Planning abilities of wild chimpanzees (Pan troglodytes troglodytes) in tool-using contexts.

Planning is a type of problem solving in which a course of future action is devised via mental computation. Potential advantages of planning for tool use include reduced effort to gather tools, closer alignment to an efficient tool design, and increased foraging efficiency. Chimpanzees (Pan troglodytes troglodytes) in the Goualougo Triangle use a variety of different types of tools. We hypothesized that procurement strategy (brought to the termite nest, manufactured or acquired at the termite nest, or borrowed from others) reflects planning for current needs, with tool transport behavior varying by tool type and by age and sex class. It is also possible that chimpanzees anticipate the need for tools at future times, which would be evidenced by transporting multiple tool types for a sequential task. One year of video recordings at termite nests were systematically screened for tool procurement; data comprised 299 tool procurement events across 66 chimpanzees. In addition, we screened video recordings of leaf sponging and honey gathering, which resulted in another 38 procurement events. Fishing probes, which are typically used during a single visit, were typically transported to termite nests, while puncturing tools, which are durable and remain on site, were more often acquired at termite nests. Most tools transported in multiples were fishing probes, perhaps in anticipation that a single probe might not last through an entire foraging bout or might be transferred to another chimpanzee. We further documented that chimpanzees transported tool sets, comprising multiple different tool types used in sequence. Mature chimpanzees transported tools more often than did immatures. These observations suggest that chimpanzees plan tool use flexibly, reflecting the availability of raw materials and the likelihood that specific tool types will be needed for particular tasks. Developmental studies and further integration of behavioral, spatial, and archaeological data will help to illuminate the decision making and time depth of planning associated with tool technologies in living primates and hominin ancestors.

Extending the conservation impact of great ape research: Flagship species sites facilitate biodiversity assessments and land preservation.

To inform regional conservation planning, we assessed mammalian and avian biodiversity in the Djéké Triangle, which is an intact forest with long-term research and tourism focused on western lowland gorillas (Gorilla gorilla gorilla). This critical region serves as a conservation conduit between the Nouabalé-Ndoki National Park (NNNP) in the Republic of Congo and the Dzanga-Ndoki National Park in Central African Republic. Wildlife inventories were conducted to determine if biodiversity in the Djéké Triangle (initially part of a logging concession) was equivalent to the NNNP. Camera traps (CTs) were deployed to estimate species richness, relative abundance, naïve occupancy, and activity patterns of medium-to-large species in mixed species and monodominant Gilbertiodendron forests that comprise the majority of regional terra firma. Species inventories were collected from CTs positioned on a grid and at termite nests throughout the Djéké Triangle and compared to CTs placed in the Goualougo Triangle located within the NNNP. From 10,534 camera days at 65 locations, we identified 34 mammal and 16 bird species. Allaying concerns of wildlife depletion, metrics of species richness in the Djéké Triangle surpassed those of the Goualougo Triangle. Many species were observed to occur across habitats, while others showed habitat specificity, with termite mounds indicated as an important microhabitat feature. Our comparisons of animal activity budgets in different habitat types provide important reference information for other populations and contexts. In conclusion, this study provided empirical evidence of the high conservation value of this region that contributed to increasing the protected status of the Djéké Triangle.

The ontogeny of termite gathering among chimpanzees in the Goualougo Triangle, Republic of Congo.

OBJECTIVES: Acquiring tool-assisted foraging skills can potentially improve dietary quality and increase fitness for wild chimpanzees (Pan troglodytes). In contrast to chimpanzees in East and West Africa, chimpanzees in the Congo Basin use tool sets and brush-tipped fishing probes to gather termites. We investigated the ontogeny of these tool skills in chimpanzees of the Goualougo Triangle, Republic of Congo, and compared it to that for chimpanzees at Gombe, Tanzania. We assessed whether chimpanzees acquired simple tool behaviors and single tool use before more complex actions and sequential use of multiple tool types. MATERIALS AND METHODS: Using a longitudinal approach, we scored remote video footage to document the acquisition of termite-gathering critical elements for 25 immature chimpanzees at Goualougo. RESULTS: All chimpanzees termite fished by 2.9 years but did not manufacture brush-tipped probes until an average of 4.3 years. Acquisition of sequential tool use extended into juvenility and adolescence. While we did not detect significant sex differences, most critical elements except tool manufacture were acquired slightly earlier by females. DISCUSSION: These findings contrast with Gombe, where chimpanzees learn to both use and make fishing probes between ages 1.5-3.5 and acquire the complete task by age 5.5. Differences between sites could reflect tool material selectivity and design complexity, the challenge of sequential tool behaviors, and strength requirements of puncturing subterranean termite nests at Goualougo. These results illustrate how task complexity may influence the timing and sequence of skill acquisition, improving models of the ontogeny of tool behavior among early hominins who likely used complex, perishable technologies.

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.

Teaching varies with task complexity in wild chimpanzees.

Cumulative culture is a transformative force in human evolution, but the social underpinnings of this capacity are debated. Identifying social influences on how chimpanzees acquire tool tasks of differing complexity may help illuminate the evolutionary origins of technology in our own lineage. Humans routinely transfer tools to novices to scaffold their skill development. While tool transfers occur in wild chimpanzees and fulfill criteria for teaching, it is unknown whether this form of helping varies between populations and across tasks. Applying standardized methods, we compared tool transfers during termite gathering by chimpanzees in the Goualougo Triangle, Republic of Congo, and in Gombe, Tanzania. At Goualougo, chimpanzees use multiple, different tool types sequentially, choose specific raw materials, and perform modifications that improve tool efficiency, which could make it challenging for novices to manufacture suitable tools. Termite gathering at Gombe involves a single tool type, fishing probes, which can be manufactured from various materials. Multiple measures indicated population differences in tool-transfer behavior. The rate of transfers and probability of transfer upon request were significantly higher at Goualougo, while resistance to transfers was significantly higher at Gombe. Active transfers of tools in which possessors moved to facilitate possession change upon request occurred only at Goualougo, where they were the most common transfer type. At Gombe, tool requests were typically refused. We suggest that these population differences in tool-transfer behavior may relate to task complexity and that active helping plays an enhanced role in the cultural transmission of complex technology in wild apes.

Tool transfers are a form of teaching among chimpanzees.

Teaching is a form of high-fidelity social learning that promotes human cumulative culture. Although recently documented in several nonhuman animals, teaching is rare among primates. In this study, we show that wild chimpanzees (Pan troglodytes troglodytes) in the Goualougo Triangle teach tool skills by providing learners with termite fishing probes. Tool donors experienced significant reductions in tool use and feeding, while tool recipients significantly increased their tool use and feeding after tool transfers. These transfers meet functional criteria for teaching: they occur in a learner's presence, are costly to the teacher, and improve the learner's performance. Donors also showed sophisticated cognitive strategies that effectively buffered them against potential costs. Teaching is predicted when less costly learning mechanisms are insufficient. Given that these chimpanzees manufacture sophisticated, brush-tipped fishing probes from specific raw materials, teaching in this population may relate to the complexity of these termite-gathering tasks.