“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.     

Successful reproduction in wild-released orphan chimpanzees (<Emphasis Type="Italic">Pan troglodytes troglodytes</Emphasis>)

We report a case of successful reproduction in wild-released orphan chimpanzees. Using non-invasive genetic analysis, we determined the paternity of an infant born to a female chimpanzee released by HELP (Habitat Ecologique et Liberté des Primates) Congo into the Conkouati-Douli National Park, Republic of Congo. The sire was a released male, thus demonstrating successful reproduction in both male and female released chimpanzees. These results provide evidence that release into the wild may be a viable response to the plight of orphan chimpanzees in Africa, and we discuss further applications of non-invasive genetic tagging to release programs. Electronic Publication     

Use of leaves as cushions to sit on wet ground by wild chimpanzees

A new type of tool use, leaf cushion, by wild chimpanzees (Pan troglodytes verus) at Bossou, Guinea, was found. We report two cases: one is indirect evidence; the other is direct observation of a chimpanzee who used the tool. Both cases indicate that chimpanzees used a set of leaves as a cushion while sitting on wet ground. Chimpanzees at Bossou show various kinds of tool use, some of which are unique to the community. Most of these behavioral patterns are subsistence tool use for obtaining food, as at other study sites. The use of leaves as a cushion adds to the few instances of nonsubsistence, elementary technology seen used by wild chimpanzees. Am. J. Primatol. 44:215–220, 1998. © 1998 Wiley-Liss, Inc.     

Fluid dipping technology of chimpanzees in Comoé National Park,Ivory Coast

Over a 6 month period during the dry season, from the end of October 2014 to the beginning of May 2015, we studied tool use behavior of previously unstudied and non‐habituated savanna chimpanzees (Pan troglodytes verus) living in the Comoé National Park, Ivory Coast (CI). We analyzed all the stick tools and leaf‐sponges found that the chimpanzees used to forage for ants, termites, honey, and water. We found a particular behavior to be widespread across different chimpanzee communities in the park, namely, dipping for water from tree holes using sticks with especially long brush‐tip modifications, using camera traps, we recorded adults, juveniles, and infants of three communities displaying this behavior. We compared water dipping and honey dipping tools used by Comoé chimpanzees and found significant differences in the total length, diameter, and brush length of the different types of fluid‐dipping tools used. We found that water dipping tools had consistently longer and thicker brush‐tips than honey dipping tools. Although this behavior was observed only during the late dry season, the chimpanzees always had alternative water sources available, like pools and rivers, in which they drank without the use of a tool. It remains unclear whether the use of a tool increases efficient access to water. This is the first time that water dipping behavior with sticks has been found as a widespread and well‐established behavior across different age and sex classes and communities, suggesting the possibility of cultural transmission. It is crucial that we conserve this population of chimpanzees, not only because they may represent the second largest population in the country, but also because of their unique behavioral repertoire.     

乌干达野生动物研究中心外来黑猩猩融入笼养群体的过程

1967至1997的30年间,生活在非洲热带地区自然栖息地的野生黑猩猩数量由60万降至不足20万,至今这个数字仍在减少,因此引起了全球的关注并急需要开展迁地保护。笼养黑猩猩与野外种群一样营群居生活,为了有效进行野生黑猩猩的迁地保护,将野外捕获的野生黑猩猩个体成功引入已在动物园的黑猩猩群体中十分必要。2004年10月至2005年3月,乌干达野生动物研究中心首先开展了这一实验。同时为更好地了解外来黑猩猩融入笼养群体的过程,2006年9月至2007年1月间收集其活动数据。选取5只黑猩猩个体并记录它们的食性、行为、体重变化及身体健康状况。除直接观察和记录外,与兽医和研究中心管理人员进行合作,以获取较多笼养个体的信息。我们发现,野生个体比笼养个体多病,因此影响了它们的取食、社会行为及活动水平。尽管笼养黑猩猩有人照料,但仍具备野生个体的行为,所以,理解外来黑猩猩融入笼养群体的过程对于黑猩猩的迁地保护和就地保护是非常重要的。     

Behavioral Adaptation of Pan troglodytes troglodytes

As wild primate populations decline, numbers of orphaned primates, sanctuaries, and attempts to release primates back to the natural environment increase. Release projects frequently are poorly documented despite IUCN guidelines recommending post-release monitoring and systematic data collection as central to the process. Since 1996, Habitat Ecologique et Liberté des Primates (HELP) has been releasing wild-born orphaned chimpanzees into natural habitat in the Conkouati-Douli National Park, Republic of Congo. HELP developed a post-release monitoring system as an integral component. We present activity budgets and diet of released chimpanzees, and compared them to those of wild chimpanzee, as primary indicators of successful release. Feeding, moving, and resting dominated activity budgets, reflecting the overall patterns in wild populations. Diet was diverse and dominated by fruit, and the released chimpanzees showed specialization on a smaller number of species, as in many wild communities. The high survival rates of the chimpanzees and overall success of the release program are attributed to careful planning and post-release support facilitated via the monitoring process. Systematic post-release data collection monitoring has confirmed that wild-born chimpanzees can adjust behaviorally and nutritionally to the wild. Survival statistics of the reintroduced chimpanzees—confirmed 56%, possible 88%— reflect the behavioral adaptability.2nd revision March 11, 2005An erratum to this article can be found at     

Polymerase chain reaction detection of Clostridium perfringens in feces from captive and wild chimpanzees, Pan troglodytes

Background For veterinary management of non‐human primates in captivity, and conservation of wild‐living primates, management of their health risks is necessary. Incidences of pathogenic bacteria in the fecal specimens are considered as one of the useful indicators for non‐invasive health monitoring. Methods We carried out the detection of Clostridium perfringens in feces from captive and wild chimpanzees by the rapid polymerase chain reaction method. Results The bacterium was detected in most fecal specimens (80%) in captive chimpanzees. Contrarily, the detection rate in the wild chimpanzees was low, with 23% (n = 12) of 53 fecal samples from the Bossou group, Guinea, and 1.2% (1/81) in the Mahale group, Tanzania. Conclusions These results show that the intestinal microflora differs between Pan populations under various living conditions, being influenced by their diet and environment.     

Chimpanzee tool technology in the Goualougo Triangle, Republic of Congo

With the exception of humans, chimpanzees show the most diverse and complex tool-using repertoires of all extant species. Specific tool repertoires differ between wild chimpanzee populations, but no apparent genetic or environmental factors have emerged as definitive forces shaping variation between populations. However, identification of such patterns has likely been hindered by a lack of information from chimpanzee taxa residing in central Africa. We report our observations of the technological system of chimpanzees in the Goualougo Triangle, located in the Republic of Congo, which is the first study to compile a complete tool repertoire from the Lower Guinean subspecies of chimpanzee (Pan troglodytes troglodytes). Between 1999 and 2006, we documented the tool use of chimpanzees by direct observations, remote video monitoring, and collections of tool assemblages. We observed 22 different types of tool behavior, almost half of which were habitual (shown repeatedly by several individuals) or customary (shown by most members of at least one age-sex class). Several behaviors considered universals among chimpanzees were confirmed in this population, but we also report the first observations of known individuals using tools to perforate termite nests, puncture termite nests, pound for honey, and use leafy twigs for rain cover. Tool behavior in this chimpanzee population ranged from simple tasks to hierarchical sequences. We report three different tool sets and a high degree of tool-material selectivity for particular tasks, which are otherwise rare in wild chimpanzees. Chimpanzees in the Goualougo Triangle are shown to have one of the largest and most complex tool repertoires reported in wild chimpanzee populations. We highlight new insights from this chimpanzee population to our understanding of ape technological systems and evolutionary models of tool-using behavior.     

Tool-set for termite-fishing and honey extraction by wild chimpanzees in the Lossi Forest, Congo

The use of perforating sticks and flexible stalks in combination for termite fishing and a complex tool-set of three components used sequentially (stout chiel, bodkin, and dip-stick) to penetrate melipone and ground-dwelling bee hives byPan troglodytes troglodytes are documented or, inferred from circumstantial evidence. Functionally, termite extraction tools were similar to other locations in west and central Africa, but the plants and the number of raw material species used were different. Tools varied in the degree of modification (fraying ends). Chimpanzees in the Lossi forest seem to be able to use the tools not in a stereotyped fashion, but in a flexible, insightful way. The extraction of Melipone honey using large pieces of wood as pounding tools has rarely been recorded elsewhere. The most impressive technological solution to the honey-getting problem by wild chimpanzees was shown by this study. This is the only known, use of a tool-set of three components in sequence to extract honey by wild chimpanzees.     

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.     

A survey of entodiniomorphid ciliates in chimpanzees and bonobos

Intestinal entodiniomorphid ciliates are commonly diagnosed in the feces of wild apes of the genera Pan and Gorilla. Although some authors previously considered entodiniomorphid ciliates as possible pathogens, a symbiotic function within the intestinal ecosystem and their participation in fiber fermentation has been proposed. Previous studies have suggested that these ciliates gradually disappear under captive conditions. We studied entodiniomorphid ciliates in 23 captive groups of chimpanzees, three groups of captive bonobos and six populations of wild chimpanzees. Fecal samples were examined using Sheather's flotation and Merthiolate‐Iodine‐Formaldehyde Concentration (MIFC) methods. We quantified the number of ciliates per gram of feces. The MIFC method was more sensitive for ciliate detection than the flotation method. Ciliates of genus Troglodytella were detected in 13 groups of captive chimpanzees, two groups of bonobos and in all wild chimpanzee populations studied. The absence of entodiniomorphids in some captive groups might be because of the extensive administration of chemotherapeutics in the past or a side‐effect of the causative or prophylactic administration of antiparasitic or antibiotic drugs. The infection intensities of ciliates in captive chimpanzees were higher than in wild ones. We suppose that the over‐supply of starch, typical in captive primate diets, might induce an increase in the number of ciliates. In vitro studies on metabolism and biochemical activities of entodiniomorphids are needed to clarify their role in ape digestion. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Demographic influences on the hunting behavior of chimpanzees

We investigated hunting in an unusually large community of wild chimpanzees at Ngogo in the Kibale National Park, Uganda. Aspects of predation were recorded with respect to the prey, the predators, and hunting episodes. During 23 months of observation, the Ngogo chimpanzees caught 128 prey items from four primate and three ungulate species. Chimpanzees preyed selectively on immature red colobus primarily during group hunts, with adult males making the majority of kills. Party size and composition were significant predictors of the probability that chimpanzees would hunt and of their success during attempts. Chimpanzees were more likely to hunt red colobus if party size and the number of male hunters were large; party size and the number of male hunters were also significantly larger in successful compared with unsuccessful hunts. The Ngogo chimpanzees did not appear to hunt cooperatively, but reciprocal meat-sharing typically took place after kills. Hunts occurred throughout the year, though there was some seasonality as displayed by periodic hunting binges. The extremely high success rate and large number of kills made per successful hunt are the two most striking aspects of predation by the Ngogo chimpanzees. We compare currently available observations of chimpanzee hunting behavior across study sites and conclude that the large size of the Ngogo community contributes to their extraordinary hunting success. Demographic differences between groups are likely to contribute to other patterns of interpopulation variation in chimpanzee predation. Am J Phys Anthropol 109:439–454, 1999. © 1999 Wiley-Liss, Inc.     

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.     

The macronutrient composition of wild and cultivated plant foods of West African chimpanzees (Pan troglodytes verus) inhabiting an anthropogenic landscape

Agricultural expansion encroaches on tropical forests and primates in such landscapes frequently incorporate crops into their diet. Understanding the nutritional drivers behind crop-foraging can help inform conservation efforts to improve human-primate coexistence. This study builds on existing knowledge of primate diets in anthropogenic landscapes by estimating the macronutrient content of 24 wild and 11 cultivated foods (90.5% of food intake) consumed by chimpanzees (Pan troglodytes verus) at Bossou, Guinea, West Africa. We also compared the macronutrient composition of Bossou crops to published macronutrient measures of crops from Bulindi, Uganda, East Africa. The composition of wild fruits, leaves, and pith were consistent with previous reports for primate diets. Cultivated fruits were higher in carbohydrates and lower in insoluble fiber than wild fruits, while wild fruits were higher in protein. Macronutrient content of cultivated pith fell within the ranges of consumed wild pith. Oil palm food parts were relatively rich in carbohydrates, protein, lipids, and/or fermentable fiber, adding support for the nutritional importance of the oil palm for West African chimpanzees. We found no differences in the composition of cultivated fruits between Bossou and Bulindi, suggesting that macronutrient content alone does not explain differences in crop selection. Our results build on the current understanding of chimpanzee feeding ecology within forest-agricultural mosaics and provide additional support for the assumption that crops offer primates energetic benefits over wild foods.     

Lateralized tool use in wild New Caledonian crows

Many vertebrate species exhibit sensory and motor asymmetries. Laterality studies of tool use have focused on primates, where hemispheric asymmetries, manifested behaviourally in hand preferences, are thought to be associated with complex motor tasks. Here we report strong individual lateralization for tool use in birds. New Caledonian crows, Corvus moneduloides, hold stick tools with their bills while foraging, often with the nonworking end laterally positioned on one side of the head and the working end possibly positioned in the binocular field. We observed four wild crows to determine whether tools were consistently held on one side. All crows showed a significant preference (two right, two left). This preference was independent of any asymmetry in tool manufacture and held for artificial holes similarly accessible for tools held on either side. This is the first demonstration of lateralized tool use in a nonprimate. In addition, all 173 tools used unilaterally were held only on a crow's preferred side. Such pronounced individual laterality for tool use in natural conditions has previously been reported only in humans and chimpanzees.     

Chimpanzees as fauna: comparisons of sympatric large mammals across long-term study sites

Although much research has shown otherwise, chimpanzees are still often classed as rainforest-dwellers. Most long-term studies of wild chimpanzees (Pan troglodytes) are not situated in evergreen, closed-canopy equatorial forests, but instead are conducted in more open habitats. This study aims to elucidate the extent of chimpanzee ecological diversity by scrutinizing (recently) sympatric mammalian fauna at established study sites. We compiled presence or absence data on large mammal species at eight sites: Assirik, Bossou, Budongo, Gombe, Kibale, Lopé, Mahale, and Tai. The sites were rank ordered on the most basic ecological variable: annual total rainfall. Only three of the 65 mammalian genera compiled were sympatric with chimpanzees at all sites: Potamochoerus (bushpig), Syncerus (buffalo), and Panthera pardus (leopard). Some subfamilies (e.g. colobines) were present at most sites, but some families (e.g. hyenids) were absent at most sites. Some taxa (e.g. suids, cercopithecines) correlated better than others (e.g. canids) with basic ecological variables. The most extreme chimpanzee study site for which data are available is Assirik, Senegal. Nowhere else are chimpanzees sympatric with Erythrocebus, Alcelaphus, Hippotragus, and Ourebia. As chimpanzees are often behavioral models for extinct hominins, these living faunal assemblages have implications for paleo-ecological reconstructions of ancestral habitats.     

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.     

Nutritional composition of actual and potential insect prey for the Kasekela chimpanzees of Gombe National Park,Tanzania

Humans, all great ape species, and some lesser apes consume insects. Insects can provide comparable nutritional yields to meat on a gram‐for‐gram basis and may serve as an important source of energy, fat, protein, minerals, and vitamins for hominoids. Although potential insect prey are abundant in ape habitats, patterns of insectivory are not consistent across species or populations. Efforts to understand these patterns are complicated by a lack of nutritional data. We collected samples of insects consumed by the Kasekela chimpanzee community of Gombe National Park, Tanzania, as well as of some insects found within the community range and ignored by these chimpanzees but known to be preyed upon by Pan elsewhere. We determined the gross energy (GE), estimated metabolizable energy (ME), fat, protein, fiber, and ash content of these samples following standard methodologies. We use these data to test the hypothesis that Kasekela chimpanzees choose insect prey (at least in part) based on energy and/or macronutrient content. On a fresh‐weight, per‐gram basis, the insect prey consumed by Kasekela chimpanzees had significantly higher fat and lower ash content than other assayed insects, and on a fresh‐weight, per‐foraging‐unit (“per‐insect,” “per‐dip,” or “per‐nest”) basis were significantly higher in GE, fat, and protein. On a per‐gram basis, the assayed insects were generally comparable in energy and macronutrients to wild vertebrate meat. We conclude that Kasekela chimpanzees do favor insects that are high in energy, fat, and protein, and that the potential macronutrient yields from some forms of insectivory are not trivial. © 2012 Wiley Periodicals, Inc.     

Drinking tools of wild chimpanzees at Bossou

Use of drinking tools by wild chimpanzees (Pan troglodytes) and the context in which the tools were used were studied at Bossou, Republic of Guinea, West Africa. During the middle to late dry season and early wet season liquids are available occasionally in the holes of trees. Chimpanzees drank water or sap using a leaf (or fiber) as a sponge or spoon. When the chimpanzees were on the ground, they tended to use one of a few kinds of soft, hairless leaves, if they were available nearby. Females, particularly juveniles and adolescents, were thought to be the main users of the drinking tool. In a few episodes, a tool set was used to procure liquid. Once a chimpanzee used a stick to push a leaf sponge into a water hole and to pull it out from the hole. In addition, three chimpanzees used a pestle to squeeze sap from an oil-palm tree before using a fiber sponge. © 1995 Wiley-Liss, Inc.