Abstract: Chimpanzees (Pan troglodytes) are the most proficient and versatile users of tools in the wild. How such skills become integrated into the behavioural repertoire of wild chimpanzee communities is investigated here by drawing together evidence from three complementary approaches in a group of oil-palm nut- ( Elaeis guineensis) cracking chimpanzees at Bossou, Guinea. First, extensive surveys of communities adjacent to Bossou have shown that population-specific details of tool use, such as the selection of species of nuts as targets for cracking, cannot be explained purely on the basis of ecological differences. Second, a 16-year longitudinal record tracing the development of nut-cracking in individual chimpanzees has highlighted the importance of a critical period for learning (3-5 years of age), while the similar learning contexts experienced by siblings have been found to result in near-perfect (13 out of 14 dyads) inter-sibling correspondence in laterality. Third, novel data from field experiments involving the introduction of unfamiliar species of nuts to the Bossou group illuminates key aspects of both cultural innovation and transmission. We show that responses of individuals toward the novel items differ markedly with age, with juveniles being the most likely to explore. Furthermore, subjects are highly specific in their selection of conspecifics as models for observation, attending to the nut-cracking activities of individuals in the same age group or older, but not younger than themselves. Together with the phenomenon of inter-community migration, these results demonstrate a mechanism for the emergence of culture in wild chimpanzees.

Abstract: 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.

Abstract: Five chimpanzees with training in counting and numerical skills selected between 2 arrays of different amounts of candy or 2 Arabic numerals. A reversed reinforcement contingency was in effect, in which the selected array was removed and the subject received the nonselected candies (or the number of candies represented by the nonselected Arabic numeral). Animals were unable to maximize reward by selecting the smaller array when candies were used as array elements. When Arabic numerals were substituted for the candy arrays, all animals showed an immediate shift to a more optimal response strategy of selecting the smaller numeral, thereby receiving the larger reward. Results suggest that a response disposition to the high-incentive candy stimuli introduced a powerful interference effect on performance, which was effectively overridden by the use of symbolic representations.

Abstract: Chimpanzees (Pan troglodytes) frequently participate in social exchange involving multiple goods and services of variable value, yet they have not been tested in a formalized situation to see whether they can barter using multiple tokens and rewards. We set up a simple barter economy with two tokens and two associated rewards and tested chimpanzees on their ability to obtain rewards by returning the matching token in situations in which their access to tokens was unlimited or limited. Chimpanzees easily learned to associate value with the tokens, as expected, and did barter, but followed a simple strategy of favoring the higher-value token, regardless of the reward proffered, instead of a more complex but more effective strategy of returning the token that matched the reward. This response is similar to that shown by capuchin monkeys in our previous study. We speculate that this response, while not ideal, may be sufficient to allow for stability of the social exchange system in these primates, and that the importance of social barter to both species may have led to this convergence of strategies.

Abstract: Summary Reliable assessment of animal populations is a long-standing challenge in wildlife ecology. Technological advances have led to widespread adoption of camera traps (CTs) to survey wildlife distribution, abundance and behaviour. As for any wildlife survey method, camera trapping must contend with sources of sampling error such as imperfect detection. Early applications focused on density estimation of naturally marked species, but there is growing interest in broad-scale CT surveys of unmarked populations and communities. Nevertheless, inferences based on detection indices are controversial, and the suitability of alternatives such as occupancy estimation is debatable. We reviewed 266 CT studies published between 2008 and 2013. We recorded study objectives and methodologies, evaluating the consistency of CT protocols and sampling designs, the extent to which CT surveys considered sampling error, and the linkages between analytical assumptions and species ecology. Nearly two-thirds of studies surveyed more than one species, and a majority used response variables that ignored imperfect detection (e.g. presence?absence, relative abundance). Many studies used opportunistic sampling and did not explicitly report details of sampling design and camera deployment that could affect conclusions. Most studies estimating density used capture?recapture methods on marked species, with spatially explicit methods becoming more prominent. Few studies estimated density for unmarked species, focusing instead on occupancy modelling or measures of relative abundance. While occupancy studies estimated detectability, most did not explicitly define key components of the modelling framework (e.g. a site) or discuss potential violations of model assumptions (e.g. site closure). Studies using relative abundance relied on assumptions of equal detectability, and most did not explicitly define expected relationships between measured responses and underlying ecological processes (e.g. animal abundance and movement). Synthesis and applications. The rapid adoption of camera traps represents an exciting transition in wildlife survey methodology. We remain optimistic about the technology's promise, but call for more explicit consideration of underlying processes of animal abundance, movement and detection by cameras, including more thorough reporting of methodological details and assumptions. Such transparency will facilitate efforts to evaluate and improve the reliability of camera trap surveys, ultimately leading to stronger inferences and helping to meet modern needs for effective ecological inquiry and biodiversity monitoring.