Intestinal parasites of sympatric Pan troglodytes and Papio Spp. at two sites: Gombe (Tanzania) and Mt. Assirik (Senegal)

Experimental and clinical parasitology need natural baselines or “controls”. We present normative data intestinal parasite loads in two genera of African primates. Wild Pan troglodytes and Papio spp. were studied at two sites: Gombe in Tanzania (P. anubis) and Mt. Assirik in Senegal (P. papio). Presence or absence of parasites, especially nematodes, was recorded from fecal specimens. Gombe's primates were more often infected than were Mt. Assirik's. At Gombe, but not at Mt. Assirik, chimpanzees seemed to have a higher incidence of infection than baboons. Comparison of three baboon troops yielded apparent differences in prevalence of infection. No differences in infection were found between the wet and dry seasons in Mt. Assirik's chimpanzees.     

Diet of wild chimpanzees (Pan troglodytes verus) at Mt. Assirik,Senegal: I. Composition

The composition of the diet of a savanna-living population of wild chimpanzees (Pan troglodytes verus) at Mt. Assirik in Senegal is presented. The study site had a prolonged dry season, high temperatures, and vegetation dominated by grasses. Data came from direct observation, fecal specimens, and feeding traces; thus, strict criteria for acceptance of the indirect data were specified. Composition of diet was given in terms of species and family of prey, parts eaten, life-form, type of habitat, and criteria for inclusion. Forty-three species of plants with 60 parts were eaten; mostly fruits, from trees, in woodland. Nine species of animal prey were eaten, mostly social insects. An additional 41 species of plants with 53 parts were classed as likely to be eaten by chimpanzees, mostly on the grounds of their being eaten by sympatric anthropoids. Overall, the diet of the apes at Mt. Assirik resembles that of this species elsewhere in Africa, but the size of the dietary repertoire seems small and the proportion of low-quality foodstuffs high. The latter are mostly time-consuming to collect or tedious to obtain or process, and include underground storage organs.     

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 in Bwindi-Impenetrable National Park, Uganda, use different tools to obtain different types of honey

Evidence of tool use for foraging for honey by chimpanzees in Bwindi-Impenetrable National Park, Uganda, is reported. These are the first records of tool use by chimpanzees in this region of the Albertine Rift. Tools of two types were found at sites of bee activity. Chimpanzees apparently use small stick tools to forage for the honey of a stingless bee [Meliponula bocandei (Trigonidae)] that nests in tree cavities and also in subterranean holes. They use significantly larger, thicker tools to assist in foraging for honey of African honeybees (Apis mellifera).     

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.     

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.     

Use of tool sets by chimpanzees for multiple purposes in Moukalaba-Doudou National Park,Gabon

We report our recent findings on the use of tool sets by chimpanzees in Moukalaba-Doudou National Park, Gabon. Direct observations and evidences left by chimpanzees showed that chimpanzees used sticks as pounders, enlargers, and collectors to extract honey from beehives of stingless bees (Meliponula sp.), which may correspond to those previously found in the same site for fishing termites and to those found in Loango National Park, Gabon. However, we observed chimpanzees using a similar set of tools for hunting a medium-sized mammal (possibly mongoose) that hid inside a log. This is the first report of hunting with tools by a chimpanzee population in Central Africa. Chimpanzees may recognize the multiple functions and applicability of tools (extracting honey and driving prey), although it is still a preliminary speculation. Our findings may provide us a new insight on the chimpanzee’s flexibility of tool use and cognitive abilities of complex food gathering.     

Subsistence Technology of Nigerian Chimpanzees

A trademark of Homo sapiens is the enormous variation in behavioral patterns across populations. Insight into the development of human cultures can be aided by studies on communities of Pan across Africa that display unique combinations of social behavior and elementary technology. Only cross-population comparisons can reveal whether the diversity reflects differential genetics, environmental constraints, or is a cultural variant. However, the recently recognized and most endangered subspecies, Pan troglodytes vellerosus, remains completely unstudied in this respect. We report first evidence from a new long-term study of Nigerian chimpanzees at Gashaka. Their dietary composition is highly varied and they have to cope with high concentrations of antifeedant defenses of plants against consumption. Gashaka chimpanzees use a varied tool kit for extractive foraging. For example, they harvest insects throughout the year, via digging sticks and probes, to obtain honey from stingless-bee and honeybee nests, dipping wands to prey on army ants, and fishing rods to eat arboreal ants. Tools appeared to be custom-made with a considerable degree of standardization in length, diameter, and preferential use of distal ends. Moreover, compared to the rainy season, tools were longer during the dry season when insects retreat further into their nests. Many of the expressions of subsistence technology seem to be environmentally constrained. Most notably, the absence of termite-eating could reflect a low abundance of mounds. Other traits may represent cultural variation. For example, the chimpanzees did not hammer open 2 types of hard-shelled nuts with tools, unlike what occurs elsewhere in West Africa. The prevalence of elementary technology may indicate that the material culture of Gashaka chimpanzees is most related to core cultural tendencies of Central African populations.     

Meat eating and ant dipping by wild chimpanzees in Sierra Leone

This report presents the first records of meat-eating and ant dipping by wild chimpanzees,Pan troglodytes, from Sierra Leone. The study was conducted in the proposed Outamba-Kilimi National Park, Northern Sierra Leone. Measurements of tools used to dip for driver ants, are compared with those from four other study sites in Africa. The results reveal some fundamentally common characteristics. From both faecal analysis and direct observation, evidence was found that the chimpanzees eat meat. These recordings indicate a varied choice of prey and add new species to those preyed upon by wild chimpanzees. These findings preliminarily support the idea that despite a wide geographical distribution throughout Africa, chimpanzees share some essential conventional behavioural patterns.     

Evidence of tool-use by chimpanzees (Pan troglodytes schweinfurthii) for digging out a bee-nest in the Kahuzi-biega national park,Zaïre

Two sticks were found near a broken bee-nest ofMeliplebeia tanganyikae aff.nigrita Alfken in the Mt. Kahuzi region of Zaïre, and were thought to have been used by a chimpanzee or perhaps several chimpanzees to dig out the subterranean nest. Honey, larvae, and most of the nest had been eaten by them. We did not find any evidence to indicate tool-use by chimpanzees in the Masisi or Itebero-Utu regions, although stingless bees were observed and honey was eaten by chimpanzees in both regions. The sticks resembled in length and diameter those known to be employed for digging termite-mounds in south-west Cameroon and Equatorial Guinea. The tool-behavior of the chimpanzees observed at Mt. Kahuzi may be similar to that of those in central Africa, rather than of those in east Africa where digging-tools have yet to be found. Another possibility is that the chimpanzees have developed the digging-tools independently, based on the need to take animal protein in the Mt. Kahuzi region, where termite-mounds are rarely observed. Instead of seeking termites, they may have a stronger motivation to seek bee larvae, especially the larvae of stingless bees beneath the ground, than to the chimpanzees inhabiting lower or drier forests.     

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.     

Prospects for Bonobo Insectivory: Lui Kotal, Democratic Republic of Congo

Chimpanzees (Pan troglodytes) are well-known to eat invertebrates, especially social insects, across Africa, but allopatric bonobos (P. paniscus) are not. Bonobo insectivory is sparsely documented and apparently sporadic. However, the availability to bonobos of social insect prey and raw materials with which to make tools to exploit them is unknown. Here, we test a set of hypotheses that relates to questions of presence, abundance, density, and distribution of taxa that Pan consume and of vegetation suitable for making extractive foraging tools. We worked at Lui Kotal, Democratic Republic of Congo, where unprovisioned bonobos live in intact forest, far from villages. We collected insect and fecal specimens, transected for prey and assessed raw materials, and monitored mounds of Macrotermes. All but 1 of the major taxa of relevant termites, ants, and (stinging) honey bees were present. The 3 main taxa of insects that chimpanzees elsewhere eat —Macrotermes (fungus-growing termites), Dorylus (Anomma; army or driver ants), and Apis (honey bees)— were abundant and widespread, and usually at densities exceeding those at well-known chimpanzee study-sites. Similarly, woody and nonwoody vegetation suitable for making fishing probes was common at mounds of Macrotermes. There is no obvious ecological reason why bonobos should not use elementary technology in extractive foraging, e.g., termite-fish, ant-fish, ant-dip, honey-dip, to obtain social insects.     

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.     

Exploitation of underground bee nests by three sympatric consumers in Loango National Park,Gabon

Honey represents a highly nutritious resource for animals, but is difficult to obtain given bees' defensive strategies. We investigated exploitation of the underground nests of stingless bees (Meliplebeia lendliana) by three sympatric consumers in Loango National Park, Gabon: the central African chimpanzee (Pan troglodytes troglodytes), forest elephant (Loxodonta cyclotis) and honey badger (Mellivora capensis). Given the differences in their respective morphological traits and sensory abilities, we hypothesized that chimpanzees would be more limited in digging out the bee nests, compared to the other two competitors, and would show behavioral strategies to overcome such constraints. Our dataset comprised camera trap footage recorded over 60 mo at 100 different bee nests. Chimpanzees visited the nests more often than the other consumers, showing a frequency of extraction success comparable to that observed in honey badgers, the most efficient digger. Both chimpanzees and honey badgers increased their extractive attempts across the dry season, whereas elephants did not. The soil hardness was greater during the dry season than the wet season and, possibly in order to compensate for this, chimpanzees showed a tendency toward digging at nests found in relatively softer soil. They also seemed to be inhibited by indirect cues left by other consumers, possibly as a risk‐avoidance strategy. Overall, chimpanzees and honey badgers extracted the underground nests of stingless bees with similar frequencies, whilst forest elephants did so only occasionally. Moreover, chimpanzees can use tools and other behavioral strategies to overcome the physical limitations that may constrain their exploitation of this resource.     

Do chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>) use cleavers and anvils to fracture <Emphasis Type="Italic">Treculia africana</Emphasis> fruits? Preliminary data on a new form of percussive technology

Wild chimpanzees (Pan troglodytes) are renowned for their use of tools in activities ranging from foraging to social interactions. Different populations across Africa vary in their tool use repertoires, giving rise to cultural variation. We report a new type of percussive technology in food processing by chimpanzees in the Nimba Mountains, Guinea: Treculia fracturing. Chimpanzees appear to use stone and wooden “cleavers” as tools, as well as stone outcrop “anvils” as substrate to fracture the large and fibrous fruits of Treculia africana, a rare but prized food source. This newly described form of percussive technology is distinctive, as the apparent aim is not to extract an embedded food item, as is the case in nut cracking, baobab smashing, or pestle pounding, but rather to reduce a large food item to manageably sized pieces. Furthermore, these preliminary data provide the first evidence of chimpanzees using two types of percussive technology for the same purpose.     

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 note on the chimpanzee ecology of the Ugalla Area,Tanzania

Results of a nine-day survey of the chimpanzee habitat in Ugalla area of western Tanzania in 1975 is reported. The most dominant vegetation of the area wasJulbernardia globiflora woodland. Evidence of chimpanzees, both indirect (such as beds, faeces, and food remnants) and direct (calls and actual observation), suggests that chimpanzees are sparsely distributed throughout the area, but that they disproportionately useBrachystegia bussei woodland along escarpments andCynometra-Albizzia riverrine forests. The hilly country of Sisegwa/Mnyangwa was the only area where chimpanzees appeared to be comparatively abundant. Mammal hair found in faeces indicates mammal-predation by the chimpanzees in this area. Hard-shelled fruits ofStrychnos were apparently bitten open without the use of extra-somatic objects. The food repertoire included fruits of the generaAzanza, Canthium, Cordia, andTamarindus.     

Miscellaneous notes on Discomycetes of the Bonin Islands,Ani-jima Island

Cup fungi of Ani-jima Island, the Bonin Islands, were collected in November 1990 and described for the first time from Ani-jima Island. Four species,Dicephalospora rufocornea, Lachnum abnormis, Lachnum pritzelianum, andOrbilia delicatula were collected from the materials in the litter layer ofLivistona chinensis var.boninensis community around Mt. Kita-hutago in Ani-jima Island.Pulvinula globifera was collected from the soil under a camellia (Schima mertensiana) community to the southeast of Mt. Kita-hutago.Lachnum pritzelianum was new to Japan.     

Tool use by wild cebus monkeys at Santa Rosa National Park,Costa Rica

Although wild cebus monkeys have been observed to use tools, this behavior has been reported only rarely. No one has systematically examined tool use in wildCebus, and it is not known how prevalent tool use is in the species' natural repertoire. During 300 hr of observation on 21 wild capuchins (Cebus capucinus imitator) at Santa Rosa National Park in Costa Rica, 31 incidents of tool use, including eight different types of tool-use behavior, were observed. These observations indicate that tool use is a notable behavior pattern in this troop. Considering these incidents of tool use in conjunction with other reports on complex food-getting and preparation behavior byCebus suggests that tool use is a manifestation ofCebus' high behavioral adaptability. Since onlyCebus and the great apes (especially chimpanzees) have been observed to show such a diverse tool-use repertoire, to use tools so frequently, or to show such complex food-getting behavior in the wild, these observations also support the notion thatCebus and the great apes have followed a parallel evolutionary development of tool-using capacity.     

Use of a tool-set by capuchin monkeys (Cebus apella)

The purpose of this study was to examine the use of a tool-set by capuchin monkeys (Cebus apella). Capuchins were presented with an apparatus designed to accommodate the use of pounding tools to crack walnuts and the use of probing tools to loosen and extract the inner meat. Three capuchins used stones and sticks sequentially for these purposes. The capuchins' behavior was similar in form and function to behavior that has been reported for chimpanzees in analogous situations. These results provide further evidence of the extensive tool-using capabilities of capuchin monkeys and are consistent with a hypothesis of cross-species continuity in the skillful use of tools by primates.