Food Preferences Among Captive Western Gorillas (Gorilla gorilla gorilla) and Chimpanzees (Pan troglodytes)

I used a zoological park setting to address food preferences among gorillas (Gorilla gorilla gorill) and chimpanzees (Pan troglodytes). Gorillas and chimpanzees are different sizes, and consequently, have been traditionally viewed as ecologically distinct. Sympatric western gorillas and chimpanzees have proved difficult to study in the wild. Limited field data have provided conflicting information about whether gorillas are fundamentally different from chimpanzees in diet and behavior. Fruit eating shapes the behavior of most apes, but it is unclear whether the large-bodied gorillas are an exception to this rule, specifically whether they are less selective and more opportunistic fruit eaters than chimpanzees are. My research provides experimental observational data to complement field data and to better characterize the diets and food preferences of the African apes. During laboratory research at the San Francisco Zoological Gardens, I examined individual and specific differences in food preferences of captive gorillas and chimpanzees via experimental paired-choice food trials with foods that varied in nutritional content. During the study, I offered 2500 paired-food choices to 6 individual gorillas and 2000 additional pairs to them as a group. I also proffered 600 food pairs to 4 individual chimpanzees. Despite expectations of the implications of body size differences for diet, gorillas and chimpanzees exhibited similar food preferences. Both species preferred foods high in non-starch sugars and sugar-to-fiber ratios, and low in total dietary fiber. Neither species avoided foods containing tannins. These data support other suggestions of African apes sharing a frugivorous adaptation.     

A Survey of Chimpanzees (Pan troglodytes troglodytes) and Gorillas (Gorilla gorilla gorilla) in the Selectively Logged Ngotto Forest,Central African Republic

Currently, the timber company Industrie Forestiére du Batalimo is selectively logging the Ngotto Forest in the Central African Republic. The forest is home to a population of chimpanzees (Pan troglodytes troglodytes) and gorillas (Gorilla gorilla gorilla) and provides the Bofi-speaking people of Grima with food, medicine, housing material, and other commodities. Over a 7-month period, the research team conducted a line-transect survey of the great ape population in the forests to the south of Grima to document their distribution. For comparison purposes, the team also surveyed a section of adjacent forest that had already been logged. Ape nests were significantly rarer in the logged forest than in the unlogged forest, and ape nests were most common in the more pristine forests to the south. This report further discusses the effects of logging and other human activities.     

Mandibular Shape, Ontogeny and Dental Development in Bonobos (Pan paniscus) and Chimpanzees (Pan troglodytes)

The postnatal ontogenetic patterns and processes that underlie species differences in African ape adult mandibular morphology are not well understood and there is ongoing debate about whether African ape faces and mandibles develop via divergent or parallel trajectories of shape change. Using three-dimensional (3D) morphometric data, we first tested when in postnatal development differences in mandibular shape are initially evident between sister species Pan troglodytes and P. paniscus. Next, we tested whether each species has a distinct and non-parallel trajectory of mandibular development. Mandibles sampled across a broad developmental range of wildshot bonobos (n = 44) and chimpanzees (n = 59) were radiographed and aged from their dental development. We then collected 3D landmark surface data from all the mandibles. A geometric morphometric analysis of size-corrected 3D data found that bonobos and chimpanzees had parallel and linear ontogenetic trajectories of mandibular shape change. In contrast, mandibular shape was statistically different between P. paniscus and P. troglodytes as early as infancy, suggesting that species shape differences are already established near or before birth. A linear and stable trajectory of shape change suggests that mandibular ontogeny in these apes is unimpacted by non-linear variation in tooth developmental timing.     

Environmental Influences on Sleep Behavior in Captive Male Chimpanzees (Pan troglodytes)

Little is known as to the influence of captivity and stressful events on sleep patterns in primates. We investigated the sleep patterns of 19 male chimpanzees living under similar conditions at the Chimpanzee Sanctuary Uto (CSU) in Kumamoto, Japan, using a behavioral sleep index. We conducted nighttime observations of all subjects during a stable period and then observed three subjects after relocation to an unfamiliar facility at CSU. We estimated length of sleep and nonsleep periods over 13-h video recordings using instantaneous sampling at 1-min intervals to record sleep, which we defined operationally as an inactive posture with the body lying down with the head on the floor or on nesting materials. The 19 subjects slept for a mean ± SEM of 11.3?±?0.26?h during the stable period, and sleep patterns varied significantly among the subjects. The three relocated subjects all showed temporarily decreased sleep duration in the post-move period but subsequently recovered to the levels observed during the stable period when habituated to the new living quarters. These results suggest that a stressful event may induce temporary sleep shortage lasting for >1?wk in captive chimpanzees. Sleep patterns may serve as a useful behavioral index of the stress response, as it is less confounded by other behaviors and the actions of human caretakers than other indices.     

The Influence of Transect Use by Local People and Reuse of Transects for Repeated Surveys on Nesting in Western Lowland Gorillas (Gorilla gorilla gorilla) and Central Chimpanzees (Pan troglodytes troglodytes) in Southeast Cameroon

Monitoring populations of endangered species over time is necessary to guide and evaluate conservation efforts. This is particularly important for nonprotected areas that ensure connectivity between protected populations but are prone to uncontrolled hunting pressure. We investigated whether use of transects by local people and transect reuse for repeated surveys influence great ape nesting and bias results. We conducted simultaneous marked nest count surveys over 12 mo on established and newly opened transects in a nonprotected area subject to traditional heavy use by local people and recorded forest composition and signs of human activity. Chimpanzee and gorilla density estimates and encounter rates per kilometer were lower on established transects than on new ones. A generalized linear model indicated that hunting activity, distance to a regularly used forest trail, and transect type (old or new) predicted chimpanzee nest abundance, and distance to the trail and transect type predicted gorilla nest abundance, with no effect of habitat type (percentage suitable habitat) for either species. We, therefore, suggest that the difference in great ape nesting is a result of high levels of hunting by local people on established transects and forest trails. Our results support the use of repeated line transect surveys for monitoring great ape populations in many circumstances, although we advocate taking precautions in nonprotected areas, to avoid the bias imposed by use of established transects for hunting.     

Allogrooming Behavior and Grooming Site Preferences in Captive Bonobos (Pan paniscus): Association with Female Dominance

I tested the utility of Seyfarth's (1977) model of rank-related attractiveness to explain the distribution of allogrooming behavior among captive bonobos (Pan paniscus). Adult female bonobos generally have high social status and may be dominant over males. As predicted by the model, I found that high-ranking adult females received most allogrooming within each of the four investigated groups. Among adult female-adult female dyads, however, allogrooming was not clearly associated with dominance rank. Contradictory to predictions of the model, the highest-ranking females were responsible for most displacements over allogrooming, and grooming competition is positively correlated with dominance rank. In the second part of this study, I investigated the social significance of allogrooming body site preferences. Bonobos direct significantly most allogrooming to the face of conspecifics, and high- and low-ranking individuals, as well as males and females, differ significantly in their preferences for certain allogrooming sites. Subordinates and males tended to avoid facial grooming and preferred the back and anogenital region, while high-ranking individuals and females directed most allogrooming to the face and head of grooming partners. Data from this study support the hypothesis that high-ranking females are the most attractive grooming partners within a female-centered bonobo society. Many other aspects of allogrooming behavior, however, are not consistent with the model of rank-related attractiveness.     

The location of DNA homologous to human satellite III DNA in the chromosomes of chimpanzee (Pan troglodytes), gorilla (Gorilla gorilla) and orang utan (Pongo pygmaeus)

Radioactive RNA with sequences complementary to human DNA satellite III was hybridised in situ to metaphase chromosomes of the chimpanzee (Pan troglodytes), the gorilla (Gorilla gorilla) and the orangutan (Pongo pygmaeus). A quantitative analysis of the radioactivity, and hence of the chromosomal distribution of human DNA satellite III equivalent sequences in the great apes, was undertaken, and the results compared with interspecies chromosome homologies based upon Giemsa banding patterns. In some instances DNA with sequence homology to human satellite III is present on the equivalent (homologous) chromosomes in identical positions in two or more species although quantitative differences are observed. In other cases there appears to be no correspondence between satellite DNA location and chromosome homology determined by banding patterns. These results differ from those found for most transcribed DNA sequences where the same sequence is located on homologous chromosomes in each species.     

Hand preferences in chimpanzees (Pan troglodytes), bonobos (Pan paniscus), and orangutans (Pongo pygmaeus) in food-reaching and other daily activities

We observed 36 captive great apes [31 chimpanzees (Pan troglodytes),2 bonobos (Pan paniscus), and3 orangutans (Pongo pygmaeus)]reaching for items of food and engaging in other spontaneous behaviors. We consider circumstantial factors that could influence the preferred choice of one hand or the other. Most of the subjects (81%) showed a significant hand preference in food reaching that is consistent in directionality with the hand preferences that they showed during other spontaneous behaviors. The results show that the postural adjustment required when carrying out an action seems to influence directly the expression of manual laterality in individuals. Moreover, the results indicate the possible existence of a relationship between maturational factors and persistence in directionality of hand preferences.     

Blood groups in the species survival plan®, European endangered species program,and managed in situ populations of bonobo (Pan paniscus), common chimpanzee (Pan troglodytes), gorilla (Gorilla ssp.), and orangutan (Pongo pygmaeus ssp.)

Blood groups of humans and great apes have long been considered similar, although they are not interchangeable between species. In this study, human monoclonal antibody technology was used to assign human ABO blood groups to whole blood samples from great apes housed in North American and European zoos and in situ managed populations, as a practical means to assist blood transfusion situations for these species. From a subset of each of the species (bonobo, common chimpanzee, gorilla, and orangutans), DNA sequence analysis was performed to determine blood group genotype. Bonobo and common chimpanzee populations were predominantly group A, which concurred with historic literature and was confirmed by genotyping. In agreement with historic literature, a smaller number of the common chimpanzees sampled were group O, although this O blood group was more often present in wild‐origin animals as compared with zoo‐born animals. Gorilla blood groups were inconclusive by monoclonal antibody techniques, and genetic studies were inconsistent with any known human blood group. As the genus and, specifically, the Bornean species, orangutans were identified with all human blood groups, including O, which had not been reported previously. Following this study, it was concluded that blood groups of bonobo, common chimpanzees, and some orangutans can be reliably assessed by human monoclonal antibody technology. However, this technique was not reliable for gorilla or orangutans other than those with blood group A. Even in those species with reliable blood group detection, blood transfusion preparation must include cross‐matching to minimize adverse reactions for the patient. Zoo Biol 30:427–444, 2011. © 2010 Wiley‐Liss, Inc.     

Survey of Grauer's Gorillas (Gorilla gorilla graueri) and Eastern Chimpanzees (Pan troglodytes schweinfurthi) in the Kahuzi-Biega National Park Lowland Sector and Adjacent Forest in Eastern Democratic Republic of Congo

We describe the distribution and estimate densities of Grauer's gorillas (Gorilla gorilla graueri) and eastern chimpanzees (Pan troglodytes schweinfurthi) in a 12,770-km ² area of lowland forest between the Lowa, Luka, Lugulu, and Oku rivers in eastern Democratic Republic of Congo, the site of the largest continuous population of Grauer's gorillas. The survey included a total of 480 km of transects completed within seven sampling zones in the Kahuzi-Biega National Park lowland sector and adjacent Kasese region and approximately 1100 km of footpath and forest reconnaissance. We estimate total populations of 7670 (4180–10,830) weaned gorillas within the Kahuzi-Biega lowland sector and 3350 (1420–5950) individuals in the Kasese survey areas. Within the same area, we estimate a population of 2600 (1620–4500) chimpanzees. Ape nest site densities are significantly higher within the Kahuzi-Biega lowland sector than in the more remote Kasese survey area in spite of a significantly higher encounter rate of human sign within the lowland sector of the park. Comparison of our data with information obtained by Emlen and Schaller during the first rangewide survey of Grauer's gorillas in 1959 suggests that gorilla populations have remained stable in protected areas but declined in adjacent forest. These findings underscore the key role played by national parks in protecting biological resources in spite of the recent political and economic turmoil in the region. We also show that forest reconnaissance is a reliable and cost-effective method to assess gorilla densities in remote forested areas.     

Interspecific and Ontogenetic Variation in Proximal Pedal Phalangeal Curvature of Great Apes (Gorilla gorilla, Pan troglodytes, and Pongo pygmaeus)

Considerable attention has been devoted to understanding phalangeal curvature in primates, particularly with regard to locomotion. Previous work has found that increased phalangeal curvature may be indicative of increased grasping during suspensory and climbing behaviors, but the details of this relationship, particularly as regards feet, is still unclear. Using behavioral studies to predict an interspecific gradient of variation in pedal phalangeal curvature, I collected digital data from the third and fifth digit proximal pedal phalanges in adult Gorilla gorilla, Pan troglodytes, and Pongo pygmaeus and calculated included angles of phalangeal curvature to assess the appropriateness of pooling digits within taxa and evaluate the association between variation in pedal phalangeal curvature and frequency of climbing behavior. I also used an ontogenetic sample of Pan troglodytes to evaluate the postnatal relationship between variation in phalangeal curvature and grasping behaviors. I found intraspecific variation in phalangeal curvature suggesting among-digit variation in grasping behaviors. Curvature of Pongo was significantly greater than of both Pan and Gorilla. In contrast, Pan was significantly more curved than Gorilla only in comparison of third digits. Ontogenetic decreases in pedal phalangeal curvature among Pan troglodytes accorded well with postnatal decreases in documented climbing frequency. These findings largely support earlier work regarding the association between arboreal grasping and phalangeal curvature, and provide a unique intraspecific analysis that illuminates a number of areas where our knowledge of the behavioral and biomechanical determinants of phalangeal curvature should be explored further, particularly with respect to the role of among-digit variation in phalangeal curvature.     

The distribution of sequences complementary to human satellite DNAs I,II and IV in the chromosomes of chimpanzee (Pan troglodytes), gorilla (Gorilla gorilla) and orang utan (Pongo pygmaeus)

Human satellite DNAs I, II and IV were transcribed to yield radioactive complementary RNAs (cRNAs). These cRNAs were hybridised to metaphase chromosomes of man, chimpanzee (Pan troglodytes), gorilla (Gorilla gorilla) and orang utan (Pongo pygmaeus). The results of this in situ hybridisation were analysed quantitatively and compared with accepted chromosome homologies based on Giemsa banding patterns. The cRNA to satellite II (cRNAII) did not hybridise to chimpanzee chromosomes, although its hybridisation to chromosomes of gorilla and orang utan yielded more autoradiograph grains than hybridisation to human chromosomes, and cRNAIV hybridised to many chromosomes of gorilla and chimpanzee but was almost entirely restricted to the Y chromosome in orang utan. Most sites of hybridisation were located on homologous chromosomes in all four species, but there were a number of sites which showed no correspondence between satellite DNA location and chromosome banding patterns, and others where a given chromosomal location hybridised with different cRNAs in each species. These results are in contrast to those found for many transcribed DNA sequences, where the same sequence is usually located at homologous chromosome sites in different species, and appear to cast doubt on many proposed models of satellite DNA function.     

Comparing the performances of apes (Gorilla gorilla, Pan troglodytes, Pongo pygmaeus) and human children (Homo sapiens) in the floating peanut task

Recently, Mendes et al. [1] described the use of a liquid tool (water) in captive orangutans. Here, we tested chimpanzees and gorillas for the first time with the same "floating peanut task." None of the subjects solved the task. In order to better understand the cognitive demands of the task, we further tested other populations of chimpanzees and orangutans with the variation of the peanut initially floating or not. Twenty percent of the chimpanzees but none of the orangutans were successful. Additional controls revealed that successful subjects added water only if it was necessary to obtain the nut. Another experiment was conducted to investigate the reason for the differences in performance between the unsuccessful (Experiment 1) and the successful (Experiment 2) chimpanzee populations. We found suggestive evidence for the view that functional fixedness might have impaired the chimpanzees' strategies in the first experiment. Finally, we tested how human children of different age classes perform in an analogous experimental setting. Within the oldest group (8 years), 58 percent of the children solved the problem, whereas in the youngest group (4 years), only 8 percent were able to find the solution.     

Hypothesis for the causes and periodicity of repetitive linear enamel hypoplasia in large, wild African (Pan troglodytes and Gorilla gorilla) and Asian (Pongo pygmaeus) apes

Repetitive linear enamel hypoplasia (rLEH) is often observed in recent large-bodied apes from Africa and Asia as well as Mid- to Late Miocene sites from Spain to China. The ubiquity and periodicity of rLEH are not understood. Its potential as an ontogenetic marker of developmental stress in threatened species (as well as their ancient relatives) makes rLEH an important if enigmatic problem. We report research designed to show the periodicity of rLEH among West African Pan troglodytes (12 male, 32 female), Gorilla gorilla (10 male, 10 female), and Bornean and Sumatran Pongo pygmaeus (11 male, 9 female, 9 unknown) from collections in Europe. Two methods were employed. In the common chimpanzees and gorillas, the space between adjacent, macroscopically visible LEH grooves on teeth with two or more episodes was expressed as an absolute measure and as a ratio of complete unworn crown height. In the orangutans, the number of perikymata between episode onsets, as well as duration of rLEH, was determined from scanning electron micrographs of casts of incisors and canines. We conclude that stress in the form of LEH commences as early as 2.5 years of age in all taxa and lasts for several years, and even longer in orangutans; the stress is not chronic but episodic; the stressor has a strong tendency to occur in pulses of two occurrences each; and large apes from both land masses exhibit rLEH with an average periodicity of 6 months (or multiples thereof; Sumatran orangutans seem to show only annual stress), but this needs further research. This is supported by evidence of spacing between rLEH as well as perikymata counts. Duration of stress in orangutans averages about 6 weeks. Finally, the semiannual stressor transcends geographic and temporal boundaries, and is attributed to regular moisture cycles associated with the intertropical convergence zone modified by the monsoon. While seasonal cycles can influence both disease and nutritional stress, it is likely the combination of seasonal variation in fruiting cycles with specific stressors (malaria and/or intestinal parasites, especially hookworm) that results in this widespread phenomenon. This seasonal stress is sufficiently common and of long duration (6 weeks on average in orangutans) that we think rLEH may reflect significant stress in recent and, inferentially, fossil apes. Increasing seasonality may have impinged negatively on later Miocene apes, especially if they lacked a clear birth peak or seasonality in their reproductive cycles.     

Sympatric Chimpanzees (<Emphasis Type=Italic>Pan troglodytes troglodytes</Emphasis>) and Gorillas (<Emphasis Type=Italic>Gorilla gorilla gorilla</Emphasis>) in Loango National Park,Gabon: Dietary Composition,Seasonality, and Intersite Comparisons

Dietary overlap of sympatric apes is complex and understudied, but its examination is essential to further our understanding of species distribution, abundance, and community ecology. Over 3 yr we studied food availability and dietary composition of central chimpanzees (Pan troglodytes troglodytes) and western gorillas (Gorilla gorilla gorilla) in Loango National Park, Gabon. We predicted that living in a habitat dominated by mature forest with sparse terrestrial herbaceous vegetation would lead to an increase in frugivory by gorillas, resulting in increased dietary overlap between the 2 ape species vs. other sites, but that chimpanzees would remain more frugivorous than gorillas. Through fecal analysis we measured overlap in fruit consumption between the 2 species on a bimonthly basis using the Renkonens method. Mean overlap was 27.5% but varied greatly seasonally, ranging between 0.3% and 69%, indicating that when examined on a finer scale, the degree of overlap appears much lower than at other study sites. In contrast to studies elsewhere, there was not a positive correlation between rainfall and fruit availability in Loango, and the long dry season was a period of high fruit production. As observed elsewhere, we found a positive correlation between fruit consumption and fruit availability for both chimpanzees and gorillas and we found that chimpanzees were more frugivorous than gorillas. A very low availability of herbs did not lead to increased frugivory by gorillas nor increased overlap between the 2 ape species vs. other field sites. We conclude that forest composition, fruit availability, and dietary variability of sympatric species can vary greatly between locations, and that chimpanzees and gorillas can adapt to primary forest with little undergrowth, where they concentrate their diet on fruit and leaves.