple high-speed cameras were used to record bouts of terrestrial locomotion of 77 habituated mountain gorillas at Bwindi Impenetrable National Park (Uganda) and Volcanoes National Park (Rwanda). RESULTS: We captured high-speed video of hand contacts in 8% of the world's population of mountain gorillas. Our results reveal that nearly 40% of these gorillas used "non-KW" hand postures, and these hand postures constituted 15% of all hand contacts. Some of these "non-KW" hand postures have never been documented in gorillas, yet match hand postures previously identified in orangutans. DISCUSSION: These results highlight a previously unrecognized level of hand postural diversity in gorillas, and perhaps great apes generally. Although present at lower frequencies than KW, we suggest that the possession of multiple, versatile hand postures present in wild mountain gorillas may represent a shared feature of the African ape and human clade (or even great ape clade) rather than KW per se.

Quantifying linear enamel hypoplasia in Virunga Mountain gorillas and other great apes.

OBJECTIVE: Linear enamel hypoplasia (LEH) is a condition marked by localized reductions in enamel thickness, resulting from growth disruptions during dental development. We use quantitative criteria to characterize the depth of LEH defects and "normal" perikymata in great apes. We test the hypothesis that mountain gorillas have shallow defects compared to other taxa, which may have led to their underestimation in previous studies. MATERIALS AND METHODS: Previous attempts to characterize LEH morphology quantitatively have been limited in sample size and scope. We generated digital elevation models using optical profilometry (Sensofar PLu Neox) and extracted 2D coordinates using ImageJ to quantify depths in canines from three great ape genera (N = 75 perikymata; 255 defects). RESULTS: All defect depths fall outside the distribution of perikymata depths. Mountain gorilla defects are significantly shallower than those of other great ape taxa examined, including western lowland gorillas. Females have significantly deeper defects than males in all taxa. The deepest defect belongs to a wild-captured zoo gorilla. Virunga mountain gorilla specimens collected by Dian Fossey exhibit deeper defects than those collected recently. DISCUSSION: Shallow defect morphology in mountain gorillas may have led to an underestimation of LEH prevalence in past studies. Defect depth is used as a proxy for insult severity, but depth might be influenced by inter- and intra-specific variation in enamel growth. Future studies should test whether severe insults are associated with deeper defects, as might be the case with Haloko, a wild-captured gorilla. Ongoing histologic studies incorporating associated behavioral records will test possible factors that underlie differences in defect morphology.