Hand Preferences for a Haptic Task in Chimpanzees (Pan troglodytes)

We tested the hand preferences of 20 chimpanzees (Pan troglodytes) for a haptic task requiring individuals to search for grapes in an opaque bucket filled with water. We compared these data to the hand preferences displayed by the same chimpanzees during reaching and bimanual feeding tasks. The chimpanzees displayed no significant hand preference for the reaching or bimanual feeding tasks, but exhibited a right-hand preference while performing the haptic task. In contrast, New and Old World monkeys display left-hand preferences for similar tasks. We discuss the relevance of these findings for the evolution of handedness in primates.     

Handedness and Grooming in Pan troglodytes: Comparative Analysis Between Findings in Captive and Wild Individuals

Grooming is a complex set of motor actions, common in highly social primates. We tested for asymmetries in hand use during unimanual and bimanual allogrooming in 215 captive chimpanzees. In addition to hand use, we coded in the ethogram whether the manual grooming action co-occurred with the use of the mouth. Overall, grooming did not elicit strong handedness at the individual level, but there is a small yet significant population-level right-hand bias for bimanual grooming. Mouth use during grooming had no influence on hand use. A comparison of the findings with previously published data on handedness for grooming in wild chimpanzees suggests that wild apes are more right-handed than captive individuals are for allogrooming. Collectively, the results suggest that role differentiation of the hands is an important factor in the assessment of handedness for grooming, and perhaps additional manual actions of chimpanzees and other primates.     

An apparatus and procedure for studying the choice component of foraging in a captive group of chimpanzees (Pan troglodytes)

An apparatus is described which was used to investigate the choice component of foraging in a captive group of chimpanzees maintained in a large, outdoor compound at the Yerkes Regional Primate Research Center Field Station. The utilization of more than one apparatus would allow the investigation of other ecological and psychological concepts in nonhuman primates housed under semi-natural conditions.     

A cross-setting study of chimpanzee (Pan troglodytes) personality structure and development: zoological parks and Yerkes National Primate Research Center

This study addressed whether personality ratings using a 43 adjective questionnaire based on the Five-Factor Model generalized from a sample of 202 zoo-housed chimpanzees to a sample of 175 chimpanzees housed in Yerkes National Primate Research Center. Mean interrater reliabilities of adjectival ratings were lower for the chimpanzees housed in Yerkes. In addition, rank order of the interrater reliabilities of items differed between settings. To compare factor structure, we first examined whether we could replicate the original six factor structure found in an earlier study of 100 zoo chimpanzees using principal factors analysis in the Yerkes sample and 102 new zoo chimpanzees. The dominance, extraversion, conscientiousness, and agreeableness factors were clearly replicated in the Yerkes sample and the 102 new zoo chimpanzees. The Neuroticism and Openness factors did not replicate in the Yerkes sample, but they also did not replicate in the new zoo chimpanzees. These findings suggest the need to sample more adjectives representing neuroticism and openness in future versions of the questionnaire. We next sought to determine whether factor structure, as determined by principal components analysis, remained invariant across the two settings. This analysis revealed dominance, extraversion, conscientiousness, and agreeableness factors in both settings and a high level of congruence between the zoo and Yerkes samples for these factors. Finally, we tested whether factor scores in the two samples were similarly related to age and sex. With the exception of differences in age effects for dominance and agreeableness, age, and sex effects were consistent across samples. These findings suggest that, whereas there may be differences in the ease with which ratings are made, personality structure, and development are largely consistent across widely differing settings.     

Using the Tools at Hand: Manual Laterality and Elementary Technology in Cebus spp. and Pan spp.

Cebus and Pan appear to be a remarkable example of evolutionary convergence in behavioral ecology. We examine their apparently analogous solutions to problems posed by laterality of hand function and elementary technology. We scrutinize appropriate published data in a meta-analysis, focusing on Cebus apella and C. capucinus and on Pan paniscus and P. troglodytes. We compare behavioral data in terms of captive versus wild, and tool use versus non-tool use, but notable gaps exist in the data, especially for bonobos. Cebus and Pan spp. are equivalent tool users in captivity, but chimpanzees are notably more extensively so in nature. For hand preference, captive bonobos and wild and captive chimpanzees show ambipreference for non-tool-use patterns. For both Cebus spp. and Pan spp., there is a tendency for individuals to be committed exclusively to one hand or the other for tool use. The data for laterality of hand function fit consistently into the five-level model proposed by McGrew and Marchant (1996).     

Peer interaction in infant chimpanzees

The peer interactions of 6 infant chimpanzees (Pan troglodytes) ranging in age from 18 to 50 months were observed in a seminatural context. The infants and their mothers lived as members of a captive social group at the Yerkes Regional Primate Research Center. An analysis of contact initiations between infants indicated that the most preferred peer interactant was the youngest and the least preferred was the oldest infant. Infants also initiated more interactions with the offspring of adults that had the closest relationships with both themselves and their mothers. These results indicate that a number of factors may influence the peer affiliations of infant chimpanzees, including the age of the infant and the mother's social relationships.     

Inhibition of social behavior in chimpanzees under high-density conditions

This is the first study to investigate the short-term effects of high population density on captive chimpanzees (Pan troglodytes). Subjects of the study were 45 chimpanzees living in five different groups at the Yerkes Regional Primate Research Center. The groups were observed under two conditions: 1) when they had access to both the indoor and outdoor sections of their enclosures; 2) during cold days when they were locked into the indoor runs, which reduced the available space by more than half. Under the high-density condition, allogrooming and submissive greetings decreased, but juvenile play increased. Remarkably, the rate of various forms of agonistic behavior, such as aggression, bluff charge, bluff display, and hooting, occurred less frequently under the high-density condition. This general decrease in adult social activity, including agonistic behavior, can be interpreted as an inhibition strategy to reduce opportunities for conflict when interindividual distances are reduced. This strategy is probably effective only in the short run, however. Behavioral indicators of anxiety, such as rough scratching and yawning, showed elevated rates, suggesting increased social tension under the high-density condition. Am. J. Primatol. 41:213–228, 1997. © 1997 Wiley-Liss, Inc.     

Personality in Captive Chimpanzees: Use of a Novel Rating Procedure

We present a new method to assess the personalities of captive chimpanzees (Pan troglodytes). Using individually generated bipolar constructs, experienced observers rated the personalities of chimpanzees on a five-point scale. Each observer chose a subset of his/her own constructs and we performed a separate factor analysis on each set of ratings. The analyses indicate four factors: dominance, sociability, machiavellianism, and anxiety. We propose that this factor structure will be useful in the development of a single rating instrument with which to assess the personalities of captive chimpanzees.     

Reproductive Parameters of Female<Emphasis Type="Italic">Pan paniscus</Emphasis> and <Emphasis Type="Italic">P. troglodytes</Emphasis>: Quality versus Quantity

We investigated intra- and interspecific differences in life history and reproductive parameters in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes). We compare the parameters of wild and captive females in order to shed light on the influence of habitat or specific differences or both on reproduction. We present new and additional information on reproductive parameters from captive bonobos and chimpanzees. Captive chimpanzees birth more live offspring and have a shorter interbirth interval, but experience higher infant mortality than captive bonobos. Although captive bonobo females tend to start reproduction at a younger age than chimpanzees, this is effectively only so for wild-born females of both species. Ultimately both species reach the same rate of production of offspring surviving to 5 yr. These results contrast with data from the wild. Wild bonobos tend to have higher reproductive success, a higher fertility rate and a shorter interbirth interval than wild chimpanzees. Reproduction is similar for wild and captive bonobos, which suggests that they are producing at their maximum under both conditions. Overall captive chimpanzees perform better than their wild conspecifics, probably because of lower feeding competition. Infant survival is the only specific difference not affected by captivity. Bonobo infants survive better, which suggests that chimpanzee infants are more at risk. We argue that the interspecific variation in reproductive parameters in captivity is related to the different influence of captivity on reproduction and different pressures of external sources of infant and juvenile mortality.     

A survey of entodiniomorphid ciliates in chimpanzees and bonobos

Intestinal entodiniomorphid ciliates are commonly diagnosed in the feces of wild apes of the genera Pan and Gorilla. Although some authors previously considered entodiniomorphid ciliates as possible pathogens, a symbiotic function within the intestinal ecosystem and their participation in fiber fermentation has been proposed. Previous studies have suggested that these ciliates gradually disappear under captive conditions. We studied entodiniomorphid ciliates in 23 captive groups of chimpanzees, three groups of captive bonobos and six populations of wild chimpanzees. Fecal samples were examined using Sheather's flotation and Merthiolate‐Iodine‐Formaldehyde Concentration (MIFC) methods. We quantified the number of ciliates per gram of feces. The MIFC method was more sensitive for ciliate detection than the flotation method. Ciliates of genus Troglodytella were detected in 13 groups of captive chimpanzees, two groups of bonobos and in all wild chimpanzee populations studied. The absence of entodiniomorphids in some captive groups might be because of the extensive administration of chemotherapeutics in the past or a side‐effect of the causative or prophylactic administration of antiparasitic or antibiotic drugs. The infection intensities of ciliates in captive chimpanzees were higher than in wild ones. We suppose that the over‐supply of starch, typical in captive primate diets, might induce an increase in the number of ciliates. In vitro studies on metabolism and biochemical activities of entodiniomorphids are needed to clarify their role in ape digestion. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Menstrual cycles continue into advanced old age in the common chimpanzee (Pan troglodytes)

A long postreproductive lifespan may distinguish women from all other female primates. A long-held consensus among reproductive scientists has been that our closest living relative, the chimpanzee (Pan troglodytes), experiences menstrual cycles until death. However, a recent study of biannual assessments of gonadotropins, but lacking observations of menstruation, concluded that menopause occurs in chimpanzees between 35 and 40 yr of age. A separate report, but on wild chimpanzees, documented fertility through the 40-44 age range in all populations studied. These contradictory reports pose questions about differences between wild and captive populations and about assessments of menopause. The present study revisits this controversy by analyzing longitudinal records of anogenital swelling and menstruation in 89 female chimpanzees aged 6 to 59 yr (n = 2386 records on cycle length), monitored for most of their adult lives at the Yerkes National Primate Research Center. Twenty of these chimpanzees were observed past 39 yr of age; all 20 displayed menstrual cycles beyond this age, as confirmed by at least two observations of menses about 35 days apart. Three of these were older than 50 yr and still displayed menstrual cycles. Only the oldest female appeared menopausal, with cycles of anogenital swelling ceasing 2 yr prior to her death at age 59. Random-effects statistical modeling reveals a slight decrease in cycle length until 20 yr of age and a slight lengthening thereafter. Mean cycle length across the lifespan is 35.4 days. Our findings, based upon actual observations of menstrual cycles, suggest that menopause in the chimpanzee is rare, occurring near the end of the lifespan.     

Chimpanzee and human grips: A new classification with a focus on evolutionary morphology

We observed grips by the hand during locomotor and manipulative behavior of captive chimpanzees to improve our ability to interpret differences between chimpanzees and humans in hand morphology that are not easily explained by current behavioral data. The study generated a new classification of grips,which takes into account three elements of precision and power gripping that appear to distinguish between the chimpanzees and humans, and which have not been explored previously in relation to hand morphology. These elements are (1) the relative force of the precision grips (pinch versus hold), (2) the relative ability to translate and rotate objects by the thumb and fingers (precision handling), and (3) the relative ability to orient a cylindrical object so that it functions effectively as an extension of the forearm (power squeeze). We recommend that this classification be incorporated into protocols for field and laboratory studies of nonhuman primate manipulative behavior, in order to test our prediction that these three elements clearly distinguish humans from chimpanzees and other nonhuman primates. The results of this test will have direct bearing upon decisions as to which grips (with their associated behaviors) are most likely to guide us through kinematic and kinetic analysis to possible explanations for morphological differences between humans and other species. These explanations, in turn, are fundamental to our ability to discern evidence for potential grips and tool behaviors in the manual morphology of fossil hominids.     

Hand Preference for a Novel Bimanual Coordinated Task During Termite Feeding in Wild Western Gorillas (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>)

Although the level of handedness in humans varies cross-culturally, humans are generally described as right-handed, which has been considered a uniquely human trait. Recently, captive chimpanzees (Pan troglodytes) have been shown to exhibit right-hand preference when performing bimanual but not unimanual tasks. Less clear is whether this pattern also occurs in wild chimpanzees and other African apes. Using videos (N = 49) of six wild western gorillas (Gorilla gorilla gorilla) feeding on termites at the Mondika Research Center (Republic of Congo), we tested whether they exhibit hand preference when performing unimanual, i.e., reaching for termite mound pieces; bimanual, i.e., “termite tapping”: rhythmically shaking a piece of termite mound with the dominant hand and collecting the termites in the other hand tasks; or hand transfer prior to bimanual tasks, i.e., transferring a piece of termite mound from one hand to the other. All individuals exhibited exclusive hand preference when performing the bimanual tasks, with five of six gorillas preferring the right hand. Conversely, most individuals did not show any manual preference during the unimanual task. In addition, hand preference during hand transfer revealed clear hand dominance of similar strength and direction of those shown for the bimanual task, suggesting that this measure is as sensitive as the bimanual task itself. Thus, we propose “termite feeding” as a novel task to be considered in future hand-preference studies in wild western gorillas. Our results are in concordance with those for chimpanzees and captive gorillas showing hemispheric specialization for bimanual actions in apes.     

Assessing the effects of cognitive experiments on the welfare of captive chimpanzees (Pan troglodytes) by direct comparison of activity budget between wild and captive chimpanzees

We investigated the effects of cognitive experiments by direct comparison of activity budgets between wild and captive chimpanzees. One goal of captive management is to ensure that the activity budgets of captive animals are as similar as possible to those of their wild counterparts. However, such similarity has rarely been achieved. We compared the activity budget among three groups of chimpanzees: wild chimpanzees in Bossou (Guinea, n = 10), and captive chimpanzees who participated in cognitive experiments (experimental chimpanzees, n = 6) or did not participate in the experiments (nonexperimental chimpanzees, n = 6) at the Primate Research Institute (Japan). The experimental chimpanzees voluntarily participated in computer‐controlled cognitive tasks and small pieces of fruits were provided as rewards. The data from captivity were obtained on the experimental days (weekdays) and nonexperimental days (weekends). In both study sites, we followed each chimpanzee from about 7 a.m. until the time when chimpanzees started to rest in the evening. The behaviors were recorded every 1 min. The results showed that on weekdays, feeding time and resting time of the experimental chimpanzees were almost the same as those of wild chimpanzees. However, for the nonexperimental chimpanzees, feeding time was significantly shorter and resting time was longer than those of the wild chimpanzees. In contrast, no difference was found in feeding time or resting time of the two groups of captive chimpanzees on weekends. The results suggested that the cognitive experiments worked as an efficient method for food‐based enrichment. Am. J. Primatol. 73:1231–1238, 2011. © 2011 Wiley Periodicals, Inc.     

Affiliation promotes the transmission of a social custom: handclasp grooming among captive chimpanzees

Handclasp grooming is a unique social custom, known to occur regularly among some, but not all populations of chimpanzees (Pan troglodytes). As with other cultural behaviors, it is assumed that this distinctive grooming posture is learned socially by one individual from another. However, statistical comparisons among factors thought to influence how a behavior spreads within a group have never, to our knowledge, been conducted. In the present study, the origination and spread of handclasp grooming in a group of captive chimpanzees was followed throughout more than 1,500 h of observation over a period of 12 years. We report on the frequency, bout duration, and number and demography of performers throughout the study period, and compare these findings to those reported for wild populations. We predicted that dyads with strong affiliative ties, measured by time spent in proximity to and grooming one another, were likely to develop a handclasp grooming partnership during the study period. A quadratic assignment procedure was used to compare correlations among observed frequencies of grooming and proximity with handclasp grooming in all possible dyads within the group. As predicted, the formation of new handclasp grooming dyads was positively correlated with the rate of overall grooming and proximity within a dyad. In addition, in nearly all dyads formed, at least one individual had been previously observed to handclasp groom. We concluded that affiliation and individual experience determines the transmission of handclasp grooming among captive chimpanzees.     

Factor Analysis of Multiple Measures of Hand Use in Captive Chimpanzees: An Alternative Approach to the Assessment of Handedness in Nonhuman Primates

We tested whether chimpanzee handedness could be characterized as either unidimensional or multidimensional when considered across multiple measures of hand use. We determined for each of 6 different tasks in a sample of 105 captive chimpanzees hand preferences, and subjected the individual hand preference scores to a factor analysis. Five of the 6 tasks loaded on two separate factors that accounted for 54% of the variance. To assess population-level handedness, we calculated handedness indices for the loadings on each factor, for the item loadings across all factors, and for all tasks including ones that did not load on any factor. There is significant population-level right handedness for all 4 indices, which suggests that chimpanzee handedness is multidimensional and not task specific.     

Genetic basis in motor skill and hand preference for tool use in chimpanzees (Pan troglodytes)

Chimpanzees are well known for their tool using abilities. Numerous studies have documented variability in tool use among chimpanzees and the role that social learning and other factors play in their development. There are also findings on hand use in both captive and wild chimpanzees; however, less understood are the potential roles of genetic and non-genetic mechanisms in determining individual differences in tool use skill and laterality. Here, we examined heritability in tool use skill and handedness for a probing task in a sample of 243 captive chimpanzees. Quantitative genetic analysis, based on the extant pedigrees, showed that overall both tool use skill and handedness were significantly heritable. Significant heritability in motor skill was evident in two genetically distinct populations of apes, and between two cohorts that received different early social rearing experiences. We further found that motor skill decreased with age and that males were more commonly left-handed than females. Collectively, these data suggest that though non-genetic factors do influence tool use performance and handedness in chimpanzees, genetic factors also play a significant role, as has been reported in humans.     

Effect of solar eclipse on the behavior of a captive group of chimpanzees (Pan troglodytes)

A captive group of chimpanzees, housed in an outdoor compound at the Yerkes Regional Primate Research Center, was observed during the annular solar eclipse of May 30, 1984. The behavior of each animal was recorded using an instantaneous scan-sampling technique (Altmann: Behaviour 49:227–265, 1974). Beginning 2 days prior to the eclipse and continuing through the day following the eclipse, data were collected from 1100 to 1300 hours daily. At 1214 hours on the day of the eclipse, when the sky began to darken and the temperature began to decrease, solitary females and females with infants moved to the top of a climbing structure. As the eclipse progressed, additional chimpanzees began to congregate on the climbing structure and to orient their bodies in the direction of the sun and moon. At 1223 hours, during the period of maximum eclipse, the animals continued to orient their bodies toward the sun and moon and to turn their faces upward. One juvenile stood upright and gestured in the direction of the sun and moon. Sunlight began to increase at 1225 hours, and as it became brighter, the animals began to descend from the climbing structure. The behaviors exhibited by the group during the period of maximum eclipse were not observed prior to or following the eclipse nor as darkness approached at normal, daily sunset. These data indicate that a solar eclipse, a rare and uncommon environmental event, can influence and modulate the behavior of chimpanzees.     

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 Faithful Record of Stressful Life Events Recorded in the Dental Developmental Record of a Juvenile Gorilla

The pattern and rate of dental development are critical components of the life history of primates. Much recent research has focused on dental development in chimpanzees and other hominoids, but comparatively little is known about dental development in Gorilla. To date, dental chronologies for Gorilla are based on a sample of 1 and information about variations in the time and timing of crown initiation and completion is lacking. We provide data on dental development in 1 captive, juvenile, female, western lowland Gorilla gorilla gorilla of known age, sex, life events, and date of death (carefully documented as part of zoo records) that experienced various physical insults during her first year of life. The perfect natural experiment allowed us to test the association of the timing of accentuated stress lines in teeth with significant physiological and psychological events during ontogeny of this juvenile gorilla. We analyzed histological sections from 14 permanent teeth (maxillary and mandibular I1-M2) and assessed crown initiation (CI) and crown formation times (CFT) using short- and long-period incremental lines in both enamel and dentine; they are advanced for all teeth compared to previously published chronology. The data suggest a relatively accelerated pace of dental development in gorillas compared to chimpanzees and fit an emerging pattern of an accelerated life history schedule in gorillas. Data on the timing of major accentuated lines in the developing dentition are tightly associated with exact dates of surgical procedures and follow-up hospital visits as recorded on zoo medical records. Our data highlight the importance of captive individuals with well-documented medical records for studying life history.