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

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American Journal of Primatology 70:402–409 (2008)
RESEARCH ARTICLE
Chimpanzees as Fauna: Comparisons of Sympatric Large Mammals
Across Long-Term Study Sites
SAMANTHA M. RUSSAK1,2 AND W.C. MCGREW1,3
1
Departments of Anthropology and Zoology, Miami University, Oxford, Ohio
2
School of Human Evolution and Social Change, Arizona State University, Tempe, Arizona
3
Leverhulme Centre for Human Evolutionary Studies, Department of Biological Anthropology, University of Cambridge,
Cambridge, United Kingdom
Although much research has shown otherwise, chimpanzees are still often classed as rainforestdwellers. Most long-term studies of wild chimpanzees (Pan troglodytes) are not situated in evergreen,
closed-canopy equatorial forests, but instead are conducted in more open habitats. This study aims to
elucidate the extent of chimpanzee ecological diversity by scrutinizing (recently) sympatric mammalian
fauna at established study sites. We compiled presence or absence data on large mammal species at
eight sites: Assirik, Bossou, Budongo, Gombe, Kibale, Lopé, Mahale, and Tai. The sites were rank
ordered on the most basic ecological variable: annual total rainfall. Only three of the 65 mammalian
genera compiled were sympatric with chimpanzees at all sites: Potamochoerus (bushpig), Syncerus
(buffalo), and Panthera pardus (leopard). Some subfamilies (e.g. colobines) were present at most sites,
but some families (e.g. hyenids) were absent at most sites. Some taxa (e.g. suids, cercopithecines)
correlated better than others (e.g. canids) with basic ecological variables. The most extreme chimpanzee
study site for which data are available is Assirik, Senegal. Nowhere else are chimpanzees sympatric
with Erythrocebus, Alcelaphus, Hippotragus, and Ourebia. As chimpanzees are often behavioral models
for extinct hominins, these living faunal assemblages have implications for paleo-ecological
c 2008 Wiley-Liss, Inc.
reconstructions of ancestral habitats. Am. J. Primatol. 70:402–409, 2008.
Key words: Pan troglodytes; faunal assemblage; biogeography; chimpanzee ecology
INTRODUCTION
Despite decades of field data to the contrary,
wild chimpanzees (Pan troglodytes) are still characterized as rain forest-dwellers in contemporary
textbooks [e.g. Relethford, 2005; p 279], and secondary reviews [e.g. Begun, 2004; p 20]. However, most
long-term studies of wild chimpanzees are not
situated in evergreen, closed-canopy equatorial forests, but instead take place in more open habitats.
Chimpanzees are found in 21 countries from far west
to east Africa, from Senegal to Tanzania, and over
201 of latitude. Sites differ greatly in annual rainfall,
lengths of dry season, and vegetation, ranging from
dry, open savanna to wet, closed rainforest to mosaic
habitats.
The biogeographical distribution of faunal communities is related to variation in climate, vegetation, and other ecological variables, with organisms
being adapted for specific environments [Fashing &
Gathua, 2004; Meiri et al., 2005; Owen-Smith, 1999].
For example, wildebeest and zebra are only found in
open grassland habitats, whereas mammals like dikdik and bushbuck are typically found in areas with
patchy vegetation and a woody overstory [McNaughton & Georgiadis, 1986]. This relationship between
r 2008 Wiley-Liss, Inc.
faunal diversity and environment of ecological
adaptation can sometimes be complicated by taxonomy; some taxon variants are distinguished on a
subspecific level (e.g. forest versus savanna buffalo,
Syncerus spp.), whereas others on a specific level (e.g.
forest versus savanna elephant, Loxodonta cyclotis,
L. africana), making levels of similarity unclear [de
Vivo & Carmignoto, 2004].
Given the strong ecological correlations noted
above, the mammalian fauna sympatric with chimpanzees should vary greatly across sites, but this
has never been elucidated. Therefore, this study
examines the extent of chimpanzee ecological diversity by scrutinizing (recently) sympatric mammalian
fauna at six established study sites. The resulting
Contract grant sponsor: Miami University Honors & Scholars
Program.
Correspondence to: Samantha M. Russak, School of Human
Evolution and Social Change, Arizona State University, P.O. Box
872402, Tempe, AZ 85287-2402.
E-mail: Samantha.Russak@asu.edu
Received 28 June 2005; revised 31 October 2007; revision
accepted 1 November 2007
DOI 10.1002/ajp.20506
Published online 1 February 2008 in Wiley InterScience (www.
interscience.wiley.com).
Chimpanzee Mammalian Faunal Communities / 403
description of the range of faunal variation at
chimpanzee study sites is an important means for
determining the environmental variation in hominin
paleo-habitats.
METHODS
Site Criteria
The following criteria were used to pick the
chimpanzee study sites: First, they ranged in location across Africa, from east to far west (Fig. 1),
covering the three recognized subspecies of
P. troglodytes. Second, we chose sites with a wide
range in mean annual rainfall, seeking the species’
limits of varying habitats, vegetation, and so,
sympatric mammalian fauna (Spearman’s rank
correlation between mean annual rainfall and dry
season length: r 5 0.83, P 5 0.05). Third, we used
sites of long-term study (i.e. four or more annual
cycles) to maximize chances of comprehensive coverage of species present. The eight sites in the study of
Whiten et al. [2001] met all of these requirements
and so were the starting point for this study.
Published lists of species of mammals found at
each site are usually limited to data on presence or
absence, and records of abundance, density, and
distribution are usually lacking. The data set
includes species of mammals seen at each chimpanzee site at any point since 1960 (the year when the
oldest field study began), but owing to increasing
environmental pressures, especially human encroachment and predation, some of these species
are now absent. The lists are inclusive for large
mammals found within the chimpanzees’ ranges, but
also include smaller mammals known to be preyed
upon by chimpanzees. Therefore, all taxa in the data
set are those that interact (directly or indirectly)
with chimpanzees. Rodents, bats, insectivores, and
elephant shrews were excluded from consideration as
not often being recorded during censuses by diurnal
primatologists.
Two sites were dropped from the analysis: Lopé
is an anomalous mosaic of rain forest and anthropogenic savanna; chimpanzees cross ‘‘empty,’’ open
grasslands from one forest fragment to another
[Tutin et al., 1997b]. Bossou is extraordinarily
depauperate (c.f. ‘‘empty forest syndrome’’) because
of its unprotected status and high-density human
population. In addition, parts of the Mahale Mountains National Park outside the chimpanzees’ range
have mammalian taxa that are not included here.
Although comparisons of taxa on the level of species
would reveal characteristics particular to a very
specific habitat, this study compares taxa on a
generic level to enable broader comparisons across
study sites so that the inherent differences in species
exhibit due to environmental conditions do not
constrain the analysis of this study.
Site Characteristics
Assirik
Mt. Assirik, in the Niokolo-Koba National Park
in southeastern Senegal, is the driest study site for
long-term studies of chimpanzees [McGrew et al.,
1981] (Table I). The area is predominantly grassland,
but also has deciduous woodland, bamboo thicket,
and gallery forest, providing a hot, dry, open savanna
habitat. Evergreen forest comprises only about 3% of
vegetation at the study site, but it is disproportionately used by chimpanzees for foraging and nesting.
This landscape resembles some stages of African
Plio-Pleistocene, and so the fauna present could
serve as important paleoecological indicators
[McGrew, 1992]. Mt. Assirik provides the most
extensive data set for this study, collected by the
Fig. 1. Locations of long-term study sites of chimpanzees in Africa [from Whiten et al., 2001].
Am. J. Primatol.
404 / Russak and McGrew
TABLE I. Sites of Long-term Study of Wild Chimpanzees [see Whiten et al., 2001]
Dominant vegetation
types
Country
coordinatesc
7 monthsa
Grassland, woodland,
bamboo thicket, gallery
foresta
Senegal
121530 N,
121460 W
SAPP 1976–1979; MAPP
2000l
1531b
6 monthse
Closed canopy forest,
Marantaceae
understory, grasslandb
Gabon
01100 S, 111350 E
Henschel, personal
communication; Tutin et al.,
p. 1192e
Gombef
1600b
6 monthsf
Evergreen riverine forest,
deciduous dry forest,
thicket, grassland,
moorlandb
Tanzania
41400 S, 291380 E
Goodall, p. 49–50f
Kibaleg
1671b
5 monthsg
Moist evergreen forest,
colonizing forestgrassland, swampb
Uganda
01320 N, 301230 E
Ghiglieri, p. 13–14m;
Struhsaker, p. 116n
Budongoh
1684a
3 monthsp
Semi-deciduous moist
rain forestb
Uganda
11440 N, 311330 E
Reynolds, p. 89–129, 251–253o
Mahalej
1774j
5–6 monthsj
Semi-evergreen forest,
miombo woodlandb
Tanzania
61070 S, 291440 E
Nishida, p. 18–19j
Taii
1800i
3 monthsi
Evergreen moist
rainforestb
Ivory Coast
51520 N, 71200 W
Boesch and BoeschAchermann, p. 280–281i
Bossouk
2230a
5 monthsq
Primary forest patches,
secondary forest,
cultivated fieldb
Guinea
71390 N, 81300 W
Humle, personal
communication
Mean annual
rainfall (mm)a
Dry season
954d
Lopee
Sites
Assirikd
Source of mammal data
a
Hunt and McGrew [2002]; bAppendix, McGrew et al. [1996]; cSarringhaus, unpubl. data; dMcGrew et al. [1981]; eTutin et al. [1997b]; fGoodall [1986];
Struhsaker [1997]; hReynolds [2005]; iBoesch and Boesch-Achermann [2000]; jNishida [1990]; kMatsuzawa [2002]; lSee site characteristics; mGhiglieri
[1984]; nStruhsaker [1975]; oReynolds [1965]; pNewton-Fisher [1999]; qTakemoto [2004].
g
Stirling African Primate Project February 1976–
December 1979 and the Miami Assirik Pan Project
February–April 2000.
Lope´
The Lopé Reserve is in central Gabon in the
Congo Basin [Tutin et al., 1997a]. The area is
characterized by evergreen forest with a dense
understory dominated by Marantaceae, and by
fire-maintained, anthropogenic savanna grassland
[White et al., 1995]. Although the amount of rainfall
is relatively low for a wet tropical rain forest,
constant cloud cover over the area causes high
humidity and moisture retention. Research on
comparative ecology of sympatric apes has been done
there since 1984, but the chimpanzees are not fully
habituated [Tutin et al., 1991].
Gombe
Gombe National Park is a small national park
(5200 ha) in western Tanzania, on the eastern shore
Am. J. Primatol.
of Lake Tanganyika [Goodall, 1986]. The mean
annual rainfall varies considerably. Gombe is a
mosaic of habitats [Collins & McGrew, 1988], but
the research area has become isolated, as the
surrounding areas have been cleared and cultivated.
In addition, several species of mammal, such as
hippopotamus and buffalo, have been hunted to
extinction since research began at Gombe in 1960.
Kibale
Kibale National Park is in western Uganda, East
Africa, and has a mosaic of habitats, including
evergreen forest, forest-grassland, and swamp communities [Struhsaker, 1997]. This diversity in vegetation types supports an array of mammalian
species. These species were threatened by intensive
logging in earlier years and continue to be endangered by human encroachment. There are two main
chimpanzee study sites at Kibale, Kanyawara
[Wrangham et al., 1991], and Ngogo [Mitani et al.,
2000], but they have similar habitats (in terms of
Chimpanzee Mammalian Faunal Communities / 405
TABLE II. Artiodactyla, Perissodactyla, and Proboscidea Sympatric with Chimpanzees at Six Long-Term
Study Sites
Genus
Alcelaphus
Ourebia
Hippotragus
Taurotragus
Tragelaphus
Potamochoerus
Syncerus
Loxodonta
Cephalophus
Phacochoerus
Hippopotamus
Sylvicapra
Kobus
Hylochoerus
Hyemoschus
Neotragus
Boocerus
Choeropsis
Total
Common name
Assirik
Hartebeest
Oribi
Roan antelope
Eland
Bush-buck
Bushpig
Buffalo
Elephant
Duiker
Warthog
Hippopotamus
Bush duiker
Waterbuck
Giant forest hog
Chevrotain
Suni/royal antelope
Bongo
Pygmy hippopotamus
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
10
Gombe
Kibale
Budongo
Mahale
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
5
9
Tai
Y
Y
Y
Y
Y
Y
Y
Y
Y
7
7
9
Study sites listed left to right in order of increasing mean annual rainfall.
both flora and fauna) and rainfall, so for this study
they have been combined [Ghiglieri, 1984].
Budongo
Also in western Uganda is the Budongo forest
[Reynolds, 1965, 2005]. Budongo is characterized by
semi-deciduous tropical rain forest. This area has
undergone intensive logging over decades but still
supports a variety of fauna [Plumptre & Reynolds,
1994]. Although extensive research has been done in
Budongo, there has been no published comprehensive list of large mammals; the data used in this
study are taken from Reynolds’ [1965, 2005] study.
Taı¨
Taı̈ National Park is located in southeastern
Ivory Coast, West Africa [Boesch & Boesch-Achermann, 2000]. It is the largest remaining block of
moist evergreen forest in West Africa, and is home to
many endemic and endangered species, such as the
pygmy hippopotamus. Several neighboring communities of chimpanzees have been studied [Herbinger
et al., 2001]. This area has remained little disturbed
by logging, but poaching has wiped out a number of
animals in the park including buffalo [Boesch
& Boesch-Achermann, 2000].
Mahale
Mahale Mountains National Park is located in
western Tanzania [Nishida, 1990]. The mountains
run in a straight line from north-west by west to
south-east by east, creating two different types of
habitats on either side of the mountain range. The
west side of the mountains (where most of the
chimpanzee research occurs) is wetter, being characterized as tropical semi-evergreen forest with
biogeographic links to the Congo basin. A rain
shadow from the mountains on the east side causes
it to be drier, with miombo woodland and open
savanna [Itani, 1979]. This study concentrates on the
mammals found in areas occupied by chimpanzees
studied long term [Uehara & Ihobe, 1998]. It is the
only study of chimpanzees where sympatric mammals have been monitored [Uehara, 2003].
Bossou
Bossou is located in southeastern Guinea, West
Africa [Matsuzawa, 2002; Sugiyama, 2004]. It is the
wettest of the eight sites used in this study and is
characterized by primary and secondary forest,
cultivated field, and savanna [Sugiyama & Koman,
1992]. The area is unprotected and human population density is high, especially after several influxes
of refugees from wars in neighboring countries. The
relict chimpanzee community at Bossou is small and
isolated, and the abundance of other species of
mammals is low, mainly because of hunting and
snaring.
RESULTS
Table II shows sympatry between chimpanzees
and three orders of large mammals: Artiodactyla,
Am. J. Primatol.
406 / Russak and McGrew
TABLE III. Carnivora Sympatric with Chimpanzees at Six Long-Term Study Sites
Genus
Lycaon
Herpestes
Atilax
Canis
Crocuta
Panthera leo
Viverra
Mungos
Felis
Genetta
Panthera pardus
Aonyx
Mellivora
Nandinia
Crossarchus
Bdeogale
Lutra
Ichneumia
Civettictis
Ictonyx
Liberiictis
Poiana
Total
Common name
Assirik
Gombe
Wild dog
Mongoose
Marsh mongoose
Jackal
Spotted hyena
Lion
Large civet
Banded mongoose
Small cat
Genet
Leopard
Clawless otter
Honey badger
Palm civet
Cusimanse
Black-footed mongoose
River otter
White-tailed mongoose
African civet
Zorilla
Liberian mongoose
Linsang
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Kibale
Budongo
Mahale
Tai
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
11
13
16
3
13
12
Study sites listed left to right in order of increasing mean annual rainfall.
Perissodactyla, and Proboscidea. These 18 genera
range from monotypic taxa, e.g. Phacochoerus
aethiopicus, Boocerus euryceros, to speciose radiations, e.g. Cephalophus, 12 spp., Tragelaphus, 4 spp
(common names for taxa listed here and below can be
found in Tables II–V). Only two genera, Potamochoerus and Syncerus, are found at all six study sites,
although three other genera, Tragelaphus, Loxodonta, and Cephalophus, lack only one site each of being
omnipresent. In contrast, half of the genera (nine of
18) occur at only one site. Of these nine genera, eight
are found at either of the two habitat extremes;
Alcelaphus, Ourebia, Hippotragus, and Taurotragus
at the dry, open habitat of Assirik and Hyemoschus,
Neotragus, Boocerus, and Choeropsis, at the wet,
closed site of Tai. Of the sites, Assirik has the most
sympatric genera, and Gombe the least.
Table III shows sympatry between chimpanzees
and 21 genera of carnivores; 15 of these genera are
monotypic, e.g. Lycaon pictus, Mellivora capensis
(note that for this study one genus, Panthera, has
been split into two, P. leo and P. panthera, given the
marked morphological, ethological, and ecological
differences between the two species). Only one taxon,
the leopard, is sympatric with the chimpanzee at all
six sites, but several other small-bodied nocturnal
carnivores (Felis, Genetta, Viverra) are found at all
but one site. Only four of the 21 taxa are found at
only one study site, and only two of these occur at the
Am. J. Primatol.
extreme sites, being Liberiictis and Poiana at Tai.
Kibale has more than five times as many carnivore
genera than does Budongo, despite their similar
annual rainfalls and geographical proximity.
Table IV shows there to be 13 genera of nonhuman primates that are sympatric with chimpanzees at the six sites. No genus is omnipresent, but
two, Cercopithecus and Papio, are found at five;
the exceptions are at the extremes: Assirik seems to
be too dry for arboreal guenons and Tai seems to be
too wet for savanna baboons. Four of the 13 genera
are sympatric with apes at only one of the six sites:
Erythrocebus only at Assirik; Lophocebus only at
Kibale; Otolemur only at Mahale; Cercocebus only at
Tai. Once again, Kibale has more than twice as many
genera than does Budongo.
Table V shows the sympatry of the four
remaining orders (Hyracoidea, Lagomorpha, Pholidota, Tubulidentata) with chimpanzees. None comes
even close to being omnipresent, but four sites had at
least one genus of pangolin (Manidae) or one of the
two genera of hyraxes (Procaviidae). Only Assirik,
with its open grasslands, had a lagomorph, Lepus.
The paucity of taxa from these orders precludes any
meaningful comparison across sites.
Of the 65 mammalian taxa found at the six study
sites, only three genera Potamochoerus, Syncerus,
and Panthera pardus, were present at all study sites.
Furthermore, the mammalian fauna assemblages
Chimpanzee Mammalian Faunal Communities / 407
TABLE IV. Primates Sympatric with Chimpanzees at Six Long-Term Study Sites
Genus
Erythrocebus
Perodicticus
Galago
Chlorocebus
Papio
Procolobus
Cercopithecus
Colobus
Euoticus
Galagoides
Lophocebus
Otolemur
Cercocebus
Common name
Assirik
Patas monkey
Potto
Lesser bushbaby
Vervet
Baboon
Colobus
Guenon
Black-and-white colobus
Needle-clawed galago
Dwarf bushbaby
Grey-cheeked mangabey
Greater bushbaby
Sooty mangabey
Y
Y
Y
Y
Y
Total
Gombe
Kibale
Budongo
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Mahale
Tai
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
5
6
9
4
7
6
Study sites listed left to right in order of increasing mean annual rainfall.
TABLE V. Hyracoidea, Lagomorpha, Pholidota, and Tubulidentata Sympatric with Chimpanzees at Six Longterm Study Sites
Genus
Lepus
Smutsia
Dendrohyrax
Phataginus
Orycteropus
Heterohyrax
Uromanis
Total
Common name
Assirik
Hare
Giant pangolin
Tree hyrax
Tree pangolin
Aardvark
Hyrax
Long-tailed pangolin
Y
Gombe
Kibale
Budongo
Mahale
Tai
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
1
1
3
1
4
4
Study sites listed left to right in order of increasing mean annual rainfall.
that are sympatric with chimpanzees are specific to
each study site and vary widely. Kibale has the most
of the 65 genera, at N 5 37, but Budongo has only 15,
whereas the other four fall in a narrower range of
25–31.
DISCUSSION
Interpreting these data is difficult because it
seems likely that not all data sets on mammals, even
at the crudest level of presence/absence are complete,
or even of comparable status (i.e. collected in the
same manner or for the same purpose). It is unlikely
that Kibale really has 2.5 times the number of
mammalian taxa as Budongo, given that the two
sites are near neighbors in western Uganda, and
have similar rainfall and vegetation. Instead, it
seems likely that the differences reflect differences
in methods of data collection across sites, e.g.
researchers who can follow apes from nest to nest
are active before sunrise and after nightfall, and so
are more likely to meet nocturnal taxa. However,
cross-site differences cannot be explained by length
of study: the site of the shortest study (Assirik, 5
years) has 27 genera, whereas the site of the longest
study (Gombe, 45 years) has 25.
If one limits comparison to large-bodied or
diurnal species, the contrasts still hold, as shown in
Tables II–IV. Hartebeest, oribi, roan antelope, and
eland are five savanna forms of ungulate, just as
chevrotain, dwarf antelopes, bongo, and pygmy
hippopotamus are evergreen closed forest taxa. In
between are a variety of bovids and suids that show
much greater habitat flexibility, including two of the
three omnipresent forms, bushpig and buffalo. In
this light, elephants and duikers (Cephalophus, spp.)
are notably absent from Gombe, but they ‘‘should’’
be there according to the ecology of the site. Gombe
is by far the smallest of the study sites, so it is not
surprising that elephants have been extinguished
there, but the absence of any Cephalophus is
surprising.
Am. J. Primatol.
408 / Russak and McGrew
Among carnivores, the small-bodied felids,
herpestids, mustelids, and viverrids are mostly
nocturnal and so easily missed by diurnal researchers. The remaining four large-bodied taxa (Lycaon,
Canis, Crocuta, Panthera) are obvious by sight,
sound, or spoor, yet their pattern of presence/
absence is confusing. Why do Assirik, Kibale, and
Mahale have Canis, but not Gombe or Budongo? (Tai
is not within the species distribution of wild Canis.)
The same question can be asked of the inconsistent
presence/absence of Crocuta. Perhaps these species
were hunted to extinction by humans; to fully
answer these questions would require a more
detailed study of the faunal and environmental
history of each site. Again, Gombe looks depauperate, lacking jackal, hyena, and lion. Only the
leopard, the carnivore species with the world’s
largest distribution, is a constant companion of the
chimpanzee among the range of carnivores.
Among the primates, the distribution of nocturnal prosimians is highly problematic; none of the six
chimpanzee study sites has been subject to intensive
study of these taxa. However, differences between
the suites of diurnal monkeys (note that all six sites
lack gorillas) are presumed to be accurate, given that
they are sought-after prey or competitors for
chimpanzees. Assirik seems to be too dry and open
to support arboreal guenons and the gallery forest
too sparse to support colobines. Tai is too wet and
closed for Papio and Chlorocebus. The two Tanzanian sites lack mangabeys altogether, though they
are found elsewhere in the country, and Gombe also
lacks any form of black and white colobus, which is
just as puzzling as Budongo’s lack of red colobus. The
stochastic vagaries of what taxa were left behind in
Pleistocene refugia during dry periods, plus the fact
that all four East African sites are on or near the
Great Rift, may account for some variation [see
Gibernau & Montuire, 1996].
Overall, the distribution patterns of the mammals at the six chimpanzee study sites are consistent
with gross biogeographical patterns for mammals
[e.g. Badgley & Fox, 2000; McNaughton & Georgiadis, 1986; Owen-Smith, 1999]. For example, herding
bovids (and the group-organized hunters that prey
upon them) require vast open spaces [de Vivo &
Carmignoto, 2004]. Three of the four genera from
Table II (Alcelaphus, Hippotragus, and Taurotragus)
that occur only at Assirik, the driest and most open
of the chimpanzee sites, are the only herding
artiodactyls of the sample, excluding the omnipresent Syncerus. The only social predators in the
sample, wild dog, hyena, and lion were also most
commonly found at the more open sites of Assirik,
Gombe, and Mahale.
de Vivo and Carmignoto [2004] showed that
large body size is highly correlated with open
landscapes. Savanna taxa are generally larger than
their rain forest counterparts. Duikers (Cephalophinae)
Am. J. Primatol.
occur at most sites, but species vary in size; the
smallest species, the blue duiker (Cephalophus
monticola) occurs at more closed sites like Tai,
Budongo, and Kibale while one of the larger species,
the red-flanked duiker (Cephalophus rufilatus) is
only found at Assirik among the six sites. When
taken to the generic level, as in this study, the
ecological correlation with body size can be lost.
Despite being classed as a rainforest-typical
species in so many secondary sources, chimpanzees
inhabit a wide range of ecotypes, most of which are
mosaics of vegetation types. These differences in
chimpanzee habitat extend to their sympatric faunal
communities. The low overlap of taxa between sites
reflects the large diversity of mammalian communities that co-occur with these apes. More studies
need to be done of P. troglodytes at sites with annual
rainfalls of below 1,500 mm or above 2,000 mm (e.g.
Fongoli, Semliki, Ugalla, or Goualougo, Kalinzu,
respectively). No single site is typical of the species
P. troglodytes; even the site richest in mammalian
diversity, Kibale, has only 57% (37 of 65) of the
genera that co-occur with chimpanzees.
Thus, chimpanzees should be considered ecological generalists. This underappreciated quality
brings them closer to humans and provides support
for their use as behavioral models for extinct
hominins. However, our results suggest that if one
seeks to compare extinct and extant faunal communities, some chimpanzee populations might be better
ecological models than others for any particular
hominin site. The apparent correlation found between fauna and one ecological variable, rainfall, has
implications for the paleo-ecological reconstruction
of these extinct sites.
ACKNOWLEDGMENTS
We thank Philipp Henschel, Tatyana Humle,
Nick Newton-Fisher, Toshisada Nishida, Caroline
Tutin, and Richard Wrangham for help with the
mammal lists; the Miami University Honors
& Scholars Program Fund for Student Initiative for
financial support.
REFERENCES
Badgley C, Fox DL. 2000. Ecological biogeography of North
American mammals: species density and ecological structure in relation to environmental gradients. J Biogeog
27:1437–1467.
Begun DR. 2004. Enhanced cognitive capacity as a contingent
fact of hominid phylogeny. In: Russon AE, Begun DR,
editors. The evolution of thought: evolutionary origins of
great ape intelligence. Cambridge: Cambridge University
Press. p 15–27.
Boesch C, Boesch-Achermann H. 2000. The chimpanzees of
the Taı̈ Forest: behavioural ecology and evolution. Oxford:
Oxford University Press. 316p.
Collins DA, McGrew WC. 1988. Habitats of three groups of
chimpanzees (Pan troglodytes) in western Tanzania compared. J Hum Evol 17:553–574.
Chimpanzee Mammalian Faunal Communities / 409
Fashing PJ, Gathua JM. 2004. Spatial variability in the
vegetation structure and composition of an East African
rain forest. Afr J Ecol 42:189–197.
Ghiglieri MP. 1984. The chimpanzees of Kibale forest. A field
study of ecology and social structure. New York: Columbia
University Press. 226p.
Gibernau M, Montuire S. 1996. Mammal diversity and
environment evolution during the Plio-Pleistocene in East
Africa. Hum Evol 11:193–204.
Goodall J. 1986. The chimpanzees of Gombe: patterns of
behavior. Cambridge, MA: Harvard University Press. 673p.
Herbinger I, Boesch C, Rothe H. 2001. Territory characteristics among three neighboring chimpanzee communities in
the Tai National Park, Cote d’Ivoire. Int J Primatol
22:143–167.
Hunt KD, McGrew WC. 2002. Chimpanzees in the dry habitats
of Assirik, Senegal and Semliki Wildlife Reserve, Uganda.
In: Boesch C, Hohmann G, Marchant LF, editors. Behavioural diversity in chimpanzees and bonobos. Cambridge:
Cambridge University Press. p 35–51.
Itani J. 1979. Distribution and adaptation of chimpanzees
in an arid area. In: Hamburg DA, McCown ER, editors.
The great apes. Menlo Park, CA: Benjamin/Cummings.
p 55–71.
Matsuzawa T, editor. 2002. Chimpanzees of Bossou and Nimba
1976–2001: collection of articles in English. Kyoto: Primate
Research Institute. 646p.
McGrew WC. 1992. Chimpanzee material culture: implications
for human evolution. Cambridge: Cambridge University
Press. 277p.
McGrew WC, Baldwin PJ, Tutin CEG. 1981. Chimpanzees in a
hot, dry and open habitat: Mt. Assirik, Senegal, West Africa.
J Hum Evol 10:227–244.
McGrew WC, Marchant LF, Nishida T, editors. 1996. Great
ape societies. Cambridge: Cambridge University Press.
328p.
McNaughton SJ, Georgiadis NJ. 1986. Ecology of African
grazing and browsing mammals. Ann Rev Ecol Systemat
17:39–65.
Meiri S, Dayan T, Simberloff D. 2005. Biogeographical
patterns in the Western Palearctic: the fasting-endurance
hypothesis and the status of Murphy’s rule. J Biogeog
32:369–375.
Mitani JC, Struhsaker TT, Lwanga JS. 2000. Primate
community dynamics in old growth forests over 23.5 years
at Ngogo, Kibale National Park, Uganda: implications for
conservation and census methods. Int J Primatol
21:269–286.
Newton-Fisher NE. 1999. The diet of chimpanzees in the
Budongo Forest Reserve, Uganda. Afr J Ecol 37:344–354.
Nishida T, editor. 1990. The Chimpanzees of the Mahale
Mountains: sexual and life history strategies. Tokyo:
University of Tokyo Press. 328p.
Owen-Smith N. 1999. Ecological links between African
savanna environments, climate change, and early hominid
evolution. In: Bromage TG, Schrenk F, editors. African
biogeography, climate change and human evolution. Oxford:
Oxford University Press. p 138–149.
Plumptre AJ, Reynolds V. 1994. The effect of selective logging
on the primate populations in the Budongo Forest Reserve,
Uganda. J Appl Ecol 31:631–641.
Relethford JH. 2005. The human species. An introduction to
biological anthropology, 6th edition. Boston: McGraw-Hill.
473p.
Reynolds V. 1965. Budongo: an African forest and its
chimpanzees. New York: Natural History Press. 253p.
Reynolds V. 2005. The chimpanzees of the Budongo forest.
Ecology, behaviour and conservation. Oxford: Oxford University Press. 352p.
Struhsaker TT. 1975. The red colobus monkey. Chicago:
University of Chicago Press. 311p.
Struhsaker TT. 1997. Ecology of an African rain forest. Logging
in Kibale and the conflict between conservation and
exploration. Gainesville: University Press of Florida. 434p.
Sugiyama Y. 2004. Demographic parameters and life history of
chimpanzees at Bossou, Guinea. Am J Phys Anthropol
124:154–165.
Sugiyama Y, Koman J. 1992. The flora of Bossou: its
utilization by chimpanzees and humans. Afr Stud Mono
13:127–169.
Takemoto H. 2004. Seasonal change in the terrestriality of
chimpanzees in relation to microclimate in the tropical
forest. Am J Phys Anthropol 124:81–92.
Tutin CEG, Fernandez M, Rogers ME, Williamson EA,
McGrew WC. 1991. Foraging profiles of sympatric lowland
gorillas and chimpanzees in the Lope Reserve, Gabon. Philo
Trans R Soc Lond B 334:179–186.
Tutin CEG, Ham RM, White LJT, Harrison MJS. 1997a. The
primate community of the Lope Reserve, Gabon: diets,
responses to fruit scarcity, and effects on biomass. Am J
Primatol 42:1–24.
Tutin CEG, White LJT, Mackanga-Missandzou A. 1997b.
The use by rain forest mammals of natural forest fragments
in an equatorial African savanna. Conserv Biol
11:1190–1203.
Uehara S. 2003. Population densities of diurnal mammals
sympatric with the chimpanzees of the Mahale Mountains,
Tanzania: comparisons between the census data of 1996 and
2000. Afr Stud Mono 24:169–179.
Uehara S, Ihobe H. 1998. Distribution and abundance of
diurnal mammals, especially monkeys at Kasoje, Mahale
Mountains, Tanzania. Anthropol Sci 106:349–369.
de Vivo M, Carmignoto AP. 2004. Holocene vegetation change
and the mammal faunas of South America and Africa. J
Biogeog 31:943–957.
White LTJ, Rogers ME, Tutin CEG, Williamson EA, Fernandez M. 1995. Herbaceous vegetation in different forest types
in the Lope Reserve, Gabon. Implications for keystone food
availability. Afr J Ecol 33:124–141.
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.
Wrangham RW, Conklin NL, Chapman CA, Hunt KD. 1991.
The significance of fibrous foods for Kibale forest chimpanzees. Philo Trans R Soc Lond B 334:171–178.
Am. J. Primatol.
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