Pan Africa News

Chief Editor:Kazuhiko Hosaka, Kamakura Womenʼs University, JapanDeputy Chief Editor:Michio Nakamura, Kyoto University, JapanAssociate Editors:Christophe Boesch, Max-Planck Institute, GermanyJane Goodall, Jane Goodall Institute, USATetsuro Matsuzawa, Kyoto University, JapanWilliam C. McGrew, University of Cambridge, UKJohn C. Mitani, University of Michigan, USAVernon Reynolds, Budongo Forest Project, UKYukimaru Sugiyama, Kyoto University, JapanRichard W. Wrangham, Harvard University, USATakeshi Furuichi, Kyoto University, JapanEditorial Secretaries:Noriko Itoh, Kyoto University, JapanKoichiro Zamma, Great Ape Research Institute, Hayashibara, JapanAgumi Inaba, Japan Monkey Centre, JapanEiji Inoue, Kyoto University, Japan

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The Newsletter of the Committee for the Care and Conservation of Chimpanzees, and the Mahale Wildlife Conservation Society

ISSN 1884-751X (print), 1884-7528 (online) mahale.main.jp/PAN/

DECEMBER 2011

Contents<NOTE>

Responses of Chimpanzees to a Python Koichiro Zamma 13

<NOTE>Chimpanzee Pith-Folding at Toro-Semliki Wildlife Reserve, Uganda

William C. McGrew & Kevin D. Hunt 15<NOTE>

Preliminary Observations of Hand-Clasp Grooming by Chimpanzees at Bulindi, Uganda

Matthew R. McLennan 18<NOTE>

Evaluating the Effectiveness of a 10-Year Old Great Ape Conservation Project in Cameroon

Nikki Tagg, Charles-Albert Petre & Jacob Willie 20<NOTE>

Chimpanzees in Bandafassi Arrondissement, Southeastern Senegal: Field Surveys as a Basis for the Sustainable Community-Based Conservation

Maja Gašperšič & Jill D. Pruetz 23<NEWS>

Memorial Service for the Late Professor Toshisada Nishida 26

<BOOK INFO>Chimpanzees of the Lakeshore 26

<NOTE>Responses of Chimpanzees to a Python

Koichiro ZammaGreat Ape Research Institute, Hayashibara Biochemical

Laboratories, Inc., Japan(E-mail: zamma@gari.jp)

INTRODUCTIONChimpanzees in captivity have been known to show

fear response to snakes1, and wild chimpanzees in Gombe,

13

Tanzania, showed not only fear and avoidance responses (e.g., leaping back, moving away) but also aggressive be-havior (e.g., hitting the snake, chasing and stamping on the ground, shaking saplings) toward live snakes (Causus rhombeatus, Philothamnus spp, etc.)2. Wild chimpanzees have also been known to express a “waa” bark2 and a “huu” (“hoo”) call2,3 when they saw snakes. However, few ob-servations of the reactions of chimpanzees to snakes have been made in the wild.

African pythons (Python sabae) have been thought to be among the potential predators of juvenile and infant chimpanzees4. Only one case involving the reactions of

P. A. N. EDITORIAL STAFF

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14 Pan Africa News, 18(2), December 2011

wild chimpanzees to a half-dead python has been report-ed under experimental conditions in Gombe2. However, no reports on encounters between wild chimpanzees and wild pythons have been published. In this study, I will re-port on the response of wild chimpanzees to a live python.

METHODSI conducted field work from November to December

2008 in the Mahale Mountains National Park in Tanzania. I observed the members of M group5, which was consisted of 59 chimpanzees during the study period. I occasion-ally used a video camera (SONY, DCR-PC105) to record behavior.

OBSERVATIONSI observed a party consisting of seven chimpanzees

(two adult females: GW, RB; two adult males: AL, PM; two young females: PF, RC; and one infant: RB07) at 12:04 on December 15, 2008. At this time, they were rest-ing in the forest near the Mpila River in the northern part of the home range of M group. At 12:07, I heard “wraa” calls from a few hundred meters east. RB then visually searched for the direction from which the “wraa” calls came and for the location of her offspring (RC and RB07); she alternated between these two search patterns. The “wraa” calls were uttered continuously by one chimpan-zee. At 12:10, GW and her adopted offspring (PF) traveled eastward, and RB followed them with her offspring. The two adult males did not move immediately; however, 2 minutes later, PM (alpha male) also went eastward, and AL (beta male) and I followed him.

At 12:18, we arrived at a bush with thick woody vines and met another seven chimpanzees (three adult females: CA, FT, LD; two juvenile females: FV, CR; and two infants: FM, LD07). The party now consisted of 14 chimpanzees, but only one chimpanzee gave “wraa” calls. Eleven chimpanzees watched the bush from a distance of 3 meters. Four juveniles (CR, FV, PF, and RC) climbed a 2-meter-high woody vine, but the adults (AL, PM, and GW) and the mothers ventrally carrying infant offspring (LD and LD07, RB and RB07) sat on the ground. These observers waited and watched events occurring near the bush. At 12:22, AL shook a woody vine quietly and leaped back, but the others did not move and did not show

surprise. Next, PM approached and sat on a woody vine about 1 meter from the bush and shook the vines both ag-gressively and gently.

At 12:25, PM stopped shaking the vines and sat on the ground; he glanced at the bush and leaped back when he noticed the head of python sticking out of the bush (Figure 1, Video 1: available online at mahale.main.jp/PAN/18_2/18(2)_01.html). The python lay motionless for several seconds but then started to come toward me. I moved away on my hands and knees, and 10 seconds later, I looked back and saw the python slithering on the ground about 5 meters away from me. The chimpanzees watched the python intently (Figure 2). We observed the python for about 1 minute until it hid in another bush. Twenty seconds later, PM tracked the python quietly on foot, but he stopped, turned to look at AL, and grimaced (Figure 3). AL immediately approached PM; RB, who was carrying her infant ventrally, and RC followed, and they all tracked the python (Figure 4). Two minutes later, GW, FV, and CA approached this group, which now surrounded the bush the python had entered. “Huu” calls were uttered by one or two chimpanzees during the 2 minutes after the python had appeared, but “wraa” calls were not heard.

At 12:35, AL stopped waiting for the python and left; PM followed him. I heard pant-hoots from their direction, and these were met by pant-hoots uttered by many other

Figure 1. A python sticking out of the bush.

Figure 2. PF watched the python intently from the woody vine.

Figure 3. PM grimaced tracking the python.

15Pan Africa News, 18(2), December 2011

members of M group, who were a few hundred meters from their location. Some of the females of the focal party remained near the bush for more than 30 minutes, but the python did not appear again.

According to estimates based on the video recording, the python was about 2.8 meters long and 0.08 meter in maximum diameter, and moved at a speed of about 0.27 m/s (= 1.0 km/h).

DISCUSSIONChimpanzees in Gombe showed fear and avoidance

responses (rushing away, climbing trees) accompanied by loud “wraa” and soft “huu” calls when they detected a nearly dead python placed within the observation area2. Chimpanzees in Mahale also expressed “wraa” and “huu” calls, but after watching the python, only “huu” calls, which express puzzlement, surprise, or slight anxiety di-rected toward such phenomena as small snakes, rustling noises made by unidentified creatures, and so on6 were heard. Chimpanzees in Mahale also showed fear and avoidance responses (leaping back, climbing on vines, grimacing) to a python, but they were thought to be more interested in the python because they spent more time waiting, watching, and tracking it. Even a mother with her infant held ventrally and a juvenile followed the python despite its status as a potential predator of juvenile and infant chimpanzees4.

Many species of primates face a risk of predation from snakes. It has been hypothesized that the need to avoid snakes shaped the evolution of the primate visual system7. This hypothesis has been supported by experi-ments on Japanese monkeys (Macaca fuscata) reared with no experience with snakes who rapidly detected a picture of a snake8. Wild chimpanzees have been shown to detect and avoid pythons, but they also expressed interest in and approached these creatures. Although many researchers have studied chimpanzees in Mahale over the course 40 years, this is the only case of an encounter between chim-panzees and a python that has been reported. Because the chimpanzees of Mahale may not have been exactly sure about the nature of a python due to their lack of experi-ence with this species, they may have been attracted to and puzzled by it.

ACKNOWLEDGEMENTSI thank TAWIRI, TANAPA and COSTECH for permission

to do the field research, and MMNP and MMWRC for logis-tic support. I also thank Mr. Hitonaru Nishie, Dr. Kazuhiko Hosaka, Dr. Akira Mori and Dr. Masami Hasegawa for valuable comments and Dr. Toshisada Nishida for his kind support. This study was financially supported by the Global Environment Research Fund of the Ministry of Environment, Japan (F-061 to T. Nishida).

REFERENCES1. Yerkes R, Yerkes AW 1936. Nature and conditions of

avoidance (fear) response in chimpanzee. J Comp Psychol 21:53–66.

2. van Lawick-Goodall J 1968. The behaviour of free-living chimpanzees in the Gombe Stream Reserve. Anim Behav Monogr 1:161–311.

3. Crockford C, Boesch C 2005. Call combinations in wild chimpanzees. Behaviour 142:397–421.

4. Izawa K, Itani J 1966. Chimpanzees in Kasakati Basin, Tanganyika (I): Ecological study in the rainy season 1963–1964. Kyoto Univ Afr Stud 1:73–156.

5. Nishida T 1990. A quarter century of research in the Mahale Mountains: An overview. In: The Chimpanzees of the Mahale Mountains, Nishida T (ed), Univ of Tokyo Press, Tokyo, pp 3–35.

6. Goodall J 1986. The Chimpanzees of Gombe: Patterns of Behavior. The Belknap Press of Harvard University Press, Cambridge.

7. Isbell L 2006. Snakes as agents of evolutionary change in primate brains. J Hum Evol 51:1–35.

8. Shibasaki M, Kawai N 2009. Rapid detection of snakes by Japanese monkeys (Macaca fuscata): An evolutionarily pre-disposed visual system. J Comp Psychol 123:131–135.

<NOTE>Chimpanzee Pith-Folding at Toro-Semliki Wildlife Reserve, Uganda

William C. McGrew1 & Kevin D. Hunt2

1 Department of Archaeology & Anthropology, University of Cambridge, UK

2 Department of Anthropology, Indiana University, USA(E-mail: wcm21@cam.ac.uk)

INTRODUCTIONChimpanzees daily make and use tools, and most

of their tool manufacturing transforms vegetation by stripping, peeling, splitting, crushing, clipping, etc. In habituated populations, behavioural data accompany the artefacts, so that observers see precisely how the tools are made and why. However, chimpanzees also modify detached vegetation when not making tools, and these ac-tivities may leave behind puzzling artefacts.

Such are the ‘wadges’ (or ‘quid’) of the pith of the wild date palm, Phoenix reclinata, which we analyse in detail here. The artefact is obvious when encountered: A straight stem is bent repeatedly to alternating sides, concertina-style, so that it has a series of folds at acute angles (see Figure 1). These objects are the spat-out prod-ucts of buccal compression, from which juices have been extracted by squeezing them between tongue and palate.

Figure 4. PM (left), AL, RB carrying her infant ventrally, and RC (right) tracked the python.

16 Pan Africa News, 18(2), December 2011

Their ‘function’ is straight-forwardly nutritional, but the puzzles exist: Why do they take this distinctive shape, and why do so many wadges have an odd number of folds while so few have an even number of folds?

The only previous report of leaf-folding comes from the chimpanzees of Bossou, Guinea, who manufacture water-extracting tools1–4. To make these tools, the apes fold sets of 1–4 leaves (mostly of Hybophrynium brau-nianum) at about 3-cm intervals, while stuffing them into the mouth. This device is inserted by hand into a tree-hole containing water, then extracted and sucked, as a source of sustenance. (Such leaf-folding differs from leaf-sponging and leaf-spooning, although all three techniques yield drinking water3.)

Wadging entails a nutritious object being manipulated by mouth but not swallowed; it seems to be a universal chimpanzee food-processing technique. Goodall5 (p. 238) described wadging at Gombe in detail, noting a variety of food-items, mostly fibrous plant-parts, e.g. figs, bark, etc. In some cases, chimpanzees add leaves to f leshy, ‘rich’-tasting foods, such as meat, eggs, honey, or over-ripe fruit, to form a compressed, homogenised bolus. This may be swallowed, but usually there is an accumulation of amorphous, jumbled fibre-spheres left on the ground. (Presumably, wadges are ejected rather than ingested, because the fibrous mass of foliage is of low quality and would take up valuable gut space.)

We have found no previous record of accordion-fold-ed wadges from any other species of non-human primate. Here we document these special wadges and seek to infer how the artefacts end up with a non-random design, that is, a prevalence of odd-numbered folds. We predicted that the number of folds is positively correlated with the length of the pithy stem involved; given a standard unit of fold-ing distance, the longer the stem, the more folds required. However, we had no clue a priori about the biased number of folds.

METHODSWe studied the wild chimpanzees (Pan troglodytes

schweinfurthii) of the Toro-Semliki Wildlife Reserve, Uganda6. If the partly-habituated chimpanzees could not be un-nested at the beginning of the day, we searched

for them, listening for calls and looking for their signs. Once contacted, we stayed with them for as long as pos-sible. These apes occupy a largely open habitat, which is a mosaic of grassland, scrub, open woodland, gallery forest, and swamp. In tracking or following chimpanzees, we noted feeding traces left by them. For wild date palm wadges, we counted the numbers in an assemblage and noted the sources from whence the piths had been de-tached. Intact and fresh (less than 24 hr old) wadges were collected in plastic ziplock bags and returned to camp for processing. In camp, WCM measured wadges to the nearest 0.5 cm, from fold to fold, and counted the number of folds. Thus, a folded-pith wadge with ‘n’ folds yielded ‘n + 1’ measureable segments.

RESULTSRemnants of the chimpanzees’ wadging of wild date

palm were found daily over May-September 2008. A typi-cal assemblage consisted of several mature fronds lying on the ground less than 5 m from the presumed source plant. A source plant was identified by freshly-damaged ends of the still-intact fronds; these were readily appar-ent because white pith revealed by the damage contrasted with the green outer colour of the foliage. Detached fronds were split length-wise and clipped cross-wise, producing segments of 8–50 cm length. All of the outer epithelium of the segments was peeled away, leaving only the grainy, moist, fibrous pith in strips of less than 1 cm diameter. Few were unused or incomplete strips of pith, by compari-son with more than 30 wadges left strewn on the ground (See Figure 1). These artefacts often were concentrated in areas of less than 50 cm diameter, sometimes in a loose pile, as if the wadger had sat still and worked through a set of wadges before moving on. Fresh wadges had a distinctive pale gold colour and were moist; older wadges turned white and shrank in size as they dried. Desiccated wadges also expanded, changing the angles of the folds from acute to obtuse, unless the fibres were tangled up with one another.

How did we know that these artefacts were made by chimpanzees, if we had not seen them being made? Several times observers were within 5 m of chimpanzees wadging palm fibre in thick undergrowth; we glimpsed fragments of their behaviour and heard the distinctive sounds of fronds being detached. We then recovered fresh artefacts from the site only minutes later. Once, a chim-panzee carried fronds into a tree and wadged arboreally. Also, we often saw the chimpanzees wadging other veg-etation, e.g. at least 4 species of tree-bark (but all of these wadges were the more typical jumbled balls). Many wild date palm wadges were recovered when we tracked chim-panzees, only minutes ahead of us. We found imprints of knuckle- and hand-prints in association with the wadges. No other animals at the study-site did such wadging, nor were there resident humans present to do so.

We measured 110 wadges to the nearest 0.5 cm. These had 1–14 folds; 56 had 3 folds, making it the modal number. Fully extended, the wadges averaged 15.9 cm long (n = 110, range 8–50, median & mode = 17.5). The mean distance between folds (i.e. length of segment) was 3.8 cm (n = 563, range = 1.5–6.5, median = 3.5, mode =

Figure 1. Folded pith wadges of Phoenix reclinata made by chimpanzees at Toro-Semliki Wildlife Reserve.

17Pan Africa News, 18(2), December 2011

3.0). There was a positive correlation (Spearman’s rho, n = 11, rs = 0.98, p < .001, two-tailed) between mean length of wadge and the number of its folds, suggesting that segment-length is relatively standardised (see Figure 2).

Many more wadges had an odd number of folds (1,3,5,7, etc.) than an even number of folds (2,4,6,8, etc). (See Figure 3) For every adjacent pairing, that is, 1 vs. 2 folds, 3 vs. 4 folds, etc., the prevalence of odd-number folds was greater. Presumably the predominance of 3–4 folds is a function of the preferred overall length divided by the typical segment length (see above). Overall, 91 (83%) wadges had 1–13 odd-numbered folds, while only 19 (17%) had 2–14 even-numbered folds (Binomial test, n = 110, z = 6.77, p < .001, two-tailed) (This was not a col-lection bias, as the number of folds was obscured in the field and could not be ascertained until the artefact was extended back in camp.)

DISCUSSIONIf the implicit interpretation of leaf-folding at Bossou

is that the optimal dimension of the fold is a function of the space available in a chimpanzee’s mouth, then it makes sense for pith-folding too. The most economical way to pack a linear object into a much smaller space is to compact it to the maximum permissible length, and the most efficient form of compaction is folding. Thus it is notable that the average length of fold of leaves at Bossou was 3 cm versus 3.8 cm for pith-folding at Semliki. So, all other things being equal, the correlation between number of folds and total length of wadge is likely to be a matter of anatomical (buccal) constraints. (Other, leafy wadges are amorphous in shape, but the stiff longitudinal fibres of the pith make folding the better alternative.)

But what about the odd-even difference? Perhaps chimpanzees consistently clipped lengths of pith that when folded to the optimal segment-length of about 3–4 cm were somehow biased toward an odd number of folds. This seems nonsensical. If an ape were sufficiently pernickety about producing exactly the right number of folds to fill up the mouth, then on average, all other things being equal, one would expect a 50:50 chance of odd or even, given individual variation in buccal volume.

A solution to the oddness riddle emerged seren-dipitously from chance, unobstructed observations of habituated chimpanzees at another Ugandan field site, Kanyawara, in Kibale National Park. There, chimpan-zees wadge the pith of papyrus (Cyperus papyrus) stems. They usually produce amorphous, jumbled-mass wadges, but they occasionally fashion concertina-shaped, folded wadges that are identical in form to the Semliki ones (Bertolani, pers. comm.; See Figure 4). The technique is as follows: After clipping the stem to the final length, the ape folds it in half, with one end in the lips and the other in one hand. She then ‘feeds’ the doubled-over stem into the mouth by the same hand, initial-fold first. Each new fold of the doubled-stem thus produces a pair of folds, which when added to the single initial fold, gives an odd-numbered total. Occasionally, the wadger does not bother with the initial fold, and just ‘feeds’ the stem into the mouth; presumably these artefacts have a 50:50 chance of ending up with an odd or even number of folds.

To what extent does pith-folding at Semliki resemble

Figure 4. Jumbled (standard) and folded pith wadges of Cyperus papyrus at Kanyawara, Kibale National Park.

Figure 3. Frequency of folded pith wadges with odd versus even number of folds.

Figure 2. Average (overall) length of folded pith wadges of Phoenix reclinata as a function of number of folds.

18 Pan Africa News, 18(2), December 2011

leaf-folding at Bossou? There are several differences3: The folded leaf is a tool, the pleats of which increase the water-holding ability of the leaves. The multi-folded pith is not a tool, and the folds seem to have no containing function. So, Semliki wadges are discarded after one use, whereas Bossou’s wadges may be re-used up to 122 times. At Semliki, only one species of plant is used for folding; at Bossou at least seven species are used. In making the wadge, at Semliki the raw material is folded over at its centre-point before being ‘fed’ into the mouth; at Bossou the raw material is stuffed directly into the mouth, start-ing at one end of the leaf. Most of these differences can be explained by the differing functions of the activities.

On the other hand, there are some similarities: Both procedures modify plant materials in order to fill up the buccal cavity, using folding as a technique to do so. The resulting pleats are evenly and similarly spaced, yielding a concertina-like artefact with alternating folds. The mak-ing of both types of artefact involves hand-mouth coordi-nation. Most of these similarities can be explained by the similar biomechanics of the tasks.

Who cares if the number of folds in a wadge is odd or even? One answer is that this is the stuff of culture, not just of gross differences across populations, but also of nuanced variation, when cultural traits are basically similar but subtly different. Such trivial variants are com-mon in human cultures, and the same may be true of non-human artefacts. This ethnographic note reminds us that culture is a layered phenomenon, and that if we operate on one level only, we may miss important features. Another answer is that researchers who work with artefacts, e.g. archaeologists, face challenges in inferring how those ar-tefacts were produced. Etho-archaeology reminds us how difficult it can be to imagine the absent processes that re-sult in material culture.

ACKNOWLEDGEMENTS We thank: Uganda Wildlife Authority and Uganda National

Council for Science and Technology for permission to work at Semliki, UWA rangers Alimosi Baluku, Patrick Biryomumsho, Charles Kasaija, Felix Anidraku, Justus Orobokiritto, and Elly Rutaro for assistance in the field; Moses Comeboy, Eriik Kasutama, Edson Katswamba, Jeremiah Nduthu for assis-tance in camp; Linda Marchant and Timothy Webster for re-search collaboration and comments on the manuscript; Lucie Salwiczek for graphics aid; Paco Bertolani for taking data at Kanyawara; an anonymous referee for helpful comments.

REFERENCES1. Biro D, Sousa C, Matsuzawa T 2006. Ontogeny and cultur-

al propagation of tool use by wild chimpanzees at Bossou, Guinea: Case studies in nut cracking and leaf folding. In: Cognitive Development in Chimpanzees. Matsuzawa T, Tomonaga M, Tanaka M (eds), Springer, Tokyo, pp 476–508.

2. Sousa C 2011. Use of leaves for drinking water. In: The Chimpanzees of Bossou and Nimba. Matsuzawa T, Humle T, Sugiyama Y (eds), Springer, Tokyo, pp 85–96.

3. Tonooka R 2001. Leaf-folding behavior for drinking water by wild chimpanzees (Pan troglodytes verus) at Bossou, Guinea. Anim Cogn 4:325–334.

4. Tonooka R, Inoue N, Matsuzawa T 1994. [Leaf-folding be-havior for drinking water by wild chimpanzees at Bossou, Guinea: A field experiment and leaf selectivity.] Primate Res 10:307–313. [Japanese, with English summary]

5. Goodall J 1986. The Chimpanzees of Gombe: Patterns of

Behavior. Harvard University Press, Cambridge, MA.6. Hunt KD, McGrew WC 2002. Chimpanzees in the dry

habitats of Assirik, Senegal and Semliki Wildlife Reserve, Uganda. In: Behavioural Diversity in Chimpanzees and Bonobos. Boesch C, Hohmann G, Marchant LF (eds), Cambridge University Press, Cambridge, pp 35–51.

<NOTE>Preliminary Observations of Hand-Clasp Grooming by Chimpanzees at Bulindi, Uganda

Matthew R. McLennanAnthropology Centre for Conservation, Environment &

Development, Oxford Brookes University, UK (E-mail: mmclennan@brookes.ac.uk)

INTRODUCTIONThe grooming hand-clasp (GHC) was the first docu-

mented social custom in wild chimpanzees1. It occurs when two chimpanzees seated opposite one another clasp hands overhead and groom each other’s underarm with their free hand. The resulting configuration is strik-ingly symmetrical. Variant forms include one partici-pant grasping their partner’s hand or wrist, and ‘wrist-to-wrist’ in which participants rest their wrists against each other’s forearm, usually one partner supporting most of the weight of both2,3. The behavior may originate from branch-clasp grooming—a universal behavior in chimpanzees4—in which participants grasp an overhead branch whilst grooming socially1. GHC has a patchy dis-tribution across Africa. It occurs in more study communi-ties than not (whether present, habitual or customary), yet is notably absent at three long-term sites: Bossou (Guinea), Gombe (Tanzania), and Budongo (Uganda)4,5. Records from new study sites are valuable because they increase our understanding of chimpanzee behavioral variation. Here I report preliminary observations of GHC in chim-panzees at Bulindi, Uganda.

STUDY SITEBulindi is a forest–agriculture ecotone, 25-km south

of Budongo Forest (Figure 1)6,7. I studied a community of ≥ 25 chimpanzees for 18 months during 2006–2008. Although chimpanzees were unhabituated to close obser-vation, from mid-2007 the community’s six adult males showed signs of semi-habituation7. Consequently, obser-vations of social behavior including grooming and domi-nance interactions increased as the study progressed.

OBSERVATIONSTwo instances of GHC were observed in which the

identity of participants was confirmed. At 0736 on 29 May 2007 we followed chimpanzee vocalizations to a clearing in heavily logged forest. Four adult males were seated on a rotting log, 27 m distant. Two males imme-diately climbed down behind the log while two (JL and MR) remained in view, staring with hair erect. Previous short-range encounters with chimpanzees generated alarm

19Pan Africa News, 18(2), December 2011

and agitation, usually accompanied by threats from adult males7. However, on this occasion the males did not threaten us. After approximately two minutes, JL and MR began self-grooming while monitoring us; a third male (JK) was peeping from behind the log. At 0808 KT climbed on the log and stood glaring at us with erect hair. When he sat JL began grooming him; after several minutes they groomed mutually. At 0818 they raised their right arms overhead and performed the wrist-to-wrist variant of GHC (Figure 2a). Video analysis indicated that JL initiated or facilitated the bout by momentarily taking hold of KT’s right fore-arm at elbow and wrist and gently pushing upwards (Figure 2b). Once extended, KT’s wrist appeared sup-ported by JL’s wrist (Figure 2a). The bout lasted 49 seconds (Video 1, available online at mahale.main.jp/PAN/18_2/18(2)_03.html).

The second instance occurred on 31 July 2007. At 0859 we observed JL, KT and MR in a Parkia filicoidea tree, 55 m distant. JL and KT were groom-ing mutually. At 0904 they raised their left arms and performed wrist-to-wrist GHC for approximately 60 seconds. On this occasion JL rested his left hand/wrist on KT’s forearm.

Two further instances of GHC were recorded. On both occasions grooming chimpanzees were observed in tree crowns from ≥ 80 m distance and identity of participants, and the form of GHC, was unconfirmed. In one case JL and KT formed a grooming clique with

an estrous female but it was unclear if the two males per-formed GHC or if one performed it with the female. In the second case two adult males partially obscured by foli-age performed GHC. Two other males in the party were identifiable (SL and MR). When the party moved off four adult males were present; the two unidentified males were almost certainly JL and KT.

Overhead branch-clasp grooming was witnessed three times: once it occurred between JL and KT, once between KT and SL, and once between an unidentified adult male and estrous female.

DISCUSSIONGHC is performed usually, but not exclusively, by

adults1,5,8. While a possible male bias was indicated in some studies2,9, adults of both sexes participate in this activity1,5. However, particular individuals9,10 and dyads5,10

may perform GHC more often than other group members. Adult females at Bulindi were shy of human observers and most grooming observations involved adult males. Thus, further study is needed to establish the occurrence of GHC in females at this site. Two well-observed GHC bouts involved the same two prime adult males (JL and KT) who were also the likely participants in two addition-al instances. These males performed GHC twice during four well-observed bouts of mutual grooming totalling 157 min. A third prime male (SL) groomed with KT dur-ing three bouts (118 min), but they did not perform GHC. JL and SL seldom interacted and were not seen grooming. Two young adult males (MR and JK) and an elderly male

Figure 2a. Adult males Julius (JL) and Keeta (KT) per forming grooming hand-clasp (May 2007); behind them a young adult male Murray (MR) is watching observers. JL and KT were the top-ranked males at Bulindi. Although in this video-still KT (facing camera) appears to rest his wrist on JL’s wrist, possibly indicating his dominance ( c f . r e f e r e n c e 2 ) , subsequent observations o f s o c i a l b e h a v i o r suggested JL was alpha male (unpubl. data).

F igure 2b . Video-st i l l showing JL (with back to camera) apparently initiating grooming hand-clasp by momentar i ly holding KT’s forearm with both hands and gently pushing upwards.

Fig.1. Map showing the location of Bulindi (at bottom) in relation to Budongo Forest in western Uganda. Gray areas represent forest. The approximate home range of Bulindi chimpanzees is encircled. The locations of the Sonso community in Budongo and nearby Kasokwa community (where grooming hand-clasp has not been seen) are also indicated. An unknown number of small chimpanzee groups utilize forest patches to the north and west of Bulindi.

20 Pan Africa News, 18(2), December 2011

(LR) were also encountered regularly but were rarely seen grooming with other males. These preliminary observa-tions suggest JL and KT—who both received submissive behavior from other males, including the largest male SL —had a GHC ‘partnership’10. More data are evidently needed, but we may tentatively classify GHC as ‘habitual’ at Bulindi4.

While GHC occurs at Bulindi, it has not been seen 25 km north in the Sonso community at Budongo11. GHC has not previously been reported present in one com-munity yet absent in another so nearby (Mahale, where it occurs, and Gombe, where it does not, are separated by 150 km; Kibale, where it occurs, and Sonso, where it does not, are 170 km apart). GHC has also not been seen in the small Kasokwa community bordering the southern edge of Budongo (Janette Wallis, pers. comm.) (see Figure 1). Bulindi chimpanzees use tools to excavate subterranean bee nests for honey—a behavior not recorded at Sonso or elsewhere in western Uganda12. The occurrence of GHC at Bulindi provides further evidence of behavioral varia-tion among chimpanzees in the Budongo region. Multiple small chimpanzee groups inhabit forests patches within the cultivated landscape south of Budongo13. Chimpanzees occur north of Bulindi, closer to Budongo’s southern border (e.g. around Kasongoire Forest Reserve). Future studies should aim to establish the status of GHC among chimpanzees in this intervening area.

GHC was first identified in 5/8 (63%) long-term study communities4. An expanded data-set reveals it occurs in 14/17 communities (82%), including two Pan paniscus communities5,8,14. If we include Bulindi and Kasokwa it occurs in 15/19 (79%). This suggests GHC is usually present in wild populations. Nevertheless, it is easier to confirm presence than absence in unhabituated or semi-habituated communities. Despite the emerging high prev-alence of GHC in wild chimpanzees, the evidence for its absence at three sites (Bossou, Gombe, Sonso) is firm— it has not been seen at these sites in decades of fieldwork. GHC has emerged spontaneously in one captive colony15 and possibly one sanctuary-released group9. This indicates GHC is a dynamic social custom that potentially emerges and disappears in local populations repeatedly over time.

ACKNOWLEDGEMENTSI thank the President’s Office, the Uganda National

Council for Science and Technology, and Uganda Wildlife Authority for permission to work in Uganda. Dan Balemesa, Gerald Mayanda, Tom Sabiiti, Moses Ssemahunge and Jane Stokoe helped in the field. Research was funded by the ESRC and NERC, and the Leverhulme Trust (project reference: F/00 382/F). The manuscript was improved by comments from W.C. McGrew.

REFERENCES1. McGrew WC, Tutin CEG 1978. Evidence for a social cus-

tom in wild chimpanzees? Man 13:234–251.2. McGrew WC, Marchant LF, Scott SE, Tutin CEG 2001.

Intergroup differences in a social custom of wild chimpan-zees: The grooming hand-clasp of the Mahale Mountains. Curr Anthropol 42:148–153.

3. Nakamura M, Uehara S 2004. Proximate factors of dif-ferent types of grooming hand-clasp in Mahale chimpan-zees: Implications for chimpanzee social customs. Curr Anthropol 45:108–114.

4. Whiten A, Goodall J, McGrew WC, Nishida T, Reynolds V, Sugiyama Y, Tutin CEG, Wrangham RW, Boesch C 2001. Charting cultural variation in chimpanzees. Behaviour 138:1481–1516.

5. Nakamura M 2002. Grooming-hand-clasp in Mahale M Group chimpanzees: Implications for culture in social behaviours. In: Behavioural Diversity in Chimpanzees and Bonobos. Boesch C, Hohmann G, Marchant LF (eds), Cambridge University Press, Cambridge, pp. 71–83.

6. McLennan MR 2010. Case study of an unusual human–chimpanzee conflict at Bulindi, Uganda. Pan Afr News 17:1–4.

7. McLennan MR, Hill CM 2010. Chimpanzee responses to researchers in a disturbed forest–farm mosaic at Bulindi, western Uganda. Am J Primatol 72:907–908.

8. Webster TH, Hodson PR, Hunt KD 2009. Grooming hand-clasp by chimpanzees of the Mugiri community, Toro-Semliki Wildlife Reserve, Uganda. Pan Afr News 16:5–7.

9. Humle T, Colin C, Raballand E 2009. Preliminary report on hand-clasp grooming in sanctuary-released chimpanzees, Haut Niger National Park, Guinea. Pan Afr News 16:7–10.

10. Bonnie KE, de Waal FBM 2006. Affiliation promotes the transmission of a social custom: Hand-clasp grooming among captive chimpanzees. Primates 47:27–34.

11. Reynolds V 2005. The Chimpanzees of the Budongo Forest. Oxford University Press, Oxford.

12. McLennan MR 2011. Tool-use to obtain honey by chim-panzees at Bulindi: New record from Uganda. Primates 52:315–322.

13. McLennan MR 2008. Beleaguered chimpanzees in the agricultural district of Hoima, western Uganda. Primate Conserv 23:45–54.

14. Fruth BI, Hohmann G, Beuerlein MM, McGrew WC 2006. Grooming hand clasp by bonobos of Lui Kotal, Democratic Republic of Congo. Pan Afr News 13:6–8.

15. de Waal FBM, Seres M 1997. Propagation of handclasp grooming among captive chimpanzees. Am J Primatol 43:339–346.

<NOTE>Evaluating the Effectiveness of a 10-Year Old Great Ape Conservation Project in Cameroon

Nikki Tagg1,2, Charles-Albert Petre3,4 & Jacob Willie1

1 Projet Grands Singes, Cameroon, Cameroon.2 Centre for Research and Conservation, Royal Zoological

Society of Antwerp, Belgium.3 Liège University, Belgium.4 Gembloux Agro-Bio Tech, Belgium.(E-mail: Nikki.tagg@kmda.org)

The world faces a massive human-accelerated bio-diversity decline. Western lowland gorillas (Gorilla go-rilla gorilla) and central chimpanzees (Pan troglodytes troglodytes) are heavily concerned by this decline, with a range-wide mean of 50% having perished1,2, due to hunt-ing for meat, disease and habitat loss and disturbance3,4,5,6. A large majority (approximately 80%) of all remaining populations of chimpanzees and gorillas live outside pro-tected areas (PAs)7,8 where human pressures are high. A serious everyday threat to great ape survival is hunting for meat3 and as a result of the poverty of an ever-growing

21Pan Africa News, 18(2), December 2011

local population with a taste for game meat, traditional bushmeat hunting has become more and more driven by economic forces9. In the Dja Biosphere Reserve (DBR) it has been noted that all large and medium-sized mam-malian species (except galagos and pottos) are hunted for human consumption10; that wildlife is the source of 98% of the animal protein consumed in villages and towns in the vicinity; and that game meat comprises close to 80% of all meat eaten by the inhabitants—a large portion of this meat acquired through poaching in the reserve11. Severe effects on animal populations can result, lead-ing to decline and extinction of even small-bodied, fast-reproducing species that were previously thought of as be-ing insusceptible to the pressures of hunting, for example some duiker species10,12. Great apes are especially vulner-able to high hunting pressures impacting negatively on their populations, as they are large-bodied animals with slow reproductive rates, and they cannot rapidly recuper-ate from losses1.

Projet Grands Singes (PGS) of the Royal Zoological Society of Antwerp (RZSA), Belgium, seeks to promote wildlife conservation and decelerate the rate of decline of species such as great apes in this region with community-based interdependent conservation and development ob-jectives13. PGS works in a non-protected forest in the buff-er zone of the Dja Biosphere Reserve (DBR; see Figure 1), which is considered as an ‘exceptional priority area’ for great ape conservation as a result of its size (5,260 km²), rich biodiversity, and estimated large population size of great apes14 but in recent years, UNESCO has threatened to declassify the reserve as a result of limited action on the ground15. Being the sole great ape conservation and research project in the northern periphery of the DBR,

PGS adopts an important multi-layered approach in its management of the site: including awareness programmes, con-trol of poaching, supply of alternative incomes and recognition of rights of local people16. In this area, rural com-munities of Baka and Bantu (Badjoue tribe) are amongst the poorest (< $1/d), least developed (infrastructure like roads, schools and health centres lacking) and least educated (literacy and rates of school attendance) in the country. PGS aids communities to de-velop participative sustainable hunting management plans to ensure liveli-hood security and an improved social and economic well-being17, as well as contributing to the maintenance of biodiversity; a method that has been suggested as being crucial, in conjunc-tion with government-led mechanisms for monitoring and law enforcement, in the maintenance of sustainability of bushmeat harvesting18. Secondly, PGS provides small-scale development training and financial aid for the local people, in an attempt to reduce pov-erty and aid in wildlife protection19,20. Finally and crucially, PGS uses scien-

tific research as a conservation tool to forge a rare and im-portant direct link between conservation and benefits for the local communities21. International researchers use es-tablished research facilities to contribute to knowledge of great apes, inarguably crucial to the conservation of any species2, while regular employment of local people in re-search activities reinforces the value of living wildlife and intact forests to the community. Furthermore, the pres-ence of the camp and staff acts as a deterrent to poachers, and in effect offers the site ‘semi-protection’22. Additional project activities including sensitisation to wildlife laws and conservation benefits, and investments in anti-poach-ing actions led by local authorities, further contribute to the better understanding and capacity of the local popula-tion to embrace great ape conservation in their forests.

However the extent and effectiveness of conservation projects such as PGS are poorly known20 and the pressing need of evaluation and evidence of success remains18,23–25. Such evaluation is crucial for planning, refinement and assessment of the effectiveness of conservation ap-proaches26. There is an urgent need to conduct extensive, regular and coordinated evaluation of all conservation ef-forts, including repeated surveys of great ape density and abundance and hunting pressures in the locality27, as well as reporting on outcomes such as technical feasibility, economic sustainability, social appropriation, and con-servation outcomes18 in order to ensure the feasibility of participative conservation actions in such non-protected buffer zones.

It has become evident in the PGS site that gun hunt-ing has dramatically increased in the locality (unpublished data), in accordance with documented trends3. This is due

Figure 1. PGS study site in the northern periphery of the Dja Biosphere Reserve, Cameroon.

22 Pan Africa News, 18(2), December 2011

to easier access to homemade cheap and effective shotguns in recent years; and better acces-sibility to rural areas as a result of an influx of cheap motorbikes allowing buyers from towns and cities to leave commands—and the ammunition required—with local people10,28. Furthermore, in accordance with this increase in gun hunting, more bushmeat being taken from the forests is being sold than in previous years; non-great ape primates are occurring more often in the bushmeat taken from the for-est; primates are mostly being caught with guns; and the majority of these primates are being sold (unpublished data). This suggests that the bush-meat markets in the region are becoming more and more commercialised9,10, in keeping with the documented in-crease in intensity and spatial extent of commercial hunt-ing which has been gradually taking hold for decades1.

This change in hunting in the region shows the seri-ousness of threats and pressures being exerted on great ape populations in non-protected forests over the last decade. It is this commercial bushmeat trade that wipes out species10. Furthermore, amongst tribes in this locality the hunting of apes is not a traditional taboo: ape meat is readily consumed and ape bones are used for traditional mystic practices. As a result, the demand for bushmeat is high and human population density continues to rise3, meaning that hunting pressure represents a serious threat to the viability of the local great ape population. Great apes produce one offspring every four to five years, when their previous offspring is weaned, so the rate of weaned individuals being killed by hunters with guns can easily be higher than the rate of replacement1. If apes in the PGS site had been subject to such intensifying hunting pres-sures since its launch 10 years ago, we might expect to see a huge decline in great ape numbers in this time, or even local extirpation. Studies in sites across the extent of great ape range have shown such declines in great ape nest en-counter rates (eg, 90% in 17 years in Ivory Coast1,14,29,30). However, encounter rates per kilometre (ERKs) of chim-panzees and gorilla nest sites in the PGS site in the buffer zone of the DBR did not significantly change between the launch of PGS in 2001 and later surveys in 2008 (paired sample t-tests: chimpanzee p = 0.579; gorilla p = 0.260; see Table 1). This implies that the abundance of chimpan-zees and gorillas has remained stable despite the increase in gun hunting in the area, suggesting that there has been a check on the negative impacts of hunting on great ape populations during this timeframe10.

This check on great ape hunting is suggested to be due to the multi-layered and long term approach of PGS. Its myriad of conservation and development activities have resulted in a local amnesty on great ape killing in the focal area and suggest that, even in non-protected and heavily-used forests, it is not too late. Such conservation projects are highly capable of having a positive effect on

the protection of biodiversity, in particular great apes, and therefore remain crucial. Stokes31 commend the effective-ness of initiatives to reduce poaching and protect habitats in the maintenance of high abundances of great apes and elephants. Others say it cannot be denied that such con-servation actions (including education, livelihoods, incen-tives and capacity-building) are important16,20,32 and that—quite simply—there is hope.

REFERENCES1. Walsh PD, Abernethy KA, Bermejo M, Beyers R, De

Wachter P, Ella Akou M, Huijbregts B, Idiata Mambounga D, Kamdem Toham A, Kilbourn AM, Lahm SA, Latour S, Maisels F, Mbina C, Mihindou Y, Ndong Obiang S, Ntsame Effa E, Starkey MP, Telfer P, Thibault M, Tutin CEG, White LJT, Wilkie DS 2003. Catastrophic ape decline in western Equatorial Africa. Nature 422:611–614.

2. Stokes EJ 2008. Conservation through scientific collabora-tion: case study-western-gorilla.org. In: Conservation in the 21st Century: Gorillas as a Case Study. Stoinski, Steklis HD, Mehlman PT (eds). Springer, New York, pp. 296–314.

3. Wilkie DS, Carpenter JF 1999. Bushmeat hunting in the Congo Basin: an assessment of impacts and options for mitigation. Biodivers Conserv 8:927–955.

4. Bassey AE, Oates JF (eds) 2001. Proceedings of the International Workshop and Conference on the Conservation of the Cross River Gorillas. Calabar, Nigeria, April 6–9, 2001.

5. Kondgen S, Kuhl H, N’Goran P, Walsh PD, Schenk S, Ernst N, Biek R, Formenty P, Matz-Rensing K, Schweiger B, Junglen S, Ellerbrok H, Nitsche A, Briese T, Lipkin WI, Pauli G, Boesch C, Leendertz FH 2008. Pandemic human viruses cause decline of endangered great apes. Curr Biol 18:1–5.

6. Bermejo M, Rodriguez-Teijeiro JD, Illera G, Barroso A, Vila C, Walsh PD 2006. Ebola outbreak killed 5000 goril-las. Science 314:1564.

7. Morgan D, Sanz C 2007. Best Practice Guidelines for Reducing the Impact of Commercial Logging on Great Apes in Western Equatorial Africa. Gland, Switzerland: IUCN SSC Primate Specialist Group.

8. Poulsen JR, Clark CJ 2004. Densities, distributions and seasonal movements of gorillas and chimpanzees in swamp forest in Northern Congo. Int J Primatol 25:285–306.

9. Chapman C, Peres C 2001. Primate conservation in the New Millenium: The role of scientists. Evol Anthropol 10:16–33.

10. Muchaal PK, Nandjui G 1999. Impact of village hunting on wildlife populations in the Western Dja Reserve, Cameroon. Conserv Biol 13:385–396.

11. Koulagna DK 2001. Country report: The issue of bushmeat in Cameroon. Bushmeat Crisis Task Force CAP Meeting Proceedings (www.bushmeat.org/may2001.htm).

Table 1. Encounter rates per kilometre of chimpanzees and gorillas in the PGS site in 2001 and 2008.

Species Year No. nest sites below cut point

No. visible nests below cut point

ERK (nest sites below cut point)

SD (nest sites below cut point)

Chimp.

2001 86 174 1.433 0.6102008 74 123 1.233 0.704

Significance (p value) 0.579a

Gorilla

2001 34 88 0.567 0.2512008 28 44 0.467 0.302

Significance(p value) 0.260a

Legend: ERK = encounter rate per kilometre (based on nest sites). The number of visible nests per kilometre, recorded along the transects. SD = standard deviation. a t-test for paired samples (N = 10, df = 9).

23Pan Africa News, 18(2), December 2011

12. Fa JE, Juste J, Perez del Val J, Castroviejo J 1995. Impact of market hunting on mammal species in Equatorial Guinea. Conserv Biol 9:1107–1115.

13. Salasfky N 2011. Integrating development with conserva-tion. Biol Conserv 144:973–978.

14. Tutin CEG, Parnell RJ, White LJT, Fernandez M 1995. Nest building by lowland gorillas in the Lopé Reserve, Gabon: environmental influences and implications for censusing. Int J Primatol 16:53–76.

15. World Heritage (2011) Convention Concerning the Protection of the World Cultural and Natural Heritage. 35Com (http://whc.unesco.org/en/decisions/4409)

16. Vermeulen C, Julve C, Doucet J-L, Monticelli D 2009. Community hunting in logging concessions: Towards a management model for Cameroon’s dense forests. Biodivers Conserv 18:2705–2718.

17. Roe D, Nelson F, Sandbrook C (eds) 2009. Community Management of Natural Resources in Africa: Impacts, Experiences and Future Directions. IIED Natural Resource Issues No. 18.

18. Van Vliet N, Milner-Gulland EJ, Bousquet F, Saqalli M, Nasi R 2010. Effect of small-scale heterogeneity of prey and hunter distributions on the sustainability of bushmeat hunt-ing. Conserv Biol 24:1327–1337.

19. Sandbrook C, Roe D 2010. Linking conservation and pov-erty alleviation: the case of great apes. An overview of current policy and practice in Africa. Conservation and Poverty Learning Group Report.

20. Brooks TM, Wright J, Sheil D 2009. Evaluating the success of conservation actions in safeguarding tropical forest bio-diversity. Conserv Biol 23:1448–1457.

21. Nepal S, Spiteri A 2011. Linking livelihoods and con-servation: An examination of local residents’ perceived linkages between conservation and livelihoods benefits around Nepal’s Chitwan National Park. Environ Manage 47:727–738.

22. Williamson EA, Usongo L 1995. I. Survey of primate populations and large mammal inventory ; II. Survey of elephants, gorillas and chimpanzees; Reserve de Faune du Dja, Cameroun. Projet ECOFAC—Composante Cameroun.

23. Ferraro PJ, Patanayak SK 2006. Money for nothing? A call for empirical evaluation of biodiversity conservation invest-ments. PLOS Biol 4:482–488; e105.

24. Pullin AS, Knight TM 2009. Doing more good than harm —Building an evidence-base for conservation and environ-mental management. Biol Conserv 142:931–934.

25. Sutherland WJ, Pullin AS, Dolman PM, Knight TM 2004. The need for evidence-based conservation. Trends Ecol Evol 19(8):305–308.

26. Matthews E, Matthews A 2004. Survey of gorillas (Gorilla gorilla gorilla) and chimpanzees (Pan troglodytes troglody-tres) in Southwestern Cameroon. Primates 45:15–24.

27. Kühl H, Maisels F, Ancrenaz M, Williamson EA 2008. Best Practice Guidelines for Surveys and Monitoring of Great Ape Populations. Gland, Switzerland: IUCN SSC Primate Specialist Group.

28. Willie J 2006. Contribution a l’evaluation de l’incidence de la chasse sur les populations de cephalophes de la periph-erie nord de la reserve de biosphere du Dja (Est-Cameroun). Master’s thesis, University of Dschang, Cameroon.

29. Campbell G, Kuhl H, N’Goran Kouame P, Boesch C 2008. Alarming decline of West African chimpanzees in Cote d’Ivoire. Curr Biol 18:R903/04.

30. Kormos R, Boesch C, Bakarr MI, Butynski TM 2003. West African Chimpanzees: Status Survey and Conservation Action Plan. IUCN/SSC Primate Specialist Group, Gland, Switzerland.

31. Stokes EJ, Strindberg S, Bakabana PC, Elkan PW, Iyenguet FC, et al. 2010. Monitoring great ape and elephant abun-dance at large spatial scales: Measuring effectiveness of a conservation landscape. PLoS ONE 5:e10294.

32. Posa MRC, Diesmos AC, Sodhi NS, Brooks TM 2008. Hope for threatened tropical biodiversity: lessons from the Philippines. BioScience 58:231–240.

<NOTE>Chimpanzees in Bandafassi Arrondissement, Southeastern Senegal: Field Surveys as a Basis for the Sustainable Community-Based Conservation

Maja Gašperšič1 & Jill D. Pruetz2

1 Department of Biology, University of Ljubljana, Slovenia2 Department of Anthropology, Iowa State University, USA(E-mail: maja.g.cisse@gmail.com)

INTRODUCTIONThe western chimpanzee (Pan troglodytes verus) is

considered as one of the most threatened ape species, fac-ing a dramatic decline over the last decade1,2. The latest conservation action plans classified Senegal as “an excep-tionally important priority area” for chimpanzee protec-tion, which demands immediate attention3. Chimpanzees have been expatriated from at least two African countries and IUCN estimated the Senegalese population to be almost extinct, numbering between 200 and 4004. Most apes range in small isolated communities in intense sym-patry with local ethnic groups. Major threats include hu-man encroachment, deforestation for crops, gold and iron digging, along with limited pet trade5. Additionally, this population lives at the northern edge of species’ distribu-tion, in extremely hot, dry and open savanna landscape that characterized an important transitional period in human evolution6. Chimpanzees though have a mythical relation with Senegalese people; therefore local folklore and taboos allows them to share space. This project is part of the investigation “Conservation of chimpanzees in south-eastern Senegal: the human element” supervised by JD Pruetz. Initially, we identified ape communities in Bandafassi Arrondissement, their ranging patterns, key water and food sources, and particularly chimpanzees’ relation to humans via an ethnoprimatological approach7.

Due to the importance of water sources with gal-lery forests, specific food-rich areas, and conflicts with humans encountered we focused on three priority field-sites: mountain slopes above Bandafassi village, where chimpanzees were known to attack goats; the stream of Angafou (12°34′N, 12°24′W), its rich gallery forest and mango orchards, being crop-raided by apes, supposedly after commercial harvesting of baobab and Saba fruits increased; and the surroundings of Nathia (12°29′N, 12°22′W), which was not identified in previous surveys2

and shows no conflict between species. Two buffer-zones are surveyed sporadically to provide basic information about the presence of chimpanzees in the remaining for-est patches between the Niokolo Koba national park and neighboring Guinea-Conakry. Several sites along Gambia River indicate various conflicts between chimpanzees and humans (e.g. palm-wine harvesting, raphia or bamboo-cutting, artisanal gold-digging sites). Chimpanzees are seen there only when people are not common, and it ap-pears that their seasonal activity affects the ranging pat-terns of apes.

24 Pan Africa News, 18(2), December 2011

METHODSHabitat composition at study sites was sampled by

walking several km in random directions and assess-ing the general habitat within a 50 m radius each 30 m. The classification of vegetation types was adapted after Bogart & Pruetz8. In March 2011 a phenology transect (1 km long, 10 m wide) was established at each study site to monitor tree production monthly throughout the year. Chimpanzee and human usage is assessed here on a total of 507 woody plants. Key food sources (e.g. Adansonia digitata, Cola cordifolia, Parkia biglobosa, etc.), and ter-mite mounds and beehives were marked as well. We also noted encounters with large mammals and humans to esti-mate the presence of predators, competitors and potential prey-species for chimpanzees. Data were systematically collected upon sightings, nesting9 and feeding sites, while signs of material culture10 were recorded opportunistical-ly. Diet composition was assessed through observations, feeding traces, and fecal-sample analyses11. Identifying ranging behavior included recording signs of chimpanzee activity and, if indices were positive (fresh debris or vo-calizations), we tried to approach them.

RESULTS From April 2010 through June 2011 we conducted

292 field surveys and identified 13 villages along several dry season water sources associated with chimpanzees. In total we recorded 157 nesting sites, 85 feeding sites, several cases of tool-use (e.g. baobab cracking, termite-fishing, ant-dipping, tuber-digging), hand-dug wells and caves/shelters. Chimpanzees of Bandafassi were encountered more than 160 times, while data on demog-raphy, activity budget and reactions to observers stems from 146 observations at three study sites.

Encounters with chimpanzeesDespite the rarity of closed-canopy

habitat12, chimpanzees were most often observed in gallery and ecotone forest (38.3% and 0.8% respectively; see Figure 1), followed by different woodland types (closed 26.3%, open 19.1% and bamboo 9.2%), on plateaus (3.5%), in grassland and along fields (1.4% both types). These results indicate chimpanzees’ preference for forested habitats, especially for nest-ing. This habitat type is also most heav-ily exploited by humans, particularly nomadic pastoralists13.

The average party size for various chimpanzee groups sighted was 5.8 (range 1–30). The largest daily parties were encountered at Nathia (mean 7.8), followed by Angafou (5.6), while the smallest groups ranged at Bandafassi (4.5). Foraging parties were on average composed of 2.4 males, 2.1 females, 1.6 juveniles, and 1.3 infants. The com-

munity at Angafou regularly nests along the stream and has indicated some acceptance of observers through time, while Nathia chimpanzees seem to have a vast home-range (> 70 km²) and use it seasonally.

As expected, apes were most often feeding (63.4% overall) upon sighting, foraging and traveling (17.9%), drinking (8.3%) resting (4.8%), nesting (2.1%) and other (in 3.4% mostly social behaviors). The most common reaction of chimpanzees in general was to leave the location (38% overall; see Figure 2), but positively they ignored us and indicated curiosity more often over time (24% and 20% respectively). However, important differences in reactions occurred among three sites with different human-chim-panzee relations. The apes indicated fear in only 14.3% of all episodes, but more significantly at Angafou. Nathia chimpanzees exhibited less fear than other communities and often ignored observers during prolonged contacts, while only apes at Bandafassi also displayed at human ob-servers (4.3%).

Competition with humans So far, chimpanzees consumed fruits (73.2%), pods

(7.6%), nuts (4.9%), pith (4.9%), bark (2.9%), leaves (1.5%; partly used for medicinal purposes) and flowers (0.4%) from 39 plant species of at least 17 families. Social insects (termites and weaver ants together in 2.6%), tubers (wild yam and unidentified underground storage organs), honey

Figure 2. Comparison of chimpanzee reactions to humans among three main sites

Figure 1. Habitat composition/available vegetation types compared with chimpanzees’ usage (nests, contacts)

25Pan Africa News, 18(2), December 2011

(each in 0.4% of cases) and traces of geophagy were de-tected in their food repertoire. Many ape food sources are also used by local humans for various purposes (Pruetz listed 17 species14), and at least five of their key feeding items are exploited commercially (e.g. Adansonia digitata, Saba senegalensis, Tamarindus indica, Parkia biglobosa, Cola cordifolia). Among tree species used for food or nest construction several were also cut by shepherds; e.g. Bombax costatum, Khaya senegalensis, Acacia spp., Ficus spp., and Zizyphus spp.13. Fulbe herders from North of the country were in fact most often encountered in the field, pruning trees for their large flocks of sheep (100–300), while harvesting of Saba fruits occurred as a key activity by local people in June.

DISCUSSION: A Vision For The Sustainable Community-Based Conservation

Chimpanzees’ responses to observers appear to be inf luenced by their experience with local people, who are the crucial element for their conservation15. Nature-culture tourism programs, including forest guards, could be an alternative solution to the complex relationship be-tween the species16, while habituating the apes is not rec-ommended17. The incentive could include local traditional customs related to primates (Bedik and Bassari initiation rites) and nature hiking with the opportunistic chance of chimpanzee viewing. This particular research project cov-ers the widest geographical area in Senegal (> 500 km²), including several chimpanzee communities in rare forest patches. As of December 2011, five field assistants, serv-ing as eco-rangers, monitor the most important chimpan-zee sites. We established a limited “surveillance system” similar to J. Carter’s in Guinea or further east in Senegal. Identified eco-guards monitor the ranging behavior of chimpanzees and at the same time limit crop-raiding and attacks on domestic animals. The two bordering-zones are important in terms of recognizing the existing forest-corridors available for possible gene-transfer within the population. It appears that with the continuous presence of researchers the apes ceased to capture goats, similarly as mango-raiding reportedly decreased. Encounters with shepherds, wine-collectors or hunters in the field are re-ported to local Eaux et forets authorities. Preliminary es-timates from surveys in Senegal by S Ndiaye in May 2011 are encouraging and indicate the Senegalese population is stronger than thought and numbers up to 500 chimpan-zees18. In fact, two of our main study areas (Angafou and Nathia) were recognized as priority sites for chimpanzee conservation18. To conclude, this project has laid solid foundations for the sustainable community-based conser-vation in Bandafassi arrondissement. However, the pros-pects for the future co-existence of savanna chimpanzees and humans depend on needed support from governmen-tal agencies as well as investment in terms of research and funding.

ACKNOWLEDGEMENTSFirst we want to thank Senegalese department of Eaux et

forêts at the Ministry of environment and the Arrondissement du Bandafassi to give us the permission for research. The project was generously supported by the Slovenian Research Agency ARRS (Z6-3676). We are sincerely grateful to local

authorities and villagers for accepting our presence in their for-ests, USAID organization Wula Nafaa for initial funding of the project, Fongoli savanna chimpanzee project for logistics and Dondo Kanté for partly managing the project. Collaboration with Janis Carter and Soulèye Ndiaye was invaluable. The research though would not be possible without dedicated field assistants / forest guardians: Gilbert Camara at Bandafassi mountains, Nambura Diallo at Nathia, Khadim Diallo at park borders, Samba Faye Kanté at Angafou, and Omar Diallo at Marewa (along with various tasks). Finally, special thanks goes to Seth Cissé for his overall help, data-collection and supervis-ing the project in my absence.

REFERENCES1. Plumptre et al. 2010. Eastern chimpanzees (Pan troglodytes

schweinfurthii): status survey and conservation action plan 2010-2020. IUCN Species Survival Commission.

2. Boesch C 2008. Why do chimpanzees die in the forest? The challenges of understanding and controlling for wild ape health. Am J Primatol 70:722–726.

3. Kormos R, Boesch C, Bakarr MI, Butynski TM (eds) 2003. West African Chimpanzees: Status Survey and Conservation Action Plan. IUCN—World Conservation Union, Gland, Switzerland.

4. Carter J, Ndiaye S, Pruetz J, McGrew WC 2003. Senegal. In: West African Chimpanzees: Status Survey and Conservation Action Plan. Kormos R, Boesch C, Bakarr MI, Butynski TM (eds), IUCN, pp. 31–39.

5. Pruetz JD, Kante D 2010. Successful return of a wild infant chimpanzee (Pan troglodytes verus) to its natal group after capture by poachers. Afr Primates 7(1):35–41.

6. Sept JM 2002. Chimpanzees on the edge: The implications of chimpanzee ecology in “savanna” landscapes for hom-inin evolution. In: Primates: evolución, cultura y diversi-dad. Contreras JM, Veá JJ (eds), CEFPSVLT, México.

7. Lee PC 2010. Sharing space: Can ethnoprimatology contrib-ute to the survival of nonhuman primates in human-domi-nated globalized landscapes? Am J Primatol 72:925–931.

8. Bogart SL, Pruetz JD 2011. Insectivory of savanna chim-panzees (Pan troglodytes verus) at Fongoli, Senegal. Am J Phys Anthropol 145:11–20.

9. Pruetz JD, Marchant LF, Arno J, McGrew WC 2002. Survey of savanna chimpanzees (Pan troglodytes verus) in southeastern Senegal. Am J Primatol 58:35–43.

10. McGrew WC, Baldwin PJ, Marchant LF, Pruetz JD, Scott SE, Tutin CEG 2003. Ethoarchaeology and elementary technology of unhabituated wild chimpanzees at Assirik, Senegal. PaleoAnthropology 2003:1–20.

11. McGrew WC, Marchant LF, Phillips CA 2009. Standardized protocol for faecal analysis. Primates 50:363–366.

12. Pruetz, JD, Bertolani P 2009. Chimpanzee (Pan troglodytes verus) behavioral responses to stresses associated with living in a savanna-mosaic environment: implications for hominin adaptations to open habitats. PaleoAnthropology 2009:252–262.

13. Massa BE 2011. Predicting Conflict over Scarce Resources: Chimpanzees ( Pan t roglody tes ver us) and Fulbe Pastoralists. Unpublished Master’s thesis, Duke University.

14. Pruetz JD 2002. Competition between savanna chim-panzees and humans in southeastern Senegal. Am J Phys Anthropol 117(S):128 (AAPA abstracts).

15. Ancrenaz M, Dabek L, O’Neil S 2007. The costs of exclu-sion: Recognizing a role for local communities in biodiver-sity conservation. PLoS Biol 5:2443–2448.

16. Hockings K, Humle T 2009. Best Practice Guidelines for the Prevention and Mitigation of Conflict between Humans and Great Apes. Occasional paper 37, IUCN Species Survival Commission.

17. Macfie EJ, Williamson EA 2010. Best Practice Guidelines for Great Ape Tourism. Occasional paper 38, IUCN Species Survival Commission.

18. Ndiaye S 2011. Conservation du chimpanzé au Sénégal. Etat des connaissances et réactualisation du plan d’actions. Rapport d’étude pour USAID/Wula Nafaa.

26 Pan Africa News, 18(2), December 2011

<NEWS>Memorial Service for the Late Professor Toshisada Nishida

The memorial service for Professor Toshisada Nishida who passed away on 7 June 2011, was held at Kyoto University and the Westin Miyako Hotel Kyoto, Japan, on 2 October 2011.

It started at three in the afternoon and was attended by more than 200 people including the colleagues, friends and students of the deceased. A brief note of his career and a copy of Pan Africa News, Special Issue Commemorating the late Professor Nishida, were given to each participant.

At a lecture hall at the Graduate School of Science, Kyoto University, Drs. Masao Kawai, Kosei Izawa, Ryutaro Ohtsuka, William McGrew, Miho Nakamura, and Kazuhiko Hosaka gave talks about Prof. Nishida’s memories in different time periods. Then the video mes-sages from his international colleagues (Drs. Richard W. Wrangham, John C. Mitani, Frans B. M. de Waal) and from Tongwe assistants were shown with Japanese subti-tles. The final speaker at this hall was Ms. Ikuko Nishida, the bereaved daughter. She shared her memories of her father at home. Each speech was really warm and remi-niscent of Professor Nishida’s frank character.

At six in the evening, the participants moved by bus to a hall of Westin Miyako Hotel Kyoto, and of-fered flowers to the altar. Dr. Akisato Nishimura made a short speech and gave a toast to the memory of Professor Nishida. Photos, videos, and some other articles left be-hind were exhibited at the hall. Participants watched them and cherished the memory of the deceased with other participants. Finally, Mrs. Haruko Nishida, the bereaved wife, made an acknowledgment to close the service at eight in the evening.

Several people kindly made donations for this serv-ice. The remaining balance will be donated to the Mahale Wildlife Conservation Society. We would like to continue our activities by following the deceased’s will to conserve chimpanzees and to promote the welfare and education of Tongwe people who have cooperated with the long-term research at Mahale.

<BOOK INFO>Chimpanzees of the Lakeshore:Natural History and Culture at MahaleBy Toshisada Nishida

Chimpanzees are humanity’s closest living relations and are of enduring interest to a range of sciences, from anthropology to zoology. In the West, many know of the pioneering work of Jane Goodall, whose studies of these apes at Gombe in Tanzania are justly famous. Less well-

known, but equally important, are the studies carried out by Toshisada Nishida on the eastern shore of Lake Tanganyika. Comparison between the two sites yields both notable similarities and startling contrasts. Nishida has written a comprehensive synthesis of his work on the behaviour and ecology of the chimpanzees of the Mahale Mountains. With topics ranging from individual devel-opment to population-specific behavioural patterns, it reveals the complexity of social life, from male struggles for dominant status to female travails in raising offspring. Richly illustrated, the author blends anecdotes with pow-erful data to explore the fascinating world of the chim-panzees of the lakeshore.

340 pages Cambridge University PressPublication date: December 2011Paperback $50.00; Hardcover $125.00ISBN: 9781107601789 (P), 9781107015784 (H)

ContentsForeword Frans de Waal; Preface; Introduction; 1. At the beginning; 2. Food and feeding behaviour; 3. Growth and development; 4. Play and exploration; 5. Communication as culture; 6. Female life histories; 7. Sexual strategies; 8. Male political strategies; 9. Culture; 10. Conservation and the future; Postscript; Acknowledgements; References; Index.

Pan Africa News, Vol. 18, No.2Published in December, 2011Address: c/o Human Evolution Studies, Dept. of Zoology, Faculty of Science, Kyoto Univ., Kyoto, 606-8502, JAPANTEL: (+81)75-753-4093FAX: (+81)75-753-4115E-mail: pan.editor@gmail.comURL: http://mahale.main.jp/PAN/ISSN: 1884-751X (Print), 1884-7528 (Online)