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: email@example.com, 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  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  Golden-handed tamarin (Saguinus midas), Guiana spider monkey (Ateles paniscus) Several species Mittermeier  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  Geissmann  Reviewed in Cormier  Mayor and Lehman  Jones et al.  Reviewed in Jones et al.  Reviewed in Kormos et al.  (e.g., Table 21.2) Inogwabini et al. ; Tashiro et al.  Wittiger and Sunderland-Groves  Ormsby [in press] Fargey ; Saj et al.  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 ; personal [in press]; personal a Classifications follow IUCN  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 . 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.  and Tooze , respectively. Our methodology was similar to that of Tooze  and Oates et al. , 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 , Oates et al. , 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  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  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  Agbo and Sokpon  Soury  2 2 1.5, 120 8, 11.5 60.8 (83.8) 9.8 (2.5) Decher  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.  Saj et al. ; 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  Kangah-Kesse et al.  Amoako-Atta  Dorm-Adzobu et al.  105.1 (216.2, 1–1,500) (0.5–2) Githitho  Riley and Brokensha  8.5 (0.7) Virtanen  Akinpelu and Areo  3.5 (1.5, 1–6) Lebbie and Guries  2.81 (3.92, 0.05–10) (0.125–50) McKone  Mwihomeke et al.  0.14 (0.06–0.26) (o1–200) Mgumia and Oba  Mwihomeke et al.  2 16, 4 10 (8.5) Banana et al.  1 5 (0.5–20 for all groves in Zanzibar) Madeweya et al.  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.  Tooze  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  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 , 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  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. REFERENCES Agbo V, Sokpon N. 1998. Forêts sacrées et patrimoine vital au Bénin. Rapport technique final. Projet CRDI (Centre de recherches pour le développement international) no. 95–8170. Université Nationale du Bénin, Faculté des Sciences Agronomiques. 32p. Akinpelu AI, Areo A. 2007. The snakes of Osun Grove: a World Heritage site in Osogbo, Nigeria. Rev Biol Trop 55:717–721. Amoako-Atta B. 1998. Preservation of sacred groves in Ghana: Esukawkaw Forest Reserve and its Anweam Sacred Grove. Paris: UNESCO. 40p. Anoliefo GO, Isikhuemhen OS, Ochije NR. 2003. Environmental implications of the erosion of cultural taboo practices in Awka—South Local Government Area of Anambra State, Nigeria: 1. Forests, trees, and water resource preservation. J Agr Environ Ethics 16:281–296. Baker LR, Olubode OS. 2008. Correlates with the distribution and abundance of endangered Sclater’s monkeys (Cercopithecus sclateri) in southern Nigeria. Afr J Ecol 46: 365–373. Banana AY, Bahati J, Gombya-Ssembajjwe W, Vogt N. 2008. Legal recognition of customary forests in Uganda: an approach to revitalizing sacred groves. In: Sheridan MJ, Nyamweru C, editors. African sacred groves: ecological dynamics and social change. Oxford: James Currey Ltd. p 195–206. Baranga D. 2004. Red-tail monkey groups in forest patches outside the protected area system in the ‘Kampala area’. Afr J Ecol 42:78–83. Am. J. Primatol. 584 / Baker et al. Barbour KM, Oguntoyinbo JS, Onyemelukwe JOC, Nwafor JC. 1982. Nigeria in maps. London: Hodder and Stoughton. 148p. Basden GT. 1966. Niger Ibos. London: Frank Cass and Co., Ltd. 456p. Bossart JL, Opuni-Frimpong E, Kuudaar S, Nkrumah E. 2006. Richness, abundance, and complementarity of fruit-feeding butterfly species in relict sacred forests and forest reserves of Ghana. Biodivers Conserv 15:333–359. Butynski TM. 1990. Comparative ecology of blue monkeys (Cercopithecus mitis) in high- and low-density subpopulations. Ecol Monogr 60:1–26. Butynski TM, Kingdon JS, Kalina J, editors. In press. The mammals of Africa. Vol. 2. Primates. Amsterdam: Academic Press. Campbell MO. 2005. Sacred groves for forest conservation in Ghana’s coastal savannas: assessing ecological and social dimensions. Singapore J Trop Geogr 26:151–169. Carter A, Carter C. 1999. Cultural representations of nonhuman primates. In: Dolhinow P, Fuentes A, editors. The nonhuman primates. Mountain View (CA): Mayfield Publishing Co. p 270–276. Chapman CA, Chapman LJ. 2000. Constraints on group size in red colobus and red-tailed guenons: examining the generality of the ecological constraints model. Int J Primatol 21:565–585. Clavette K. 2003. Conservation through folklore: ethnoprimatology in southeastern Sénégal [MA thesis]. Flagstaff (AZ): Northern Arizona University. 98p. Colding J, Folke C. 1997. The relations among threatened species, their protection, and taboos. Conserv Ecol 1:6 [Internet]. Available at: http://www.consecolorg/vol1/iss1/art6/. Colding J, Folke C. 2001. Social taboos: invisible systems of local resource management and biological conservation. Ecol Appl 11:584–600. Cords M, Rowell TE. 1986. Group fission in blue monkeys of the Kakamega Forest, Kenya. Folia Primatol 46:70–82. Cormier L. 2006. A preliminary review of neotropical primates in the subsistence and symbolism of indigenous lowland South American peoples. Ecol Environ Anthropol 2:14–32. Decher J. 1997. Conservation, small mammals, and the future of sacred groves in West Africa. Biodivers Conserv 6: 1007–1026. Dorm-Adzobu C, Ampadu-Agyei O, Veit PG. 1991. Religious beliefs and environmental protection: the Malshegu Sacred Grove in northern Ghana. Washington, D.C. and Nairobi: Center for International Development and Environment (World Resources Institute) and African Centre for Technology Studies. 35p. Ejidike OM. 1999. Human rights in the cultural traditions and social practice of the Igbo of south-eastern Nigeria. J Afr Law 43:71–98. Fa JE, Seymour S, Dupain J, Amin R, Albrechtsen L, Macdonald D. 2006. Getting to grips with the magnitude of exploitation: bushmeat in the Cross–Sanaga rivers region, Nigeria and Cameroon. Biol Conserv 129:497–510. Fargey PJ. 1991. Assessment of the conservation status of the Boabeng-Fiema Monkey Sanctuary. Final report to the Flora & Fauna Preservation Society. Kumasi (Ghana): University of Science and Technology. 73p. Geissmann T. 2007. First field data on the Laotian black crested gibbon (Nomascus concolor lu) of the Nam Kan area of Laos. Gibbon J 3:56–65. Geomatics International Inc., Beak Consultants Ltd., Unilag Consult of Nigeria. 1998. The assessment of vegetation and land use changes in Nigeria between 1976/78 and 1993/95. Abuja (Nigeria): Forestry Management, Evaluation, and Coordinating Unit (FORMECU) and Nigerian Federal Department of Forestry (under the World Bank Environmental Management Project). Am. J. Primatol. Githitho A. 2004. The coastal terrestrial forests of Kenya: a report on resources, threats, and investments. Nairobi: World Wildlife Fund. 29p. Glenn ME, Bensen KJ, Matsuda R. In press. Mona monkey (Cercopithecus mona). In: Butynski TM, Kingdon JS, Kalina J, editors. The mammals of Africa. Vol. 2. Primates. Amsterdam: Academic Press. Gonedelé Bi S, Sangaré A, N’Goran E, Zinner D, Siedel H, Vigilant L, Boesch C. 2007. Mitochondrial DNA shows three cryptic populations of Campbell’s and Lowe’s monkeys (Cercopithecus campbelli, Waterhouse, 1838) in Côte d’Ivoire. In: Hazelhurst S, Ramsay M, editors. Proceedings of the first Southern African Bioinformatics Workshop. Johannesburg: University of the Witwatersrand. p 59–62. Gonedelé Bi S, Bitty A, Gnangbé F, Bené JC, Koné I, Sangaré A, Zinner D. In press. Conservation status of Geoffroy’s pied colobus monkey (Colobus vellerosus, Geoffroy 1834) has dramatically declined in Côte d’Ivoire. Afr Primates. Gornitz V, NASA. 1985. A survey of anthropogenic vegetation changes in West Africa during the last century—climatic implications. Climatic Change 7:285–325. Green Shepherd Ghana. Monkey Sanctuary. Available at: http://greenshepherdghana.org/monkey.html. Herbinger I, Boesch C, Tondossama A, McManus E. 2005. Republic of Côte d’Ivoire. In: Caldecott J, Miles L, editors. World atlas of great apes and their conservation. Berkeley: University of California Press. p 328–332. Horowitz LS. 1998. Integrating indigenous resource management with wildlife conservation: a case study of Batang Ai National Park, Sarawak, Malaysia. Hum Ecol 26:371–403. Hubert J. 1994. Sacred beliefs and beliefs of sacredness. In: Carmichael DL, Hubert J, Reeves B, Schanche A, editors. Sacred sites, sacred places. London and New York: Routledge. p 9–19. Indrawan M, Supriyadi D, Supriatna J, Andayani N. 1995/1996. Javan gibbon surviving at a mined forest in Gunung Pongkor, Mount Halimun National Park, West Java: considerable toleration to disturbances? Asian Primates 5:11–13. Inogwabini BI, Matungila B, Mbende L, Abokome M, Tshimanga TW. 2007. Great apes in the Lake Tumba landscape, Democratic Republic of Congo: newly described populations. Oryx 41:1–7. IUCN. 2008. IUCN Red List of Threatened Species. Available at: http://www.iucnredlist.org. Jarman P, Smith AP, Southwell C. 1996. Complete counts. In: Wilson DE, Cole FR, Nichols JD, Rudran R, Foster MS, editors. Measuring and monitoring biological diversity: standard methods for mammals. Washington, D.C.: Smithsonian Institution Press. p 192–193. Jell-Bahlsen S. 1997. Eze mmiri di egwu, the water monarch is awesome: reconsidering the mammy water myths. Ann NY Acad Sci 810:103–134. Jones JPG, Andriamarovololona MM, Hockley N. 2008. The importance of taboos and social norms to conservation in Madagascar. Conserv Biol 22:976–986. Kangah-Kesse L, Attuquayefio D, Owusu E, Gbogbo F. 2007. Bird species diversity and abundance in the Abiriw Sacred Grove in the Eastern Region of Ghana. West Afr J Appl Ecol 11:38–44. Kankam BO. 1997. The population of black-and-white colobus (Colobus polykomos) and the mona monkeys (Cercopithecus mona) at the Boabeng-Fiema Monkey Sanctuary and surrounding villages [BSc thesis]. Kumasi (Ghana): University of Science and Technology. 71p. Kavanagh M. 1980. Invasion of the forest by an African savannah monkey: behavioural adaptations. Behaviour 73:238–260. Kingdon JS. 1980. The role of visual signals and face patterns in African forest monkeys (guenons) of the genus Cercopithecus. Trans Zool Soc Lond 35:425–475. Sacred Monkeys in Nigeria / 585 Kormos R, Boesch C, Bakarr MI, Butynski TM, editors. 2003. West African chimpanzees. Status survey and conservation action plan. IUCN/SSC Primate Specialist Group. Gland (Switzerland) and Cambridge: IUCN. ix1p 219. Available at: http://www.primate-sg.org/PDF/WACAP.English.pdf. Kortlandt A. 1986. The use of stone tools by wild-living chimpanzees and earliest hominids. J Hum Evol 15:77–132. Lebbie A, Guries RP. 2008. The role of sacred groves in biodiversity conservation in Sierra Leone. In: Sheridan MJ, Nyamweru C, editors. African sacred groves: ecological dynamics and social change. Oxford: James Currey Ltd. p 42–61. Madeweya KH, Oka H, Matsumoto M. 2004. Sustainable management of sacred forests and their potential for ecotourism in Zanzibar. Bull Forestry Forest Prod Res Inst 3:33–48. Mayor M, Lehman SM. 1999. Conservation of Perrier’s sifaka (Propithecus diadema perrieri) in Analamera Special Reserve, Madagascar. Lemur News 4:21–23. McKone D. 1995. A brief survey of the traditional forest reserves of Rungwe District, Mbeya Region, Tanzania. Unpublished draft report. Government of Tanzania/EEC Agroforestry, Soil and Water Conservation Project, Mbeya and District Forestry Office, Rungwe District. Available at: http://www.mckone.org/tfrrung.html. Mgumia FH, Oba G. 2003. Potential role of sacred groves in biodiversity conservation in Tanzania. Environ Conserv 30:259–265. Mittermeier RA. 1987. Effects of hunting on rain forest primates. In: Marsh CW, Mittermeier RA, editors. Primate conservation in the tropical rain forest. New York: Alan R. Liss, Inc. p 109–146. Mwihomeke ST, Msangi TH, Mabula CK, Ylhaisi J, Mndeme KCH. 1998. Traditionally protected forests and nature conservation in the North Pare Mountains and Handeni District, Tanzania. J East Afr Nat Hist 87:279–290. National Population Commission. 2007. Schedule II. Fed Rep Nigeria Official Gazette 94:B179–B198. Oates JF. 1989. A survey of primates and other forest wildlife in Anambra, Imo and Rivers States, Nigeria. Report to National Geographic Society (USA), Nigerian Conservation Foundation, Nigerian Federal Department of Forestry, and Governments of Anambra, Imo and Rivers States. 41p. Oates JF. 1996. Habitat alteration, hunting and the conservation of folivorous primates in African forests. Aust J Ecol 21:1–9. Oates JF, Anadu PA. 1988. The status of two guenon monkeys (Cercopithecus erythrogaster and Cercopithecus sclateri) in Nigeria. Report to World Wildlife Fund (USA) and Nigerian Conservation Foundation. 40p. Oates JF, Anadu PA. 1989. A field observation of Sclater’s guenon (Cercopithecus sclateri Pocock, 1904). Folia Primatol 52:93–96. Oates JF, Anadu PA, Gadsby EL, Werre JL. 1992. Sclater’s guenon—a rare Nigerian monkey threatened by deforestation. Natl Geogr Res Explor 8:476–491. Oates JF, Bergl RA, Linder JM. 2004. Africa’s Gulf of Guinea forests: biodiversity patterns and conservation priorities. Washington, D.C.: Conservation International, Center for Applied Biodiversity Sciences. 91p. Okpoko PU. 2001. Harnessing the tourism potentials of sacred groves and shrines in southeast Nigeria. West Afr J Archaeol 31:93–114. Olaniran OJ, Sumner GN. 1989. A study of climatic variability in Nigeria based on the onset, retreat, and length of the rainy season. Int J Climatol 9:253–269. Ormsby A. In press. Cultural and conservation values of sacred forests in Ghana. In: Pungetti G, Oviedo G, Hooke D, editors. Sacred species and sites: advances in biocultural conservation. Cambridge: Cambridge University Press. Osemeobo GJ. 1994. The role of folklore in environmental conservation: evidence from Edo State, Nigeria. Int J Sust Dev World Ecol 1:48–55. Pocock RI. 1904. Description of a new species of spot-nosed monkey of the genus Cercopithecus. Proc Zool Soc Lond 1:433–436. Redford KH. 1992. The empty forest. BioScience 42:412–422. Riley BW, Brokensha D. 1988. The Mbeere in Kenya. Vol. 1. Changing rural ecology. Lanham (MD): University Press of America. 365p. Riley EP. 2005. Ethnoprimatology of Macaca tonkeana: the interface of primate ecology, human ecology, and conservation in Lore Lindu National Park, Sulawesi, Indonesia [PhD dissertation]. Athens (GA): University of Georgia. 189p. Rudran R, Fernandez-Duque E. 2003. Demographic changes over thirty years in a red howler population in Venezuela. Int J Primatol 24:925–947. Sabater Pi J, Groves C. 1972. The importance of higher primates in the diet of the Fang of Rio Muni. Man 7:239–243. Saj TL, Sicotte P. 2007. Predicting the competitive regime of female Colobus vellerosus from the distribution of food resources. Int J Primatol 28:315–336. Saj TL, Teichroeb JA, Sicotte P. 2005. The population status of the ursine colobus (Colobus vellerosus) at Boabeng-Fiema, Ghana. In: Paterson JD, Wallis J, editors. Commensalism and conflict: the human-primate interface. Norman (OK): American Society of Primatologists. p 350–375. Saj TL, Mather C, Sicotte P. 2006. Traditional taboos in biological conservation: the case of Colobus vellerosus at the Boabeng-Fiema Monkey Sanctuary, Central Ghana. Soc Sci Inform 45:285–310. Sicotte P, Macintosh AJ. 2004. Inter-group encounters and male incursions in Colobus vellerosus in Central Ghana. Behaviour 141:533–553. Sicotte P, Teichroeb JA, Saj TL. 2007. Aspects of male competition in Colobus vellerosus: preliminary data on male and female loud calling, and infant deaths after a takeover. Int J Primatol 28:627–636. Sokpon N, Ago EE. 2001. Sacralisation et niveau de maturation des forets denses semi-decidues du plateau Adja au sudouest du Benin. J Recherche Scientifique l’Universite de Lome 5:319–331. Soury A. 2007. Sacred forests: a sustainable conservation strategy? The case of sacred forests in the Ouémé Valley, Benin [MSc thesis]. Wageningen: Wageningen University. 141p. Southwick CH, Malik I, Siddiqi MF. 2005. Rhesus commenalism in India: problems and prospects. In: Paterson JD, Wallis J, editors. Commensalism and conflict: the humanprimate interface. Norman (OK): American Society of Primatologists. p 240–257. Struhsaker TT, Leland L. 1988. Group fission in redtail monkeys (Cercopithecus ascanius) in the Kibale Forest, Uganda. In: Gautier-Hion A, Bourliere F, Gautier JP, Kingdon J, editors. A primate radiation: evolutionary biology of the African guenons. Cambridge: Cambridge University Press. p 364–388. Tashiro Y, Idani G, Kimura D, Bongori L. 2007. Habitat changes and decreases in the bonobo population in Wamba, Democratic Republic of the Congo. Afr Study Monogr 28:99–106. Teichroeb JA, Saj TL, Paterson JD, Sicotte P. 2003. Effect of group size on activity budgets in Colobus vellerosus in Ghana. Int J Primatol 24:743–758. Tooze Z. 1994. Does sacred mean secure? Investigation of a sacred population of Sclater’s guenon (Cercopithecus sclateri) in southeast Nigeria. Report to the Wildlife Conservation Society, New York. 14p. Tooze Z. 1995. Update on Sclater’s guenon Cercopithecus sclateri in southern Nigeria. Afr Primates 1:38–42. Am. J. Primatol. 586 / Baker et al. Ubah CN. 1982. The supreme being, divinities, and ancestors in Igbo traditional religion: evidence from Otanchara and Otanzu. Africa 52:90–105. Uchendu VC. 1965. The Igbo of southeast Nigeria. New York: Holt, Rinehart, and Winston. 111p. Virtanen P. 2002. The role of customary institutions in the conservation of biodiversity: sacred forests in Mozambique. Environ Values 11:227–241. Wheatley BP. 1999. The sacred monkeys of Bali. Prospect Heights (IL): Waveland Press, Inc. 189p. Whittaker DJ. 2006. A conservation action plan for the Mentawai primates. Primate Conserv 20:95–105. Windfelder TL, Lwanga JS. 2002. Group fission in red-tailed monkeys (Cercopithecus ascanius) in Kibale National Park, Uganda. In: Glenn ME, Cords M, editors. The guenons: diversity and adaptation in African Am. J. Primatol. monkeys. New York: Kluwer Academic/Plenum Publishers. p 147–159. Wittiger L, Sunderland-Groves JL. 2007. Tool use during display behavior in wild Cross River gorillas. Am J Primatol 69:1307–1311. Wolfe LD. 2002. Rhesus macaques: a comparative study of two sites, Jaipur, India, and Silver Springs, Florida. In: Fuentes A, Wolfe LD, editors. Primates face to face: the conservation implications of human–nonhuman primate interconnections. Cambridge: Cambridge University Press. p 310–330. Wong SNP, Sicotte P. 2006. Population size and density of Colobus vellerosus at the Boabeng-Fiema Monkey Sanctuary and surrounding forest fragments in Ghana. Am J Primatol 68:465–476. Wong SNP, Sicotte P. 2007. Activity budget and ranging patterns of Colobus vellerosus in forest fragments in Central Ghana. Folia Primatol 78:245–254.