Tactics to obtain a hidden food item in chimpanzee pairs (Pan troglodytes)

Five dyads of chimpanzees were tested in a competitive situation, as a pilot study to examine chimpanzees' understanding of conspecifics' knowledge. A human experimenter baited one of five containers in an outdoor enclosure. Chimpanzee A (witness) could see where the food was hidden, while chimpanzee B (witness-of-witness) could not see the baited place but could observe the chimpanzee A watching the food being hidden. Then the two were released into the enclosure. This procedure was repeated for a certain number of days along with a control condition in which neither could see the baited location. The witness-of-witness developed tactics to forestall the witness in two pairs. The witness misled the witness-of-witness by taking a route to an empty container in several cases. These episodes might represent examples of deception. Tactics and counter-tactics thus developed through the interaction between the witness and the witness-of-witness, illustrating the high social intelligence of chimpanzees. An examination of the changes in tactics suggests a possibility that the witness-of-witness understands the witness's knowledge of the location of hidden food. Accepted after revision: 22 May 2001 Electronic Publication     

Forest chimpanzees (Pan troglodytes verus) remember the location of numerous fruit trees

It is assumed that spatial memory contributes crucially to animal cognition since animals’ habitats entail a large number of dispersed and unpredictable food sources. Spatial memory has been investigated under controlled conditions, with different species showing and different conditions leading to varying performance levels. However, the number of food sources investigated is very low compared to what exists under natural conditions, where food resources are so abundant that it is difficult to precisely identify what is available. By using a detailed botanical map containing over 12,499 trees known to be used by the Taï chimpanzees, we created virtual maps of all productive fruit trees to simulate potential strategies used by wild chimpanzees to reach resources without spatial memory. First, we simulated different assumptions concerning the chimpanzees’ preference for a particular tree species, and, second, we varied the detection field to control for the possible use of smell to detect fruiting trees. For all these assumptions, we compared simulated distance travelled, frequencies of trees visited, and revisit rates with what we actually observed in wild chimpanzees. Our results show that chimpanzees visit rare tree species more frequently, travel shorter distances to reach them, and revisit the same trees more often than if they had no spatial memory. In addition, we demonstrate that chimpanzees travel longer distances to reach resources where they will eat for longer periods of time, and revisit resources more frequently where they ate for a long period of time during their first visit. Therefore, this study shows that forest chimpanzees possess a precise spatial memory which allows them to remember the location of numerous resources and use this information to select the most attractive resources.     

Chimpanzees (Pan troglodytes) recognize spatial and object correspondences between a scale model and its referent

In the present study, the contributions of spatial and object features to chimpanzees' comprehension of scale models were examined. Seven chimpanzees that previously demonstrated the ability to use a scale model as an information source for the location of a hidden item were tested under conditions manipulating the feature correspondence and spatial-relational correspondence between objects in the model and an outdoor enclosure. In Experiment 1, subjects solved the task under two conditions in which one object cue (color or shape) was unavailable, but positional cues remained. Additionally, performance was above chance under a third condition in which both types of object cues, but not position cues, were available. In Experiment 2, 2 subjects solved the task under a condition in which shape and color object cues were simultaneously unavailable. The results suggest that, much like young children, chimpanzees are sensitive to both object and spatial-relational correspondences between a model and its referent.     

Chimpanzees (Pan troglodytes) respond to nonvisible sets after one-by-one addition and removal of items

Two chimpanzees (Pan troglodytes) made numerousness judgments of nonvisible sets of items. In Experiment 1, 1-10 items were dropped 1 at a time into an opaque cup, and then an additional 1-10 items were dropped 1 at a time into another opaque cup. The chimpanzees' performance levels were high and were more dependent on factors indicative of an analogue-magnitude mechanism for representation of set size than on an object file mechanism. In Experiment 2, a 3rd visible set was made available after the sequential presentation of the first 2 sets. The chimpanzees again performed at high levels in selecting the largest of the 3 sets. In Experiment 3, 1 of the 2 initially presented sets was reduced in number by the sequential removal of 1, 2, or 3 items. Both chimpanzees performed above chance levels for the removal of 1, but not more than 1, item.     

Erratum to: Chimpanzees (Pan troglodytes) accurately compare poured liquid quantities

Although many studies have shown that nonhuman animals can choose the larger of two discrete quantities of items, less emphasis has been given to discrimination of continuous quantity. These studies are necessary to discern the similarities and differences in discrimination performance as a function of the type of quantities that are compared. Chimpanzees made judgments between continuous quantities (liquids) in a series of three experiments. In the first experiment, chimpanzees first chose between two clear containers holding differing amounts of juice. Next, they watched as two liquid quantities were dispensed from opaque syringes held above opaque containers. In the second experiment, one liquid amount was presented by pouring it into an opaque container from an opaque syringe, whereas the other quantity was visible the entire time in a clear container. In the third experiment, the heights at which the opaque syringes were held above opaque containers differed for each set, so that sometimes sets with smaller amounts of juice were dropped from a greater height, providing a possible visual illusion as to the total amount. Chimpanzees succeeded in all tasks and showed many similarities in their continuous quantity estimation to how they performed previously in similar tasks with discrete quantities (for example, performance was constrained by the ratio between sets). Chimpanzees could compare visible sets to nonvisible sets, and they were not distracted by perceptual illusions created through various presentation styles that were not relevant to the actual amount of juice dispensed. This performance demonstrated a similarity in the quantitative discrimination skills of chimpanzees for continuous quantities that matches that previously shown for discrete quantities.     

Spatial memory and monitoring of hidden items through spatial displacements by chimpanzees (Pan troglodytes)

This study examined chimpanzee (Pan troglodytes) short-term memory for food location in near space. In Experiments 1 and 2, either 1 or 2 items (chocolate pieces) were hidden in an array of 3 or 5 containers that either remained stationary or were rotated 180 degrees or 360 degrees. When the array remained stationary, the chimpanzees remembered both item locations. When arrays were rotated, however, chimpanzees found only 1 item. In Experiment 3, 2 items were hidden in an array of 7 cups. Both items were found at levels significantly better than chance. Ninety percent of errors were made after the 1st item was found, and errors reflected memory failure rather than a failure of inhibitory control.     

Chimpanzees (Pan troglodytes) transfer tokens repeatedly with a partner to accumulate rewards in a self-control task

There has been extensive research investigating self-control in humans and nonhuman animals, yet we know surprisingly little about how one’s social environment influences self-control. The present study examined the self-control of chimpanzees in a task that required active engagement with conspecifics. The task consisted of transferring a token back and forth with a partner animal in order to accumulate food rewards, one item per token transfer. Self-control was required because at any point in the trial, either chimpanzee could obtain their accumulated rewards, but doing so discontinued the food accumulation and ended the trial for both individuals. Chimpanzees readily engaged the task and accumulated the majority of available rewards before ending each trial, and they did so across a number of conditions that varied the identity of the partner, the presence/absence of the experimenter, and the means by which they could obtain rewards. A second experiment examined chimpanzees’ self-control when given the choice between immediately available food items and a potentially larger amount of rewards that could be obtained by engaging the token transfer task with a partner. Chimpanzees were flexible in their decision-making in this test, typically choosing the option representing the largest amount of food, even if it involved delayed accumulation of the rewards via the token transfer task. These results demonstrate that chimpanzees can exhibit self-control in situations involving social interactions, and they encourage further research into this important aspect of the self-control scenario.     

Size matters: impact of item size and quantity on array choice by chimpanzees (Pan troglodytes)

The authors previously reported that chimpanzees (Pan troglodytes) showed a striking bias to select the larger of 2 candy arrays, despite a reversed reward contingency in which the animals received the smaller, nonselected array as a reward, except when Arabic numerals were used as stimuli. A perceptual or incentive-based interference occurred that was overcome by symbolic stimuli. The authors of the present study examined the impact of element size in choice arrays, using 1 to 5 large and small candies. Five test-sophisticated chimpanzees selected an array from the 2 presented during each trial. Their responses were not optimal, as animals generally selected arrays with larger total mass; thus, they received the smaller remaining array as a reward. When choice stimuli differed in size and quantity, element size was more heavily weighted, although choices reflected total candy mass. These results replicate previous findings showing chimpanzees' difficulties with quantity judgments under reverse reward contingencies and also show that individual item size exerts a more powerful interference effect.     

Unprompted recall and reporting of hidden objects by a chimpanzee (Pan troglodytes) after extended delays

The ability to recall features of environments not present to the senses is important in human thinking, planning, and communication, but to date there are almost no data on recall capabilities in nonverbal animals. In this study, the author used symbol knowledge as a tool to study chimpanzee memory. An 11-year-old female chimpanzee (Pan troglodytes) that had already learned a large number of arbitrarily designated geometric forms (lexigrams) watched as an experimenter hid an object in the woods outside her outdoor enclosure. The type and location of the object varied across trials. After an imposed delay of up to 16 h, the chimpanzee could interact indoors with a person who did not know that an object had been hidden, let alone the type or location of the object. A keyboard in the indoor cage displayed 256 lexigrams. From Trial 1, the chimpanzee seemed to do whatever it took to catch the person's attention and then touched the lexigram corresponding to the type of object hidden, pointed outdoors, went outdoors (if followed), and continued to point manually toward the object and vocalize until the person found the object. The subject indicated nonfood objects as well as more than 20 food types.     

Chimpanzees (Pan troglodytes) and the question of cumulative culture: an experimental approach

There is increasing evidence for cultural variations in behaviour among non-human species, but human societies additionally display elaborate cumulative cultural evolution, with successive generations building on earlier achievements. Evidence for cumulative culture in non-human species remains minimal and controversial. Relevant experiments are also lacking. Here we present a first experiment designed to examine chimpanzees' capacity for cumulative social learning. Eleven young chimpanzees were presented with a foraging device, which afforded both a relatively simple and a more complex tool-use technique for extracting honey. The more complex 'probing' technique incorporated the core actions of the simpler 'dipping' one and was also much more productive. In a baseline, exploration condition only two subjects discovered the dipping technique and a solitary instance of probing occurred. Demonstrations of dipping by a familiar human were followed by acquisition of this technique by the five subjects aged three years or above, whilst younger subjects showed a significant increase only in the elements of the dipping technique. By contrast, subsequent demonstrations of the probing task were not followed by acquisition of this more productive technique. Subjects stuck to their habitual dipping method despite an escalating series of demonstrations eventually exceeding 200. Supplementary tests showed this technique is within the capability of chimpanzees of this age. We therefore tentatively conclude that young chimpanzees exhibit a tendency to become 'stuck' on a technique they initially learn, inhibiting cumulative social learning and possibly constraining the species' capacity for cumulative cultural evolution.     

Chimpanzees (Pan troglodytes) can wait,when they choose to: a study with the hybrid delay task

Self-control has been studied in nonhuman animals using a variety of tasks. The inter-temporal choice (ITC) task presents choices between smaller–sooner (SS) and larger–later (LL) options. Using food amounts as rewards, this presents two problems: (a) choices of the LL option could either reflect self-control or instead result from animals’ difficulty with pointing to smaller amounts of food; (b) there is no way to verify whether the subjects would not revert their choice for the LL option, if given the opportunity to do so during the ensuing delay. To address these problems, we have recently introduced a new protocol, the hybrid delay task, which combines an initial ITC with a subsequent accumulation phase in which selection of the SS option leads to its immediate delivery, but choice of the LL option then leads to one-by-one presentation of those items that continues only as long as the subject does not eat any of the accumulated items. The choice of the LL option therefore only reflects self-control when the number of items obtained from LL choices during the accumulation phase is higher than what could be received in the SS option. Previous research with capuchin monkeys demonstrated that their apparent self-control responses in the ITC task may have overestimated their general self-control abilities, given their poor performance in the hybrid delay task. Here, chimpanzees instead demonstrated that their choices for the LL option in the ITC phase of the hybrid delay task were confirmed by their ability to sustain long delays during accumulation of LL rewards.     

Inferences about the location of food in the great apes (Pan paniscus, Pan troglodytes, Gorilla gorilla, and Pongo pygmaeus)

Bonobos (Pan paniscus; n = 4), chimpanzees (Pan troglodytes; n = 12), gorillas (Gorilla gorilla; n = 8), and orangutans (Pongo pygmaeus; n = 6) were presented with 2 cups (1 baited) and given visual or auditory information about their contents. Visual information consisted of letting subjects look inside the cups. Auditory information consisted of shaking the cup so that the baited cup produced a rattling sound. Subjects correctly selected the baited cup both when they saw or heard the food. Nine individuals were above chance in both visual and auditory conditions. More important, subjects as a group selected the baited cup when only the empty cup was either shown or shaken, which means that subjects chose correctly without having seen or heard the food (i.e., inference by exclusion). Control tests showed that subjects were not more attracted to noisy cups, avoided shaken noiseless cups, or learned to use auditory information as a cue during the study. It is concluded that subjects understood that the food caused the noise, not simply that the noise was associated with the food.     

Maintenance of Self-Imposed Delay of Gratification by Four Chimpanzees (Pan troglodytes) and an Orangutan (Pongo pygmaeus)

Delay maintenance, which is the continuance over time of the choice to forgo an immediate, less preferred reward for a future, more preferred reward, was examined in 4 chimpanzees (Pan troglodytes) and 1 orangutan (Pongo pygmaeus). In the 1st experiment, the apes were presented with 20 chocolate pieces that were placed, one at a time, into a bowl that was within their reach. The apes could consume the available chocolate pieces at any time during a trial, but no additional pieces would be given. The total length of time taken to place the 20 items into the bowl ranged from 60 s to 180 s. All 5 apes delayed gratification on a majority of trials until all 20 chocolate pieces were presented. Unlike in most experiments with human children using this test situation, attention by the apes to the reward was not detrimental to delay maintenance. In a 2nd experiment with the chimpanzees only, 4 foods of differing incentive value were presented in different trials in the same manner as in Experiment 1. The chimpanzees were highly successful in obtaining all food pieces, and there was no difference in performance as a function of food type.     

Acquisition and Transmission of Tool Making and Use for Drinking Juice in a Group of Captive Chimpanzees (Pan troglodytes)

The present study examined the acquisition and transmission of tool making and use in a group of chimpanzees. We set up a piece of apparatus that provided orange juice in an outdoor compound for a group of nine chimpanzees. Although they could reach the juice with their hands, eight of the nine subjects used tools. Fifteen kinds of tools in total were used, such as straw, twigs, and some kinds of leaves. The chimpanzees showed high selectivity with regard to tool type. They preferred to use Thuja occidentalis as a tool although there were 28 species of tree and several kinds of grass available in the compound. Two females initiated the use of the Thuja tool. Since then, five other individuals have begun to use it selectively. Before making the tools by themselves, these five chimpanzees first watched others using the Thuja tool for drinking juice, and then used the Thuja tool which had been used and left by another chimpanzee.     

Executive Function in Young Children and Chimpanzees (Pan troglodytes): Evidence From a Nonverbal Dimensional Change Card Sort Task

In this article the authors compared chimpanzees’ executive function with that of children. They developed a nonverbal dimensional change card sorting task, which indexed the development of executive function. Three pairs of mother and offspring chimpanzees and 30 typically developed 5-year-old children were presented with 2 target stimuli and a test stimulus comprising 2 dimensions (size and shape) on a display; they were required to sort the test stimulus according to 1 dimension (e.g., shape). After 5 consecutive correct trials, the participants had to sort the test stimulus according to the other dimension (e.g., size). The results showed that the chimpanzees often failed to sort the test stimuli according to the first and reversed dimensions. On the other hand, the children were correctly able to use both dimensions. These results indicate that chimpanzees may have less developed executive skills than children.     

Facial perception of conspecifics: chimpanzees (Pan troglodytes) preferentially attend to proper orientation and open eyes

This paper reports on the use of an eye-tracking technique to examine how chimpanzees look at facial photographs of conspecifics. Six chimpanzees viewed a sequence of pictures presented on a monitor while their eye movements were measured by an eye tracker. The pictures presented conspecific faces with open or closed eyes in an upright or inverted orientation in a frame. The results demonstrated that chimpanzees looked at the eyes, nose, and mouth more frequently than would be expected on the basis of random scanning of faces. More specifically, they looked at the eyes longer than they looked at the nose and mouth when photographs of upright faces with open eyes were presented, suggesting that particular attention to the eyes represents a spontaneous face-scanning strategy shared among monkeys, apes, and humans. In contrast to the results obtained for upright faces with open eyes, the viewing times for the eyes, nose, and mouth of inverted faces with open eyes did not differ from one another. The viewing times for the eyes, nose, and mouth of faces with closed eyes did not differ when faces with closed eyes were presented in either an upright or inverted orientation. These results suggest the possibility that open eyes play an important role in the configural processing of faces and that chimpanzees perceive and process open and closed eyes differently.     

Recognition of Line-Drawing Representations by a Chimpanzee (Pan troglodytes)

The ability of a chimpanzee to recognize individuals portrayed in line drawings was evaluated. A 12-year-old female chimpanzee with extensive prior experience in the use of visual symbols matched the line drawings of chimpanzees, humans, and an orangutan with a specific letter of the alphabet. When a line drawing of a familiar individual was presented on the computer screen, the chimpanzee responded by punching a key with the letter of the alphabet that corresponded to the individual's name. Results indicate that the chimpanzee is able to categorize individuals from novel line-drawing representations.     

Chimpanzees remember the results of one-by-one addition of food items to sets over extended time periods

Four chimpanzees were highly accurate in selecting the larger of two concurrent accumulations of bananas in two opaque containers over a span of 20 min. One at a time, bananas were placed into the containers, which were outside the chimpanzees' cages. The chimpanzees never saw more than one banana at a time, and there were no cues indicating the locations of the bananas after they were placed into the containers. The performance of these animals matched that of human infants and young children in similar tests. The chimpanzees were successful even when the sets to be compared were sufficiently large (5 vs. 8, 5 vs. 10, and 6 vs. 10) to cast doubt on the possibility that the chimpanzees were using an object file mechanism. These chimpanzees are the first nonhuman animals to demonstrate extended memory for accumulated quantity.     

Chimpanzee (Pan troglodytes) spatial problem solving with the use of mirrors and televised equivalents of mirrors

Two adult male chimpanzees reached through a hole in the wall of their home cage and, by tracking the images of their hands and of an otherwise hidden target object in a mirror or closed-circuit television picture, moved their hands in whichever direction was necessary to make contact with the target object. They discriminated between live video images and tapes and performed effectively when the target objects were presented in novel locations and when the video picture was presented at random in different orientations. There was thus no consistent relation between the location of images on the monitor and the location of their real-world counterparts. Comparable performances in monkeys and nonprimates seem unlikely.