Eastern Lowland Gorillas Research Articles

Historical Genomes Reveal the Genomic Consequences of Recent Population Decline in Eastern Gorillas

Many endangered species have experienced severe population declines within the last centuries [1, 2]. However, despite concerns about negative fitness effects resulting from increased genetic drift and inbreeding, there is a lack of empirical data on genomic changes in conjunction with such declines [3-7]. Here, we use whole genomes recovered from century-old historical museum specimens to quantify the genomic consequences of small population size in the critically endangered Grauer's and endangered mountain gorillas. We find a reduction of genetic diversity and increase in inbreeding and genetic load in the Grauer's gorilla, which experienced severe population declines in recent decades. In contrast, the small but relatively stable mountain gorilla population has experienced little genomic change during the last century. These results suggest that species histories as well as the rate of demographic change may influence how population declines affect genome diversity.

Phylogenetic and environmental effects on limb bone structure in gorillas.

The effects of phylogeny and locomotor behavior on long bone structural proportions are assessed through comparisons between adult and ontogenetic samples of extant gorillas. A total of 281 wild-collected individuals were included in the study, divided into four groups that vary taxonomically and ecologically: western lowland gorillas (G. g. gorilla), lowland and highland grauer gorillas (G. b. graueri), and Virunga mountain gorillas (G. b. beringei). Lengths and articular breadths of the major long bones (except the fibula) were measured, and diaphyseal cross-sectional geometric properties determined using computed tomography. Ages of immature specimens (n = 145) were known or estimated from dental development. Differences between groups in hind limb to forelimb proportions were assessed in both adults and during development. Diaphyseal strength proportions among adults vary in parallel with behavioral/ecological differences, and not phylogeny. The more arboreal western lowland and lowland grauer gorillas have relatively stronger forelimbs than the more terrestrial Virunga mountain gorillas, while the behaviorally intermediate highland grauer gorillas have intermediate proportions. Diaphyseal strength proportions are similar in young infants but diverge after 2 years of age in western lowland and mountain gorillas, at the same time that changes in locomotor behavior occur. There are no differences between groups in length or articular proportions among either adults or immature individuals. Long bone diaphyseal strength proportions in gorillas are developmentally plastic, reflecting behavior, while length and articular proportions are much more genetically canalized. These findings have implications for interpreting morphological variation among fossil taxa.

Population-level assessment of genetic diversity and habitat fragmentation in critically endangered Grauer's gorillas.

The critically endangered Grauer's gorilla (Gorilla beringei graueri) has experienced an estimated 77% population decline within a single generation. Although crucial for informed conservation decisions, there is no clear understanding about population structure and distribution of genetic diversity across the species' highly fragmented range. We fill this gap by studying several core and peripheral Grauer's gorilla populations throughout their distribution range. We generated genetic profiles for a sampling of an unstudied population of Grauer's gorillas from within the species' core range at 13 autosomal microsatellite loci and combined them with previously published and newly generated data from four other Grauer's gorilla populations, two mountain gorilla populations, and one western lowland gorilla population. In agreement with previous studies, the genetic diversity of Grauer's gorillas is intermediate, falling between western lowland and mountain gorillas. Among Grauer's gorilla populations, we observe lower genetic diversity and high differentiation in peripheral compared with central populations, indicating a strong effect of genetic drift and limited gene flow among small, isolated forest fragments. Although genetically less diverse, peripheral populations are frequently essential for the long-term persistence of a species and migration between peripheral and core populations may significantly enrich the overall species genetic diversity. Thus, in addition to central Grauer's gorilla populations from the core of the distribution range that clearly deserve conservation attention, we argue that conservation strategies aiming to ensure long-term species viability should include preserving peripheral populations and enhancing habitat connectivity.

Extant ape dental topography and its implications for reconstructing the emergence of early Homo

Dental topography reflects diet accurately in several extant and extinct mammalian clades. However, dental topographic dietary reconstructions have high success rates only when closely related taxa are compared. Given the dietary breadth that exists among extant apes and likely existed among fossil hominins, dental topographic values from many species and subspecies of great apes are necessary for making dietary inferences about the hominin fossil record. Here, we present the results of one metric of dental topography, Dirichlet normal energy (DNE), for seven groups of great apes (Pongo pygmaeus pygmaeus, Pan paniscus, Pan troglodytes troglodytes and schweinfurthii, Gorilla gorilla gorilla, Gorilla beringei graueri and beringei). Dirichlet normal energy was inadequate at differentiating folivores from frugivores, but was adequate at predicting which groups had more fibrous diets among sympatric African apes. Character displacement analyses confirmed there is substantial dental topographic and relative molar size (M1:M2 ratio; length, width, and area) divergence in sympatric apes when compared to their allopatric counterparts, but character displacement is only present in relative molar size when DNE is also considered. Presence of character displacement is likely due to indirect competition over similar food resources. Assuming similar ecological conditions in the Plio-Pleistocene, the derived masticatory apparatuses of the robust australopiths and early Homo may be due to indirect competition over dietary resources between the taxa, causing dietary niche partitioning. Our results imply that dental topography cannot be used to predict dietary categories in fossil hominins without consideration of ecological factors, such as dietary and geographic overlap. In addition, our results may open new avenues for understanding the community compositions of early hominins and the formation of specific ecological niches among hominin taxa.

Whole mitochondrial genome capture from faecal samples and museum-preserved specimens.

Population-scale molecular studies of endangered and cryptic species are often limited by access to high-quality samples. The use of noninvasively collected samples or museum-preserved specimens reduces the pressure on modern populations by removing the need to capture and handle live animals. However, endogenous DNA content in such samples is low, making shotgun sequencing a financially prohibitive approach. Here, we apply a target enrichment method to retrieve mitochondrial genomes from 65 museum specimens and 56 noninvasively collected faecal samples of two endangered great ape species, Grauer's gorilla and the eastern chimpanzee. We show that the applied method is suitable for a wide range of sample types that differ in endogenous DNA content, increasing the proportion of target reads to over 300-fold. By systematically evaluating biases introduced during target enrichment of pooled museum samples, we show that capture is less efficient for fragments shorter or longer than the baits, that the proportion of human contaminating reads increases postcapture although capture efficiency is lower for human compared to gorilla fragments with a gorilla-generated bait, and that the rate of jumping PCR is considerable, but can be controlled for with a double-barcoding approach. We succeed in capturing complete mitochondrial genomes from faecal samples, but observe reduced capture efficiency as sequence divergence increases between the bait and target species. As previously shown for museum specimens, we demonstrate here that mitochondrial genome capture from field-collected faecal samples is a robust and reliable approach for population-wide studies of nonmodel organisms.

Catastrophic Decline of World's Largest Primate: 80% Loss of Grauer's Gorilla (Gorilla beringei graueri) Population Justifies Critically Endangered Status.

Grauer’s gorilla (Gorilla beringei graueri), the World’s largest primate, is confined to eastern Democratic Republic of Congo (DRC) and is threatened by civil war and insecurity. During the war, armed groups in mining camps relied on hunting bushmeat, including gorillas. Insecurity and the presence of several militia groups across Grauer’s gorilla’s range made it very difficult to assess their population size. Here we use a novel method that enables rigorous assessment of local community and ranger-collected data on gorilla occupancy to evaluate the impacts of civil war on Grauer’s gorilla, which prior to the war was estimated to number 16,900 individuals. We show that gorilla numbers in their stronghold of Kahuzi-Biega National Park have declined by 87%. Encounter rate data of gorilla nests at 10 sites across its range indicate declines of 82–100% at six of these sites. Spatial occupancy analysis identifies three key areas as the most critical sites for the remaining populations of this ape and that the range of this taxon is around 19,700 km2. We estimate that only 3,800 Grauer’s gorillas remain in the wild, a 77% decline in one generation, justifying its elevation to Critically Endangered status on the IUCN Red List of Threatened Species.

New survey reveals dramatic decline of Grauer's gorilla

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The evolutionary origin and population history of the grauer gorilla.

Gorillas living in western central Africa (Gorilla gorilla) are morphologically and genetically distinguishable from those living in eastern central Africa (Gorilla beringei). Genomic analyses show eastern gorillas experienced a significant reduction in population size during the Pleistocene subsequent to geographical isolation from their western counterparts. However, how these results relate more specifically to the recent biogeographical and evolutionary history of eastern gorillas remains poorly understood. Here we show that two rare morphological traits are present in the hands and feet of both eastern gorilla subspecies at strikingly high frequencies (>60% in G. b. graueri; ∼28% in G. b. beringei) in comparison with western gorillas (<1%). The intrageneric distribution of these rare traits suggests that they became common among eastern gorillas after diverging from their western relatives during the early to middle Pleistocene. The extremely high frequencies observed among grauer gorillas-which currently occupy a geographic range more than ten times the size of that of mountain gorillas-imply that grauers originated relatively recently from a small founding population of eastern gorillas. Current paleoenvironmental, geological, and biogeographical evidence supports the hypothesis that a small group of eastern gorillas likely dispersed westward from the Virungas into present-day grauer range in the highlands just north of Lake Kivu, either immediately before or directly after the Younger Dryas interval. We propose that as the lowland forests of central Africa expanded rapidly during the early Holocene, they became connected with the expanding highland forests along the Albertine Rift and enabled the descendants of this small group to widely disperse. The descendant populations significantly expanded their geographic range and population numbers relative to the gorillas of the Virunga Mountains and the Bwindi-Impenetrable Forest, ultimately resulting in the grauer gorilla subspecies recognized today. This founder-effect hypothesis offers some optimism for modern conservation efforts to save critically endangered eastern gorillas from extinction.

The geographic distribution of genetic diversity within gorillas.

Gorillas, like all non-human great apes, are endangered. Understanding the distribution of genetic diversity across their range is important because low diversity may arise in small populations through increased inbreeding, and, by reducing reproductive fitness, may lead to decreased chances of persistence of a given population. Previous studies found higher genetic diversity in the western (Gorilla gorilla) than in the eastern gorillas (Gorilla beringei), but rarely employed individuals of known geographic origin to investigate the distribution of diversity across multiple populations. The present study fills that gap by analyzing 1,161 individuals from nine sites across all four currently recognized Gorilla subspecies. Genetic diversity at each site was estimated using published data from seven highly-variable microsatellite loci. We found that the small and fragmented populations of Cross River gorillas, eastern lowland gorillas and mountain gorillas were less diverse than any of the five analyzed western lowland gorilla populations. The higher levels of genetic variation within the western lowland gorillas might be best explained by the facts that they (i) exhibit larger present and past effective population sizes than the other subspecies and (ii) maintain higher rates of gene flow through the existence of largely continuous habitat within their range. With regard to conservation, the high genetic diversity within western lowland gorillas is encouraging and retention of dispersal corridors between already protected areas is essential. Am. J. Primatol. 77:974-985, 2015. © 2015 Wiley Periodicals, Inc.

Lack of Correlation Between Fluctuating Asymmetry and Morphological Masculinity/Femininity in Primate Skulls

As both degree of masculinity/femininity in sexually dimorphic organisms and developmental instability are put forward as indicators of individual quality, they are expected to correlate positively. However, the results in the literature are equivocal. One reason may be that most research has been performed in populations exposed to relatively low levels of parasitism. This article studies associations between the degree of masculinity/femininity and the fluctuating asymmetry (a measure of developmental instability) of skulls of three primate species [baboon (Papio anubis), eastern lowland gorilla (Gorilla beringei graueri), and chimpanzee (Pan troglodytes)] collected in the wild. The exposure to infection and parasites is likely to be higher in these individuals compared to captive primates and humans, and therefore likely to increase the levels of association. While the degree of individual masculinity/femininity and levels of fluctuating asymmetry were quantified accurately, I found no evidence of an association between them. My results thus are in line with the conclusion of a recent meta-analysis, that there is little evidence for associations between masculinity/femininity and developmental instability.

Inference of gorilla demographic and selective history from whole-genome sequence data.

Although population-level genomic sequence data have been gathered extensively for humans, similar data from our closest living relatives are just beginning to emerge. Examination of genomic variation within great apes offers many opportunities to increase our understanding of the forces that have differentially shaped the evolutionary history of hominid taxa. Here, we expand upon the work of the Great Ape Genome Project by analyzing medium to high coverage whole-genome sequences from 14 western lowland gorillas (Gorilla gorilla gorilla), 2 eastern lowland gorillas (G. beringei graueri), and a single Cross River individual (G. gorilla diehli). We infer that the ancestors of western and eastern lowland gorillas diverged from a common ancestor approximately 261 ka, and that the ancestors of the Cross River population diverged from the western lowland gorilla lineage approximately 68 ka. Using a diffusion approximation approach to model the genome-wide site frequency spectrum, we infer a history of western lowland gorillas that includes an ancestral population expansion of 1.4-fold around 970 ka and a recent 5.6-fold contraction in population size 23 ka. The latter may correspond to a major reduction in African equatorial forests around the Last Glacial Maximum. We also analyze patterns of variation among western lowland gorillas to identify several genomic regions with strong signatures of recent selective sweeps. We find that processes related to taste, pancreatic and saliva secretion, sodium ion transmembrane transport, and cardiac muscle function are overrepresented in genomic regions predicted to have experienced recent positive selection.

Healed mid-shaft fracture of an Early Roman bovine femur

We evaluated a healed fracture that was observed in the shaft of a bovine femur from an Early Roman settlement context at the site of Wange, Belgium. Using external observation and X-ray images, the specimen is described, paying particular attention to displacement of the bone ends, shortening of the bone, and callus formation. Similar fractures seen in femora of a modern Derby's antelope (Taurotragus derbianus) and of a modern eastern lowland gorilla (Gorilla gorilla graueri) are described, and we discuss why the incidence of healed femur fractures is so limited in large mammals.

Recent divergences and size decreases of eastern gorilla populations.

Compared with other African apes, eastern gorillas (Gorilla beringei) have been little studied genetically. We used analysis of autosomal DNA genotypes obtained from non-invasively collected faecal samples to estimate the evolutionary histories of the two extant mountain gorilla populations and the closely related eastern lowland gorillas. Our results suggest that eastern lowland gorillas and mountain gorillas split beginning some 10 000 years ago, followed 5000 years ago by the split of the two mountain gorilla populations of Bwindi Impenetrable National Park and the Virungas Massif. All three populations have decreased in effective population size, with particularly substantial 10-fold decreases for the mountain gorillas. These dynamics probably reflect responses to habitat changes resulting from climate fluctuations over the past 20 000 years as well as increasing human influence in this densely populated region in the last several thousand years.

Brain organization of gorillas reflects species differences in ecology.

Gorillas include separate eastern (Gorilla beringei) and western (Gorilla gorilla) African species that diverged from each other approximately 2 million years ago. Although anatomical, genetic, behavioral, and socioecological differences have been noted among gorilla populations, little is known about variation in their brain structure. This study examines neuroanatomical variation between gorilla species using structural neuroimaging. Postmortem magnetic resonance images were obtained of brains from 18 captive western lowland gorillas (Gorilla gorilla gorilla), 15 wild mountain gorillas (Gorilla beringei beringei), and 3 Grauer's gorillas (Gorilla beringei graueri) (both wild and captive). Stereologic methods were used to measure volumes of brain structures, including left and right frontal lobe gray and white matter, temporal lobe gray and white matter, parietal and occipital lobes gray and white matter, insular gray matter, hippocampus, striatum, thalamus, each hemisphere and the vermis of the cerebellum, and the external and extreme capsules together with the claustrum. Among the species differences, the volumes of the hippocampus and cerebellum were significantly larger in G. gorilla than G. beringei. These anatomical differences may relate to divergent ecological adaptations of the two species. Specifically, G. gorilla engages in more arboreal locomotion and thus may rely more on cerebellar circuits. In addition, they tend to eat more fruit and have larger home ranges and consequently might depend more on spatial mapping functions of the hippocampus.

The complete mitochondrial genome of eastern lowland gorilla, Gorilla beringei graueri, and comparative mitochondrial genomics of Gorilla species

In this study, we determined the complete mitochondrial (mt) genome of eastern lowland gorilla, Gorilla beringei graueri for the first time. The total genome was 16,416 bp in length. It contained a total of 13 protein-coding genes, 22 transfer RNA genes, 2 ribosomal RNA genes and 1 control region (D-loop region). The base composition was A (30.88%), G (13.10%), C (30.89%) and T (25.13%), indicating that the percentage of A+T (56.01%) was higher than G+C (43.99%). Comparisons with the other publicly available Gorilla mitogenome showed the conservation of gene order and base compositions but a bunch of nucleotide diversity. This complete mitochondrial genome sequence will provide valuable genetic information for further studies on conservation genetics of eastern lowland gorilla.

The interpretive power of infraorbital foramen area in making dietary inferences in extant apes.

The infraorbital foramen (IOF) is located below the orbit and transmits the sensory infraorbital nerve (ION) to mechanoreceptors located throughout the maxillary region. The size of the IOF correlates with the size of the ION; thus, the IOF appears to indicate relative touch sensitivity of maxillary region. In primates, IOF size correlates well with diet. Frugivores have relatively larger IOFs than folivores or insectivores because fruit handling/processing requires increased touch sensitivity. However, it is unknown if the IOF can be used to detect subtle dietary differences among closely related hominoid species. Hominoids are traditionally grouped into broad dietary categories, despite the fact that hominoid diets are remarkably diverse. This study examines whether relative IOF size is capable of differentiating among the dietary preferences of closely related species with overlapping, yet divergent diets. We measured IOF area in Hylobates lar, Symphalangus syndactulus, Pongo pygmaeus spp., Pan troglodytes, Gorilla gorilla, Gorilla beringei graueri, and Gorilla beringei beringei. We classified each species as a dedicated folivore, mixed folivore/frugivore, soft object frugivore, or hard object frugivore. The IOF is documented to be larger in more frugivorous species and smaller in more folivorous taxa. Interestingly, G.b. beringei, had the largest relative IOF of any gorilla, despite being a dedicated folivore. G.b. beringei does have unique food processing behavior that relies heavily on maxillary mechanoreception, thus this finding is not entirely unsupported behaviorally. The results of this study provide evidence that the IOF is an informative feature in interpretations of fossil apes.

Asymmetry of the midfacial skeleton of eastern lowland gorillas (Gorilla beringei graueri) and potential association with frontal lobe asymmetries

In 1973, Groves and Humphrey reported facial asymmetry in eastern gorillas and suggested that it may be indicative of asymmetrical brain function. Being cautious, the authors favored the explanation that facial asymmetry was likely due to asymmetrical chewing behavior rather than anatomical asymmetry of the cerebral hemispheres. The nature of the association between lateralized functions (e.g., handedness, mastication) and brain asymmetries in hominoids represents an active field of research. There has been both support (Hoogmartens and Caubergh, 1987; Nisan et al., 2004) and lack of support (Martinez-Gomis et al., 2009) for an association between chewing side preference and handedness in humans. In gorillas, contradictory results can be found on the question of handedness, varying from no handedness (Harrison and Nystrom, 2010) to equal proportion of right and left (Fagot and Vauclair, 1988; Byrne and Byrne, 1991; Parnell, 2001; Lambert, 2012) to true handedness (Meguerditchian et al., 2010). Cerebral anatomical asymmetries, and particularly fronto-occipital petalia, have been found to be related to handedness in modern humans (LeMay, 1976, 1977; Galaburda et al., 1978; LeMay and Kido, 1978; LeMay et al., 1982; Kertesz et al., 1986, 1990; but see Chiu and Damasio, 1980), and more generally to specific aspects of human cognition (Hecaen and Albert, 1978; Nass and Gazzaniga, 1987; Bryden, 1988; Gazzaniga, 1995a, b; Harrington, 1995). The existence of asymmetrical patterns for the frontal and occipital petalias in great apes, including Gorilla beringei graueri, have recently been reported (Balzeau and Gilissen, 2010; Balzeau et al., 2012). However, the potential association between facial and cerebral anatomical asymmetries in great apes and its functional and evolutionary implications remain unclear. Moreover, the original paper of Groves and Humphrey (1973) has been over-interpreted, as several authors refer to it as a claim of cerebellar asymmetry in gorillas (e.g., Balonov et al., 1984; Hopkins and Marino, 2000; Holloway et al., 2004). Here, we propose to quantify facial asymmetry in adult eastern lowland gorillas (Gorilla beringei graueri), determine whether or not directional asymmetry (DA) is present in our sample, and subsequently investigate the potential relationship between facial and frontal lobe asymmetries using data previously published by Balzeau and Gilissen (2010).

The Status of Conservation Education in Africa

There are 5 global eco-regions regarded as top conservation priorities in Africa, and each one has a center for conservation studies. Established in 1962, the College of Wildlife Management at Mweka, Tanzania, has trained more than 5000 wildlife managers from 52 countries till date in the Eastern Arc Montane eco-region. The Upper and Lower Guinea eco-region is represented by the Sapo Conservation Center (SCC) in the eponymous National Park in Liberia and the A.P. Levenits Ornithological Research Institute (APLORI), Jos, Nigeria. The SCC offers training programs of up to 2 weeks duration, 2 to 3 times a year for students and academic instructors from the University of Liberia, Monrovia, Forestry Training Institute, Tubmanburg and Nimba County Community College, Nimba. The APLORI, founded in 2002, is a premier institute of ornithological research and conservation training in West Africa, and 66 graduates have left its portals to take up crucial responsibilities in the government, civil society and academia in Africa. The Cape Fynbos eco-region famed for its restioid, ericoid and proteoid components hosts the Southern African Wildlife College (SAWC) for training managers and conservationists in diverse fields such as such as nature-based tourism, trans-frontier conservation area management and capacity building at the community level. SAWC is situated in South Africa in a natural low veld environment, 10km west of the Orpen Gate of the Kruger National Park. The Albertine Rift Montane eco-region with flagship endangered species such as the Mountain and Eastern Lowland gorillas, saw the launch of a conservation primary school in Ilima, Democratic Republic of Congo (DRC) in September 2013; building 14 other unique primary schools over a period of 10 years in key landscapes in Africa is a commitment of the African Wildlife Foundation (AWF). Last but not the least, the Ethiopian highlands eco-region, houses species like the Ethiopian wolf and is served by the Wondo Genet College of Forestry in southern Ethiopia to train conservation professionals in the Horn of Africa region and beyond.

Complete mitochondrial genome sequence of the Eastern gorilla (Gorilla beringei) and implications for african ape biogeography.

The Western and Eastern species of gorillas (Gorilla gorilla and Gorilla beringei) began diverging in the mid-Pleistocene, but in a complex pattern with ongoing gene flow following their initial split. We sequenced the complete mitochondrial genomes of 1 Eastern and 1 Western gorilla to provide the most accurate date for their mitochondrial divergence, and to analyze patterns of nucleotide substitutions. The most recent common ancestor of these genomes existed about 1.9 million years ago, slightly more recent than that of chimpanzee and bonobo. We in turn use this date as a calibration to reanalyze sequences from the Eastern lowland and mountain gorilla subspecies to estimate their mitochondrial divergence at approximately 380000 years ago. These dates help frame a hypothesis whereby populations became isolated nearly 2 million years ago with restricted maternal gene flow, followed by ongoing male migration until the recent past. This process of divergence with prolonged hybridization occurred against the backdrop of the African Pleistocene, characterized by intense fluctuations in temperature and aridity, while at the same time experiencing tectonic uplifting and consequent shifts in the drainage of major river systems. Interestingly, this same pattern of introgression following divergence and discrepancies between mitochondrial and nuclear loci is seen in fossil hominins from Eurasia, suggesting that such processes may be common in hominids and that living gorillas may provide a useful model for understanding isolation and migration in our extinct relatives.

Tooth cusp sharpness as a dietary correlate in great apes

Mammalian molars have undergone heavy scrutiny to determine correlates between morphology and diet. Here, the relationship between one aspect of occlusal morphology, tooth cusp radius of curvature (RoC), and two broad dietary categories, folivory and frugivory, is analyzed in apes. The author hypothesizes that there is a relationship between tooth cusp RoC and diet, and that folivores have sharper teeth than frugivores, and further test the correlation between tooth cusp RoC and tooth cusp size. Eight measures of tooth cusp RoC (two RoCs per cusp) were taken from 53 M(2) s from four species and subspecies of frugivorous apes (Pongo pygmaeus, Pan troglodytes troglodytes, Pan troglodytes schweinfurthii, and Gorilla gorilla gorilla) and two subspecies of folivorous apes (Gorilla beringei beringei, and Gorilla beringei graueri). Phylogenetically corrected ANOVAs were run on the full dataset and several subsets of the full dataset, revealing that, when buccolingual RoCs are taken into account, tooth cusp RoCs can successfully differentiate folivores and frugivores. PCAs revealed that folivores consistently had duller teeth than frugivores. In addition, a weak, statistically significant positive correlation exists between tooth cusp size and tooth cusp RoC. The author hypothesizes differences in tooth cusp RoC are correlated with wear rates, where, per vertical unit of wear, duller cusps will have a longer length of exposed enamel ridge than sharper cusps. More data need to be gathered to determine if the correlation between tooth cusp RoC and tooth cusp size holds true when small primates are considered.