A comparative study of the variety and complexity of object manipulation in captive chimpanzees (Pan troglodytes) and bonobos (Pan paniscus)

Four types of specific objects: wooden spoons, metal bowls, plastic boxes, and cotton towels were introduced in a similar setting to two captive groups of different species in the genusPan, the bonobo and the chimpanzee. In total, 582 unique manipulation forms were distinguished by a set of variables: types of objects, motor patterns, body-parts used, the number of objects manipulated, and types of orienting manipulation. In sum, chimpanzees and bonobos were not so different in the variety and the complexity of object manipulation forms. However, comparison of the two species revealed significant differences as follows: (1) chimpanzees preferred to use only one hand during manipulation of both single- and multiple-objects, whereas in the case of multiple-objects bonobos used both hands significantly more often; (2) chimpanzees performed more orienting manipulations in single-object manipulations than did bonobos, whereas the reverse was the case in multiple-object manipulations; and (3) chimpanzees' object manipulations were overall more substrate-oriented than were bonobos'. The factors producing these differences are discussed in relation to positional behaviors and habitual tool use in the two species.     

How does stone-tool use emerge? Introduction of stones and nuts to naïve chimpanzees in captivity

Nut-cracking behavior has been reported in several communities in West Africa but not in East and Central Africa. Furthermore, even within nut-cracking communities, there are individuals who do not acquire the skill. The present study aimed to clarify the cognitive capability required for nut-cracking behavior and the process through which the the nut-cracking behavior emerges. To examine emergence, we provided three naïve adult chimpanzees with a single opportunity to observe human models. A human tester demonstrated nut-cracking behavior using a pair of stones and then supplied stones and nuts to the chimpanzee subjects. Two out of three chimpanzees proceeded to hit a nut on an anvil stone using a hammer stone, one of whom succeeded in cracking open the nuts during the first test session. The third chimpanzee failed to crack open nuts. We used four variables (object, location, body part used, and action) to describe stone/nut manipulation in order to analyze further the patterns of object manipulation exhibited by the subjects. The analysis revealed that there were three main difficulties associated with nut-cracking behavior. (1) The chimpanzee who failed at the task never showed hitting action. (2) The chimpanzee who failed at the task manipulated nuts but rarely stones. (3) The combination of three objects was not commonly observed in the three chimpanzees. We also discuss our results with reference to the effect of enculturation in captivity and the social context of learning in the wild.     

Behavioral and functional strategies during tool use tasks in bonobos

Different primate species have developed extensive capacities for grasping and manipulating objects. However, the manual abilities of primates remain poorly known from a dynamic point of view. The aim of the present study was to quantify the functional and behavioral strategies used by captive bonobos (Pan paniscus) during tool use tasks. The study was conducted on eight captive bonobos which we observed during two tool use tasks: food extraction from a large piece of wood and food recovery from a maze. We focused on grasping postures, in‐hand movements, the sequences of grasp postures used that have not been studied in bonobos, and the kind of tools selected. Bonobos used a great variety of grasping postures during both tool use tasks. They were capable of in‐hand movement, demonstrated complex sequences of contacts, and showed more dynamic manipulation during the maze task than during the extraction task. They arrived on the location of the task with the tool already modified and used different kinds of tools according to the task. We also observed individual manual strategies. Bonobos were thus able to develop in‐hand movements similar to humans and chimpanzees, demonstrated dynamic manipulation, and they responded to task constraints by selecting and modifying tools appropriately, usually before they started the tasks. These results show the necessity to quantify object manipulation in different species to better understand their real manual specificities, which is essential to reconstruct the evolution of primate manual abilities.     

Sex Differences in Object Manipulation in Wild Immature Chimpanzees (Pan troglodytes schweinfurthii) and Bonobos (Pan paniscus): Preparation for Tool Use?

Sex differences in immatures predict behavioural differences in adulthood in many mammal species. Because most studies have focused on sex differences in social interactions, little is known about possible sex differences in ‘preparation’ for adult life with regards to tool use skills. We investigated sex and age differences in object manipulation in immature apes. Chimpanzees use a variety of tools across numerous contexts, whereas bonobos use few tools and none in foraging. In both species, a female bias in adult tool use has been reported. We studied object manipulation in immature chimpanzees at Kalinzu (Uganda) and bonobos at Wamba (Democratic Republic of Congo). We tested predictions of the ‘preparation for tool use’ hypothesis. We confirmed that chimpanzees showed higher rates and more diverse types of object manipulation than bonobos. Against expectation, male chimpanzees showed higher object manipulation rates than females, whereas in bonobos no sex difference was found. However, object manipulation by male chimpanzees was play-dominated, whereas manipulation types of female chimpanzees were more diverse (e.g., bite, break, carry). Manipulation by young immatures of both species was similarly dominated by play, but only in chimpanzees did it become more diverse with age. Moreover, in chimpanzees, object types became more tool-like (i.e., sticks) with age, further suggesting preparation for tool use in adulthood. The male bias in object manipulation in immature chimpanzees, along with the late onset of tool-like object manipulation, indicates that not all (early) object manipulation (i.e., object play) in immatures prepares for subsistence tool use. Instead, given the similarity with gender differences in human children, object play may also function in motor skill practice for male-specific behaviours (e.g., dominance displays). In conclusion, even though immature behaviours almost certainly reflect preparation for adult roles, more detailed future work is needed to disentangle possible functions of object manipulation during development.     

Developing classification in action: II. Young chimpanzees (Pan troglodytes)

The development of spontaneous object manipulation in 5 chimpanzees (Pan troglodytes) from ages 15 to 54 months was investigated, focusing on formal properties of subjects’ acts and the objects they manipulated. Young chimpanzees’ manipulation progress from serial one-at-a-time acts on one object to parallel two-at-a-time acts on two or more objects. With age, simultaneous acts become increasingly transformational and identical or reciprocal to each other. Moreover, the class properties of objects manipulated simultaneously change. When presented with objects belonging to two different classes, subjects shift, with age, from manipulating different objects to manipulating identical or similar objects. In all these respects young chimpanzee’ development is similar to human infants’. In others it differs. Most especially, the onset age is later and the development is slower as well as less structurally complex.     

Precision grips in young chimpanzees

A precision grip, thumb-finger opposition, has been regarded as an uniquely human trait. Napier's conclusion that chimpanzees were incapable of precision grip was based on two subjects and prehension of a single object (i.e., a grape). The purpose of the present study was to specify grip type and hand use by 13 young chimpanzees to prehend three different-sized food objects. The subjects were laboratory raised (eight males and five females) and ranged in age from 27 to 58 months. An ethogram was devised that comprised 43 different grip types: ten configurations of precision grips were found, in addition to imprecise or inefficient grip types (nine types), thumb-to-finger opposition (10 types), power grips (two types), and a variety of other grips (12 types). Subjects most often prehended were very small-sized (5 mm × 5 mm × 3 mm) or small-sized (10 mm × 10 mm × 3 mm) food objects with precision and imprecise grips. An analysis of latency to prehend, i.e., efficiency, revealed (1) precision grips were equally efficient for all object sizes; (2) power grips were most efficient with the largest object (a grape); (3) with imprecise grips, the left hand was more efficient than the right with small objects, and with power grips the right hand was more efficient than the left for medium-sized objects. No population handedness was observed, but individual handedness was seen in nine subjects for some grip types and some object sizes. This study provides evidence that young chimpanzees preferentially use a true precision grip to prehend small and very small objects. © 1996 Wiley-Liss, Inc.     

Perspectives on object manipulation and action grammar for percussive actions in primates

The skill of object manipulation is a common feature of primates including humans, although there are species-typical patterns of manipulation. Object manipulation can be used as a comparative scale of cognitive development, focusing on its complexity. Nut cracking in chimpanzees has the highest hierarchical complexity of tool use reported in non-human primates. An analysis of the patterns of object manipulation in naive chimpanzees after nut-cracking demonstrations revealed the cause of difficulties in learning nut-cracking behaviour. Various types of behaviours exhibited within a nut-cracking context can be examined in terms of the application of problem-solving strategies, focusing on their basis in causal understanding or insightful intentionality. Captive chimpanzees also exhibit complex forms of combinatory manipulation, which is the precursor of tool use. A new notation system of object manipulation was invented to assess grammatical rules in manipulative actions. The notation system of action grammar enabled direct comparisons to be made between primates including humans in a variety of object-manipulation tasks, including percussive-tool use.     

Development of the visual preference of chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) for photographs of primates: effect of social experience

In a study by Tanaka (2003) five captive chimpanzees preferred photographs of humans to those of chimpanzees. All the subjects were raised by humans and lived in captivity for many years. This suggests their preference might have developed through social experience. In this study examined this hypothesis by using three young chimpanzees raised by their mothers in a captive chimpanzee community. The young chimpanzees were tested four times before six years of age. I also tested eight adult chimpanzees that had been in captivity for more than 20 years. Each subject was presented with digitized color photographs of different species of primates on a touch-sensitive screen. The subjects received a food reward when they touched a photograph, irrespective of which photograph they touched. All the adult chimpanzees touched photographs of humans more frequently than those of any other species of primate. Two of the young chimpanzees showed no species preference before reaching 5 years of age, when they started to show preference for humans. The remaining young chimpanzee consistently preferred chimpanzees. These results suggest that development of visual preference of chimpanzees is affected by social experience during infancy.     

Advanced age influences chimpanzee behavior in small social groups

Management strategies for captive chimpanzees (Pan troglodytes) must begin to take into account the increasing age of the captive chimpanzee population. This study represents a baseline assessment of the relationship between advancing age and behavior among male and female chimpanzees living in pairs and trios in indoor/outdoor runs. Data collected on 14 old individuals (30–44 years old) and 20 younger adult individuals (11–22 years old) totaled 240 hours. Levels of agonistic and affiliative social behavior, non‐social activity, abnormal behaviors, and behavioral indicators of anxiety were evaluated. In the same captive setting, the behavior of old chimpanzees was significantly different from younger chimpanzees. Old chimpanzees showed less aggression and moved about their enclosures less. Old females behaved submissively more often than younger adult females; the reverse was found among males. However, affiliative social behavior occurred at similar levels in old and younger adult chimpanzees, implying continued need for social housing with advancing age. The effect of enrichment devices may differ for aged female chimpanzees, given their submissiveness and the lower levels of object manipulation found in aged subjects. These results suggest that aging in chimpanzees may be accompanied by altered patterns of social interaction, requiring careful attention to the compatibility of social partners. Zoo Biol 19:111–119, 2000. © 2000 Wiley‐Liss, Inc.     

Hierarchical complexity of combinatorial manipulation in capuchin monkeys (Cebus apella)

The purpose of this study was to examine the hierarchical complexity of combinatorial manipulation in capuchin monkeys (Cebus apella). Two experiments were conducted. In Experiment 1 capuchins were presented with an apparatus designed to accommodate the use of probing tools. In Experiment 2 the same capuchins were presented with sets of nesting containers. Five of the ten subjects used probing tools and seven subjects placed objects in the containers. The capuchins' behavior reflected three hierarchically organized combinatorial patterns displayed by chimpanzees and human infants. Although the capuchins sometimes displayed the two more complex patterns (“pot” and “subassembly”), their combinatorial behavior was dominated by the simplest pattern (“pairing”). In this regard capuchins may not attain the same grammar of manipulative action that has been reported for chimpanzees and young human children. © 1994 Wiley-Liss, Inc.     

Use of novel objects to enhance environments of captive chimpanzees

A small group of chimpanzees was allowed to manipulate different quantities of novel objects in order to enhance their captive conditions. This procedure led to a general increase in the manipulation frequency by the apes. It decreased their inactivity and their self-grooming, and almost eliminated the abnormal behaviors displayed by one individual. The distribution of novel objects did not affect the total frequency of social interactions and did not increase the aggressivity of the animals, even though dominant individuals secured most of the objects when only a small number of them were available. Following their familiarization with the novel objects, the chimpanzees' manipulation frequency decreased whereas self-grooming and abnormal behaviors were increased. Suggestions are made on how to maximize the use of a limited bank of toys to occupy captive chimpanzees.     

Chimpanzee vocal signaling points to a multimodal origin of human language

The evolutionary origin of human language and its neurobiological foundations has long been the object of intense scientific debate. Although a number of theories have been proposed, one particularly contentious model suggests that human language evolved from a manual gestural communication system in a common ape-human ancestor. Consistent with a gestural origins theory are data indicating that chimpanzees intentionally and referentially communicate via manual gestures, and the production of manual gestures, in conjunction with vocalizations, activates the chimpanzee Broca's area homologue--a region in the human brain that is critical for the planning and execution of language. However, it is not known if this activity observed in the chimpanzee Broca's area is the result of the chimpanzees producing manual communicative gestures, communicative sounds, or both. This information is critical for evaluating the theory that human language evolved from a strictly manual gestural system. To this end, we used positron emission tomography (PET) to examine the neural metabolic activity in the chimpanzee brain. We collected PET data in 4 subjects, all of whom produced manual communicative gestures. However, 2 of these subjects also produced so-called attention-getting vocalizations directed towards a human experimenter. Interestingly, only the two subjects that produced these attention-getting sounds showed greater mean metabolic activity in the Broca's area homologue as compared to a baseline scan. The two subjects that did not produce attention-getting sounds did not. These data contradict an exclusive "gestural origins" theory for they suggest that it is vocal signaling that selectively activates the Broca's area homologue in chimpanzees. In other words, the activity observed in the Broca's area homologue reflects the production of vocal signals by the chimpanzees, suggesting that this critical human language region was involved in vocal signaling in the common ancestor of both modern humans and chimpanzees.     

Multiple classification performance of juvenile chimpanzees,normal children,and retarded children

The cognitive capacities of juvenile chimpanzees, normal children, and retarded children were evaluated by using a nonverbal, Piagetian-type multiple classification task. The three groups of subjects were tested with the same two by two stimulus matrix, which was formed by combining two different colors with two different shapes. On each problem the subjects were required to select one of four stimulus items from the response tray and place it inside the empty cell of the stimulus matrix. It was found that the human and nonhuman primates tested were able to select objects with correct color and shape cues (C⁺/S⁺)significantly more often than the other objects with only color cues correct (C⁺/S^-), only shape cues correct (C^-/S⁺), or neither cue correct (C^-/S^-). Although the two groups of human children were able to select the C⁺/S⁺ objects about 100% of the time by the end of testing, the normal children required significantly fewer problems than the retarded children. The juvenile chimpanzees needed significantly more problems than the human subjects before they consistently selected the C⁺/S⁺ object on the first trial and attained a level of correct responding that was above chance. Moreover, their level of performance did not exceed 70% correct. These data suggest that the human children (both normal and retarded) used a conceptual strategy, while the juvenile chimpanzees employed a perceptual strategy to solve the multiple classification problems. The relationship between language and conceptual problem-solving strategies for Piagetiantype tasks is discussed. Portions of the data reported in this study were presented at the Annual Meeting of the Southern Psychological Association, Atlanta, Georgia, March 15–18, 1978.     

Spatial aspects of spontaneous object grouping by young chimpanzees(Pan troglodytes)

I report spontaneous spatial object grouping in five chimpanzees (Pan troglodytes)ranging from 1 to 4 years of age. I recorded subjects' spontaneous interactions with groups of objects, noting the spatial relations in the groupings and the constructive processes that the subjects adopted. Though one subject developed vertical stacks, none of them realized horizontal alignments or spatial correspondences between groups. All subjects showed consistent trends in their constructive processes toward manipulating objects in relation to the body: they increasingly manipulated objects globally, that is to say, they moved or held objects together with the same part of the body. They also increasingly placed objects on the body or in correspondence to their symmetrical body parts. The increasing importance assigned by chimpanzees to body- object relations over object- object relations contrasts with human developmental trends in the domain of manipulative space.     

Use of temporary and semipermanent enrichment objects by five chimpanzees

At the Chimpanzee and Human Communication Institute, caregivers provide the 5 chimpanzees who reside there with many different forms of social, food, habitat, and object enrichment. In this study, we examined the chimpanzees' use of both semipermanent and temporary objects. Semipermanent objects included cargo nets, climbing structures, a treat mound, and other objects that were present at the chimpanzees' enclosure throughout the duration of this study. Each day, 50 temporary objects were placed in the chimpanzees' outdoor enclosure or indoor exercise rooms. Frequency of use was examined in 2 conditions: rotated and same. In the rotated condition, temporary objects were replaced with different temporary objects after 3 hr. In the same condition, temporary objects were presented for the entire day. Focal and scan sampling were used to record the chimpanzees' use of enrichment objects. Observers collected focal sample data to record the chimpanzees' initial reaction to objects when entering the indoor exercise rooms, outdoor enclosures, or both at 9:00 a.m. and 1.00 p.m. A total of 35 hr of focal data and 156 hr of scan data were collected over an 8-week period. Temporary object rotation in- creased the overall frequency of temporary object use both in the initial 15 min of focal sample data and during the following 6 hr of scan sample data for 4 of the chimpanzees. All of these chimpanzees used both semipermanent and temporary objects throughout the day. Each chimpanzee's pattern of use was unique. The results of this study emphasize the importance of temporary object rotation and presentation of both temporary and semipermanent objects to captive chimpanzee environments.     

Dexterity and technique in termite fishing by chimpanzees (Pan troglodytes troglodytes) in the Goualougo Triangle,Republic of Congo

Although the phenomenon of termite fishing by chimpanzees (Pan troglodytes) has historical and theoretical importance for primatology, we still have a limited understanding of how chimpanzees accomplish this activity, and in particular, about details of skilled actions and the nature of individual variation in fishing techniques. We examined movements, hand positions, grips, and other details from remote video footage of seven adult and subadult female chimpanzees using plant probes to extract Macrotermes muelleri termites from epigeal nests. Six chimpanzees used exclusively one hand (left or right) to grip the probe during termite fishing. All chimpanzees used the same repertoire of actions to insert, adjust, and withdraw the probe but differed in the frequency of use of particular actions. Chimpanzees have been described as eating termites in two ways—directly from the probe or by sweeping them from the probe with one hand. We describe a third technique: sliding the probe between the digits of one stationary hand as the probe is extracted from the nest. The sliding technique requires complementary bimanual coordination (extracting with one hand and grasping lightly with the other, at the same time). We highlight the importance of actions with two hands—one gripping, one assisting—in termite fishing and discuss how probing techniques are correlated with performance. Additional research on digital function and on environmental, organismic, and task constraints will further reveal manual dexterity in termite fishing.     

Video-task acquisition in rhesus monkeys (Macaca mulatta) and chimpanzees (Pan troglodytes): A comparative analysis

This study describes video-task acquisition in two nonhuman primate species. The subjects were seven rhesus monkeys (Macaca mulatta) and seven chimpanzees (Pan troglodytes). All subjects were trained to manipulate a joystick which controlled a cursor displayed on a computer monitor. Two criterion levels were used: one based on conceptual knowledge of the task and one based on motor performance. Chimpanzees and rhesus monkeys attained criterion in a comparable number of trials using a conceptually based criterion. However, using a criterion based on motor performance, chimpanzees reached criterion significantly faster than rhesus monkeys. Analysis of error patterns and latency indicated that the rhesus monkeys had a larger asymmetry in response bias and were significantly slower in responding than the chimpanzees. The results are discussed in terms of the relation between object manipulation skills and video-task acquisition.     

Mortality rates among wild chimpanzees

In order to compare evolved human and chimpanzees' life histories we present a synthetic life table for free-living chimpanzees, derived from data collected in five study populations (Gombe, Ta?, Kibale, Mahale, Bossou). The combined data from all populations represent 3711 chimpanzee years at risk and 278 deaths. Males show higher mortality than females and data suggest some inter-site variation in mortality. Despite this variation, however, wild chimpanzees generally have a life expectancy at birth of less than 15 years and mean adult lifespan (after sexual maturity) is only about 15 years. This is considerably lower survival than that reported for chimpanzees in zoos or captive breeding colonies, or that measured among modern human hunter-gatherers. The low mortality rate of human foragers relative to chimpanzees in the early adult years may partially explain why humans have evolved to senesce later than chimpanzees, and have a longer juvenile period.     

Captive Bottlenose Dolphins (Tursiops truncatus) Spontaneously Using Water Flow to Manipulate Objects

Several terrestrial animals and delphinids manipulate objects in a tactile manner, using parts of their bodies, such as their mouths or hands. In this paper, we report that bottlenose dolphins (Tursiops truncatus) manipulate objects not by direct bodily contact, but by spontaneous water flow. Three of four dolphins at Suma Aqualife Park performed object manipulation with food. The typical sequence of object manipulation consisted of a three step procedure. First, the dolphins released the object from the sides of their mouths while assuming a head-down posture near the floor. They then manipulated the object around their mouths and caught it. Finally, they ceased to engage in their head-down posture and started to swim. When the dolphins moved the object, they used the water current in the pool or moved their head. These results showed that dolphins manipulate objects using movements that do not directly involve contact between a body part and the object. In the event the dolphins dropped the object on the floor, they lifted it by making water flow in one of three methods: opening and closing their mouths repeatedly, moving their heads lengthwise, or making circular head motions. This result suggests that bottlenose dolphins spontaneously change their environment to manipulate objects. The reason why aquatic animals like dolphins do object manipulation by changing their environment but terrestrial animals do not may be that the viscosity of the aquatic environment is much higher than it is in terrestrial environments. This is the first report thus far of any non-human mammal engaging in object manipulation using several methods to change their environment.     

Juvenile Sandhill Cranes exhibit wider ranging and more exploratory movements than adults during the breeding season

Sandhill Cranes Antigone canadensis exhibit delayed sexual maturity and breeding, and therefore juvenile Cranes searching for suitable territories to occupy have different ecological constraints on movements than adults, which must defend a territory and raise young. We used fine-scale GPS telemetry data to characterize and compare movements of adult and juvenile Cranes near the boundary between two populations in Minnesota, USA, from arrival on natal areas in the spring until staging prior to autumn migration. Juvenile and adult Cranes had marked differences in movement patterns throughout the breeding season. Juveniles were more likely than adults to display long-distance movements: they revisited areas less frequently throughout the breeding season and they had lower average residence times.