Flexible and Persistent Tool-using Strategies in Honey-gathering by Wild Chimpanzees

Several populations of wild chimpanzees use tools to raid bee nests, but preliminary observations of chimpanzees in the Congo Basin indicate that they may have developed sophisticated technical solutions to gather honey that differ from those of apes in other regions. Despite the lack of habituated groups within the range of the central subspecies, there have been several reports of different types of tools used by chimpanzees to open beehives and gather honey. Researchers have observed some of these behaviors (honey dipping) in populations of western and eastern chimpanzees, whereas others (hive pounding) may be limited to this region. Toward evaluating hypotheses of regional tool using patterns, we provide the first repeated direct observations and systematic documentation of tool use in honey-gathering by a population of Pan troglodytes troglodytes. Between 2002 and 2006, we observed 40 episodes of tool use in honey-gathering by chimpanzees in the Goualougo Triangle, Republic of Congo. Pounding was the most common and successful strategy to open beehives. Chimpanzees at this site used several tools in a single tool-using episode and could also attribute multiple functions to a single tool. They exhibited flexibility in responses toward progress in opening a hive and hierarchical structuring of tool sequences. Our results support suggestions of regional tool using traditions in honey-gathering, which could be shaped by variation in bee ecology across the chimpanzee range. Further, we suggest that these chimpanzees may have an enhanced propensity to use tool sets that could be related to other aspects of their tool repertoire. Clearly, there is still much to be learned about the behavioral diversity of chimpanzees residing within the Congo Basin.     

Chimpanzees prey on army ants with specialized tool set

Several populations of chimpanzees have been reported to prey upon Dorylus army ants. The most common tool‐using technique to gather these ants is with “dipping” probes, which vary in length with regard to aggressiveness and lifestyle of the prey species. We report the use of a tool set in army ant predation by chimpanzees in the Goualougo Triangle, Republic of Congo. We recovered 1,060 tools used in this context and collected 25 video recordings of chimpanzee tool‐using behavior at ant nests. Two different types of tools were distinguished based on their form and function. The chimpanzees use a woody sapling to perforate the ant nest, and then a herb stem as a dipping tool to harvest the ants. All of the species of ants preyed upon in Goualougo are present and consumed by chimpanzees at other sites, but there are no other reports of such a regular or widespread use of more than one type of tool to prey upon Dorylus ants. Furthermore, this tool set differs from other types of tool combinations used by chimpanzees at this site for preying upon termites or gathering honey. Therefore, we conclude that these chimpanzees have developed a specialized method for preying upon army ants, which involves the use of an additional tool for opening nests. Further research is needed to determine which specific ecological and social factors may have shaped the emergence and maintenance of this technology. Am. J. Primatol. 72:17–24, 2010. © 2009 Wiley‐Liss, Inc.     

Design complexity in termite-fishing tools of chimpanzees (Pan troglodytes)

Adopting the approach taken with New Caledonian crows (Corvus moneduloides), we present evidence of design complexity in one of the termite-fishing tools of chimpanzees (Pan troglodytes) in the Goualougo Triangle, Republic of Congo. Prior to termite fishing, chimpanzees applied a set of deliberate, distinguishable actions to modify herb stems to fashion a brush-tipped probe, which is different from the form of fishing tools used by chimpanzees in East and West Africa. This means that ‘brush-tipped fishing probes’, unlike ‘brush sticks’, are not a by-product of use but a deliberate design feature absent in other chimpanzee populations. The specialized modifications to prepare the tool for termite fishing, measures taken to repair non-functional brushes and appropriate orientation of the modified end suggest that these wild chimpanzees are attentive to tool modifications. We also conducted experimental trials that showed that a brush-tipped probe is more effective in gathering insects than an unmodified fishing probe. Based on these findings, we suggest that chimpanzees in the Congo Basin have developed an improved fishing probe design.     

Chaînes opératoires and resource-exploitation strategies in chimpanzee (Pan troglodytes) nut cracking

We apply archaeological methods to extend our knowledge of chimpanzee material culture. The cha?ne opératoire conceptual framework, as introduced by ethnography, established technology as a phased process. Prehistoric archaeology adopted this concept to elucidate technological variability in tool-making procedures, based on knowledge of tool functions or subsistence patterns. We focused on the detection of operational sequences by wild chimpanzees (Pan troglodytes verus) when nut cracking with lithic implements at the sites of Bossou and Diecké, Guinea, West Africa. Thus, while it has recently been claimed that chimpanzees leave behind recognizable assemblages of stone hammers that can be morphologically distinguished from Oldowan hammers, this is the first study to focus specifically on the existence of operational sequences during the utilization of stone tools by wild chimpanzees. By combining primatological and archaeological methods and examining ecological areas inhabited by different chimpanzee groups, we sought technological variability and identified variables influencing regional diversity in tool typology and technology. We compared three case studies: (1) Bossou-direct recording of experimental nut-cracking sessions; (2) Bossou- direct and indirect monitoring of nut-cracking sites in the wild; (3) Diecké-indirect monitoring of nut-cracking sites in the wild. Results suggest that chimpanzees perform sequences of repeated tool transport and nut cracking. Data show discrimination of tool functions based on tool features. We identified the most technologically complex tool for nut cracking, which was composed of four stones. We found regional diversity in chimpanzee stone assemblages. Raw-material type and tool mobility constrain technological development in human and nonhuman primates. Spatial analysis of tool distribution indicates a pattern of resource-exploitation strategy, revealing affinities with Oldowan.     

Ape Abundance and Habitat Use in the Goualougo Triangle, Republic of Congo

Chimpanzee (Pan troglodytes troglodytes) and western gorilla (Gorilla gorilla gorilla) populations in central Africa are rapidly declining as a result of disease epidemics, commercial bushmeat hunting, and habitat destruction. Our main objective was to estimate the absolute abundance and habitat utilization of chimpanzees and gorillas in the intact forests of the Goualougo Triangle in the Republic of Congo, and in an adjacent area in which selective logging will take place in the near future. The estimates provide a unique baseline for apes inhabiting an undisturbed environment. A second objective was to compare estimates of abundance and patterns of habitat utilization generated by different techniques: 1) distance sampling of individual ape nests and nest sites along line transects, 2) direct observations of apes during reconnaissance surveys, and 3) observations of ape traces during reconnaissance surveys. We completed a total of 222 km of line transect surveys in 4 sampling areas, resulting in overall density estimates of 1.53 chimpanzees/km² and 2.34 gorillas/km² from nest sites. We generated a density estimate of 2.23 chimpanzees/km² from direct observations during reconnaissance surveys of a semihabituated community in 1 of the 4 sampling areas. Habitat use profiles that nest surveys depicted on transects differed from those of direct observations and traces we encountered on reconnaissance surveys. We found the highest overall abundance of chimpanzee nests in monodominant Gilbertiodendron forest, whereas our direct observations showed that chimpanzees preferred mixed species forest. Transects that traversed the core area of the community range had the highest encounter rates of chimpanzee nests and nest sites. Gorilla nests on transects showed a preference only for mixed species forest with an open canopy, but direct observations and traces on reconnaissance surveys clearly indicated that gorillas use several habitat types. We conclude by evaluating the precision of these nest surveys and our ability to detect future trends in ape densities in the Goualougo Triangle.

Pestle-pounding behavior of wild chimpanzees at Bossou,Guinea: A newly observed tool-using behavior

A new type of tool-using behavior was observed in a group of wild chimpanzees (Pan troglodytes verus) at Bossou, Guinea. The chimpanzees used the leaf-petiole of oil-palm trees (Elaeis guineensis) as a pounding tool to deepen a hole in the oil-palm crown which appeared after the chimpanzees had pulled out the central young shoots. Finally, the chimpanzees extracted and ate the apical meristem or apical bud of the oil-palm tree which is edible but inaccessible without such tool use. The motor pattern which the chimpanzees employed is similar to that used for termite-nest digging but it is more exaggerated and requires great force. The behavior is reminiscent of pestlepounding. The chimpanzees exploit substantial amounts of food with this tool-using skill, compensating for insufficient fruit foods in the primary forest. This tool-using behavior was first observed in 1990 and, to date, almost half of the group members have been confirmed to use the pestle tool. It appears that this tool-using behavior was invented recently and has since spread widely throughout the group as a habitual one.     

Complex tool sets for honey extraction among chimpanzees in Loango National Park, Gabon

Homo faber was once proposed as a label for humans specifically to highlight their unique propensity for tool use. However, new observations on complex tool use by the chimpanzees of Loango National Park, Gabon, expand our knowledge about tool-using abilities in Pan troglodytes. Chimpanzees in Loango, when using tools to extract honey from three types of bee nests, were observed to regularly use three- to five-element tool sets. In other words, different types of tools were used sequentially to access a single food source. Such tool sets included multi-function tools that present typical wear for two distinct uses. In addition, chimpanzees exploited underground bee nests and used ground-perforating tools to locate nest chambers that were not visible from the ground surface. These new observations concur with others from Central African chimpanzees to highlight the importance of honey extraction in arguments favoring the emergence of complex tool use in hominoids, including different tool types, expanded tool sets, multifunction tools, and the exploitation of underground resources. This last technique requires sophisticated cognitive abilities concerning unseen objects. A sequential analysis reveals a higher level of complexity in honey extraction than previously proposed for nut cracking or hunting tools, and compares with some technologies attributed to early hominins from the Early and Middle Stone Age. A better understanding of similarities in human and chimpanzee tool use will allow for a greater understanding of tool-using skills that are uniquely human.     

Gene flow in wild chimpanzee populations: what genetic data tell us about chimpanzee movement over space and time

The isolation of phylogenetically distinct primate immunodeficiency viruses from at least seven wild-born, captive chimpanzees indicates that viruses closely related to HIV-1 may be endemic in some wild chimpanzee populations. The search for the chimpanzee population or populations harbouring these viruses is therefore on. This paper attempts to answer the question of whether or not such populations of chimpanzees are likely to exist at all, and, if so, where they are likely to be found. We summarize what is known about gene flow in wild populations of chimpanzees, both between major phylogeographical subdivisions of the species, and within these subdivisions. Our analysis indicates that hitherto undocumented reproductively isolated chimpanzee populations may in fact exist. This conclusion is based on the observation that, despite limited geographical sampling and limited numbers of genetic loci, conventional notions of the nature and extent of chimpanzee gene flow have recently been substantially revised. Molecular genetic studies using mitochondrial DNA sequences and hypervariable nuclear microsatellite markers have indicated the existence of heretofore undocumented barriers to chimpanzee gene flow. These studies have identified at least one population of chimpanzees genetically distinct enough to be classified into a new subspecies (Pan troglodytes vellerosus). At the same time, they have called into question the long-accepted genetic distinction between eastern chimpanzees (Pan troglodytes schweinfurthii) and western equatorial chimpanzees (Pan troglodytes troglodytes). The same studies have further indicated that gene flow between local populations is more extensive than was previously thought, and follows patterns sometimes inconsistent with those documented through direct behavioural observation. Given the apparently incomplete nature of the current understanding of chimpanzee gene flow in equatorial Africa, it seems reasonable to speculate that a chimpanzee population or populations may exist which both harbour the putative HIV-1 ancestor, and which have remained reproductively isolated from other chimpanzee populations over the time-scale relevant to the evolution of the SIVcpz-HIV-1 complex of viruses. Continued extensive sampling of wild chimpanzee populations, both for their genes and their viruses, should be performed quickly considering the high probability of extinction that many wild chimpanzee populations face today. The history of human-chimpanzee contacts is discussed.     

Naïve Encounters with Chimpanzees in the Goualougo Triangle, Republic of Congo

We describe the behavior of an unhabituated population of chimpanzees in the Goualougo Triangle, Republic of Congo. We encountered chimpanzee parties on 218 occasions during two field seasons (February 1999–December 1999, June 2000–June 2001). Overall contact rate was 0.63 contacts per day in the field (n = 347). During the first 5 min of observation, we recorded individual responses as curious, ignore, hide, or depart. In contrast to other unhabituated chimpanzees, curiosity was the most common response (84%) of individuals in the Goualougo Triangle. However, the responses were deeply integrated in the group's reaction to our arrival and behavior throughout an encounter. Based on the behavior of the majority of individuals in a group, we categorized entire contact events as naïve, ignore, nervous, or depart. Naïve contacts accounted for 69% of all encounters. Other contacts types occurred much less frequently: nervous (12%), depart (11%), ignore (8%). Naïve contacts were characterized by chimpanzees that continued to exhibit curiosity throughout the encounter, the arrival of other individuals at the contact location, and relatively prolonged contact with observers (average duration: 136 min). It is likely that the high frequency of curious responses and naïve contacts are due to the remote location of the Goualougo Triangle and the chimpanzees's lack of experience with humans. Documentation of this naïve phenomenon has been successfully used to lobby for the protection of the chimpanzees and their habitat.     

“Stepping-sticks” and “seat-sticks”: New types of tools used by wild chimpanzees (Pan troglodytes) in Sierra Leone

In Tenkere, Sierra Leone, a community of wild chimpanzees (Pan troglodytes verus) spent long hours eating the fruits and flowers of the Kapok (Ceiba pentandra) tree. The branches of this species are covered in sharp thorns which make movement in their high canopies problematic for the chimpanzees. In an apparent attempt to increase their mobility and to ease the discomfort of lengthy bouts of eating in these trees, some of the Tenkere chimpanzees have been observed using stick tools as foot (“stepping-sticks”) and body (“seat-sticks”) protection against the painful thorns. This form of tool-using is culturally unique to the Tenkere chimpanzees, as at other sites where these apes have been observed eating parts of kapok trees, there are no published records of this tool technology. In three of the stepping-stick tool use incidents, the chimpanzee used the tool(s), held between their greater and lesser toes, in locomotion. This form of tool use is the first recorded case of habitually used tools that can be justifiably categorized as being “worn” by any known wild population of Pan troglodytes. Am J Primatol 41:45–52, 1997. © 1997 Wiley-Liss, Inc.     

Ecological and social correlates of chimpanzee tool use

The emergence of technology has been suggested to coincide with scarcity of staple resources that led to innovations in the form of tool-assisted strategies to diversify or augment typical diets. We examined seasonal patterns of several types of tool use exhibited by a chimpanzee (Pan troglodytes) population residing in central Africa, to determine whether their technical skills provided access to fallback resources when preferred food items were scarce. Chimpanzees in the Goualougo Triangle exhibit a diverse repertoire of tool behaviours, many of which are exhibited throughout the year. Further, they have developed specific tool sets to overcome the issues of accessibility to particular food items. Our conclusion is that these chimpanzees use a sophisticated tool technology to cope with seasonal changes in relative food abundance and gain access to high-quality foods. Subgroup sizes were smaller in tool using contexts than other foraging contexts, suggesting that the size of the social group may not be as important in promoting complex tool traditions as the frequency and type of social interactions. Further, reports from other populations and species showed that tool use may occur more often in response to ecological opportunities and relative profitability of foraging techniques than scarcity of resources.     

Tool-use to obtain honey by chimpanzees at Bulindi: new record from Uganda

Honey-gathering from bee nests has been recorded at chimpanzee (Pan troglodytes) study sites across tropical Africa. Different populations employ different strategies, ranging from simple ‘smash-and grab’ raids to use of sophisticated tool-sets, i.e., two or more types of tool used sequentially in a single task. In this paper I present evidence of tool-use, and the probable use of a tool-set, for honey-gathering by unhabituated chimpanzees at Bulindi, a forest–farm mosaic south of the Budongo Forest in Uganda. Between June and December 2007, 44 stick tools were found in association with 16 holes dug in the ground, corresponding to the period when stingless bees (Meliponula sp.) appeared in chimpanzee dung. In 11 cases the confirmed target was a Meliponula ground nest. Two potential tool types were distinguished: digging sticks encrusted with soil, and more slender and/or flexible sticks largely devoid of soil that may have functioned to probe the bees’ narrow entry tubes. Reports of chimpanzees using tools to dig for honey have been largely confined to Central Africa. Honey-digging has not previously been reported for Ugandan chimpanzees. Similarly, use of a tool-set to obtain honey has thus far been described for wild chimpanzee populations only in Central Africa. Evidence strongly suggests that Bulindi chimpanzees also use sticks in predation on carpenter bee (Xylocopa sp.) nests, perhaps as probes to locate honey or to disable adult bees. These preliminary findings from Bulindi add to our understanding of chimpanzee technological and cultural variation. However, unprotected forests at Bulindi and elsewhere in the region are currently severely threatened by commercial logging and clearance for farming. Populations with potentially unique behavioral and technological repertoires are being lost.     

Army ant prey availability and consumption by chimpanzees (Pan troglodytes vellerosus) at Gashaka (Nigeria)

Army ant predation by chimpanzees has been studied as an intriguing example of tool use and a possible case of cultural variation. However, the importance of army ant prey in chimpanzee diet and feeding ecology is still only poorly understood. We studied the availability and consumption of army ants in a population of the chimpanzee subspecies Pan troglodytes vellerosus in Nigeria. Army ants were collected from nests and trails (workers) and near artificial light sources (males). Three potential prey species were found: Dorylus rufescens, Dorylus gerstaeckeri and Dorylus kohli . Dorylus rufescens was by far more abundant than the other two species. Only remains from D. rufescens were present in chimpanzee faeces. This is the first report of consumption of this ant species by chimpanzees. However, because of the low availability of the other two species, it is unclear whether this pattern reflects a preference for D. rufescens . Although D. rufescens ' availability varied with weather conditions, the occurrence as well as the absolute and relative numbers of Dorylus fragments in faeces did not. This finding, together with the considerable difficulties encountered by human observers in their efforts to locate nests by following trails, suggests that the chimpanzees in this population do not harvest army ants from trails and do not use trails to locate nests. The overall occurrence of army ant fragments in 42.3% of all faecal samples is the highest ever recorded in any chimpanzee population. This indicates that in this chimpanzee population, army ant prey is not a fallback during periods of sparse availability of plant food, but quantitatively important throughout the year. Future studies will be needed to clarify which cues and strategies chimpanzees use to locate army ant nests and to assess the role of myrmecophagy with respect to macro- or micronutrient demands.     

Density and distribution of western chimpanzees around a bauxite deposit in the Boé Sector,Guinea‐Bissau

The Boé sector in southeast Guinea‐Bissau harbors a population of western chimpanzees (Pan troglodytes verus) that inhabits a mosaic of forest and savanna. The Boé sector contains a substantial bauxite deposit in a region called Ronde Hill, and there are plans for the construction of a mine, which may endanger the chimpanzee population. In 1‐week survey in May 2013, we used the standing crop nest counts method to obtain the number of chimpanzee nests and from that estimate the density and abundance of chimpanzees. We carried out five 1‐km line transects that covered the bauxite deposit and surrounding valleys. We used density surface modeling to analyze habitat preferences, then predicted chimpanzee nest density and distribution based on environmental variables. We found the projected location of the mine partially coincides with an area of high predicted abundances of chimpanzee nests and is surrounded by highly suitable areas for chimpanzees (northeast and southwest). We conclude the mine could have significant direct and indirect effects on this population of chimpanzees whose impacts must be carefully considered and properly mitigated if the mine is built.     

Use of a tool-set by Pan troglodytes troglodytes to obtain termites (Macrotermes) in the periphery of the Dja Biosphere Reserve, southeast Cameroon

At the northern periphery of the Dja Biosphere Reserve (southeastern Cameroon) we recorded a new use of a tool-set by Pan troglodytes troglodytes to prey on Macrotermes muelleri, M. renouxi, M. lilljeborgi, and M. nobilis. We recovered 79 puncturing sticks and 47 fishing probes at 17 termite nests between 2002 and 2005. The mean length of the puncturing sticks (n = 77) and fishing probes (n = 45) was 52 cm and 56 cm, respectively, and the mean diameter was 9 mm and 4.5 mm, respectively. Sixty-eight percent of 138 chimpanzee fecal samples contained major soldiers of four Macrotermes species. The chimpanzees in southeastern Cameroon appeared to be selective in their choice of plant material to make their tools. The tools found at our study site resemble those from other sites in this region. However, in southeastern Cameroon only one tool-set type was found, whereas two tool-set types have been reported in Congo. Our study suggests that, along with the different vegetation types and the availability of plant material around termite nests, the nest and gallery structure and foraging behavior of the different Macrotermes spp. at all Central African sites must be investigated before we can attribute differences in tool-use behavior to culture.     

Individual acquisition of “stick pounding” behavior by naïve chimpanzees

Many studies investigating culture in nonhuman animals tend to focus on the inferred need of social learning mechanisms that transmit the form of a behavior to explain the population differences observed in wild animal behavioral repertoires. This research focus often results in studies overlooking the possibility of individuals being able to develop behavioral forms without requiring social learning. The disregard of individual learning abilities is most clearly observed in the nonhuman great ape literature, where there is a persistent claim that chimpanzee behaviors, in particular, require various forms of social learning mechanisms. These special social learning abilities have been argued to explain the acquisition of the relatively large behavioral repertoires observed across chimpanzee populations. However, current evidence suggests that although low‐fidelity social learning plays a role in harmonizing and stabilizing the frequency of behaviors within chimpanzee populations, some (if not all) of the forms that chimpanzee behaviors take may develop independently of social learning. If so, they would be latent solutions—behavioral forms that can (re‐)emerge even in the absence of observational opportunities, via individual (re)innovations. Through a combination of individual and low‐fidelity social learning, the population‐wide patterns of behaviors observed in great ape species are then established and stably maintained. This is the Zone of Latent Solutions (ZLS) hypothesis. The current study experimentally tested the ZLS hypothesis for pestle pounding, a wild chimpanzee behavior. We tested the reinnovation of this behavior in semi‐wild chimpanzees at Chimfunshi Wildlife Orphanage in Zambia, Africa, (N = 90, tested in four social groups). Crucially, all subjects were naïve to stick pounding before testing. Three out of the four tested groups reinnovated stick pounding—clearly demonstrating that this behavioral form does not require social learning. These findings provide support for the ZLS hypothesis alongside further evidence for the individual learning abilities of chimpanzees.     

Female Coalitions Against Male Aggression in Wild Chimpanzees of the Budongo Forest

In the wild, female chimpanzees (Pan troglodytes) are subject to male aggression that at times can be prolonged or particularly violent. There are no reports of cooperative retaliation to such aggression, a strategy observed in the congeneric Pan paniscus, from the wild despite >4 decades of detailed behavioral study across a number of populations and its occurrence among captive female chimpanzees. If the reports from captivity represent an inherent capacity, then the absence of similar reports from wild populations suggests that females may be able to form coalitions only under appropriate ecological and demographic conditions. During a study of male and female aggressive interactions among chimpanzees of the Budongo Forest, Uganda, wild adult female chimpanzees sometimes formed coalitions with one another to retaliate against male aggression. This may be possible because these females tend to be more gregarious than in other populations of East African chimpanzees, as other studies of the same population have suggested; the extent and variation of female chimpanzee social strategies may, therefore, need reconsideration. Further, my observations strengthen the argument that at least some of the differences between chimpanzees and bonobos may be more of degree than of kind.     

Skeletal pathology in Pan troglodytes schweinfurthii in Kibale National Park, Uganda

The ecological pressures shaping chimpanzee anatomy and behavior are the subject of much discussion in primatology and paleoanthropology, yet empirical data on fundamental parameters including body size, morbidity, and mortality are rare for wild chimpanzees. Here, we present skeletal pathology and body size data for 20 (19 crania, 12 postcrania) chimpanzees (Pan troglodytes schweinfurthii) from Kibale National Park, Uganda. We compare these data with other East African populations, especially Gombe National Park. Estimated body size for Kibale chimpanzees was similar to other East African populations and significantly larger than Gombe chimpanzees. The high rates of trauma and other skeletal pathology evident in the Kibale chimpanzee skeletons were similar to those in the Gombe skeletal sample. Much of the major skeletal trauma in the Kibale skeletons was attributable to falls, although other pathologies were noted as well, including apparent injuries from snares, degenerative arthritis, and minor congenital abnormalities.