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Distribution and abundance of sacred monkeys in Igboland southern Nigeria.

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American Journal of Primatology 71:574–586 (2009)
RESEARCH ARTICLE
Distribution and Abundance of Sacred Monkeys in Igboland, Southern Nigeria
LYNNE R. BAKER1, ADEBOWALE A. TANIMOLA2, OLUSEUN S. OLUBODE3, AND DAVID L. GARSHELIS1,4
1
Conservation Biology Program, Department of Fisheries, Wildlife, and Conservation Biology, University of Minnesota,
St. Paul, Minnesota
2
Department of Crop and Soil Sciences, Faculty of Agriculture, University of Port Harcourt, Port Harcourt,
Rivers State, Nigeria
3
Department of Crop Protection and Environmental Biology, University of Ibadan, Ibadan, Nigeria
4
Minnesota Department of Natural Resources, Grand Rapids, Minnesota
Although primates are hunted on a global scale, some species are protected against harassment and
killing by taboos or religious doctrines. Sites where the killing of sacred monkeys or the destruction of
sacred groves is forbidden may be integral to the conservation of certain species. In 2004, as part of a
distribution survey of Sclater’s guenon (Cercopithecus sclateri) in southern Nigeria, we investigated
reports of sacred monkeys in the Igbo-speaking region of Nigeria. We confirmed nine new sites where
primates are protected as sacred: four with tantalus monkeys (Chlorocebus tantalus) and five with
mona monkeys (Cercopithecus mona). During 2004–2006, we visited two communities (Akpugoeze and
Lagwa) previously known to harbor sacred populations of Ce. sclateri to estimate population abundance
and trends. We directly counted all groups and compared our estimates with previous counts when
available. We also estimated the size of sacred groves and compared these with grove sizes reported in
the literature. The mean size of the sacred groves in Akpugoeze (2.06 ha, n 5 10) was similar to others in
Africa south of the Sahel, but larger than the average grove in Lagwa (0.49 ha, n 5 15). We estimated a
total population of 124 Sclater’s monkeys in 15 groups in Lagwa and 193 monkeys in 20 groups in
Akpugoeze. The Akpugoeze population was relatively stable over two decades, although the proportion
of infants declined, and the number of groups increased. As Sclater’s monkey does not occur in any
official protected areas, sacred populations are important to the species’ long-term conservation.
Despite the monkeys’ destruction of human crops, most local people still adhere to the custom of not
killing monkeys. These sites represent ideal locations in which to study the ecology of Sclater’s monkey
and human–wildlife interactions. Am. J. Primatol. 71:574–586, 2009.
r 2009 Wiley-Liss, Inc.
Key words: Cercopithecus sclateri; Cercopithecus mona; Chlorocebus tantalus; conservation;
Nigeria; taboo
INTRODUCTION
Hunting of primates is common worldwide.
Hunting may be more detrimental to primate
survival than habitat destruction and can locally
extirpate populations even where suitable habitat
remains [Mittermeier, 1987; Oates, 1996; Redford,
1992]. Primates are hunted for a variety of reasons:
to eat, sell, or keep as pets; for use in medicines, in
rituals, or as ornaments; or because they are
regarded as crop raiders [Mittermeier, 1987].
Sites where humans do not hunt primates are
less common, although examples occur globally
(Table I). Reasons for not hunting primates are
varied, but are frequently based on religion. Islam,
Hinduism, and Buddhism have various restrictions
or beliefs that prevent followers from eating or
killing primates [Mittermeier, 1987]. A well-known
example is the Hindu-based protection of monkeys in
parts of Asia, such as long-tailed macaques (Macaca
r 2009 Wiley-Liss, Inc.
fascicularis) in Bali, Indonesia [Wheatley, 1999], and
rhesus macaques (M. mulatta) and gray langurs
(Semnopithecus spp.) in India and the neighboring
Contract grant sponsors: The American Society of Primatologists; CENSHARE (University of Minnesota-UMN); Doctoral
Dissertation International Research Grant (UMN); Interdisciplinary Center for the Study of Global Change (UMN); Lincoln
Park Zoo; Margot Marsh Biodiversity Foundation; National
Science Foundation; Rufford Small Grants Foundation; Sigma Xi.
Oluseun S. Olubode’s current address is Plant Science and
Biotechnology Program, Department of Biological Sciences and
Biotechnology, Caleb University, Lagos, Nigeria
Correspondence to: Lynne R. Baker, 2713 Chimney Hill
Drive, Waco, TX, 76708. E-mail: bake0389@umn.edu, sclateri@
yahoo.com
Received 29 October 2008; revised 19 February 2009; revision
accepted 19 March 2009
DOI 10.1002/ajp.20690
Published online 30 April 2009 in Wiley InterScience (www.
interscience.wiley.com).
Sacred Monkeys in Nigeria / 575
TABLE I. Examples of Sites where Primates are Protected as Sacred or by Local Taboos
Location
Asia
Lindu Highlands, Lore Lindu NP,
Sulawesi, Indonesia
Mount Halimun NP, Java, Indonesia
Batang Ai NP, Sarawak, Malaysia
Mentawai Islands, Indonesia
Nam Kan Valley, Laos
South America
Parts of Suriname
Several sites/countries
Madagascar
Analamera Special Reserve
Fianarantsoa Province
(Ranomafana NP)
Several sites
Africa
Several sites in West Africa
Malebo and Wamba, Democratic
Republic of Congo
Kagwene Mountain, Cameroon
Speciesa
Source
Tonkean macaque (Macaca tonkeana)
Riley [2005]
Silvery gibbon (Hylobates moloch)
Bornean orangutan (Pongo
pygmaeus)
Kloss’s gibbon (Hylobates klossii)b
Laotian black-crested gibbon
(Nomascus concolor lu)
Indrawan et al. [1995/1996]
Horowitz [1998]
Golden-handed tamarin (Saguinus
midas), Guiana spider monkey
(Ateles paniscus)
Several species
Mittermeier [1987]
Perrier’s sifaka (Propithecus perrieri)
Eastern avahi (Avahi laniger), MilneEdwards’ sifaka (Propithecus
edwardsi)
Verreaux’s sifaka (Propithecus
verreauxi), Ring-tailed lemur
(Lemur catta), Decken’s sifaka
(Propithecus deckenii), Goldencrowned sifaka (Propithecus
tattersalli), Indri (Indri indri)
Western chimpanzee (Pan troglodytes
verus)
Bonobo (Pan paniscus)
Guelitapia, Côte d’Ivoire
Cross River gorilla (Gorilla gorilla
diehli)
Mona monkey (Cercopithecus mona)
White-thighed colobus (Colobus
vellerosus), Lowe’s monkey
(Cercopithecus lowei)
Eastern lesser spot-nosed monkey
(Cercopithecus petaurista
petaurista), olive colobus
(Procolobus verus), Ce. lowei
Green monkey (Chlorocebus sabaeus),
Ce. p. petaurista, Co. vellerosus,
Ce. lowei
P. verus, Ce. p. petaurista, Ce. lowei
Dinaoudi, Côte d’Ivoire
P. verus, Co. vellerosus
Tafi Atome, Ghana
Boabeng-Fiema, Ghana
Duasidan, Ghana
Soko, Côte d’Ivoire
Whittaker [2006]
Geissmann [2007]
Reviewed in Cormier [2006]
Mayor and Lehman [1999]
Jones et al. [2008]
Reviewed in Jones et al. [2008]
Reviewed in Kormos et al. [2003]
(e.g., Table 21.2)
Inogwabini et al. [2007];
Tashiro et al. [2007]
Wittiger and Sunderland-Groves
[2007]
Ormsby [in press]
Fargey [1991]; Saj et al. [2006]
Green Shepherd Ghanac; P. Sicotte,
personal communication
Gonedelé Bi et al.
S. Gonedelé Bi,
communication
Gonedelé Bi et al.
S. Gonedelé Bi,
communication
Gonedelé Bi et al.
S. Gonedelé Bi,
communication
[2007, in press];
personal
[2007];
personal
[in press];
personal
a
Classifications follow IUCN [2008] and Butynski et al. [in press] for African primates.
Author notes that Mentawais have largely abandoned this taboo.
Personal communication with GSG executive director (E. Abugbila).
b
c
region [Carter & Carter, 1999; Southwick et al.,
2005; Wolfe, 2002].
Primates may also be protected by social
taboos [Colding & Folke, 2001]. Some chimpanzee
(Pan troglodytes) populations are not hunted
because of their physical similarity to humans or
folklore regarding an ancestral relationship with
humans [Clavette, 2003; Herbinger et al., 2005;
Kortlandt, 1986; Osemeobo, 1994; Sabater Pi &
Groves, 1972].
In Nigeria, it is not unusual to find communities
(village-groups) or villages that hold animals, trees,
Am. J. Primatol.
576 / Baker et al.
forests, or streams sacred. There is much variation
across cultures and languages as to the meaning of
‘‘sacredness’’ [Hubert, 1994]. Here we use sacred to
refer to something that is worshipped, respected,
undisturbed, or specially managed or used owing to its
association with a religion or local belief system. In
this sense, sacred species are associated with taboos
that prohibit a group of people from engaging in
harmful behaviors toward these species, such as
hunting or eating them [Colding & Folke, 1997, 2001].
Examples of sacredness occur throughout the
Igbo-speaking region of Nigeria (Igboland) [e.g.,
Anoliefo et al., 2003; Oates et al., 1992; Okpoko,
2001]. Igboland includes the predominantly Igbo
states of Abia, Anambra, Ebonyi, Enugu, and Imo
(Fig. 1), as well as parts of Delta and Rivers States.
Many Igbo people, for example, adhere to a taboo
against harming the royal python (Python regius)
[Jell-Bahlsen, 1997; Okpoko, 2001; Ubah, 1982]. In
Igbo sites where monkeys are considered sacred, they
are often associated with a shrine (deity) and
unharmed wherever they travel within a community
[Oates et al., 1992]. In other cases, the monkeys
themselves are not sacred, but are unharmed when
they occur inside sacred forests. Elsewhere in southeastern Nigeria, hunting is widespread [Fa et al.,
2006; Oates et al., 2004]. Igboland has no official
protected areas, such as wildlife sanctuaries or
national parks.
As part of a survey in 2004 on the distribution of
Sclater’s guenon (Cercopithecus sclateri), which is
endemic to southern Nigeria, we investigated reports
of sacred monkeys in Igboland. We identified nine
previously unreported communities with either
sacred tantalus monkeys (Chlorocebus tantalus) or
mona monkeys (Cercopithecus mona). We also conducted censuses to estimate the abundance of two
previously known sacred populations of Sclater’s
Fig. 1. Igboland’s five main states: Abia, Anambra, Ebonyi,
Enugu, and Imo.
Am. J. Primatol.
monkeys in 2004 and again in 2005–2006. In one of
these sites, previous estimates of abundance were
available, which enabled us to assess trends in the
population and evaluate the effectiveness of the
monkeys’ sacred status. To estimate available forest
habitat for monkeys in both sites, we measured
sacred groves and compared them with groves
elsewhere in Africa south of the Sahel.
METHODS
Study Subject
In 1902, Sclater’s monkey was described and
named in honor of zoologist Philip Lutley Sclater
[Pocock, 1904]. It is recognized as a distinct species
and member of the cephus superspecies [Kingdon,
1980]. Investigations of its distribution were not
initiated until 1987 [Oates & Anadu, 1989; Oates
et al., 1992]. These surveys found that the species is
restricted to isolated populations between the Niger
and Cross Rivers in southern Nigeria. Long considered endangered, this species is now listed as
vulnerable [IUCN, 2008] because of recent evidence
of its adaptability and persistence in degraded,
fragmented forests across its range [Baker &
Olubode, 2008].
Study Region
We conducted a distribution survey in the range
of Sclater’s monkey [Baker & Olubode, 2008]. Most
of the study area is low-lying (o300 m) with little
relief [Barbour et al., 1982]. Southern Nigeria
has high mean annual rainfall: from 1,500 mm in
the northerly portion to 4,000 mm near the coast
[Barbour et al., 1982]. The rainy season is generally
from late March to early November [Olaniran &
Sumner, 1989], but can last more than 10 months
in the Niger Delta and along the coast [Barbour
et al., 1982].
The five Igbo states have high average human
density: 595 individuals/km2 (range 5 328–878) [Geomatics International Inc. et al., 1998; National
Population Commission, 2007]. Vegetation in the
northern part of Igboland is generally classified as
forest-savanna mosaic and Guinea savanna woodland, whereas oil palm (Elaeis guineensis), farmbush (newly regenerating vegetation), and secondary
forest predominate elsewhere [Barbour et al., 1982;
Gornitz & NASA, 1985]. The region is largely
cultivated. In the mid-1990s, 73–89% of land use in
these states was classified as ‘‘intensive (crop)
agriculture’’ [Geomatics International Inc. et al.,
1998].
As a result of intense human use, Igboland is
believed to have long been devoid of game species.
Writing about Igboland in the early 20th century,
G.T. Basden noted: ‘‘There has been, since the
introduction of firearms, an unrestricted slaughter
Sacred Monkeys in Nigeria / 577
of bird and beast until they have been reduced
almost to vanishing point. y This lack of game y is
also influenced by the density of the population,
particularly in the southern districts. There is little
country where wild animals can roam undisturbed
y ’’ [Basden, 1966, p 117].
We conducted censuses of Sclater’s monkey in
Akpugoeze, Enugu State, and Lagwa, Imo State.
Vegetation in these communities generally comprises
farm-bush and active farmland. Oil palm is especially
common throughout Lagwa; also present are Raphia
spp. palms, which are planted and tapped for palm
wine. Small forest patches are present in both
communities, particularly in Akpugoeze, and are
often protected as sacred groves. With limited
natural food and cover, monkeys invade farms and
gardens and run on rooftops; they are widely viewed
as pests and usually driven away [Oates et al., 1992;
Tooze, 1994; L. R. Baker, unpublished data].
The settlement of Akpugoeze (45 km2) comprises three autonomous communities, which include seven villages. Two of these villages consider
monkeys sacred, although some residents of the
other five villages do not kill or eat monkeys.
Monkeys are also generally unharmed anywhere in
Akpugoeze when found in sacred groves. Lagwa
(8.5 km2) comprises seven villages and one newly
autonomous community, Umunokwu. The entirety
of Lagwa–Umunokwu protects monkeys. In both the
sites, only Sclater’s monkey is known to occur,
although the mona monkey occurs just a few
kilometers to the east of Akpugoeze in Inyi, where
it is considered sacred [Oates, 1989].
In Akpugoeze, monkeys are protected because
they are considered the property of two shrines
(deities). In Lagwa, two stories exist regarding the
origin of the monkeys’ sacredness. Monkeys are
associated with a deity of the traditional religion, and
they are said to have been friends of Lagwa since the
community was founded. One common variation of
the latter story is that monkeys dropped food from
trees for the hungry, pregnant (or nursing) wife of
Agwa, the founder of Lagwa, and he subsequently
decreed that they should never be harmed.
In Igboland, shrines may be housed within or
nearby a sacred grove. Many groves have sacred
status because of their associated shrines and are
thus named after them [Anoliefo et al., 2003;
Okpoko, 2001]. Shrines in Lagwa are physical
structures (small buildings or altars) that may reside
within or near a sacred grove. Few shrines contain
structures in Akpugoeze; instead, a shrine is usually
part of a sacred grove, which is maintained free of
vegetative undergrowth and debris by one or more
shrine priests. Some groves contain pots, drums, or
other symbolic items.
Land in Igbo society usually belongs to a lineage,
through which an individual gains access to parcels
of land (secures tenure) and thus rights to plant
crops or construct buildings [Ejidike, 1999; Uchendu,
1965]. Sacred groves are generally considered the
property of their associated shrines and are under
the purview and control of the shrines’ chief priests.
No one can farm or cut trees in the groves, or even
enter them, without permission from the chief priest
[Anoliefo et al., 2003; Tooze, 1994]. The priesthood of
a shrine usually stays within the priest’s family
(inherited from one’s father or uncle, for example)
[Ubah, 1982; L. R. Baker, unpublished data]. If a
shrine priest abandons his duties and no longer cares
for the shrine and grove, or if he dies and no one
takes over, the shrine may lose its status, and
protection of the grove may be at risk.
Distribution Survey
In 2004, we visited villages and forest sites in
southern Nigeria to confirm the presence of Sclater’s
monkey; methods of this study are detailed in Baker
and Olubode [2008]. When local people reported the
presence of sacred monkeys elsewhere, we traveled
to these sites and generally spent a few hours
attempting to visually identify species by investigating areas where monkeys were typically seen.
When we could not obtain visual sightings,
owing to logistical and time constraints associated
with our main study, we conducted informal interviews with elders, leaders, or other available community members and accepted reliable reports as
confirmation. Reliable reports were those where local
people (usually Z four people) correctly described a
species by physical traits or vocalizations and
selected the described species from among several
photos. In addition, we visited six communities
previously known to harbor sacred monkeys (three
with Sclater’s monkeys and three with mona
monkeys).
Population Censuses
In Lagwa, we counted monkeys in April–May
2004 and October–November 2005. In 2004, a local
political conflict limited our census to six of Lagwa’s
eight villages. In Akpugoeze, we conducted counts in
June 2004 and April–May 2006. Most groups in
Lagwa and Akpugoeze are habituated to humans,
and thus we attempted to make total counts
(censuses) [Jarman et al., 1996]. Because of the
paucity of forest, groups were generally located in or
around sacred groves or tree patches. Four main
observers (L. R. Baker and three trained assistants)
worked independently to maximize area coverage
and made repeated visits to these areas.
We counted all monkeys in each group, using
binoculars when needed, and distinguished dependent
infants by their size and behavior (young that were
nursing or clung to mothers). For final group counts,
we used the maximum reliable count or the most
consistent count of each group. To assess trends in
Am. J. Primatol.
578 / Baker et al.
Akpugoeze, we compared our counts with those made
in January 1989 and March–April 1994 by Oates et al.
[1992] and Tooze [1994], respectively. Our methodology was similar to that of Tooze [1994] and Oates
et al. [1992], although the latter study provided fewer
details about group size and composition.
Sacred Groves
In 2005–2006, we estimated the size of sacred
groves. We measured only those groves that had
relatively clear boundaries and for which we had the
necessary permission (Lagwa: n 5 15, Akpugoeze:
n 5 10). In Akpugoeze, we included ‘‘bad bush,’’ a
forest patch formerly used for disposing of unwanted
twin infants [Tooze, 1994], because it had remained
off-limits to human use. Some grove borders were
difficult to distinguish because of reported boundary
discrepancies and connectivity with farm-bush. We
attempted to exclude such vegetation, although some
sacred groves, particularly in Lagwa, comprised
primarily shrubs and a few trees. We slowly walked
the perimeter of each grove in clear weather and
calculated area using Global Positioning System
tracking technology. For 17 groves, we made two
perimeter walks and calculated the mean.
We conducted a literature search to compare our
estimates with sizes reported for sacred groves
elsewhere in Africa south of the Sahel. The literature
on sacred groves is vast, and sizes are reported
infrequently. We could not find or obtain all possible
sources; hence our aim was to compile a representative list. Our list should be viewed as thorough, but
not comprehensive. When possible, we calculated
summary statistics from raw data presented in
sources. Using two-tailed Mann–Whitney U tests,
we determined whether grove sizes differed among
Akpugoeze, Lagwa, and those reported in the
literature (n 5 74; Table II, excluding the particularly
large 2,000 ha grove). For Akpugoeze, we considered
only those sacred groves that were within or bordered
the two villages that protect monkeys, as these forest
patches were most often used by monkeys.
All research was conducted with permission of
the Imo and Enugu Forestry Departments and local
communities and adhered to the legal requirements
of Nigeria.
RESULTS
informants reported that protection was conferred
specifically on one monkey (Ce. mona), whereas in
the neighboring communities of Amiri and Awomamma, we confirmed the presence of just Ch.
tantalus, although all monkey types were said to be
protected. Reports were unclear in some sites, such
as Awka, where we observed only mona monkeys,
but other species may receive protection in this
community.
We also determined the continued presence of
sacred monkeys at six of seven sites where they had
been reported previously. In Akpugoeze, Lagwa,
and Ikot Uso Akpan, we observed only Sclater’s
monkeys, as reported previously by Oates [1989],
Oates et al. [1992], and R. King and E. Egwali
[personal communication], respectively. In Inyi and
Imerienwe, we reaffirmed, through local reports,
previous findings that only mona monkeys were
present [Oates, 1989; Oates & Anadu, 1988]. Oates
[1989] found that Ce. mona was sacred in
Ejemekwuru and suggested that Ch. tantalus may
also be present. During our visit there, a community
leader reported that just one type of monkey
(Ce. mona) was protected by a particular deity;
correspondingly, we observed only mona monkeys
there.
Anecdotal information and our own observations indicated that the belief systems that protect
monkeys were breaking down in some sites, although
we did not formally evaluate this. In Awomamma
and Amiri, protection of monkeys was now limited to
two villages, one in each community. We also saw in
Amiri a recently captured young tantalus monkey,
which was said to be destined for the dinner table
once it grew bigger. In Abacha, local people reported
that the belief system protecting monkeys was now
very weak. In Lagwa, the shrine dedicated to the
monkeys’ deity was demolished by a Christian
association in 2000.
In this study, we also confirmed another protected
(though not sacred) Ce. sclateri population on the
forested grounds of Nekede Zoo in Imo State (Fig. 2).
The zoo covers 22 ha, of which 15 ha are forested and
referred to as a free-range zone. Here we observed
eight individuals: six adults, a juvenile, and an infant.
We were informed that the monkeys have always been
wild, but are now considered part of the zoo. Although
most of the free-range area is fenced, people were said
to hunt and set traps there on rare occasions.
Distribution Survey
We located nine previously unreported communities that harbor sacred monkeys: five with mona
monkeys and four with tantalus monkeys (Fig. 2).
Three were confirmed by visual observations and six
by reliable reports. We did not confirm more than
one sacred species per site, although some residents
indicated that other types of monkeys were present
(Fig. 2). In Eluama and Isuochi, for example,
Am. J. Primatol.
Population Censuses
The Sclater’s monkey population in Akpugoeze
changed little from 1989 to 2006, remaining at
just under 200 individuals (including dependent
infants; Table III). However, we observed an
increase in the number of groups and a decrease in
mean group size. Tooze [1994] recorded 199 individuals in 12 groups, averaging 16.6 individuals per
Sacred Monkeys in Nigeria / 579
TABLE II. Examples of Sacred-Grove Sizes Reported for African Countries South of the Sahel
Location/grove(s)
Number
Benin
Adja Plateau
Benin (countrywide)
13
2,940
Ouémé Valley
52
Ghana
Accra Plains/Adumanya, Pinkwae
Ashanti Region/Kagyase; Jachie,
Jachie-Pramso
Ashanti Region, Kumasi Area
Brong Ahafo Region/Boabeng-Fiema
Area (ha)a
Mean area (ha)
(SD, range)b
2.41 (5.75, 0.06–21)
r1 (n 5 2,040);
41o5
(n 5 538);
Z5 (n 5 362)
o1 (n 5 25);
1–5 (n 5 20);
45 (n 5 7)
Source
Sokpon and Ago [2001]
Agbo and Sokpon [1998]
Soury [2007]
2
2
1.5, 120
8, 11.5
60.8 (83.8)
9.8 (2.5)
Decher [1997]
GACONc
4
1
6, 8, 11.5, 259
192
71.1 (125.3, 6–259)
Brong Ahafo Region/Duasidan
Brong Ahafo Region/Mprisi, Buoyem;
Tano, Tano Boase
Coastal Savanna
1
2
22
369.4, 121.5
Bossart et al. [2006]
Saj et al. [2006];
P. Sicotte, personal
communication
Green Shepherd Ghana
GACONc
4
Eastern Region/Abiriw
Eastern Region, Esukawkaw Forest
Reserve/Anweam
Northern Region/Malshegu
Kenya
Southern Coast (kayas)
1
1
0.08, 0.10, 0.48,
16.01
16
2,000
1
0.8
Mbeere, Embu District
Mozambique
Chôa Highlands, Barué District/
Chinda, Mungwa
Nigeria
Osun State/Osun-Osogbo
Sierra Leone
Moyamba District
Tanzania
Mbeya Region, Rungwe District
North Pare Mountains, Mwanga
District
146
Tabora Region
Tanga Region, Handeni District
Uganda
Mpigi District/Magezigoomu, Mukasa
Zanzibar
Jambiani Village/Kuumbi
58d
2
8, 9
1
75
14
15e
230
8
499
o0.5 (26%);
0.5o2 (46%);
2–5 (24%);
45 (4%)
o2 (26%);
2–5 (39%);
45–10 (18%);
410 (17%)
245.5 (175.3)
4.17 (7.90,
0.08–16.01)
Campbell [2005]
Kangah-Kesse et al. [2007]
Amoako-Atta [1998]
Dorm-Adzobu et al. [1991]
105.1 (216.2,
1–1,500)
(0.5–2)
Githitho [2004]
Riley and Brokensha [1988]
8.5 (0.7)
Virtanen [2002]
Akinpelu and Areo [2007]
3.5 (1.5, 1–6)
Lebbie and Guries [2008]
2.81 (3.92, 0.05–10)
(0.125–50)
McKone [1995]
Mwihomeke et al. [1998]
0.14 (0.06–0.26)
(o1–200)
Mgumia and Oba [2003]
Mwihomeke et al. [1998]
2
16, 4
10 (8.5)
Banana et al. [2008]
1
5
(0.5–20 for all
groves in
Zanzibar)
Madeweya et al. [2004]
a
Areas of individual groves are listed when the source reports five or fewer groves.
Summary statistics are listed if reported in, or could be calculated from, source data.
GACON: Ghana Association for the Conservation of Nature (Y. B. Agyeman, personal communication).
d
Excluded from data analysis as estimates include some non-forest area and so are not directly comparable (A. Githitho, personal communication).
e
Includes only sites visited by author.
b
c
Am. J. Primatol.
580 / Baker et al.
Fig. 2. Communities that hold monkeys sacred in Igboland (except ]2, which is an Ibibio community), listed by the species confirmed as
protected and survey reference. In some sites (), reports indicated that other monkey species could also be present. (1) Azumini
(population is not fully protected, as both Ibibio and Igbo people reside there, and only Igbos consider monkeys sacred [Tooze, 1995]); (2)
Ikot Uso Akpan; (3) Lagwa; (4) Imerienwe; (5) Nekede Zoo (monkeys are not sacred, but are not hunted here); (6) Ejemekwuru; (7)
Awomamma; (8) Amiri; (9) Akatta; (10) Umuihi; (11) Eluama; (12) Isuochi; (13) Ihe; (14) Inyi; (15) Akpugoeze; (16) Awka; and (17)
Abacha.
group, whereas our most recent estimate was 193
individuals in 20 groups, with an average size of 9.7
(Table III). We also observed about half the number
of dependent infants as reported in 1994.
We were unable to determine absolute density
because it was unclear how far monkeys ranged in
and around Akpugoeze. Rarely did we observe
monkeys in the five villages that do not protect
them, and then, usually near the boundaries of
‘‘safe’’ areas. Local people, however, reported that
monkeys occurred in these five villages, where there
are several forest patches, and complained of
monkeys raiding their crops. Also, we previously
observed Ce. sclateri in riverine forest 2 km from
Akpugoeze’s western boundary [Baker & Olubode,
2008]; monkeys may disperse between these sites.
During our survey, however, the groups we counted
occurred within a region of 6.5–7 km2. Using this
area, density would be 28–30 individuals/km2.
In Lagwa, we counted 83 individuals in 10
groups in 2004 and 124 individuals in 15 groups in
2005 (Table III). Our 2004 count was low because a
local political conflict restricted our survey area. All
villages within Lagwa protect monkeys, and local
Am. J. Primatol.
people indicated that monkeys rarely ventured out of
the community boundaries into areas where they
could be hunted. Using the approximate area of
Lagwa (8.5 km2), density was 14.6 individuals/km2,
or about half that of Akpugoeze. A neighboring
village that borders one Lagwa village was reported
to protect monkeys; we could not confirm this, but if
so, the range of one or more monkey groups may
include this area.
Monkey groups were occasionally wary and hid or
fled during surveys. Although they are habituated to
humans, some individuals and groups seemed unaccustomed to people stopping and watching them.
Local people generally ignored the monkeys, except
when they were near their homes or gardens or raided
their crops, at which point the monkeys were usually
chased away (sometimes with sticks or rocks).
Sacred Groves
In Akpugoeze, we measured 10 groves that
ranged from 0.24 to 9.60 ha (x ¼ 2:06, SD 5 2.84,
median 5 0.93). These groves were usually conspicuous forest patches, with some individual trees
–
1.0 (0.8, 0–2)
–
1:7.3
This study
This study
>30 m high. Some of the larger emergent species
included Milicia excelsa, Piptadeniastrum africanum, Triplochiton scleroxylon, Ceiba pentandra,
and Khaya spp. We observed African pied hornbills
(Tockus fasciatus), gray parrots (Psittacus erithacus), and white-crested hornbills (Tropicranus albocristatus) in the groves. Several sacred groves also
occurred within the villages that do not protect
monkeys; many were small, although two that we
measured were each 2 ha.
Based on local reports, most of Akpugoeze’s
shrine groves still exist, but have diminished in size
as a result of encroachment from agriculture or
construction. Few were currently maintained by
shrine priests; several priests had died and had not
been replaced. Informants reported that the priest of
one of the two shrines that confer protection on the
monkeys died several years ago, and his family had
not appointed another priest and was not interested
in maintaining the shrine.
In Lagwa, we measured 15 sacred groves that
ranged from 0.01 to 2.45 ha (x ¼ 0:49, SD 5 0.74,
median 5 0.16). They were significantly smaller than
those in Akpugoeze (U 5 25.0, Z 5 2.774, P 5 0.006)
and generally comprised a few trees and shrubs. Local
reports indicated that most groves had been greatly
reduced in size, and many community shrines had been
demolished or long abandoned. The sacred forests of
Akpugoeze and particularly Lagwa were smaller than
those reported elsewhere south of the Sahel (x ¼ 18:98,
SD 5 58.69, median 5 2.00, range 5 0.05–369.4, n 5 74).
This difference was statistically significant for Lagwa
(U 5 218.0, Z 5 3.695, P 5 0.000), but not for Akpugoeze (U 5 322.0, Z 5 0.663, P 5 0.507).
–: Not reported/available.
a
IND: Independents (adults and juveniles); DI: Dependent infants.
b
Owing to a local political conflict, census was limited and is thus an underestimate.
c
Excludes one solitary individual.
–
109
Lagwa
April–May 2004
October–November 2005
83b
124
10b
15
9.1c (6.4, 3–24)
8.3 (3.3, 4–14)
–
15
–
2.8 (1.4, 0–5)
–
0.8 (0.8, 0–3)
–
33
–
16
–
166
–
177
– (15–30)
16.6 (6.6, 8–28)
13.4 (5.6, 5–25)
9.7 (4.2, 4–19)
Akpugoeze
January 1989
March–April 1994
June 2004
April–May 2006
180–225
199
187
193
Z9
12
14
20
]INDa
Mean group size
(SD, range)
]Groups
]Individuals
Month/year
Community
TABLE III. Population Estimates of Cercopithecus sclateri in Akpugoeze and Lagwa
]DIa
Mean ]DI/group
(SD, range)
–
1:5.0
–
1:11.1
Ratio DI:IND
Source
Oates et al. [1992]
Tooze [1994]
This study
This study
Sacred Monkeys in Nigeria / 581
DISCUSSION
Distribution of Sacred Monkeys
The discovery of nine sacred-monkey sites in
Igboland is not surprising because sacred animals
and forests occur throughout the region. That each
of these sites harbored only a single monkey species
was also not unexpected, given that nearly all
previously described sacred-monkey sites in Nigeria
contained a single species (although Tooze [1995]
found both Sclater’s and mona monkeys in one
location) (Fig. 2). Possible reasons for this pattern
include: (1) differing ecological flexibility among
species; (2) doctrines of the belief system or religion;
(3) species-specific habitat preferences; (4) historic
distribution and abundance; and (5) inter-specific
competition. The first reason seems least likely
because all three species found in this region—Ce.
sclateri, Ce. mona, and Ch. tantalus—are considered
ecologically flexible and known to persist in modified
environments [Baker & Olubode, 2008; Glenn et al.,
in press; Kavanagh, 1980].
Doctrines are probably most important. If a
belief system specifically confers protection on one
Am. J. Primatol.
582 / Baker et al.
monkey, others may be hunted and eventually
extirpated. In Ejemekwuru, for example, a community leader said that mona monkeys are the only
species that belongs to a local deity, and thus other
monkey types can be killed. Hunting pressure from
neighboring communities may further encourage
sacred species to restrict their movements. Presently
in Akpugoeze, monkeys occur primarily in a limited
area where they are not hunted, even though this
area is human-dense, and several other forest
patches exist in the vicinity.
Certain doctrines may have stemmed from
ecological factors that initially favored a single
species, which then received special status because
of its relative commonness. Doctrines may have also
changed over time to single out species that were less
destructive to crops or less dangerous. Multiple
species should have historically existed in these sites,
but this situation predated the memories of most
people. Many elders and leaders in Lagwa, for
example, claimed to know of the existence of only
one type of monkey, although a 60-year-old informant recalled the presence of another monkey that
he said had not been seen for at least 30 years. In
some communities with one sacred species, informants reported formerly or occasionally seeing other
monkeys in the surrounding area; these were often
described as patas monkeys (Erythrocebus patas) or
olive baboons (Papio anubis).
Where doctrines do not specify a certain species,
such factors as habitat, historic distribution and
abundance, and competition may individually or in
combination explain which species (and why only one
species) presently occurs in an area. Tantalus
monkeys are savanna-woodland dwellers and less
arboreal than Ce. mona and Ce. sclateri; they may
have traditionally occurred or been more common in
areas less suitable for Cercopithecus monkeys.
Limited resources and space may also have encouraged inter-specific competition and favored larger or
more aggressive monkeys. Baranga [2004], for
example, found that the smaller red-tailed monkeys
(Cercopithecus ascanius) were displaced by Ch.
tantalus (revised taxonomy [IUCN, 2008]) in forest
fragments.
The region where most sacred-monkey populations occur overlaps the northern portion of Sclater’s
range, which is largely forest-savanna mosaic. This is
atypical habitat for Sclater’s monkey, and the species
has not been widely recorded here [Baker & Olubode,
2008]. This, combined with Sclater’s monkey being
the smallest diurnal primate in the region, may
explain why the majority of sacred-monkey sites do
not harbor this species.
Censuses of Sclater’s Monkeys
Our comparison of census results showed that
the Akpugoeze population has remained relatively
Am. J. Primatol.
constant. We recognize that censuses of sacred
monkeys may result in underestimates [Wong &
Sicotte, 2006]. Detectability, even with habituated
monkeys in sparse vegetation, was unlikely to be
perfect, as some groups and individuals were wary
and difficult to count. However, we used similar
methods as previous investigators, as well as the
same observers in 2004 and 2006; hence these counts
should be comparable.
The stability of the Akpugoeze population is
likely owing in large part to the continued respect by
most residents for the taboo against harming
monkeys. That the population has not increased
may reflect the site’s carrying capacity. At the
Boabeng-Fiema Monkey Sanctuary (192 ha) in
Ghana, where a local belief system confers protection
on two primates (Colobus vellerosus and Cercopithecus lowei), the Ce. lowei population increased in a
few years from 216 [Fargey, 1991] to 342 [Kankam,
1997] and has continued to increase [B. Kankam,
personal communication]. The Co. vellerosus population also grew about 1.8 over 13 years; however,
in smaller-sized forest fragments surrounding the
sanctuary, Co. vellerosus abundance decreased
slightly [Wong & Sicotte, 2006].
The decline in the proportion of dependent infants
in Akpugoeze between the 1994 and 2006 surveys may
reflect poor reproductive success. It may also be a result
of a natural or periodic fluctuation in the age structure
or reproductive output of the population [e.g., Rudran
& Fernandez-Duque, 2003]. Notably, however, both
surveys were conducted at similar times of the year,
which may rule out a seasonal effect.
We observed a greater number of groups in
Akpugoeze than in previous studies, but only slightly
more individuals in 2006 than in 2004. This suggests
a degree of fluidity in group composition in this
population. One explanation is that groups undergo
fission–fusion, and that some observed groups were
subgroups or smaller foraging parties. Alternately,
average group size may have decreased. Permanent
fission, although uncommon in Cercopithecus species, has been observed when group sizes reached
23–50 individuals [Butynski, 1990; Cords & Rowell,
1986; Struhsaker & Leland, 1988; Windfelder &
Lwanga, 2002]. Struhsaker and Leland [1988]
posited that larger groups may undergo fission when
female reproductive success is low, which may have
been the case in Akpugoeze, given the relatively
small proportion of infants.
Access to and competition for food and other
resources may also limit group size [Chapman &
Chapman, 2000]. In the unique village environments
of Akpugoeze and Lagwa, monkeys exploit limited
areas by raiding the gardens and crops of residents
and often using human structures (rooftops, compound walls, etc.) to travel and rest. In such
anthropogenic sites, there may be an added benefit
for monkeys to live and forage in smaller parties:
Sacred Monkeys in Nigeria / 583
Smaller groups are more likely to go unnoticed or be
more tolerated by local residents, who regularly
chase and throw objects at garden-raiding monkeys.
Conservation Implications
Religious doctrines and species-specific taboos
can be important in the conservation of declining or
threatened species [Colding & Folke, 1997; Table I].
Of the 24 primates listed in Table I, three are
critically endangered (Nomascus concolor lu, Propithecus perrieri, and Gorilla gorilla diehli); eight are
endangered (Hylobates moloch, Pongo pygmaeus,
Hylobates klossii, Propithecus edwardsi, Propithecus
tattersalli, Indri indri, Pan troglodytes verus, and Pan
paniscus); and five are vulnerable (Macaca tonkeana,
Ateles paniscus, Propithecus verreauxi, Propithecus
deckenii, and Co. vellerosus) [IUCN, 2008].
Lagwa and Akpugoeze are two of just three
known communities across the range of Sclater’s
monkey where it is considered sacred and thus
protected (Sclater’s monkeys are also relatively
protected at Nekede Zoo and receive partial protection at Azumini). These sites represent the species’
only ‘‘protected areas.’’ In all sites, however, available natural habitat (forest) is scant and patchy, as
evidenced by the relatively small size of the sacred
groves. Lagwa’s groves are particularly small and
probably insignificant to the monkeys’ survival.
Monkeys regularly sought food in people’s gardens
and farms, and residents were increasingly annoyed
with them [Oates et al., 1992; Tooze, 1994; L. R.
Baker, unpublished data]. In both Akpugoeze and
Lagwa, we received reports of outsiders occasionally
being permitted to kill monkeys, as well as locals
killing them surreptitiously.
The long-term survival of these monkeys largely
depends on the willingness of the local people to
continue to adhere to the taboo against harming
monkeys, despite depredation of their crops and
apparent erosion of the religious beliefs protecting
the monkeys. Residents were generally unaware of
the uniqueness and overall rarity of Sclater’s
monkey, the role of their community in its conservation, or any benefit that its presence provides or
could provide in the future (e.g., ecotourism).
This situation contrasts with Boabeng-Fiema in
Ghana, where most local people considered the
monkeys an important part of their cultural heritage
and believed that the monkeys’ presence could
promote development of their villages [Fargey,
1991]. A presently strong adherence to the hunting
taboo is partly attributed to a growth in tourism
revenue, the success of which has even encouraged
neighboring communities to stop hunting monkeys
(particularly colobus) and seek involvement in the
program [Saj et al., 2005]. For several years BoabengFiema has also had a steady influx of researchers, who
are responsible for a growing literature on this site
and species [e.g., Saj & Sicotte, 2007; Saj et al., 2006;
Sicotte & Macintosh, 2004; Sicotte et al., 2007;
Teichroeb et al., 2003; Wong & Sicotte, 2006, 2007].
Boabeng-Fiema is an example of the potential
that sacred-monkey sites have for research and
ecotourism. Similarly, Lagwa and Akpugoeze are
unique sites for the study of Sclater’s monkey
(ecology, behavior, etc.), human–wildlife interactions
(disease transmission, etc.), and the conservation
role of local belief systems. Both communities have
relatively high primate densities. Monkeys occur in
human-dense settings with sparse forest cover,
which makes visual observations relatively easy.
Additional
research
is
needed
on
this
under-studied species, to monitor population and
behavioral changes with respect to environmental
and human-related factors, especially potential
changes in the monkeys’ sacred status.
ACKNOWLEDGMENTS
The authors thank the communities of LagwaUmunokwu and Akpugoeze and the Imo and Enugu
State Forestry Departments for their assistance and
permission to conduct research, which adhered to
Nigeria’s legal requirements. For their support in
Nigeria, much appreciation goes to Rose Bassey,
Jennifer Seale, Mobil Producing Nigeria, Chief
Assam Assam, Zena Tooze, CERCOPAN, and Eli
Khawaja. Helpful comments on the manuscript were
provided by John Oates, Janette Wallis, Douglas
Hawkins, Zena Tooze, and Tamara Giles-Vernick.
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