Pneumatic processes in the temporal bone of chimpanzee (Pan troglodytes) and gorilla (Gorilla gorilla)

The ontogeny of human temporal bone pneumatization has been well studied from both comparative and clinical perspectives. While a difference in the extent of air cell distribution has been noted in our closest living relatives, chimpanzees and gorillas, the processes responsible have been relatively unexplored. To examine these processes, a large, age‐graded series of hominoid skulls was radiographed and the progress of pneumatization recorded. Additionally, a subsample of 30 chimpanzees and 12 gorillas was subjected to high‐resolution CT scanning. Neonatal specimens show a well‐developed mastoid antrum, as well as a capacious hypotympanum extending into the petrous apex. In African apes, as in humans, the mastoid antrum serves as the focus for air cell expansion into the mastoid and immediately adjacent areas. In chimpanzees and gorillas, however, a pronounced lateral structure, described as the squamous antrum, serves as the focus of pneumatization for anterior structures such as the squamous and zygomatic. The diminution of this structure in Homo sapiens explains the difference in air cell distribution in these regions. J. Morphol. 241:127–137, 1999. © 1999 Wiley‐Liss, Inc.     

Nationwide census of gorilla (gorilla g. gorilla) and chimpanzee (Pan t. troglodytes) populations in Gabon

A census was made of gorilla and chimpanzee populations throughout Gabon between December 1980 and February 1983. The aim of the census was to estimate the total numbers of both species and describe their distributions. The method was based on nest counts from line transects which allowed the calculation of population densities of all individuals except suckling infants. Fifteen types of habitat were recognized and defined in terms of their structural features. In the initial phase of the study we did transects in each habitat-type and computed mean densities for each species in each habitat-type. In the second phase of the study we estimated the sizes of gorilla and chimpanzee populations throughout the country by extrapolation from these population density values. We did transects in all areas of the country and conducted interviews to check the accuracy of the population totals obtained by extrapolation. Corrections were made to the extrapolated totals to take into account different levels of hunting pressure and other human activities found to modify ape population densities. Total populations of 34,764 gorillas and 64,173 chimpanzees were estimated. An error of ± 20% was associated with the estimated population totals, which allows the conclusion that Gabon contains 35,000 ± 7,000 gorillas and 64,000 ± 13,000 chimpanzees. The figure for gorillas is much larger than previous estimates. This seems to be because (1) gorillas occur in almost all types of forest and are not restricted to man-made secondary forest as had been though; and (2) the geographical distribution of gorillas in Gabon is wider than previously believed. Gabon's large areas of undisturbed primary forest offer exceptional potential for conservation, not only of gorillas and chimpanzees, but also of the intact tropical rain forest ecosystems which they inhabit.     

Behavioral Responses of Gorillas to Habituation in the Dzanga-Ndoki National Park,Central African Republic

We monitored the impact of habituation for tourism through changes in gorillas' behavior during the habituation process at Bai Hokou (Dzanga-Ndoki National Park, Central African Republic) from August 1996 to December 1999. From August 1998 onwards we focused on one gorilla group: the Munye. During the habituation process, it became increasingly easier to locate and to remain with the gorillas. Their initial reactions of aggression, fear and vocalization upon contact were replaced increasingly by ignoring us. Curiosity appeared to be an intermediate stage in the process. The way in which contacts with the Munye ended became more subdued over time. Regular daily contact is important in promoting habituation. Likewise, contacting gorillas while they are in a tree or in dense forest provides positive results compared to open habitat. Contacts within 10 m and contacts without forewarning the gorillas of observer presence, e.g., via tongue clacking, should be avoided. As of December 1999, habituation had progressed well; habituation of western gorillas is feasible. However, the gorillas experience negative effects during the habituation process, showing, for example, an increase in daily path length, and reactions of aggression, fear and vocalization upon contact. Impacts diminish over time. Given these and other potentially negative effects, the decision to begin habituation should not be taken lightly.     

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.     

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.     

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.     

Brain structure variation in great apes, with attention to the mountain gorilla (Gorilla beringei beringei)

This report presents data regarding the brain structure of mountain gorillas (Gorilla beringei beringei) in comparison with other great apes. Magnetic resonance (MR) images of three mountain gorilla brains were obtained with a 3T scanner, and the volume of major neuroanatomical structures (neocortical gray matter, hippocampus, thalamus, striatum, and cerebellum) was measured. These data were included with our existing database that includes 23 chimpanzees, three western lowland gorillas, and six orangutans. We defined a multidimensional space by calculating the principal components (PCs) from the correlation matrix of brain structure fractions in the well-represented sample of chimpanzees. We then plotted data from all of the taxa in this space to examine phyletic variation in neural organization. Most of the variance in mountain gorillas, as well as other great apes, was contained within the chimpanzee range along the first two PCs, which accounted for 61.73% of the total variance. Thus, the majority of interspecific variation in brain structure observed among these ape taxa was no greater than the within-species variation seen in chimpanzees. The loadings on PCs indicated that the brain structure of great apes differs among taxa mostly in the relative sizes of the striatum, cerebellum, and hippocampus. These findings suggest possible functional differences among taxa in terms of neural adaptations for ecological and locomotor capacities. Importantly, these results fill a critical gap in current knowledge regarding great ape neuroanatomical diversity.     

An unusual incident of adoption in a wild chimpanzee (Pan troglodytes) population at Gombe National Park

Allomothering and adoption are well documented across primate species. Multiple hypotheses have been proposed to explain the evolution of such behavior according to the costs and benefits to the caregiver, mother, and infant. Permanent adoptions and allomothering have been observed in chimpanzees, but they typically involve the infants' siblings or nulliparous females. Here, I report a unique incident of adoption where an infant was adopted by its grandmother without the death of its mother. I conclude by considering how the adoption may have benefited the grandmother, mother, and infant.     

Ranging Patterns of a Western Gorilla Group During Habituation to Humans in the Dzanga-Ndoki National Park,Central African Republic

The ranging patterns of western gorillas have been described in relation to their more frugivorous diet as compared to mountain gorillas. However, the lack of data from habituated groups prevents a full understanding of their ranging behavior and its implications for western gorilla social behavior. Unhabituated gorillas often flee from approaching researchers, possibly resulting in longer daily ranges and home range expansions and/or shifts. I report on the influence of habituation on the ranging patterns of a western gorilla group. I analyzed progress in habituation through changes in gorilla reactions, displayed upon detecting the observers. I measured daily path length and home range while following the group's trails and monitored diet via fecal analyses and direct observations. During the study period, habituation progressed and the gorillas increasingly ignored the arrival of observers. As a result, daily path lengths were longer during the early stages of habituation, when the gorillas were avoiding observers. There was no range expansion or shift, though the measure is more difficult to investigate, being subject to error due to variation in sample size. We recorded seasonal changes in the consumption of fruit by gorillas, but no direct effect on ranging patterns is detectable with the data available, which is likely a consequence of the combined effect of habituation and seasonality on group movements. The findings are not only important for understanding the extent of human influence while conducting research on wild gorillas but also suggest the need for caution when interpreting results from non-habituated gorillas.     

Insect-eating by sympatric Lowland gorillas (Gorilla g. gorilla) and chimpanzees (Pan t. troglodytes) in the Lopé Reserve,Gabon

Sympatric populations of lowland gorillas (Gorilla gorilla gorilla) and chimpanzees (Pan troglodytes troglodytes) in the Lopé Reserve in central Gabon consumed insects at similar average frequencies over a 7-year period (30% versus 31% feces contained insect remains). Data came mostly from fecal analysis supplemented by observation and trail evidence. The weaver ant (Oecophylla longinoda) was the species eaten most frequently by both gorillas and chimpanzees. Other species of insects wore eaten but there was virtually no overlap: Chimpanzees used tools to eat Apis bees (and their honey) and two large species of ants; gorillas ate three species of small ants. Thus, despite their shared habitat, the esources utilized were not identical as gorillas do not show the tool-use “technology” of chimpanzees. The frequency of insect-eating by both species of ape varied seasonally and between years but in different ways. This variation did not seem to be related to the ratio of fruit to foliage in their diets. Gorillas of all age-classes ate insects at similar rates. Comparisons with insectivory by other populations of gorillas indicate differences exist. Mountain gorillas (Gorilla g. beringei) in the Virunga Volcanoes, Rwanda, consume thousands of invertebrates daily, eating them inadvertently with handfuls of herbaceous foods but they deliberately ingest insect-foods only rarely. Lowland gorillas at Lopé habitually ate social insects, and their selective processing of herbaceous foods probably minimizes inadvertent consumption of other invertebrates. Gorillas at Belinga in northeastern Gabon, 250 km from Lop6, ate social insects at similar rates but ignored weaver ants in favor of Cubitermes sulcifrons, a small species of termite that occurs at Lopé but was not eaten by gorillas. This indicates that local traditions similar to those reported for chimpanzees also exist amongst populations of gorillas. © 1992 Wiley-Liss, Inc.     

Effects of group dynamics and diet on the ranging patterns of a western gorilla group (Gorilla gorilla gorilla) at Bai Hokou, Central African Republic

This study describes how group dynamics and diet have influenced the ranging patterns of a western gorilla group at Bai Hokou, Central African Republic. The results are compared with those from an earlier study [Cipolletta, International Journal of Primatology, 2003], when the same group was larger and undergoing the process of habituation to humans. Data were obtained from maps of the gorillas' travel routes, direct observations, and analysis of fecal samples. Through the years, the group has experienced a decrease in size, from eight to three individuals, with periods of membership fluctuation. The male's search for new mates resulted in a larger home range than was recorded when the group consisted of more individuals. Moreover, despite an average group size of three throughout this study, the monthly range and mean daily path length (DPL) were also larger when the group was acquiring/losing members in new areas, than when no new members joined or left the group. Fruit was consumed year-round, although more heavily so during wet months. The influence of fruit consumption on the ranging patterns was concealed initially by the effect of habituation [Cipolletta, International Journal of Primatology, 2003], and later (at least partially) by the male's search for new mates. In the last 14 months of the study, when the group numbered only three individuals and was ranging in a restricted area, the average DPL, but not the monthly range, increased when the gorillas were consuming more fruit.     

Bipedality in chimpanzee (Pan troglodytes) and bonobo (Pan paniscus): testing hypotheses on the evolution of bipedalism

A host of ecological, anatomical, and physiological selective pressures are hypothesized to have played a role in the evolution of hominid bipedalism. A referential model, based on the chimpanzee (Pan troglodytes) and bonobo (Pan paniscus), was used to test through experimental manipulation four hypotheses on the evolution of hominid bipedalism. The introduction of food piles (Carry hypothesis) increased locomotor bipedality in both species. Neither the introduction of branches (Display hypothesis) nor the construction of visual barriers (Vigilance hypothesis) altered bipedality in either species. Introduction of raised foraging structures (Forage hypothesis) increased postural bipedality in chimpanzees. These experimental manipulations provided support for carrying of portable objects and foraging on elevated food-items as plausible mechanisms that shaped bipedalism in hominids.     

Morphological analyses and 3D modeling of the tongue musculature of the chimpanzee (Pan troglodytes)

Knowledge of the comparative anatomy of tongue musculature is crucial to the discussion of the origin and the evolution of speech because of the indispensable role played by this organ in speech. However, the tongue musculature of primates has rarely been studied. In a previous study, the author analyzed human tongue musculature and developed a 3D model of this organ [Takemoto, Journal of Speech, Language, and Hearing Research 44:95-107, 2001]. In this study, the tongue musculature of chimpanzees was examined using methods similar to those used for humans. Results showed that tongue musculature was topologically the same for both humans and chimpanzees. As in humans, the tongue musculature of chimpanzees consisted of inner and outer regions. The inner musculature was composed of serial "structural units," made up of two types of laminae whose fibers were perpendicular to the tongue surface. The outer musculature was a thin layer of fibers oriented parallel to the surface and superficial to the inner musculature. Although the tongue musculature of humans and chimpanzees is similar, the external shapes differ: the chimpanzee tongue is flat, whereas the human tongue is round. Applying the muscular hydrostat theory to the external shape of the tongue suggests that the primary actions of the chimpanzee tongue are protrusion and retrusion, whereas the human tongue can be deformed in the oral cavity with a high degree of freedom. It is hypothesized that the evolution of the external shape of the tongue is one of the factors that led to the development of human speech. The results of this study suggest that modeling based on muscular hydrostatic theory of the effects of changes in external tongue shape on articulatory movements should be included in discussions on the origin of speech.     

Parasitological analyses of the male chimpanzees (Pan troglodytes schweinfurthii) at Ngogo, Kibale National Park, Uganda

Numerous intestinal parasites identified in populations of wild nonhuman primates can be pathogenic to humans. Furthermore, nonhuman primates are susceptible to a variety of human pathogens. Because of increasing human encroachment into previously nonimpacted forests, and the potential for disease transmission between human and nonhuman primate populations, further detailed investigations of primate ecological parasitology are warranted. For meaningful comparisons to be made, it is important for methods to be standardized across study sites. One aspect of methodological standardization is providing reliable estimates of parasite prevalence and knowing how many samples are needed to adequately estimate an individual's parasite prevalence. In this study the parasitic fauna of 37 adult, adolescent, and juvenile male chimpanzees from the Ngogo group, Kibale National Park, Uganda, were assessed from 121 fecal samples collected over a 3-month period. Twelve taxa of intestinal species (five helminth and seven protozoan) were recovered from the samples. The four most prevalent species were Troglodytella abrassarti (97.3%), Oesophagostomum sp. (81.1%), Strongyloides sp. (83.8%), and Entamoeba chattoni (70.3%). No one species was found in all samples from any one animal, and Troglodytella abrassarti, the most common intestinal organism, was found in all of the serial samples of only 69.4% of the chimpanzees. The cumulative species richness for individuals significantly increased for every sequential sample (up to three to four samples) taken per animal during this study. The results indicate that to accurately diagnose total intestinal infection and evaluate group prevalence, three to four sequential samples from each individual must be collected on nonconsecutive days. This conclusion applies only to short study periods in which possible seasonal effects are not taken into consideration. Validation of these results at different study sites in different regions with different climatic patterns is needed.     

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.     

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.