Primates

DOI 10.1007/s10329-011-0287-x

ORIGINAL ARTICLE

Plant-food and tool transfer among savanna chimpanzees

at Fongoli, Senegal
Jill D. Pruetz Stacy Lindshield

Received: 10 October 2010 / Accepted: 31 October 2011

Japan Monkey Centre and Springer 2011

Abstract Transferring food is considered a defining

characteristic of humans, as such behavior is relatively
uncommon in other animal species save for kin-based
transfer. Chimpanzees (Pan troglodytes) are one exception,
as they commonly transfer meat among nonrelatives but
rarely transfer other resources. New observations at Fongoli, Senegal, show habitual transfer of wild-plant foods
and other non-meat resources among community members
beyond transfers from mother to offspring. We explore
various explanations for these behaviors with a focus on
age- and sex-class patterns in transfer events. In a total of
27 of 41 cases, male chimpanzees at Fongoli transferred
wild-plant foods or tools to females. Most other cases
involved transfer among males or males taking food from
females. In light of malefemale transfer patterns at Fongoli, we examine four hypotheses that have been applied to
food transfer in apes: (1) testing for male-coercive tendency (van Noordwijk and van Schaik, Behav Ecol Sociobiol 63:883890, 2009), (2) costly signaling (Hockings
et al. PLoS ONE 2:e886, 2007), (3) food-for-sex (Gomes
and Boesch, PLoS ONE 4:5116, 2009), and (4) sharingunder-pressure (Gilby, Anim Behav 71:953963, 2006).
We also consider hypotheses posed to explain transfer
among callitrichids, where such behavior is more common
(Ruiz-Miranda et al. Am J Primatol 48:305320, 1999).
Finally, we examine variables such as patch and food size
and food transport. We discuss our findings relative to
J. D. Pruetz (&)
S. Lindshield
Department of Anthropology, Iowa State University,
324 Curtiss Hall, Ames, IA 50011, USA
e-mail: pruetz@iastate.edu
S. Lindshield
Ecology and Evolutionary Biology, Iowa State University,
Ames, IA 50011, USA

general patterns of non-meat transfer in Pan and examine

them in the context of chimpanzee sociality in particular.
We then contrast chimpanzee species and subspecies in
terms of non-meat food and tool transfer and address the
possibility that a savanna environment contributes to the
unusual pattern observed at Fongoli.
Keywords Sharing
Food transfer
Chimpanzee

Savanna
Pan troglodytes verus

Introduction
The transfer of resources other than meat, such as plant
foods, is uncommon among most nonhuman primates but
common in humans and, to some degree, in bonobos (Pan
paniscus) compared to the other great apes (Hohmann and
Fruth 1996). In addition, callitrichids frequently share plant
foods, and these transfers are mostly associated with
cooperative-breeding behavior (Feistner and McGrew
1989; Huck et al. 2004). Reviews of food-transfer behavior
can be found in Feistner and McGrew (1989) and Stevens
and Gilby (2004). Although food transfer among chimpanzees is frequent between mothers and offspring (Nishida and Turner 1996), and among nonkin when meat is
involved (Gomes and Boesch 2009), it is otherwise rare
(McGrew 1975; Bethell et al. 2000; Nakamura and Itoh
2001; Slocombe and Newton-Fisher 2005; Hockings et al.
2007).
In chimpanzees, when nonrelatives transfer plant foods,
the transfer often involves human-provisioned foods that
are large and energy-rich, such as bananas at Gombe,
Tanzania (McGrew 1975; Goodall 1986), sugar cane at
Mahale (Nishida 1970), or cultivars, such as papaya, at
Bossou, Guinea (Hockings et al. 2007). In general,

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Primates
Table 1 Reports of (non-meat) food and tool transfers at wild Pan study sites
Species and study
site

Events
(N)a

Items shared

Reference (duration of study)

Pan troglodytes verus

Fongoli, Senegal

41

FL, FR, HN, IN, LV, PI, TL (Piliostigma thonningii,

Oncoba spinosa, Ficus sp., Hymenocardia acida,
Adansonia digitata, Oxytenanthera)

Present study (34 months)

Bossou, Guinea

59

FR, LV, TB, PI (Carica papayab, Citrus aurantifoliab,

Ananas comosusb, Zea maysb, Manihot esculentab,
Theobroma cacaob, Saccharum officinarumb, Ficus
exasperata)

Hockings et al. (2007) (12 months)

Tai Forest, Ivory

Coast

N/A

FR (Treculia africana, Coula edulis, Panda oleosa)

Boesch and Boesch-Achermann (2000); Boesch (2009)

(N/A; habituated communities studied since 1984)

FR (Strychnos, Sterculia quinqueloba)

Wrangham (1975) (12 months)

P. t. schweinfurthii
Gombe, Tanzania

457

FR (Musa sp.)

McGrew (1975) (21 months)

Budongo, Uganda

FR, PI, IN (Cubitermes termites, Monodora myristica,

Ficus mucuso, Desplatsia dewevrei, Treculia
africana, Raphia farinifera)

Bethell et al. (2000) (N/A); Newton-Fisher (1999)

(16 months); Reynolds (2005) (N/A; habituated
community studied since 1990); Slocombe and
Newton-Fisher (2005) (1 month)

Mahale, Tanzania

FR, TL (Citrus sp.b, Voacanga africana)

Nishida and Hiraiwa (1982) (22.5 months); Nakamura

and Itoh (2001) (N/A; habituated communities
studied since mid-1960s)

76

PI (Saccharum officinarum)c

Nishida (1970) (13 months)

TB (Clematis sp.)

Lanjouw (2002) (36 months)

Lomako,
Democratic
Republic of
Congo (DRC)

24

FR (Treculia, Carpodinus)

White (1994) (36 months)

Lomako, DRC
(Eyengo)

80

FR (Treculia, Anonidium)

Hohmann and Fruth (1996); Fruth and Hohmann

(2002) (46 months)

Wamba, DRC

175

FR (Anonidium mannii, Treculia africana, Antrocaryon

micraster, Baissea thollonii, Cuervea macrophylla,
Dialium excelsum, Dialium zenkeri, Irvingia
gabonensis, Musanga cecropioides, Pancovia
laurentii)

Kano (1980) (13 months)

Wamba, DRC

40

FR, HN (Treculia, Anonidium, Dialium sp., Ficus,

Gilbertiodendron, Landolphia, Manniophyton,
Musanga, Synsepalum, Tabernaemontana)

Kuroda (1984) (8 months)

Tongo, DRC
P. paniscus

FL Flower, FR fruit, HN honey, IN insect, LV leaf, PI pith, TB tuber, TL tool, N/A not available
a

Due to variations in sampling methods, observations may refer to number of exchanges, accounts, or shared fruits

Non-provisioned and domesticated plant foods that were cultivated (Hockings et al. 2007) or naturally dispersed (Nakamura and Itoh 2001)

Provisioned and domesticated-plant foods

wild-plant food and tool transfer is rare (Table 1; Humle

et al. 2008). We present new observations of habitual
transfer of tools and foods other than meat in the Fongoli
chimpanzee community. Given that chimpanzees share
meat at every site where it is eaten but wild-plant food is
rarely transferred among individuals, we intentionally
excluded vertebrate prey-sharing here. Such a difference in
food transfer patterns suggests that different ultimate and
proximate explanations exist for meat versus non-meat
transfer. Here, we refer only to transfers of tools and foods

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other than meat. While meat transfer at Fongoli appears to

be similar to that in chimpanzees elsewhere, the prevalence
of wild-plant and tool transfer at this site may differ. We
considered variables such as characteristics of the transferred resources (patch size, food size, and availability),
male rank, and female reproductive state, which might
elicit or affect transfer behavior in Fongoli chimpanzees.
Because our data showed a pattern of male chimpanzees
as owners and females as recipients during transfer, we
examined hypotheses set forth to explain resource transfer

Primates

in terms of specific age- and sex-class representation.

These hypotheses include: (1) females taking resources as a
test of males coercive tendencies (van Noordwijk and van
Schaik 2009), (2) food transfer representing a costly signal for males (Hockings et al. 2007), (3) food-for-sex
reciprocity eliciting food transfer in return for mating
opportunity (Gomes and Boesch 2009), and (4) sharingunder-pressure, in which active begging behavior provokes
an owner to give up some part of a resource as a means to
reduce harassment (Gilby 2006). Additionally, we explored
hypotheses that have been posed in large part to explain
food transfer in callitrichids (Ruiz-Miranda et al. 1999).
These hypotheses predict food transfers (1) of items individuals cannot get themselves (foraging assistance
hypothesis: Price and Feistner 1993), (2) as a means to
teach conspecifics (dietary breadth hypothesis), (3) of items
that are rare but nutritionally valuable (rare item hypothesis), and (4) in order to maintain social relationships
(social benefits hypothesis).

Methods
The Fongoli chimpanzee community ranges within the
Kedougou (formerly Tambacounda) region in southeastern
Senegal (1240 N, 1213 W). The savanna environment
here is a mosaic of woodland, grassland, bamboo, and
gallery forest habitats (Pruetz 2006). The extensive dry
season lasts for more than 7 months, and rainfall averages
less than 1000 mm annually (Pruetz and Bertolani 2009;
Pruetz, unpublished data). Maximum temperature in the
dry season exceeds 40C. Although apes here are sympatric
with humans who grow corn and millet predominantly,
Fongoli chimpanzees do not crop raid. The Fongoli
chimpanzee community varied annually between 30 and 36
individuals from 2005 to 2011, averaging 31 individuals
during this 6-year period. From January 2008 to August
2011 the community numbered 2834 individuals and was
comprised of eight adult females, 811 adult males, and
1315 immature individuals. Some immature individuals
advanced to older age classes during the study period (e.g.,
from infant to juvenile). We defined adults as older than
13 years, unless they had not integrated into the adult
social dominance hierarchy (in the case of males), in which
case they were termed subadults. We defined adolescents
as 913 years, juveniles as 49 years (or younger if
weaned earlier than age 4), and infants up to 4 years of age,
unless they were not yet weaned (after Baldwin 1979, with
slight modification).
Most Fongoli community members were habituated in
2005, but some adult females remained semi-habituated, in
that they exhibited signs of nervousness around observers
when adult males were absent. Consequently, adult females

were not followed apart from when they were in mixed-sex

parties, but all-occurrence samples of behavior (Altmann
1974) could be reliably collected on females in the presence of males. Chimpanzee parties that included adult male
focal subjects were followed from nest to nest between
March 2005 and August 2011 for a minimum of 20 days
per month. Adult male ranks were identified by the direction of pant-grunt greetings. Rank reordering frequently
occurred during the study. The study periods included here
are September 2007September 2009; FebruaryMarch,
JuneJuly, OctoberDecember 2010; and MayAugust
2011 (approximately 34 months). Before January 2008,
data were collected ad libitum. Later, data were collected
on transfer events among individuals on an all-occurrence
basis.
A transfer event was defined as the exchange of a
resource (e.g., fruit, tool) between one dyad within an
uninterrupted feeding or foraging bout. Multiple transfers
of the same resource between a single dyad within a bout
were considered one event. We rated transfer events on a
scale originally developed by Boesch and Boesch
(1989:551) to classify meat-sharing behavior, but we used
the letter T for transfer to label these categories (for
additional scoring approaches, see Nishida 1970; McGrew
1975) (Table 1). Theft (T1) occurred when the receiver
used force or aggression to take a resource from the owner,
and the owner protested the receivers behavior; theft is the
only obvious agonistic exchange in this classification system. Recovery (T2) occurred when the receiver took an
item dropped or placed on the ground by the owner, and the
owner tolerated the receivers actions. We modified this
category by stipulating that the food must be within an
arms length of the owner to distinguish from scrounging
of foods that had been distinctly abandoned. In passive
transfer (T3), the receiver took an item held by the owner,
and the owner passively tolerated the receivers behavior.
Activepassive transfer (T4) differed from T3 in that the
owner moved to facilitate the exchange. Active transfer
(T5) occurred when the owner actively divided the item so
that the recipient could easily take a portion, or presented a
portion to the recipient, but kept a majority of the item for
him/herself. Finally, giving (T6) occurred when the owner
allotted the majority of an item to the recipient.
We considered aspects of transferred resources, male
rank, and female reproductive state as variables that might
elicit or affect transfer behavior in Fongoli chimpanzees.
To provide some indication of food availability, we
described food patches according to their size (small,
medium, large) relative to the potential number of chimpanzee co-feeders (after Pruetz 2009) in an average party at
Fongoli (mean party size 15 individuals: Pruetz and Bertolani 2009). A patch was defined as an area in which an
individual could continuously move and eat, usually a tree

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Primates

crown (after Pruetz 2009). Patch distribution was scored as

scarce, abundant, or ubiquitous, relative to other plant
foods within the home range of Fongoli chimpanzees. Food
item size was based on length, width, or diameter
measurements.
To take into account the possible influence of reproductive condition on transfer, we scored adolescent and
adult females according to the appearance of their estrus
swelling. We used a four-point scale, with 0 indicating a
fully detumescent female and 3 indicating a fully tumescent female. Chimpanzee females can exhibit genital
swellings that do not correspond with true estrus cycles
(Wallis and Lemmon 1986). Therefore, we indicated
whether females were cycling regularly or showed acyclic
swelling, because male chimpanzees may be attracted to
females with genital swelling regardless of their reproductive state.
In order to place our observations into a comparative
context, we reviewed the literature for studies of plant-food
and tool transfer behaviors in Pan (Table 2). In our most
specific comparison, we focused on studies enabling us to
compare transfer events as defined here, and to identify the
age and sex classes of individuals (Table 3). We excluded
instances of direct provisioning by humans and cases of
crop raiding in order to identify features of food transfer
that did not involve artificially abundant food patches. This
comparison provides the most appropriate context in which
to identify evolutionarily important proximal causes of
transfer. We included two sites where researchers had
provided detailed descriptions of all transfer events (Budongo, Uganda: Bethell et al. 2000; Reynolds 2005; Mahale,
Tanzania: Nakamura and Itoh 2001). To further examine
transfer behaviors in the variety of contexts in which they
have been reported, we subsequently broadened our criteria
to include instances of provisioning (Mahale: Nishida
1970) and crop raiding (Bossou, Guinea: Hockings et al.
2007).

Results
Observations of food and tool transfer
Plant-food transfer
Female recipients Twenty-seven cases of plant-food
transfer were recorded, with 15 involving four different
adult females and three juvenile or infant females as
recipients (Table 2). All the adult females were cycling or
swelling during the observation periods, but not all females
were tumescent during the transfer events.
Of the seven transfer cases involving young adult female
TM, six occurred following detumescence (estrus score 0).

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TM is estimated to be [15 years of age and gave birth to

her first infant in December 2008, although she exhibited
an estrus swelling typical of an adult for several years
preceding this. Case 1: on 2 March 2008, TM took an
Oncoba spinosa fruit from adult male SIs hand (T3). This
fruit is 57 cm in diameter and grows on a shrub-like tree
distributed in locally abundant but small patches relative to
average chimpanzee party size. Case 2: on this same day,
TM took a branch with small Ficus sp. fruits from the
mouth of adult male DF who had been carrying it in his
mouth (T3). These small fruits (\5 cm diameter) are distributed in relatively scarce, small- to medium-sized patches relative to chimpanzee party size. Case 3: on 9 March
2008, TM approached adult male BI, who had five Piliostigma erinaceus pods on the ground in front of him. He
quickly grabbed two of them, while TM retrieved the
remaining three (T2). Piliostigma trees are abundantly
scattered throughout the chimpanzees range but occur in
small- to medium-sized patches relative to chimpanzee
party size. The seedpods are approximately 1020 cm in
length and\5 cm in width. Case 4: that same day, TM took
a Piliostigma seedpod from FR, an adolescent male (T3).
Case 5: on 7 March 2009, TM took one of adult male SIs
Piliostigma pods after he groomed her (T2). SI moved
away and ate his remaining seedpods. Case 6: on 27
November 2010, TM (estrus score 2) ate Ficus sp. fruits
(similar to those described above) from a branch held by
adult male DV. DV had carried the branch approximately
5 m from the tree. It initially held more than 10 figs, but
DV was left with four figs after TM moved away. TM
peered at DV for approximately 30 s before taking the figs
from him, and DV made no attempt to stop her (T3). Case
7: on 15 July 2011, TM approached adolescent female LL
and took a branch with figs (Ficus sp.) on it from her
mouth. LL watched TM with a fear grin but made no move
to stop her (T1).
Four plant-food sharing events involved the cycling,
multiparous, adult female FA. FA was detumescent when
two events occurred and was at peak estrus when the other
two were recorded. Case 8: on 28 May 2009, FA approached adult male KL, who had broken off several Ficus
branches and taken them approximately 25 m from the
shrub-sized tree to feed on them. FA took a branch of Ficus
leaves from the ground at KLs side and began feeding on
them (T2). KL made no move to stop FA from taking the
branch but immediately took the other branches from the
ground at his side and moved away approximately 15 m.
FA was fully detumescent. Case 9: on 8 June 2009, FAs
adolescent son FR broke off a small Hymenocardia acida
branch (\1 m in length), covered in flowers, and carried it
more than 10 m to the vine clump below which other party
members rested. This shrub-like tree is relatively abundant
but occurs in small patches. FA joined him, feeding

Primates
Table 2 Summary of transfer events at Fongoli, Senegal (present study)
Case #

Resource exchanged

Transfer scorea

Ownerb

Recipientb

Swelling scorec

Oncoba (FR)

AM (10)

AF

Ficus (FR)

AM (5)

AF

Piliostigma (FR)

AM (6)

AF

Piliostigma (FR)

am

AF

Piliostigma (FR)

AM (10)

AF

Ficus (FR)

AM (45 tie)

AF

Ficus (FR)

af

AF

Ficus (LV)

AM (3)

AF

Hymenocardia (FL)

am

AF&if

10

Piliostigma (FR)

AM (5)

AF&if

11
12

Piliostigma (FR)
Piliostigma (FR)

2
2

AM (10)
AM (5)

AF
AF

3
0

13

Adansonia (FR)

AM (46 tie)

AF

14

Adansonia (FR)

sam

jf

15

Adansonia (FR)

AM (1)

jf

16

Beetle larvae (IN)

AM (56)

if

17

Adansonia (FR)

N/A

AM (1)

AM (3)

18

Piliostigma (FR)

AM (10)

AM (4)

19

Adansonia (FR)

AF

AM (3)

23

20

Adansonia (FR)

am

AM (3)

21

Adansonia (FR)

jf

am

22

Adansonia (FR)

AM (7)

AM (3)

23

Adansonia (FR)

AM (1)

AM (7)

24

Hymenocardia (FL)

am

AM (1)

25

Oxytenanthera (PI)

N/A

AM (2)

AM (7)

26

Oxytenanthera (PI)

am

AM (56 tie)

27
28

Adansonia (FR)
Honey

1
5

AM (46 tie)
AM (3)

AM (8)
jm

29

Honey

AM (3)

AM (4)

30

Honey

AM (7)

AF

23

31

Soil/clay

am

AF

32

Galago hunting (TL)

am

af

23

33

Galago hunting (TL)

am

af

34

Termite fishing (TL)

34

AM (9)

af

23

35

Termite fishing (TL)

AM (5)

AF

36

Termite fishing (TL)

AM (3)

AF

37

Termite fishing (TL)

AM (10)

AF

38

Termite fishing (TL)

am

AF

39

Termite fishing (TL)

AM (7)

AF

40

Ant dipping (TL)

im

AF

41

Termite fishing (TL)

AM

af

FR fruit, LV leaf, FL flower, IN insect, PI pith, TL tool, AF Adult female, af adolescent female, jf juvenile female, if infant female, AM adult
male, am adolescent male, jm juvenile male, im infant male
a

Resource-transfer scores are described in the Methods section

Adult-male ranks are shown in parentheses; immature males are subordinate to adult males; adult-female ranks are not known

Female state of anogenital swelling was scored on a four-point scale, where 0 is maximally detumescent and 3 is maximally tumescent

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Primates
Table 3 Number of owners and recipients by agesex class for food (all sites) and tool (Fongoli only) transfers in Pan troglodytes
Site

Events (N)

Ownera
AM

Receivera
AF

Fongoli

41b

22 (54)

1 (2)

Budongo

7b

6 (86)

1 (14)

Mahale
Mahale

5b
70c

2 (40)
53 (76)

2 (40)
10 (14)

Bossou

38d,e

35 (92)

3 (8)

Source

imt

AM

AF

14 (34)

11 (27)

22 (54)

8 (20)

Present study

2 (29)

2 (29)

3 (42)

Newton-Fisher (1999); Bethell et al. (2000); Reynolds

(2005)

3 (60)
19 (27)

2 (40)
46 (66)

0
5 (7)

Nishida and Hiraiwa (1982); Nakamura and Itoh (2001)

Nishida (1970)

1 (3)

25 (66)

12 (31)

0
1 (20)
7 (10)
0

imt

Hockings et al. (2007)

Motherinfant transfers are excluded. Numbers in parentheses are percentages

a
imt: immature males and females; note that Hockings et al. (2007) include immatures less than 8 years old and do not present information on
adolescents; for all other studies, immatures include all non-adult individuals; see Table 2 for other agesex abbreviations
b

Non-provisioned foods from wild sources

Provisioned and domesticated foods

Non-provisioned and domesticated foods

Excludes motheroffspring transfers

simultaneously on the flowers on the branch (T3). FAs

unweaned, 5-year-old infant daughter FN fed briefly on the
same branch. FR stopped feeding after several minutes, and
FA abandoned the branch after 4 min. FA was completely
detumescent. Case 10: on 24 February 2010, FA (at peak
estrus) approached and copulated with adult male DF who
had approached the tree she was in less than 2 min previously. DF had carried 34 Piliostigma seedpods in his
mouth and began eating them before FA descended and
presented to him for copulation. Following copulation, FA
reached behind DF and retrieved two of the Piliostigma
fruits from the ground where he had placed them, and she
began feeding on them. Her unweaned daughter FN
retrieved a seedpod DF had been feeding on earlier (T2).
Case 11: on 5 March 2010, FA (fully tumescent) approached and took a Piliostigma seedpod from adult male SI.
After FA took a seedpod from his pile of several fruits, he
moved approximately 4 m from her and resumed eating.
Once finished with the seedpod she took, FA approached SI
once again and took two seedpods from his pile, less than
1 m from him. SI made no attempt to stop FA either time,
other than moving a short distance away from her following the transfers (T2).
Two adult females were involved in one plant-food
transfer case each. Both of these females are timid in
respect to observers, relative to other females in the group.
One case of plant-food transfer involved the cycling,
multiparous female LU who was anestrous when the event
was observed. Case 12: on 14 March 2010, LU approached
and took two or three Piliostigma seedpods from adult
male BI. BI had carried at least five seedpods from a
feeding tree to a shady spot where he and LU fed on them.
He made no attempt to stop LU from taking the seedpods,
although he did move one closer to him as she finished one

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seedpod and before she reached past him to retrieve

another (T2). Another case involved the cycling, primiparous adult female NA, who exhibited a low estrus score at
the time. Case 13: on 4 November 2010, adult male BI
allowed NA to take a piece of baobab (Adansonia digitata)
fruit from his hand (T3). BI then broke off another piece
for himself. Adansonia trees are sparsely scattered
throughout the chimpanzees range but occur in mediumto large-sized patches relative to chimpanzee party size.
These fruits vary in size from 7.5 to 54 cm in length and
7.520 cm in diameter (Sidibe and Williams 2002).
Three cases of plant-food transfer involved juvenile or
infant females taking food from adult or subadult males.
Case 14: on 21 November 2010, juvenile female SO ate
Adansonia fruit from subadult male BOs hand and then
from fruit he held in his foot while both sat in the trees
crown (T3). Initially, BO showed no inclination to stop SO,
but 2 min later he jerked the Adansonia fruit away to keep
her from taking more. SO left to recover an abandoned
Adansonia fruit, as the alpha male approached them. Case
15: on 5 December 2010, juvenile female FN followed the
alpha male, LP, along with her mother, FA. LP initially
avoided FNs mother by moving about 1 m from her
approach, but FN twice took Adansonia fruit from his hand
(T3). FN screeched between transfers; LP was out of sight
but may have threatened FN in some way. FN moved on to
take Adansonia fruit from her mother. Case 16: on 7
August 2011, infant female AM approached and took
unidentified beetle larvae from adult male BIs feeding site
(T2). BI had pulled a large piece of bark from a dead tree to
harvest these larvae. The bark was apparently very difficult
to detach, based on the fact that adult females, including
AMs mother (TI), were repeatedly unable to pull off
pieces of bark at various sites.

Primates

Male recipients The 11 remaining cases of plant-food

transfer involved adult males. In six of these cases, adult
males transferred food to one another. Five cases involved
the same adult male (KL), who was responsible for two of
the five cases in which a dominant individual forcibly took
food from a subordinate individual. Case 17: on 17
November 2008, KL approached alpha male YO, who had
been eating Adansonia fruit, and KL subsequently moved
away with fruit. The exchange was out of view, but KL did
not have fruit immediately prior to interacting with YO
(sharing score not available [N/A]). Case 18: on 27 February 2010, KL retrieved an uneaten Piliostigma fruit from
the pile of three or more fruits that subordinate adult male
SI was feeding on, after he had carried these fruits from a
nearby tree to the ground (T2). KL had been resting less
than 1 m from where SI was feeding. Case 19: on 20
October 2010, KL took adult female NNs unripe Adansonia fruit, which she had carried with her from a tree to
the ground (T3). KL groomed NN before taking her fruit,
although he also groomed with another adult male. NN
grunted then left with another fruit she had also carried
with her. NN exhibited a swelling score of 23. NN is an
older female and, due to an illness that has affected her
genital swelling, she is almost continuously partly swollen,
but males rarely copulate with her. Case 20: on 23
November 2011, KL took an Adansonia fruit from adolescent male DW, who screamed (T1). Case 21: DW
immediately took an Adansonia fruit from juvenile female
FN, who also protested by screaming (T1).
Three cases of plant-food transfer involved LP, who
eventually became the alpha male. In two cases, he transferred food with the same subordinate, adult male, BN. Case
22: on 24 November 2008, BN approached and pant-grunted to LP. BN had been feeding on Adansonia fruit and
moved towards LP with a fruit in his mouth. LP took the
fruit from BNs mouth during the greeting (T3). Case 23: on
5 December 2010, LP (who by now had become the alpha
male) allowed BN to take an Adansonia fruit he had placed
on the ground in front of him (T2). LP had carried four fruits
approximately 17 m from a baobab tree. BN, who had been
resting and self-grooming, approached LP and grunted as he
took the fruit from in front of LP. LP made no move to stop
BN, nor did he otherwise indicate any resistance. However,
after approximately 1 min, LP moved up into vines several
meters away. BN did not immediately begin eating the fruit
but sat and self-groomed for 7 min, finally leaving with the
fruit. Case 24: on 29 May 2010, LP approached and took a
branch of Hymenocardia flowers from adolescent male LT,
who made no attempt to stop him.
Case 25: on 3 April 2009 adult male MM broke open a
stalk of bamboo (Oxytenanthera abyssinica) and began
eating the pith. Subordinate adult male BN approached

MM and took some of his pith. It is not known whether

MM had been holding the bamboo or had placed it on the
ground when BN took it (sharing score N/A). Bamboo is
found in large patches relative to average chimpanzee party
size and is locally ubiquitous but patchy within the communitys home range.
Case 26: on 16 November 2010, subordinate adult male
SI approached dominant adult male DV, who had just
descended an Adansonia tree with four fruits. SI attempted
to take one or more of DVs fruits, but DV resisted SIs
attempt with an aggressive stare and posture. SI directed a
waa bark threat at DV. DV charged past SI and hit him.
SI responded by chasing DV as other chimpanzees waa
barked. DV ultimately dropped all of his fruits. SI returned
and recovered three fruits, while DV returned to the
Adansonia tree to eat more fruit (T1).
Case 27: on 10 August 2011, adult male DF approached
and took a stalk of bamboo from late-adolescent male LT,
who protested with a fear grin but moved away and fed on
bamboo less than 5 m from DF (T1).
Honey transfer
Three instances of honey transfer were observed during the
study period. Two cases were among males during the
same feeding bout, and one was between a malefemale
dyad. Case 28: on 14 July 2009, adult male LP grabbed a
handful of honeycomb from an active hive and began
eating it after leaving the hive area. LP handed honeycomb
to juvenile male DW 7 min later (T5). DW is thought to be
LPs younger brother (mother disappeared and presumed
deceased), based on patterns of strong affiliation and the
fact that they share the same mtDNA haplotype (Stewart
et al., unpublished data). Case 29: on this same day, subordinate adult male MM, who had been traveling with LP,
approached and took a piece of honeycomb from him
9 min later, with no resistance from LP (T3). Case 30: on
13 February 2010, adult female NN recovered pieces of
honeycomb on the ground where they had fallen from adult
male BNs hands. BN was feeding while sitting within
arms reach of NN (T2). An adolescent male sitting within
arms reach of BN did not recover the fallen honeycomb
pieces, although he sat and peered at BN eating honey
during this event.
Soil/clay transfer
One case of soil/clay transfer was observed when a chimpanzee party engaged in geophagy. Case 31: on 19 August
2010, adult female TM approached and took a piece of clay
from the hand of adolescent male JM, who had carried the
clay from inside a small cave (T3).

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Primates

Tool transfer
Ten cases of tool sharing were recorded between females
and males. During six of these events, the females were at
or close to peak tumescence. Two incidents involved
cycling female TM, who was detumescent at the time. Case
32: on 26 September 2007, subadult female TM was seen
jabbing a stick tool into a tree cavity in an action typical of
Galago hunting (Pruetz and Bertolani 2007), while adolescent male JM fashioned a tool beside her. After her tool
broke, TM took JMs tool and resumed jabbing into the
cavity (T3). TM later abandoned the site, but JM remained
to hunt. Case 33: on 7 October 2007, TM was termite
fishing when adult male KM sat down beside her with a
tool in his mouth. After her tool became ineffective, she
took KMs tool and resumed fishing. Next, KM made
another tool and waited beside her with his tool in his
mouth. Within minutes, TM took the second tool from
KMs mouth (T3 or T4). KM made a third tool and waited
beside TM until she left the termite mound; he made no
attempt to termite-fish afterwards.
Case 34: on 9 June 2009, adult male DF and adult
female TI termite fished at the same mound, approximately
1 m from one another. TI then moved 1.5 m around and
behind DF to take one of two tools he had made and placed
on the ground at his side (T2). TI returned to her same
fishing site and grunted to DF as she passed. DF looked at
TI, looked for the tool she took from the ground at his side,
and resumed fishing. DF was the only adult male with this
small party on this day, and TI was fully tumescent. DF
copulated with TI once before and three times after the
transfer on this day. DF exhibited possessiveness with TI
(sensu Tutin 1979), as he was attentive to and solicitous of
her movement, as well as attentive to her interaction with
other individuals. TI ignored sexual solicitation by adolescent males in the same party on this day.
Four cases of tool transfer involved multiparous adult
female FA, who was cycling during half of the events. Case
35: on 19 April 2009, FA (fully detumescent but cycling)
approached adult male KL and took the fishing tool he was
using out of his hand. KL showed no resistance and briefly
groomed FA while she fished for termites (T3). Case 36: on
26 March 2010, FA was again detumescent when she took
the tool adult male SI was using to termite fish (T3) and
supplanted him from his fishing site. SI made another tool
and moved to a different spot to termite-fish. In the final
two cases, FA was not cycling, haven given birth 8 months
earlier. Case 37: on 25 June 2011, FA took a termite fishing
tool from adult male BN after approaching him and panting, then peering at his termite fishing for approximately
1015 s. After FA took BNs tool and began fishing, he
inspected her genitals and left the mound (T3). Case 38: on
11 July 2011, FA approached adolescent male MI, where

123

he sat termite fishing and took his tool from his hand while
he had it inserted into the termite mound. FA then moved
MI out of the way with her hand and began termite fishing
with his tool at his site (T3). MI was dominant to FA and
all other females at this point. Case 39: on 27 November
2010, infant male CY (TMs son) retrieved an abandoned
ant-dipping tool from near an army-ant nest. CY attempted
to insert the tool, which was over 1 m in length, into a tree
root. Adult female TI arrived and took the tool from CY. TI
moved to ant dip but then abandoned the tool in the nest
and fled with a handful of ants. TI had ant-dipped at this
nest approximately 5 min earlier with a similarly-sized
tool, and it is likely that the tool she took from CY was the
same one she had made and used previously. TI was at
peak estrus on this day (T3).
Two cases of tool transfer involved recent immigrant,
adolescent female LL. Case 40: on 4 July 2011, LL (at peak
estrus) approached adolescent male LT, who was using a
tool to jab into a cavity in an action typical of Galago
hunting. LL took LTs tool out of his hand and from the
cavity, sniffed the tool, and began using the tool in the
manner of Galago hunting in the same cavity (T3). LT
made no move to stop LL, and he was dominant to LL and
to adult females at this point. Case 41: on 7 July 2011, LL
termite fished at the same mound as adult male BN. LL was
at peak estrus and, after she and BN had copulated, LL took
BNs tool from him and returned to continue termite fishing at the same site (T3).
Unsuccessful attempts at food transfer
Finally, although the following four cases are not examples
of successful transfers, we believe they warrant description,
as they illustrate the tendency for subordinate Fongoli
chimpanzees outside of the motheroffspring pair to
attempt to engage in plant-food transfer. Case U1: on 23
February 2010, FA (fully tumescent) approached subadult
male BO and attempted to take a Ficus wadge from his
mouth twice with her hand. She was unsuccessful, as BO
pulled his head away each time and grunted once. BO had
twice copulated with FA earlier this day. BO was dominant
to all adult females, including FA. Case U2: on 16
November 2010, FA approached her adult son DV and
pant-grunted to him. FA exhibited no estrus swelling, as
she had recently given birth. As she arrived, DV picked up
an Adansonia fruit he had placed on the ground and moved
away, with FA following. FA put her hand to his mouth
and touched the fruit, but DV turned away. FA then left his
immediate vicinity. Eighteen minutes later, DV left the
fruits to support FA in an agonistic encounter. FA returned
to retrieve DVs food while the agonism continued. DV
then returned and repeatedly displayed past FA, who
avoided, then approached DV and pant-grunted. DV

Primates

ultimately redirected aggression at adolescent male MI,

and FA continued feeding. Case U3: on 18 November
2010, juvenile female SO attempted to take bamboo stalks
from adult male BN. BN did not allow SO to take the
bamboo, and she responded by screaming. She then fed on
stalks BN had previously fed on, approximately 50 cm
from BN. Case U4: on 28 July 2011, infant male CY
attempted to take a bamboo stalk from adult female NI,
who carried it with her. NI presented sexually to CY, who
mounted NI but attempted to take the bamboo once again.
NI kept the stalk out of CYs reach.
Patterns of transfer behavior at Fongoli and other sites
Most food and tool transfers at Fongoli involved recovery
or passive tolerance of food and tool taking (Table 2). Of
the 39 scored transfers, most belonged to the recovery
(T2 = 28%) or passive (T3 = 55%) categories. Less than
13% of transfers involved taking food while the owner
protested (T1). The fewest exchanges were activepassive
(T4 = 1%) and active (T5 = 3%). No instances of giving
(T6) were recorded. Fruits were most often transferred
(51.0% of all events), while transfers of honey (7.0%),
flowers (5.0%), pith (5.0%), leaves (2.5%), insects (2.5%),
and soil (2.5%) were rare. The fruits with the largest
dimensions, Adansonia and Piliostigma, comprised 40% of
all exchanges and 76% of all fruit exchanges. Tools used to
fish for termites accounted for 17% of all transfers, while
galago-hunting tool (5%) and ant-dipping tool (2%)
transfers were rare. Most foods transferred were rare (61%)
and found in patches that were small or small to medium in
size (58%) relative to average chimpanzee party size.
Most transfers in this study were to females. Table 2
summarizes information on agesex classes, adult males
ranks, and females sexual swelling states for transfer
events. Adult females received the most resources (54% of
all transfers), and adult males were the owners in most
exchanges (54% of all transfers). Middle- and lowerranking-adult males, as well as immature males, most often
transferred resources to females. Only two transfers
involved males taking foods from females, and no transfers
between adult females were observed. The presence of a
sexual swelling was not associated with transfer, as female
recipients exhibited tumescence in only one-third of all
transfers with adult males. However, 78% of females
involved in transfers were cycling. Less than one-third of
all transfers were among males (N = 11). All rank classes
(high, middle, low) were represented among recipients and
owners in malemale transfer, but most transfers were from
higher-ranking owners to lower-ranking recipients (71% of
exchanges between adult males). Furthermore, high-ranking males transferred resources more often to middle- and
lower-ranking males than to adult females. In general,

dominant individuals transferred resources to subordinate

individuals (68% of cases).
Fongoli chimpanzees appear to transfer wild foods and
tools at a higher frequency than other chimpanzee communities studied thus far (Tables 1, 2, 3). However, because
methodologies were not standardized across studies in
regard to defining independent events and reporting kin
transfers, we were limited to describing general similarities
and differences among chimpanzee communities. Table 3
gives the number of these transfers according to the agesex
class of owners and receivers. Adult females at Fongoli
received more wild resources (54%) than adult females at
Budongo (29%) and Mahale (40%), but small sample sizes
at the latter study sites undermined our confidence in this
comparison. To remedy this problem, we included transfers
of provisioned sugar cane at Mahale (Nishida 1970) and
non-provisioned cultivars at Bossou (Hockings et al. 2007).
When transfers of cultivars and provisioned foods were
considered, most chimpanzee communities exhibited a
predominance of adult-male owners and adult-female
receivers (Fig. 1; Table 3). Fongoli and Mahale diverged in
adult-female and immature ownership, however (Table 3).
Immature owners were more than three times as common
at Fongoli compared to other sites, and adult-female
owners were relatively rare at Fongoli and Bossou. Hockings et al. (2007) define immatures in their Bossou study
as individuals less than 8 years old and do not explicitly
report transfer behavior in adolescents or subadults. It is
possible that older immatures were part of other transfer
categories (e.g., motheroffspring: Hockings et al. 2007)
excluded from our study, however. Our results on sexdifferences in ownership should be interpreted with caution
because Fongoli females were not focal subjects, and
female owners may therefore have been underrepresented
in our sample. We have greater confidence in our

Fig. 1 Percentage of transfers by adult owner-adult recipient at

Fongoli (present study), Mahale (Nishida 1970), and Bossou (Hockings et al. 2007), excluding femalefemale transfers (see text for
details). F female, M male

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Primates

observations of female recipients of male transfer at Fongoli because females are mostly tolerant of observers when
adult males are present.
Recipients at Fongoli were adult females twice as often
as adult males. Fongoli and Mahale transfer recipients were
similar in terms of agesex classes. At Bossou, adult males
rarely received food (Table 3) and did not take food from
females (Fig. 1), although the latter exchange was rare in
general at other sites. Relative to Bossou and Mahale, there
were more transfers between males and fewer transfers
from males to females among adults at Fongoli (Fig. 1).
This pattern held after combining wild-plant food and tool
transfers at Mahale (Nishida and Hiraiwa 1982; Nakamura
and Itoh 2001), and/or excluding all agonistic transfers
(Nishida 1970; present study).

Discussion
In the following discussion, we compare transfer behavior
among chimpanzee communities, and place these behaviors into an explanatory framework. However, we consider
this discussion to be preliminary in nature for many reasons. We were unable to compare transfer rates across sites
because study intervals were not clearly defined, or
descriptions did not include the quantity of transferred
resources. In addition, even though most studies excluded
motherinfant transfers, it was not possible to determine
the effect of kinship on food-transfer behavior, because
relatedness indices were not available. Future food-transfer
studies can benefit from standardizing methods and
assessing genetic relatedness among individuals.
Although wild-plant-food transfer is relatively rare,
Fongoli chimpanzees transfer food at frequencies previously unreported for chimpanzees. At Fongoli, foods that
were transferred had almost always been transported, but
they varied in size and availability relative to chimpanzee
party size. The chimpanzees most commonly transferred
wild-plant foods, but they also transferred tools, honey, and
soil. More than one-third of the plant foods transferred
were Adansonia or baobab, a relatively large fruit that
occurred in medium- to large-sized patches relative to
average chimpanzee party size. Similarly, wild and reintroduced golden lion tamarins (Leontopithecus rosalia)
frequently transferred large, hard-to-handle fruits, although
prey items were most often transferred (Ruiz-Miranda et al.
1999). The majority of foods transferred by Fongoli
chimpanzees were rare and found in patches that were
relatively small compared to average chimpanzee party
size, conditions predicted to provoke contest competition
(van Schaik 1989). The transfer of smaller foods seen at
Fongoli is uncommon elsewhere (Table 2), although

123

bonobos also engage in such sharing to some extent

(Kuroda 1984). Foods transferred at Fongoli were often
those that were harvested during the dry season (21 of 31
cases), which is also the period of greatest fruit abundance
(Pruetz 2006; Bogart and Pruetz 2011), and owners often
moved to consume them in shaded areas, where other
individuals subsequently took them.
In general, most descriptions of transfer behavior in wild
Pan can be classified as recovery or passive sharing (sensu
Boesch and Boesch 1989), and Fongoli chimpanzee food
transfer also fits this pattern, with females commonly taking food from males. Fongoli chimpanzees transfer plant
foods among most agesex classes, but the majority of
transferred resources were from dominant to subordinate
recipients. More than three-quarters of all transfers
between males and females followed this pattern. Although
some adult females are still timid in the presence of
observers, we recorded males transferring food or tools to
five of the seven to eight adult females in their community
during the present study. This suggests that a pattern of
male tolerance of female assertiveness characterizes Fongoli chimpanzees. To determine whether our findings were
typical of chimpanzees, we compared our results to those
of other studies describing wild-plant-food and tool transfer. Fongoli females took more resources from males than
Budongo and Mahale females (Table 3), but this trend
disappeared when we incorporated domesticated-plant
transfers at Mahale and Bossou (Fig. 1), highlighting the
common problem of small sample sizes for studies of food
transfer in wild primates. The latter comparison indicates
that male-to-female transfers are common in chimpanzees.
Explanations for the widespread pattern of male-tofemale food transfer in chimpanzees invoke female choice
as the mechanism that shapes such behavior, and ultimately
shapes male mating strategies. Male affiliation with
females may be a form of mating strategy (Gomes and
Boesch 2009), which could explain the transfer behavior at
Fongoli. Specifically, the relatively small number of
females in the Fongoli community may be an incentive for
males to tolerate female assertiveness in the form of foodand tool-taking, as the average adult male-to-adult female
sex ratio was 1.3:1 during the study period. This also fits
the prediction of the hypothesis that food transfer in callitrichids maintains social relationships within a group
(Ruiz-Miranda et al. 1999). On the other hand, reciprocal
exchange might explain male tolerance for females foodtaking behavior. Because females rarely shared with males,
strict reciprocity (food for food) can be ruled out. Although
tumescent and detumescent females were equally likely to
receive resources from males, and thus immediate mating
was not an incentive for at least half of these transfers, an
analysis of mating behavior over a longer time interval

Primates

(e.g., months) might be needed to detect associations

between food transfers and mate choice (Gomes and
Boesch 2009). Alternatively, relatively high rates of foodtaking by females from adult male orangutans (Pongo sp.)
prompted van Noordwijk and van Schaik (2009) to propose
that females were testing for male coercive tendency. Few
cases of unsuccessful or protested transfer occurred in the
Fongoli sample, however, indicating this explanation is
unlikely. Finally, female assertiveness may place more
pressure on Fongoli males to give up valued resources.
Gilby (2006) found that meat-sharing by males at Gombe
could be explained by this sharing-under-pressure
hypothesis, an explanation that may initially seem likely at
Fongoli, but extensive begging did not characterize the
food- and tool-taking behavior seen in our study relative to
that characterizing meat-sharing at Fongoli (Pruetz,
unpublished data).
Hockings et al. (2007) invoke the costly-signaling
hypothesis to explain the transfer of mainly cultivars at
Bossou. Although relinquishing food to another individual
results in caloric loss, costly signaling involves additional
risks to the owner, such as possessing a high-value and rare
food (Bossou: Hockings et al. 2007) or losing energy from
an unsuccessful hunt (Gilby and Wrangham 2007). Relative to crop raiding or chasing down prey, the risk of
harvesting wild foods is probably lower at Fongoli relative
to contexts in which the costly-signaling hypothesis has
been supported for chimpanzees. Moreover, adult females
and immatures were equally likely to forage for the same
plant foods that were transferred at Fongoli, unlike findings
for cultivars (Hockings et al. 2007) and meat (Stanford
1998; Boesch and Boesch-Achermann 2000). Therefore,
the costly-signaling hypothesis is an unlikely explanation
of food transfer at Fongoli.
A number of hypotheses proposed to explain food
transfer in primates invoke specific characteristics of the
items shared. The rare-item and foraging-assistance
hypotheses posed to explain callitrichid food transfer
involve rare and difficult-to-process foods, respectively
(Price and Feistner 1993; Ruiz-Miranda et al. 1999). Fongoli data support these hypotheses to some degree, with the
transfer of Adansonia fruit being mainly responsible for
this pattern. However, the hypothesis that food transfer is a
means of teaching young individuals about new food items
is not supported by the Fongoli data. While adults of both
sexes in callitrichids commonly transfer foods to immatures (Price and Feistner 1993), in terms of transfer outside
of the motheroffspring pair, relatively few cases at Fongoli involved adult chimpanzees transferring to immature
individuals.
Given our small sample size, we cannot yet adequately
test the various hypotheses put forth to explain the cause of
food and tool transfer behavior at Fongoli, although a

pattern emerges suggesting that male chimpanzees at

Fongoli share mostly rare, hard-to-process fruits, as well as
tools, with subordinates, mainly females. We can, then,
look to the larger social system in attempting to put the
tolerance and transfer behavior observed at Fongoli in
context. In general, patterns of transfer at Fongoli fit
expectations for this relatively cohesive community.
Chimpanzees at Fongoli range in large, mixed parties,
making this community more cohesive than those studied
elsewhere (Pruetz and Bertolani 2009). Average party size
is 15 individuals, so that more than 40% of the community
is together at any one time (Pruetz and Bertolani 2009).
Such cohesiveness has been attributed to the combined
effects of savanna resource availability, a relatively large
home range ([85 km2; Pruetz and Wessling, unpublished
data), and chimpanzee sociality (Pruetz and Bertolani
2009). Relatively high tolerance for food-taking behavior
could therefore be attributed to the maintenance of affiliative and cohesive social relationships. In addition to
malefemale transfer, Fongoli adult males transferred plant
foods among each other more often than Mahale and
Bossou males (Fig. 1), indicating that Fongoli males are
more tolerant of each other while feeding. Although
motivations for malemale transfer can differ from motivations for malefemale transfer (Mitani and Watts 2001),
the small sample of malemale transfers limits our ability
to explore these differences. However, West African
chimpanzees at Tai Forest, Ivory Coast, typically range in
large, mixed malefemale parties, which have been
described as bisexually-bonded (Lehmann and Boesch
2005). West African chimpanzee societies are also somewhat reminiscent of bonobo societies (Tai: Lehmann and
Boesch 2005, 2008; Bossou: Yamakoshi 2004; and see
Doran et al. 2002), where the social unit is more cohesive
(Boesch 2009), females are dominant to males in many
situations (Parish 1994), and wild plant-foods are transferred among nonrelatives (Kano 1992). Such similarities
warrant investigation of the ultimate causes of this convergent behavior. To some degree, our findings, therefore,
support the hypothesis that West African chimpanzees are
more similar to bonobos than the East African subspecies
in certain aspects of their social behavior (Yamakoshi
2004). However, we also explore the possibility that the
savanna environment at Fongoli contributes to the pattern
of food transfer.
As the only habituated community of chimpanzees living in a savanna environment, the Fongoli community
provides detailed information on the effect of ecological
variables on social behavior and organization in this great
ape under selective pressures associated with an open, dry,
and hot environment. Alternatively, the tendency for
habitual food transfer and high social cohesion at Fongoli
may be explained by savanna living. Savanna chimpanzees

123

Primates

at Fongoli exhibit significant differences in behavior

compared to chimpanzees studied in forested habitats.
These include habitually using tools to hunt (Pruetz and
Bertolani 2007), using caves as shelter (Pruetz 2007), more
cohesive grouping patterns (Pruetz and Bertolani 2009),
and relying heavily on insect prey (Pruetz 2006; Bogart and
Pruetz 2008, 2011). Each of these differences can be linked
to the environment at the site. Possible sources of a
relatively high degree of tolerance at Fongoli include the
relative costs and benefits of food transfer (sensu Stevens
and Gilby 2004) related to the savanna habitat at Fongoli or
a general propensity for affiliation that has been genetically
or socially inherited. The tendency for Fongoli chimpanzees to transport foods to shaded areas for consumption
during the dry season likely increases the value of these
foods. Rather than abandon a shady area to forage, individuals may occasionally attempt to take transported foods
from owners. This suggests that the environment at Fongoli
may contribute to the relatively high degree of wild-plant
food transfer.
Finally, in consideration of recent studies noting the
propensity for chimpanzee plant-food sharing when conditions allow (Hockings et al. 2007), we propose that the
focus on meat-sharing among wild chimpanzees should be
broadened to include possible plant-food transfer, as well
as females supplanting males at food sites. For example, at
Fongoli, we have observed 11 instances in which adult
females supplanted adult (N = 9) or subadult males
(N = 2) at termite-fishing sites and one case in which the
alpha male shared his termite-fishing site with a female
infant, even though these same males have been observed
to dominate these females in most contexts (Pruetz,
unpublished data). Observations of malefemale foodsharing in captive chimpanzees (S. Lambeth, personal
communication) and a recent experimental study of sharing
behavior in Pan paniscus and P. troglodytes, Jaeggi et al.
(2010), showing that captive chimpanzees shared more
frequently and were more egalitarian relative to bonobos,
support our suggestion that observers of wild chimpanzees
place renewed focus on this behavior.
Acknowledgments We thank the Republic of Senegal, Department
of Eaux et Forets and the Arrondissement du Bandafassi for permission to work in Senegal. Funding was provided by Iowa State
University, Great Ape Trust of Iowa, National Science Foundation,
National Geographic Society, Leakey Foundation, Primate Conservation Inc., American Society of Primatologists, Wenner-Gren
Foundation, and the U.S. Fish and Wildlife Society. We thank
K. Boyer, J. Marshack, P. Bertolani, M. Gaspersic, S. Bogart,
E. Wessling, D. Kante, M. Sahdjakho, and M. Camara for assistance
in the field. T.C. LaDuke, K. Klag, K. Walkup, and E. OtarolaCastillo made helpful comments on the manuscript. We thank
Dr. M. Nakamura and one anonymous reviewer and would like to
especially acknowledge the contribution of the late Dr. T. Nishida in
significantly improving this manuscript.

123

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