Phylogenetic differences in the morphology and shape of the central sulcus in great apes and humans: implications for the evolution of motor functions.

The central sulcus divides the primary motor and somatosensory cortices in many anthropoid primate brains. Differences exist in the surface area and depth of the central sulcus along the dorso-ventral plane in great apes and humans compared to other primate species. Within hominid species, there are variations in the depth and aspect of their hand motor area, or knob, within the precentral gyrus. In this study, we used post-image analyses on magnetic resonance images to characterize the central sulcus shape of humans, chimpanzees (Pan troglodytes), gorillas (Gorilla gorilla), and orangutans (Pongo pygmaeus and Pongo abelii). Using these data, we examined the morphological variability of central sulcus in hominids, focusing on the hand region, a significant change in human evolution. We show that the central sulcus shape differs between great ape species, but all show similar variations in the location of their hand knob. However, the prevalence of the knob location along the dorso-ventral plane and lateralization differs between species and the presence of a second ventral motor knob seems to be unique to humans. Humans and orangutans exhibit the most similar and complex central sulcus shapes. However, their similarities may reflect divergent evolutionary processes related to selection for different positional and habitual locomotor functions.

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.

Trabecular architecture of the distal femur in extant hominids.

Extant great apes are characterized by a wide range of locomotor, postural and manipulative behaviours that each require the limbs to be used in different ways. In addition to external bone morphology, comparative investigation of trabecular bone, which (re-)models to reflect loads incurred during life, can provide novel insights into bone functional adaptation. Here, we use canonical holistic morphometric analysis (cHMA) to analyse the trabecular morphology in the distal femoral epiphysis of Homo sapiens (n = 26), Gorilla gorilla (n = 14), Pan troglodytes (n = 15) and Pongo sp. (n = 9). We test two predictions: (1) that differing locomotor behaviours will be reflected in differing trabecular architecture of the distal femur across Homo, Pan, Gorilla and Pongo; (2) that trabecular architecture will significantly differ between male and female Gorilla due to their different levels of arboreality but not between male and female Pan or Homo based on previous studies of locomotor behaviours. Results indicate that trabecular architecture differs among extant great apes based on their locomotor repertoires. The relative bone volume and degree of anisotropy patterns found reflect habitual use of extended knee postures during bipedalism in Homo, and habitual use of flexed knee posture during terrestrial and arboreal locomotion in Pan and Gorilla. Trabecular architecture in Pongo is consistent with a highly mobile knee joint that may vary in posture from extension to full flexion. Within Gorilla, trabecular architecture suggests a different loading of knee in extension/flexion between females and males, but no sex differences were found in Pan or Homo, supporting our predictions. Inter- and intra-specific variation in trabecular architecture of distal femur provides a comparative context to interpret knee postures and, in turn, locomotor behaviours in fossil hominins.

Changes in movement patterns in relation to sun conditions and spatial scales in wild western gorillas.

For most primates living in tropical forests, food resources occur in patchworks of different habitats that vary seasonally in quality and quantity. Efficient navigation (i.e., spatial memory-based orientation) towards profitable food patches should enhance their foraging success. The mechanisms underpinning primate navigating ability remain nonetheless mostly unknown. Using GPS long-term tracking (596 days) of one group of wild western lowland gorillas (Gorilla gorilla gorilla), we investigated their ability to navigate at long distances, and tested for how the sun was used to navigate at any scale by improving landmark visibility and/or by acting as a compass. Long episodic movements ending at a distant swamp, a unique place in the home range where gorillas could find mineral-rich aquatic plants, were straighter and faster than their everyday foraging movements relying on spatial memory. This suggests intentional targeting of the swamp based on long-distance navigation skills, which can thus be efficient over a couple of kilometres. Interestingly, for both long-distance movements towards the swamp and everyday foraging movements, gorillas moved straighter under sunlight conditions even under a dense vegetation cover. By contrast, movement straightness was not markedly different when the sun elevation was low (the sun azimuth then being potentially usable as a compass) or high (so providing no directional information) and the sky was clear or overcast. This suggests that gorillas navigate their home range by relying on visual place recognition but do not use the sun azimuth as a compass. Like humans, who rely heavily on vision to navigate, gorillas should benefit from better lighting to help them identify landmarks as they move through shady forests. This study uncovers a neglected aspect of primate navigation. Spatial memory and vision might have played an important role in the evolutionary success of diurnal primate lineages.

Sexual dimorphism in the cranium and endocast of the eastern lowland gorillas (Gorilla beringei graueri).

Sexual dimorphism of the nervous system has been reported for a wide range of vertebrates. However, understanding of sexual dimorphism in primate cranial structures and soft tissues, and more particularly the brain, remains limited. In this study, we aimed to investigate the external and internal (i.e., endocast) cranial differences between male and female eastern lowland gorillas (Gorilla beringei graueri). We examined the differences in the size, shape, and disparity with the aim to compare how sexual dimorphism can impact these two structures distinctively, with a particular focus on the endocranium. To do so, we reconstructed gorilla external crania and endocasts from CT scans and used 3D geometric morphometric techniques combined with multivariate analyses to assess the cranial and endocranial differences between the sexes. Our results highlighted sexual dimorphism for the external cranium and endocast with regard to both size and shape. In particular, males display an elongated face accompanied by a pronounced sagittal crest and an elongated endocast along the rostroposterior axis, in contrast to females who are identified by a more rounded brain case and endocast. Males also show a significantly larger external cranium and endocast size than females. In addition, we described important differences for the posterior cranial fossae (i.e., the position of the cerebellum within the brain case) and olfactory bulb between the two sexes. Particularly, our results highlighted that, relatively to males, females have larger posterior cranial fossae, whereas males have been characterized by a larger and rostrally oriented olfactory bulb.

Chemical contraceptive impacts on cyclic progesterone and sexual behavior in zoo-housed Western lowland gorillas.

For regularly cycling primates in captivity, chemical contraceptives can be used to reduce the likelihood of unintentional offspring, but the effects on cyclic progesterone and sexual behavior have not been well studied in gorillas (Gorilla gorilla gorilla). As zoos are invested in maintaining naturalistic behaviors and physiology, this is an important aspect of gorilla physical and mental health. We examined how exposure to combination and progestin-only oral birth control pills altered cyclic progesterone and timing of sexual behaviors in four female gorillas at four institutions compared to cycling controls. Hormonal data were collected using fecal steroid metabolites as detected by enzyme immunoassay. Behavioral data were collected via one-zero sampling done by keeper staff. All methods of oral contraceptive eliminated the ovarian cycle, with fecal progestogens being maintained below baselines in cycling control individuals. There was also no evidence of any cyclic patterning in endogenous progesterone, suggesting that the methods tested do not allow for detection of a recovery of endogenous progesterone during a short no-pill period. We also found that the oral contraceptives altered the timing of behavioral displays when compared to cycling control individuals but was not directly related to any fluctuations in endogenous progesterone. This study reveals a need for additional research on contraceptive use in gorillas and reinforces that sexual behavior in gorillas is both hormonally and socially mediated.

Variation in Milk Proteins Across Lactation in Pongo pygmaeus and Gorilla gorilla.

Milk is a critical nutrition source for all neonatal mammals. In addition to nutrition, milk contains a multitude of bioactive molecules that likely affect neonatal physiology, metabolism, and immune function. We suggest that changes in the milk proteome across lactation reflect the changing need of the neonate and juvenile offspring. We used mass spectrometry to characterize the milk proteomes from a Pongo pygmaeus (12 samples) and a Gorilla gorilla (6 samples) housed at the Smithsonian's National Zoo and Conservation Biology Institute and trained to give milk samples. We found a total of 454 proteins from P. pygmaeus and 428 proteins from G. gorilla. We specifically characterized changes across lactation in 13 proteins representing multiple compartments of milk, including the milk fat globule membrane and whey. Additionally, we characterized changes in various immunoglobulin types, finding similarities to previously published studies on primate milks. Despite broad similarities between the milk proteomes of these two apes, we demonstrated that proteomes from samples from 8 to 12 months clustered by species/individual and were distinct. Samples from more individuals are required to distinguish whether our result demonstrates species differences or individual differences. This study represents a baseline study that other zoo-based milk studies can build from. All RAW data, MetaMorpheus search results, and PAW_BLAST results are available on MassIVE at ftp://massive.ucsd.edu/MSV000089723/.

A comparison of faecal glucocorticoid metabolite concentration and gut microbiota diversity in bonobos (Pan paniscus).

Sex, age, diet, stress and social environment have all been shown to influence the gut microbiota. In several mammals, including humans, increased stress is related to decreasing gut microbial diversity and may differentially impact specific taxa. Recent evidence from gorillas shows faecal glucocorticoid metabolite concentration (FGMC) did not significantly explain gut microbial diversity, but it was significantly associated with the abundance of the family Anaerolineaceae. These patterns have yet to be examined in other primates, like bonobos (Pan paniscus). We compared FGMC to 16S rRNA amplicons for 202 bonobo faecal samples collected across 5 months to evaluate the impact of stress, measured with FGMC, on the gut microbiota. Alpha diversity measures (Chao's and Shannon's indexes) were not significantly related to FGMC. FGMC explained 0.80 % of the variation in beta diversity for Jensen-Shannon and 1.2% for weighted UniFrac but was not significant for unweighted UniFrac. We found that genus SHD-231, a member of the family Anaerolinaceae had a significant positive relationship with FGMC. These results suggest that bonobos are relatively similar to gorillas in alpha diversity and family Anaerolinaceae responses to FGMC, but different from gorillas in beta diversity. Members of the family Anaerolinaceae may be differentially affected by FGMC across great apes. FGMC appears to be context dependent and may be species-specific for alpha and beta diversity but this study provides an example of consistent change in two African apes. Thus, the relationship between physiological stress and the gut microbiome may be difficult to predict, even among closely related species.

What's not to like about Likert? Developing a nonverbal animal preference scale (NAPS).

Rating scales, such as Likert scales, are incredibly flexible and intuitive tools for measuring individuals' rating of agreement with or relative preference for many types of stimuli. For humans, this typically involves ratings of agreement between end points representing distinct attitudes or beliefs; For example, strongly disagree to strongly agree. Nonverbal versions of Likert scales have also been presented to children, allowing them to indicate their degree of preference, pain, or happiness. However, before the current study, no known efforts had been made to develop a nonverbal rating scale for use with nonhuman animals. Such a scale would be a useful welfare tool, allowing nonverbal individuals to indicate not just relative preferences between pairs of items but their degree of liking for individual items. I present an outline of the steps taken to create such a scale for use with three zoo-housed gorillas. Two gorillas succeeded in associating preferred and less preferred foods with different response buttons but none of the gorillas were able to effectively use the neutral response button. It is possible that limits in gorillas' capacity for conditional discriminations and/or conceptualization of constructs as abstract as "liking" impeded training. These data are relevant for understanding gorilla cognition and can inform continued efforts to create a tool for nonhumans to communicate their preferences to human caregivers in a more nuanced way than is currently possible.

Prevalence of regurgitation and reingestion and occurrence of coprophagy in the North American AZA Gorilla (Gorilla gorilla gorilla) population.

The aims of this study were to determine the current prevalence of regurgitation and reingestion (R/R) and gain an insight on the occurrence of coprophagy in the Association of Zoos & Aquariums (AZA) gorilla (Gorilla gorilla gorilla) population. To do this, an online survey was sent to all AZA accredited gorilla institutions in 2018. This survey collected information on the severity of R/R at the individual level and the occurrence of coprophagy at the zoo level. Previous research suggests more than half of zoo-housed gorillas in the North American population engaged in R/R and coprophagy to some extent. The current study had a 100% response and completion rate; data were collected on all 296 noninfant gorillas in the population. Responses indicated that 60% of the AZA gorilla population engages in R/R to some degree and 24% of the population are observed exhibiting coprophagy on a weekly basis. With this preliminary information on coprophagy, and the realization that the Zoo community has been unsuccessful at eliminating R/R over the past 30 years, the next steps should be aimed at empirically assessing the potential drivers of these behaviors. Continuing efforts to further understand and eliminate R/R and coprophagy in zoo-housed primates is important not only because these behaviors are unsightly to visitors, but more importantly, R/R and coprophagy may signal deficiencies in current practices surrounding animal care.

Exploring the functional morphology of the Gorilla shoulder through musculoskeletal modelling.

Musculoskeletal computer models allow us to quantitatively relate morphological features to biomechanical performance. In non-human apes, certain morphological features have long been linked to greater arm abduction potential and increased arm-raising performance, compared to humans. Here, we present the first musculoskeletal model of a western lowland gorilla shoulder to test some of these long-standing proposals. Estimates of moment arms and moments of the glenohumeral abductors (deltoid, supraspinatus and infraspinatus muscles) over arm abduction were conducted for the gorilla model and a previously published human shoulder model. Contrary to previous assumptions, we found that overall glenohumeral abduction potential is similar between Gorilla and Homo. However, gorillas differ by maintaining high abduction moment capacity with the arm raised above horizontal. This difference is linked to a disparity in soft tissue properties, indicating that scapular morphological features like a cranially oriented scapular spine and glenoid do not enhance the abductor function of the gorilla glenohumeral muscles. A functional enhancement due to differences in skeletal morphology was only demonstrated in the gorilla supraspinatus muscle. Contrary to earlier ideas linking a more obliquely oriented scapular spine to greater supraspinatus leverage, our results suggest that increased lateral projection of the greater tubercle of the humerus accounts for the greater biomechanical performance in Gorilla. This study enhances our understanding of the evolution of gorilla locomotion, as well as providing greater insight into the general interaction between anatomy, function and locomotor biomechanics.

Fiber-type phenotype of the jaw-closing muscles in Gorilla gorilla, Pan troglodytes, and Pan paniscus: A test of the Frequent Recruitment Hypothesis.

Skeletal muscle fiber types are important determinants of the contractile properties of muscle fibers, such as fatigue resistance and shortening velocity. Yet little is known about how jaw-adductor fiber types correlate with feeding behavior in primates. Compared with chimpanzees and bonobos, gorillas spend a greater percentage of their daily time feeding and shift to herbaceous vegetation when fruits are scarce. We thus used the African apes to test the hypothesis that chewing with unusually high frequency is correlated with the expression in the jaw adductors of a high proportion of type 1 (slow, fatigue-resistant) fibers at the expense of other fiber types (the Frequent Recruitment Hypothesis). We used immunohistochemistry to determine the presence and distribution of the four major myosin heavy chain (MHC) isoforms in the anterior superficial masseter (ASM), superficial anterior temporalis, and deep anterior temporalis of four Gorilla gorilla, two Pan paniscus, and four Pan troglodytes. Serial sections were stained against slow (MHC-1/-α-cardiac) and fast (MHC-2/-M) fibers. Fibers were counted and scored for staining intensity, and fiber cross-sectional areas (CSAs) were measured and used to estimate percentage of CSA of each MHC isoform. Hybrid fibers accounted for nearly 100% of fiber types in the masseter and temporalis of all three species, resulting in three main hybrid phenotypes. As predicted, the gorilla ASM and deep anterior temporalis comprised a greater percentage of CSA of the slower, fatigue-resistant hybrid fiber type, significantly so for the ASM (p = 0.015). Finally, the results suggest that fiber phenotype of the chewing muscles contributes to behavioral flexibility in ways that would go undetected in paleontological studies relying solely on morphology of the bony masticatory apparatus.

Variability of weaning age in mountain gorillas (Gorilla beringei beringei).

OBJECTIVES: Weaning is a key life history milestone for mammals that represents both the end of nutritional investment from the perspective of mothers and the start of complete nutritional independence for the infants. The age at weaning may vary depending on ecological, social, and demographic factors experienced by the mother and infant. Bwindi mountain gorillas live in different environmental conditions and have longer interbirth intervals than their counterparts in the Virunga Volcanoes, yet other life history characteristics of this population remain less well known. We use long-term data from Bwindi Impenetrable National Park, Uganda to examine factors related to weaning age. MATERIALS AND METHODS: We analyzed data on infants born in four mountain gorilla groups in Bwindi to quantify their age of weaning (defined as last nipple contact) and to test if the sex of offspring, parity, and dominance rank of mother influences age of weaning. We also compared the age at weaning and time to conception after resumption of mating in Bwindi and Virunga gorillas. RESULTS: Bwindi gorillas were weaned at an average age of 57.5 months. No difference was found between age of weaning for primiparous and multiparous mothers, nor did maternal dominance rank influence age of weaning, but sons were weaned at a later age than daughters. The majority of Bwindi mothers were still suckling when they resumed mating and mothers generally conceived before they weaned their previous offspring. The age of weaning was significantly later in Bwindi than in Virunga gorillas. After mothers resumed mating, the time to conceiving the next offspring was not significantly longer for Bwindi females than Virungas females (6 vs. 4 months). DISCUSSION: Later weaning age for sons than daughters is similar to findings of other studies of great apes. Bwindi mountain gorillas are weaned at approximately the same age as western gorillas and chimpanzees, which is more than a year later than Virunga mountain gorillas. The results of this study suggest that variation in ecological conditions of populations living in close geographic proximity can result in variation in life history patterns, which has implications for understanding the evolution of the unique life history patterns of humans.

Gorilla calcaneal morphological variation and ecological divergence.

OBJECTIVES: The primate foot has been extensively investigated because of its role in weight-bearing; however, the calcaneus has been relatively understudied. Here we examine entire gorilla calcaneal external shape to understand its relationship with locomotor behavior. MATERIALS AND METHODS: Calcanei of Gorilla gorilla gorilla (n = 43), Gorilla beringei graueri (n = 20), and Gorilla beringei beringei (n = 15) were surface or micro-CT scanned. External shape was analyzed through a three-dimensional geometric morphometric sliding semilandmark analysis. Semilandmarks were slid relative to an updated Procrustes average in order to minimize the bending energy of the thin plate spline interpolation function. Shape variation was summarized using principal components analysis of shape coordinates. Procrustes distances between taxa averages were calculated and resampling statistics run to test pairwise differences. Linear measures were collected and regressed against estimated body mass. RESULTS: All three taxa exhibit statistically different morphologies (p < .001 for pairwise comparisons). G. g. gorilla demonstrates an anteroposteriorly elongated calcaneus with a deeper cuboid pivot region and mediolaterally flatter posterior talar facet. G. b. beringei possesses the flattest cuboid and most medially-angled posterior talar facets. G. b. graueri demonstrates intermediate articular facet morphology, a medially-angled tuberosity, and an elongated peroneal trochlea. DISCUSSION: Articular facet differences separate gorillas along a locomotor gradient. G. g. gorilla is adapted for arboreality with greater joint mobility, while G. b. beringei is adapted for more stereotypical loads associated with terrestriality. G. b. graueri's unique posterolateral morphology may be due to a secondary transition to greater arboreality from a more terrestrial ancestor.

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.

Morphometric analysis of the hominin talus: Evolutionary and functional implications.

The adoption of bipedalism is a key benchmark in human evolution that has impacted talar morphology. Here, we investigate talar morphological variability in extinct and extant hominins using a 3D geometric morphometric approach. The evolutionary timing and appearance of modern human-like features and their contributions to bipedal locomotion were evaluated on the talus as a whole, each articular facet separately, and multiple combinations of facets. Distinctive suites of features are consistently present in all fossil hominins, despite the presence of substantial interspecific variation, suggesting a potential connection of these suites to bipedal gait. A modern human-like condition evolved in navicular and lateral malleolar facets early in the hominin lineage compared with other facets, which demonstrate more complex morphological variation within Homininae. Interestingly, navicular facet morphology of Australopithecus afarensis is derived in the direction of Homo, whereas more recent hominin species such as Australopithecus africanus and Australopithecus sediba retain more primitive states in this facet. Combining the navicular facet with the trochlea and the posterior calcaneal facet as a functional suite, however, distinguishes Australopithecus from Homo in that the medial longitudinal arch had not fully developed in the former. Our results suggest that a more everted foot and stiffer medial midtarsal region are adaptations that coincide with the emergence of bipedalism, whereas a high medial longitudinal arch emerges later in time, within Homo. This study provides novel insights into the emergence of talar morphological traits linked to bipedalism and its transition from a facultative to an obligate condition.

Fruit-feeding and activity patterns of mountain gorillas (Gorilla beringei beringei) in Bwindi Impenetrable National Park, Uganda.

OBJECTIVES: Availability of fruit is an important factor influencing variation in great ape foraging strategies and activity patterns. This study aims to quantify how frugivory influences activity budgets across age-sex classes of mountain gorillas in Bwindi Impenetrable National Park, Uganda. MATERIALS AND METHODS: Daily proportions of fruit-feeding and activity budgets were calculated using 6 years of observational data on four habituated groups. We fitted generalized linear mixed models to test for age-sex differences in the amount of fruit-feeding, and to test whether these factors influence the proportion of time spent feeding, resting, and traveling. RESULTS: Bwindi mountain gorillas spent on average 15% of feeding time consuming fruit, with monthly variation ranging from 0 to 70%. Greater amounts of fruit-feeding were associated with more time feeding and traveling, and less time resting. Immatures tended to spend more feeding time on fruit than adults, but less overall time feeding and more time traveling. There were no significant differences in the amount of fruit-feeding and overall feeding time between adult females and silverback males, despite differences in body size. DISCUSSION: This study confirms that gorillas are frugivorous, and only the Virunga mountain gorilla population can be characterized as highly folivorous. Along with other frugivorous great apes, Bwindi mountain gorillas alter their activity patterns in response to varying amounts of fruit in their diet. A better understanding of how variable ecological conditions can drive diversity even within a subspecies has important implications for understanding relationships between ecology, body size, and foraging strategies in great apes.

Brief communication: Maximum ingested bite size in captive western lowland gorillas (Gorilla gorilla gorilla).

OBJECTIVES: Previously, we found that maximum ingested bite size (Vb ), the largest piece of food an animal can consume without biting it into smaller pieces first, isometrically scales relative to body size in strepsirrhines and with negative allometry in anthropoids. In the current study, we rectify the omission of great apes from the earlier sample to now characterize the Vb of the entire size-range of the order. MATERIALS AND METHODS: Five gorillas (Gorilla gorilla gorilla-G. g. gorilla) were studied to ascertain Vb in relation to the mechanical properties of five foods. RESULTS: Gorilla Vb ranged from 166.38 cm3 (for the least obdurate food: watermelon) to 8 cm3 (for the most obdurate food: turnip), with an average Vb of 33.50 cm3 across all food types. CONCLUSIONS: When these data were compared to those from our previous studies, we found that gorillas consumed relatively slightly smaller volumes of food compared to the trend found across primates. However, because the more frugivorous gorillas consumed relatively larger pieces of food than the large folivorous monkeys previously studied, including the gorilla data increased the slope of the linear regression between body mass and Vb in anthropoids. Thus, the addition of the largest living primate brings the anthropoid Vb trend closer to the Vb trend of the order. Notwithstanding, there is still negative allometry in anthropoid Vb , in contrast with the isometry in strepsirrhine Vb . Future research should include species with body masses between the smaller anthropoids and gorillas by studying the Vb of large papionids and the other great apes.

Ecological correlates of space use patterns in wild western lowland gorillas.

The distribution of resources is a crucial determinant of animals' space use (e.g., daily travel distance, monthly home range size, and revisitation patterns). We examined how variation in ecological parameters affected variability in space use patterns of western lowland gorillas, Gorilla gorilla gorilla. They are an interesting species for investigating this topic because key components of their diet are nonfruit items (herbaceous vegetation and tree leaves) that occur at low density and are sparsely distributed, and fruits, which show high spatiotemporal variation in availability. We estimated how availability of nonfruit foods and fruit, frugivory (proportion of feeding time consuming fruit), and swamps in areas used by the gorillas influenced daily travel distance, monthly home range size, and revisit frequency to grid cells in the home range of one habituated gorilla group in Loango National Park, Gabon. Using location data from 2015 to 2018, we found that the gorillas decreased their daily travel distance as both the density of nonfruit foods and the proportion of swamps in areas used increased. Daily travel distances were shorter when both frugivory and availability of fruit were higher, yet, daily travel distances were longer when availability of fruit was low but frugivory was still high. Furthermore, monthly home range size increased as frugivory increased and monthly revisit frequencies to an area increased as fruit availability of an area increased. In conclusion, the availability of both nonfruit and fruit influenced the gorillas' space use patterns. Gorillas decreased foraging effort when food availability was high but were willing to incur increasing foraging costs to feed on fruit when availability was low. This study highlights how animals have to adjust their space use with changing resource availability and it emphasizes the value of examining multiple parameters of space use.

Effects of a recurring late-night event on the behavior and welfare of a population of zoo-housed gorillas.

The impact of visitors on the well-being of captive animals presents both positive and potentially negative consequences. While some amount of novelty through visitor stimulation offers the opportunity for a more complex captive environment, anecdotal evidence from primate staff observations at the Smithsonian's National Zoo and Conservation Biology Institute suggested that gorillas exhibited increased restlessness during the annual month-long late night ZooLights (ZL) event than before it. The current study compared activity budgets, aggression (interactions involving contact between conspecifics and displays toward visitors), and abnormal behaviors in two groups of socially housed gorillas for 1-month periods before, during, and after the 2015 ZL event. We also compared the fecal glucocorticoid metabolite profiles of all six gorillas across these same observation periods. Physiologically, most individuals appeared to cope appropriately with the increased visitor presence during the event. We saw little difference in contact aggression; however, abnormal behavior was observed in some gorillas during and after the event, which highlights the importance of individual analysis and data interpretation. As predicted, we found that the majority of gorillas rested less during ZL than during other observation periods, particularly adult females in the mixed sex troop. Preliminary results of this study aided the decision of zoo management to close initially the Great Ape House and subsequently most animal buildings during future ZL events to avoid the potential disruption of normal activity patterns. While the findings of this study were mixed as to the impact of the event on these gorillas, the results suggest that zoos should carefully consider the possible ramifications of keeping great ape buildings open for lengthy multi-night events.