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This cultural expression is part of the Guineans culture and it is still practised nowadays, involving however a more reduced presence in the “Sacred Woods” (about three months). It’s a collective ceremony and aims at the transmission of reserved knowledge, fortifying, as well, the cohesion and community unity (Campredon 1997). 3 The armed conflict begun in the south, in January of 1963 (Simpósio 2008). 4 “Nhinte Camatchol” is a Nalu sculpture used at fanado. It represents a bird’s head with a human face, being the main message of this ritual to the participants that they must consider true brotherhood, a collective interest affirmation, above individual or familiar interest. 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Using Genetics as a Tool in Primate Conservation | Learn Science at Scitable 12/11/28 20:59 | Lead Editor: Sigurdur Greipsson CONSERVATION AND RESTORATION Using Genetics as a Tool in Primate Conservation By: Maria J. Ferreira da Silva (Cardiff University, CIBIO-Porto University), Tania Minhos (Cardiff University, CESAM-Lisbon University), Rui M. Sa (Cardiff University, New University of Lisbon, CRIA) & Michael W. Bruford (Cardiff University) © 2012 Nature Education Citation: Ferreira da Silva, M. J., Minhos, T., Sa, R. M. & Bruford, M. W. (2012) Using Genetics as a Tool in Primate Conservation. Nature Education Knowledge 3(10):89 "Here I am, in the middle of this forest, and I can't see the primates? How can I learn more about the species I am studying?" This question can be answered with the help of recent advances in non-invasive molecular genetics. The current conservation status and decline of nonhuman primates is alarming (Norconk et al. 2011) due to factors such as disease, habitat destruction, hunting, illegal trade and climate change (WWF 2010). Unfortunately, their restricted geographical range, resource requirements, long lifespan and slow reproduction rate, dispersal needs, and degree of specialization, make primates more susceptible to extinction than many other species (Cowlishaw & Dunbar 2000, Harcourt et al. 2002, Gibbons & Harcourt 2009). Moreover, the geographical range of threatened primate species often overlaps with areas of high human density (Harcourt & Parks 2003). Recent studies conclude that 48% of primate species are in danger of becoming extinct, making well-informed conservation measures crucial for ensuring their long-term survival (Mittermeier et al. 2009). Aside from their contribution as models for human evolutionary research, their value to ecotourism, and their charisma, primate species play important ecological roles, especially as seed dispersers (Chapman & Russo 2007). Some studies suggest that maintaining this role could be important for ecosystem resilience because monkeys and apes visit trees in social groups where they tend to stay longer during feeding periods than other mammals or birds (Lambert 2011). http://www.nature.com/scitable/knowledge/library/using-genetics-as-a-tool-in-primate-59120330 Page 1 of 11 Using Genetics as a Tool in Primate Conservation | Learn Science at Scitable 12/11/28 20:59 Figure 1: Elusive animals. Guinea baboons (Papio hamadryas papio) live in dense forested habitats and are very hard to observe. By using molecular tools, primatologists can gather information about this elusive species. © 2012 Nature Education Photo courtesy of Chimbo Foundation. All rights reserved. Non-Invasive Genetic Analysis The relatively recent development of non-invasive genetics has allowed primatologists to better understand the population and group dynamics of wild primates, simply by the fact that it is now possible to obtain genetic information by extracting DNA from by-products such as feces, shed hair, and urine. The first study of this kind was in chimpanzees (Pan troglodytes) in which Morin et al. (1993) analyzed patterns of gene flow in the Gombe chimpanzee community (Tanzania). Since then, major technical improvements in non-invasive genetics have greatly expanded our capacity to address a wide range of questions about the structure of primate populations, their evolutionary histories, and adaptation, while allowing the study of wild populations without direct contact with the animals (Charpentier et al. 2007, Tung et al. 2008). More importantly, the combination of genetics with long-term socio-ecological data has enabled comprehensive analysis at an individual and social group level for a wide range of primate species. DNA obtained non-invasively can be analyzed in individuals and populations for a wide range of molecular genetic markers such as microsatellites, minisatellites, mitochondrial DNA, amplified fragment length polymorphism (AFLP) and the major histocompatibility complex (MHC). By using a variety of software to analyze the genetic data produced, primatologists can now obtain information on effective population size, parentage, relatedness, sex, dispersal, population structure, population assignment and gene flow. This is crucial if we are to fully understand population dynamics at a local scale and evaluate the threats and suggest appropriate conservation measures (Goossens & Bruford 2009). Threats to Primate Conservation Habitat fragmentation Habitat fragmentation can influence several key features of primate populations: 1) their diet, 2) the social group size or density, and 3) the dispersal and gene flow between social groups or subpopulations (Marsh 2003, Frankham et al. 2002). Consequently, the capacity for populations to persist in fragmented landscapes is related to a species' particular characteristics (Marsh 2003) (Figure 2). It is theoretically possible for primate populations to increase in size within fragments-for example, if natural predators disappear (Strier 2007)-but more frequently, they decrease or become extinct. This can be due either to direct mortality, caused by an increased hunting pressure, since isolated areas become accessible for humans (Marsh 2003, Strier 2007), or due to genetic changes (Frankham et al. 2002) (Figure 3). In the long term, fragmentation can lead to a reduction in genetic diversity and increased genetic differentiation. This results from the decrease of gene flow between breeding groups and the action of random genetic drift and/or inbreeding (Frankham et al. 2002). http://www.nature.com/scitable/knowledge/library/using-genetics-as-a-tool-in-primate-59120330 Page 2 of 11 Using Genetics as a Tool in Primate Conservation | Learn Science at Scitable 12/11/28 20:59 Figure 2: Behavioral adaptation to habitat fragmentation. Black-and-white colobus are almost exclusively arboreal. Fragmentation may force the adaptation to new habitats or result in local extinctions. © 2012 Nature Education All rights reserved. Figure 3: Habitat loss and deforestation. Deforestation is happening at an accelerated rate and can be caused by logging, collection of non-timber forest products, and fires. It not only promotes isolation of populations but also increases hunting pressure and contributes to climate change. © 2012 Nature Education All rights reserved. Each population fragment may show different levels of genetic diversity and significantly different allele frequencies from the other fragments. The risk of inbreeding depression is increased if the population is smaller and isolated, with lower genetic diversity. Migration of individuals between fragments and subsequent reproduction will introduce new alleles into the population (increasing genetic diversity) and it will counterbalance the effects of genetic drift and inbreeding, preventing complete fixation of alleles (Frankham et al. 2002). By using non-invasive genetic methods it is possible to identify the genetic structure of a fragmented population, and levels of gene flow between units, and determine whether ecological corridors should be created/maintained, or individuals should be translocated (e.g., Bruford et al. 2010). http://www.nature.com/scitable/knowledge/library/using-genetics-as-a-tool-in-primate-59120330 Page 3 of 11 Using Genetics as a Tool in Primate Conservation | Learn Science at Scitable 12/11/28 21:00 In the fragmented range of the Cross River Gorilla (Gorilla gorilla diehli) three sub-populations have been uncovered using microsatellite markers (Bergl & Vigilant 2007). Although this genetic structure corresponds broadly to the pattern of habitat fragmentation, migrants between fragments could be identified. Since different levels of genetic diversity were found between the sub-populations, it was suggested that the conservation of the most genetically diverse sub-population should be prioritized. Also, habitat corridors between fragments, along with measures to control hunting in areas between fragments, were recommended (Bergl et al. 2008). The Bornean orang-utan (Pongo pymaeus), living in forest fragments of the Lower Kinabatangan flood plain in Sabah, Malaysia, shows a different pattern: high levels of heterozygosity within fragments, with a relative scarcity of rare alleles, suggesting that this population was large in the past and has suffered a recent major reduction (Goossens et al. 2005). Goossens et al. (2006), using extensive non-invasive sampling across the area and 14 microsatellite loci, showed that the Bornean orang-utans population has decreased in size by 95% over the last decades or centuries, due to anthropogenetic fragmentation of the habitat. Therefore, the high genetic diversity found is transitory and may disappear if forest corridors alongside the riverbank are not established (Bruford et al. 2010). Hunting The impact of hunting pressure on primate populations is often difficult to evaluate. Although information on the amount of harvested primates can be obtained by counting carcasses in urban bushmeat markets, morphological identification can be hindered if a carcass has been processed, or if the meat has been smoked (Figure 4). Figure 4: Bushmeat markets. Hunting of primates is occurring at very high rates. Primate meat is consumed in rural areas for subsistence and in urban centers as a delicacy. It is the result of an illegal, organized trade. © 2012 Nature Education All rights reserved. Primatologists can use molecular PCR-based tools to taxonomically identify unknown specimens. After extracting DNA and amplifying a specific DNA fragment, these fragments can then be compared with other DNA fragments obtained from specimens of known species. The comparison can also be accomplished by verifying the presence/size of the fragment after PCR: for a review of the techniques see Fajardo et al. (2010). In many cases however, the researcher might not have access to specimens of known species. To overcome this difficulty, it is necessary to amplify a standard gene fragment that can be compared with fragments from voucher species deposited in public databases. A fragment of 648bp from the mitochondrial cytochrome c oxidase (COI) gene was proposed by Hebert et al. (2003) as a standard fragment for DNA barcoding, the data for which are deposited in the Barcode of Life Database (http://www.barcodeoflife.org/) as well as in public databases such as GenBank. Lorenz et al. (2005) tested the use of this mitochondrial DNA region to identify the species of primate samples. All samples, representing 56 primate species, amplified with at least one of the 3 different primers used and, with few exceptions, the fragments obtained clustered together with sequences retrieved from GenBank (Figure 5). More recently, Rönn et al. (2009) proposed the use of a micro-array system to assign samples of primates to the genus level, using both http://www.nature.com/scitable/knowledge/library/using-genetics-as-a-tool-in-primate-59120330 Page 4 of 11 Using Genetics as a Tool in Primate Conservation | Learn Science at Scitable 12/11/28 21:00 nuclear and mitochondrial genes. This technique uses 111 diagnostic nucleotide positions to perform a hierarchical assignment of samples. This method can be used to process a large number of samples at a relatively low cost, and 45 out of the 64 samples were correctly assigned to their Primates genus. http://www.nature.com/scitable/knowledge/library/using-genetics-as-a-tool-in-primate-59120330 Page 5 of 11 Using Genetics as a Tool in Primate Conservation | Learn Science at Scitable 12/11/28 21:00 Figure 5: Molecular identification of bushmeat. Morphological identification can be difficult if carcasses have been processed. Molecular identification is the easiest and most reliable tool available. © 2012 Nature Education All rights reserved. Diseases Disease is another important aspect for primate conservation. The Ebola and anthrax outbreaks that have occurred in Central Africa in recent decades caused a dramatic decline in gorilla and chimpanzee populations (Leendertz et al. 2006, Bermejo et al. 2007, Campbell et al. 2008). Additionally, recent studies on parasite infection dynamics have demonstrated an association with hunting, human population growth, and fragmentation in wild primates (Gillespie & Chapman 2006, Goldberg et al. 2007, Gillespie et al. 2008, Riley & Fuentes 2011). With the incorporation of molecular approaches to epidemiology, Johnston et al. (2010) have demonstrated cross species transmission of Giardia duodenalis between humans, livestock, and wild primates in Western Uganda. Likewise, Goldberg et al. (2009) discovered three novel retroviruses in red colobus monkeys, shedding light on the dynamics of primate retroviral transmission. More recently, Yildirim et al. (2010) unveiled the gut microbial community of three nonhuman primate species by sequencing the small subunit rRNA unit from fecal samples, allowing future analysis on comparative and evolutionary studies of human gut microbes and other primates. Furthermore, using an innovative method that combines a single-genome amplification of Plasmodium sp. recovered non-invasively from fecal material of great apes, Liu et al. (2010) inferred that the origin of the human malignant malaria Plasmodium falciparum is gorilla-derived. This result argues against the previous study from Prugnolle et al. (2009) that showed that P. falciparum emerged from P. reichenowi by a single transfer from chimpanzees. Similarly, HIV/AIDS is the result of a cross-species transmission event of simian immunodeficiency virus (SIV) to humans from non-human African primates, and much attention has been paid to the understanding of the evolutionary history of these emerging infection diseases (Gao et al. 1999, Damond et al. 2004, Liu et al. 2008). By using a molecular dating technique, Wertheim and Worobey (2009) estimated a surprisingly recent common ancestor of infectious SIV in chimpanzees (between 1266 to 1685 years ago) and sooty mangabeys (between 1729 to 1875 years ago), the reservoirs of HIV-1 and HIV-2, respectively. Conversely, human transmitted pathogens to great apes such as bacteria (e.g., Streptococcus pneumoniae) or viruses (e.g., human metapneumovirus) are causing fatal respiratory outbreaks (Chi et al. 2007, Kaur et al. 2008, Köndgen et al. 2008, 2011, Palacios et al. 2011) and to mitigate the risk of disease transmission the use of face masks by researchers, tourists and staff is advocated as a good practice (Macfie & Williamson 2010). These studies emphasize the fact that there is much to be learned concerning disease transmission and its implications for wild primates using molecular tools. Figure 6: Primate parasites. a) Entamoeba sp. and b) Strongylida fam. gen. sp. (Hookworms) are gastrointestinal parasites that can cause severe infections and weaken the health of the host. © 2012 Nature Education All rights reserved. Applying Conservation Genetics Primate census The abundance and density of wild primate populations are key parameters for assessing their conservation status and management (Arandjelovic et al. 2010). Biomonitoring and molecular censusing allows the determination of population size, as well as individual movements in the landscape (Storfer et al. 2007, Vigilant & Guschanski 2009). Guschanski et al. (2009), using a panel of 16 microsatellite loci, estimated that the population size of the endangered mountain gorillas (Gorilla beringei beringei) was http://www.nature.com/scitable/knowledge/library/using-genetics-as-a-tool-in-primate-59120330 Page 6 of 11 Using Genetics as a Tool in Primate Conservation | Learn Science at Scitable 12/11/28 21:00 10% less when compared to the classical nest-count methods. All molecular census estimates in primates have shown a population size smaller than previously accessed by traditional methods. In contrast, Zhan et al. (2006), comparing the numbers of traditional survey methods with molecular censusing, demonstrated that the DNA-based estimate for a well-studied giant panda (Ailuropoda melanoleuca) population was more than double the ecological estimate.When using a capture-recapture analysis for census purposes, Arandjelovic et al. (2010) recommended that three times more samples should be collected than the predicted population size for apes when assuming a closed population model. Therefore, molecular surveys provide a complementary method to more traditional census approaches. "Evolutionary significant units" (ESUs) and "management units" (MUs) ESUs and MUs are two types of conservation units described using genetic information: ESUs have been defined as needing to be reciprocally monophyletic mitochondrial lineages (i.e., occupying different branches in a phylogenetic tree) and requires long-term historical population differentiation, whereas MUs are identified based on current demographic isolation (i.e., no current or recent gene-flow), evidenced by differences in allele frequency distributions and significantly different frequencies for both mitochondrial and nuclear loci (Moritz 1994). Although the criteria to identify these units have been subject to debate (e.g., see Paektau 1999), such definitions can be key indicators to preserve genetic distinctiveness (evolutionary heritage, genetic diversity and differentiation). For example, Kanthaswamy et al. (2006) suggested that the Bornean and Sumatran orangutans should be considered two distinct MUs based on the analysis of mtDNA and microsatellite loci, and consequently the authors advised against the inter-island translocation of animals. Population and habitat viability analysis Population and habitat viability analysis (PHVA) evaluates the risk of extinction within a certain period of time (e.g., 100 or 200 years) and identifies which factors play a major role in the extinction process. PHVA relies on stochastic modeling by using simulation software, such as VORTEX (Miller & Lacy 2005), and requires the input of parameters on the ecology and life history of the species (e.g., population size, mortality and birth rates, sex ratio, dispersal rates, and main threats to the habitat) to be able to simulate (by Monte Carlo iterations) species responses that are realistic. Molecular census and genetic data can also be very important parameters for PHVA. Moreover, it allows the introduction of different and combined management measures (e.g., ecological corridors, reintroduction, translocations, habitat rehabilitation) and simulates the evolution of the species under such interventions. This tool allows conservationists to detect the major threats for rare and endangered species and thereby help implement the most long- term viable conservation actions. Bruford et al. (2010) incorporated the genetic data of 200 orangutans from the Kinabatangan floodplain in Sabah, Malaysia, to study the implications of non-intervention, translocation, corridor establishment, and a combination of the latter two measures, on the future genetic diversity of this highly fragmented population. They found that non- intervention would result in the extinction of some of the subpopulations within five generations, and that translocation or corridor establishment alone would not be sufficient to prevent high levels of inbreeding. Instead, a combination of the two measures would retain the demographic stability of even the most isolated subpopulations and constrain localized inbreeding to a sustainable threshold (Figure 7). http://www.nature.com/scitable/knowledge/library/using-genetics-as-a-tool-in-primate-59120330 Page 7 of 11 Using Genetics as a Tool in Primate Conservation | Learn Science at Scitable 12/11/28 21:00 Figure 7: Population viability analysis. The table shows the result of demographic and genetic parameters of the vortex simulation for 25 to 250 years under a non-intervention scenario. Note the mean extinction time (column TE) for most of the sub-populations (PSU) is less than 250 years. P(E): probability of extinction; Pop: population; GD: genetic diversity (from Bruford et al. 2010). © 2012 Nature Education All rights reserved. Summary The extensive use of molecular techniques as tools has provided new opportunities to better understand the mechanisms underlying the evolution and adaptation of primates (Figure 8). By integrating genetic and ecological data into simulation models, conservation predictions will be more accurate, and long-term conservation strategies will be more effective. http://www.nature.com/scitable/knowledge/library/using-genetics-as-a-tool-in-primate-59120330 Page 8 of 11 Using Genetics as a Tool in Primate Conservation | Learn Science at Scitable 12/11/28 21:00 Figure 8: Survival of endangered species. Interspecific association between western Red and Black-and-white colobus (Cantanhez National Park, Guinea Bissau). Prospects for the survival of these endangered species will benefit from the use of non-invasive genetic tools. © 2012 Nature Education All rights reserved. All authors have contributed equally to this article. References and Recommended Reading Arandjelovic M. et al. 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Outline | Keywords Explore This Topic BASIC INTERMEDIATE Conservation of Biodiversity Population Viability Analysis: Origins Conservation Biology and Contributions Restoration Ecology Applications of Decision Theory to Conservation Planning and Conservation and Restoration Management Introduction Notes from the Field: A Phytoremediation Primatologist's Point of View ADVANCED Using Genetics as a Tool in Primate Conservation http://www.nature.com/scitable/knowledge/library/using-genetics-as-a-tool-in-primate-59120330 Page 11 of 11 S H O R T C O M M U N I C A T I O N 36.513 km2, which corresponds to 33 individuals (Sousa, 2009). However, the exact number of individuals and communities for the whole CNP and the rest of the country remains unclear; with the aid of a molecular census, however, it will be possible to infer its effective population size (Sá et al., 2009). Anthropogenic disturbances such as habitat loss and fragmentation (e.g. logging activities and shifting land occupation for the purposes of agriculture and food production, e.g. cashew nuts), the hunting of infant animals for the pet trade, and casual deaths from crop raiding allied to extrinsic factors such as disease, are the main threats, not only to Chimpanzees but to all non- human primates in Guinea-Bissau (Gippoliti et al., 2003; DAVID LAWSON / WWF-UK / LAWSON DAVID Casanova and Sousa, 2007; Brugière et al., 2009). The The Trade and Ethnobiological Use of species is classified by IUCN as Endangered, and listed in CITES Appendix I, and is also protected in Guinea- Chimpanzee Body Parts in Guinea-Bissau: Bissau. Even though most primate species in Guinea- Implications for Conservation Bissau are traded for meat consumption, there is no evidence that this is the case for Chimpanzees (Minhos Rui Miguel Moutinho Sá, Maria Ferreira da Silva, et al., in prep.). Fernando Miguel Sousa and Tânia Minhós This paper reports on the use and trade of Chimpanzee body parts in Guinea-Bissau for traditional practices (e.g. for nutritional, medicinal or ritual purposes, or “animistic INTRODUCTION myths”). Informal interviews were conducted and observations made with a view to providing insight into uinea-Bissau represents the western-most limit of how these human traditions and myths might pose an the endangered West African Chimpanzee Pan additional threat. troglodytes verus (Sousa et al., 2005). During G ETHODS the 1980s, Chimpanzees were erroneously considered to M be extinct in the country due to a total absence of information owing largely to political and civil unrest Seven visits, of approximately four hours each, were (Lee et al., 1988). In 1994, a preliminary survey was made to Bandim market, the largest market in Bissau, the conducted and the presence of Chimpanzees was capital, during two weeks in September 2008 and a reconfirmed (Gippoliti and Dell’ Omo, 1995; 1996). similar period in June 2010. Some 10–15 men were More recently, research has been carried out in co- found to be offering wild animal body parts for sale (e.g. operation with national and local authorities, establishing skin, bones, teeth, horns and scales). Where possible, a system for the systematic monitoring and management morphological identification of the specimens viewed of this great ape (Casanova and Sousa, 2007). Within the was made and photographs taken. country, Chimpanzees are distributed across the south of An ethnoprimatological approach (i.e. the study of the Corubal River. Their presence is confirmed in two human and non-human primate interactions) aims to protected areas—Cantanhez National Park (CNP) and understand the incorporation of non-human primates into Cufada Lagoons Natural Park (CLNP) in the folklore, myths, the hunting of non-human primates for southwestern region—and in the eastern region of Boé food, keeping non-human primates as pets, indigenous (Casanova and Sousa 2007; Brugière et al., 2009). knowledge of non-human primate behaviour, among Due to high levels of exploitation, loss of habitat and others (Wolfe and Fuentes, 2007; Fuentes and Hockings, habitat quality as a result of human activities, this 2010). In this study, the authors were interested in subspecies is estimated to have experienced a significant understanding and placing into context the social population reduction in the past 20 to 30 years (IUCN, inclusion of Chimpanzee body parts for human traditional 2011). However, no recent data are available to allow for practices using informal interviews and ethnographic an estimation of rates of decline (IUCN, 2011). The most observations, although not enough data were collected to recent figures available, from 1996 (Gippoliti et al., provide an in-depth analysis for such an approach. 2003), estimate that the number of Chimpanzees in Most of the vendors encountered were male. Five Guinea-Bissau ranges from between 600 and 1000 urban vendors in Bandim market and 17 rural informants individuals. It is estimated that Chimpanzee density in in villages in the CNP and the Boé region were informally the southern area of CNP is of 2.34 nest builders/km2 in a interviewed following an unstructured script, in order to total area of 17.225 km2, corresponding to 40 individuals document the geographical origin and use of Chimpanzee (Sousa et al., 2011), while in the neighbouring area east body parts, prices and the scale of the trade, i.e. whether of Gadamael, just outside the CNP area, this value at a national, regional, or transnational level. Direct decreases to 0.89 nest builders/km2 in a total area of observations of the trade were conducted in the market TRAFFIC Bulletin Vol. 24 No. 1 (2012) 31 S H O R T C O M M U N I C A T I O N 1. Leopard Panthera pardus (1a complete). CITES Appendix I/IUCN Near Threatened; 2. Chimpanzee Pan troglodytes verus. CITES Appendix I/IUCN Endangered; 3. Nile Crocodile Crocodylus niloticus. CITES Appendix I/II/IUCN Lower Risk/Least Concern. 4. Guinea Baboon Papio papio. CITES Appendix II/IUCN Near Threatened. 5. Possibly Lion Panthera leo. CITES I/IUCN Vulnerable. 6. Possibly African Civet Civettictis civetta. CITES Appendix III/IUCN Least Concern. 7. Antelope horns (species not identified). Fig. 1. All animal-derived products for human traditional R.M. SÁ, M. FERREIRA DA SILVA, F.M. SOUSA AND T. MINHÓS T. AND SOUSA F.M. SILVA, DA R.M. SÁ, M. FERREIRA purposes in Bandim market, Bissau, Guinea-Bissau. and field notes were taken. Informants were assured that All the species mentioned above are reported as the purpose of the work was not to condemn or report occurring in Guinea Bissau except for Olive Baboons, their practices to the local authorities. Every observation whose western limit of distribution is reported to be in heard and/or seen was recorded and notes/interviews Mali and the Republic of Guinea (IUCN, 2011). The organized into social demographic categories (e.g. urban Olive Baboon skin seen was morphologically quite traders, local villagers, gender, ethnic group). Only different to the Guinea baboon skins found at the market. information relevant to the research topic was assigned to While Guinea Baboon skins present red/brownish these categories (Rubin and Rubin, 1995). coloration, the Olive Baboon skin had a green hue, typical Only pieces of animal skins were seen for sale during of what has been described for the subspecies (Groves, the surveys (which could have derived from one or more 2001). specimens). As the vendors were reluctant to answer questions related to the animal numbers involved in the Costs, origin and scale trade, it was not possible to estimate the number of skins being offered for sale for each species recorded. This paper Interviews with urban traders revealed that the cost of a therefore focuses on the morphological identification of the piece of Chimpanzee skin was relatively high, ranging from species and not to the number of skins traded. XOF1500 (CFA Francs) to XOF90 000 (USD2.9 to USD173.96, based on an exchange rate in 2008 of RESULTS AND DISCUSSION XOF460.77 to USD1). The average monthly wage in 2008 was XOF40 000 (approximately USD88.00) (UNDP, 2010). Traded species All urban vendors reported that the Chimpanzee and other animal body parts (apart from the elephant hide During visits to Bandim market, morphologically seen) originated from the “southern part”, and frequently identified dried Chimpanzee skins were found being sold mentioned the regions of Cantanhez and Gabú specif- for traditional medicinal purposes. Additionally, dried skins ically. Vendors considered the “southern part” every from Temminck’s Red Colobus monkeys Procolobus location south of Bissau. The authors were told that the badius temminckii, Guinea Baboons Papio papio and Olive elephant hide had come from Senegal. According to Blanc Baboons Papio anubis were also found. The authors also et al., 2007, at least one, and at most 10 elephants remain detected trade in dried skins of several non-primate species in Senegal. Most vendors said that consumers were of such as Leopard Panthera pardus, Nile Crocodile both sexes, different ethnic groups and social status. Crocodylus niloticus, African Civet Civettictis civetta, It was apparent to the authors that witchdoctors are elephant Loxodonta sp., hare Lepus sp., African Buffalo not the only people to buy animal-derived products for Syncerus caffer, Spotted Hyaena Crocuta crocuta and traditional medicine or protection fetishes. For example, several species of antelopes, snakes and lizards, as well as according to statements from three vendors: skins alleged to be of Wild Dog Lycaon pictus and Lion “All sort of people buy. Men and women, poor or Panthera leo (Fig. 1). Other animal body parts observed rich… Fulas, Pepel, Balanta, even Europeans. Every included bones, Crested Porcupine Hystrix cristata spines, kind. Not only djamba kuss [witchdoctors] to please the teeth, antelope horns, pangolin Manis sp. scales, mollusc irans [magical and religious entities].” shells, fish bones and feathers. Morphologically specific According to Robillard, in litt. to TRAFFIC, July identification was not possible in most cases due to the 2011, it is common practice in Africa for people who are similarity of those body parts to other species, as well as to unwell to buy their own products based on a list provided their condition. A few sellers mentioned that some of the by the traditional doctor. See also Marshall (1998). bones being offered for sale were from primates. 32 TRAFFIC Bulletin Vol. 24 No. 1 (2012) S H O R T C O M M U N I C A T I O N Two of the vendors also mentioned that individuals Chimpanzee body parts in birth and circumcision rituals from neighbouring countries such as Senegal, Guinea or (Mallart Guimera, 1981). The Yoruba people of south- Gambia are involved in the trade within the country: western Nigeria believe in the magical properties of “Other foreigners also buy and sell their own plants, Chimpanzee body parts in appeasing witches and fortune shells or skins”. tellers (Dedeke and Aboyami, 2006). However, it is not easy for people to obtain these remedies or to gain access Symbolic and medicinal use to these animals. In Central Africa, the consumption of Chimpanzee meat is taboo for young men, pregnant women Most male informants in rural CNP and Boé villages and children (Robillard, in litt. to TRAFFIC, July 2011). associated the use of Chimpanzee-derived products with The presence in Bandim market of the skin of an Olive the needs of women, as revealed by one elder Fula Baboon suggests a foreign origin for some of the animal respondent in Béli, Boé: body parts being offered for sale. While the distribution “Dári [chimpanzee] is mezinho [traditional medicine] area for this species (Papio anubis)includes neighbouring of women.” Guinea and Mali, it does not occur in Guinea-Bissau Three Balanta women in CNP confirmed that (Soewu, 2008). The Guinea Baboon Papio papio is the Chimpanzee skin is used to: “prepare a cleansing mixture only baboon species reported and observed in the country against hideousness when they are pregnant or their (IUCN, 2011). There are striking differences in children are still babies in the event they see a lonely morphology between both baboon species (namely coat chimpanzee cross their way”. Likewise, another woman coloration (Groves, 2001)), which enables a distinction to said that “the leaves of the nest where a menstruating female be made based on their skins. Furthermore, in Colobane chimpanzee sleeps can be applied to heal mental problems”. and Boucotte markets in Senegal (in Dakar and One informant admitted that he uses a stitched amulet Ziguinchor, respectively), several species of reptiles and made of chimpanzee body parts to help provide awareness mammals, including primate species (data not shown) to protect him and his friends while in the bush (Rui Sá, were found in trade for use in traditional medicinal pers. obs., 2008). practices and/or magic ceremonies (Fernando Sousa, pers. obs., 2008). According to information provided by the Guinea-Bissau in the context of previous studies sellers, those animal body parts were brought from Niger, Nigeria, Ivory Coast and Mali. Chimpanzee skins were One possible explanation for the lack of information also found in these Senegalese markets (Fernando Sousa, on magic practices and traditional medicines using animal pers. obs., 2008). The respondents pinpointed Cassamance body parts in Guinea-Bissau is the difficulty in collecting (on the border between Senegal and Guinea-Bissau) as the information on such an undisclosed subject, as well as putative origin of Chimpanzee skins at Boucotte market, both a lack of interest and in-depth study of such and Guinea-Bissau and the Republic of Guinea as the practices. As a result, the authors’ observations are possible origin of the Chimpanzee skins being sold at opportunistic. However, the use of animals’ body parts Colobane market. The possibility that the Chimpanzee for medicinal purposes could seriously threaten the skins found in Bandim market could also be from the biodiversity of Guinea-Bissau and, in particular, Republic of Guinea cannot be excluded since sellers constitutes an additional and significant threat to mentioned the “south” as the origin but not specifically Chimpanzee populations already menaced by habitat loss the south of Guinea-Bissau. and fragmentation, the pet trade and crop-raiding conflicts. Therefore, this phenomenon deserves to be Implications for conservation thoroughly investigated (Cá, 2008). Although not previously reported for Guinea-Bissau, The suggested transnational interest for Guinea-Bissau the use of non-human primate body parts in traditional Chimpanzee skins may constitute an even bigger threat medicine is not unusual elsewhere in the world (Alves et for the conservation of this population. Since Chimpanzee al., 2010; Leypey and Fomine, 2010). In a recent review, populations are declining in West African countries Alves et al. (2010) reported the use of 101 species of (IUCN, 2010), foreign hunters could be attracted to primates in folk/magic-religious practices, most Guinea-Bissau and the hunting of Chimpanzees could frequently in Africa, Latin America and Asia. Although therefore increase in the near future. Biodiversity Cercopithecidae species are the most affected, management authorities in Guinea-Bissau (IBAP and Chimpanzees are also referred to as a remedy for diseases Direcção Geral de Florestas e Fauna) have introduced and for use in folk medicine (Alves et al., 2010). In new laws to regulate the trade in wild meat (e.g. recently, Nigeria, Mali, Sierra Leone, Congo and Guinea, the hunting of primates throughout the country was Chimpanzee body parts are used to cure male impotency, prohibited (Anon., 2011). However, the lack of resources epilepsy, bone fractures and infertility in women (Dedeke and lack of awareness of management authorities and and Aboyami, 2006). In Cameroon, the Bakweri people politicians is hindering law enforcement in the country. believe that by using the liquid derived from boiled At the international level, conservation agencies should Chimpanzee bones, the bones of children or babies will re-examine their strategies to mitigate this trade, and, at become stronger (Leypey and Fomine, 2010). the national level, specific programmes should be Additionally, in the forested areas, people use designed and applied to empower all actors involved (e.g. TRAFFIC Bulletin Vol. 24 No. 1 (2012) 33 S H O R T C O M M U N I C A T I O N park rangers, Customs officers, the military, police, etc.), Gippoliti, S., and Dell’Omo, G. (1995). Status and conservation complemented at the same time by provision of environ- of the chimpanzee Pan troglodytes verus in Guinea-Bissau. mental education for the local communities. African Primates 1:3–5. Further work by the authors will include the molecular Gippoliti, S., and Dell’Omo, G. (1996). Primates of the Cantanhez Forest and the Cacine Basin, Guinea-Bissau. determination of the origin of the skins observed in the Oryx 30:74–80. markets and of the species involved. This will assist in Gippoliti, S., Embalo, D., and Sousa, C. (2003). Guinea-Bissau. evaluating the scale of the trade. Finally, an ethnographic In: West African Chimpanzees: Status Survey and Conser- study specifically centred on the use of non-human vation Action Plan. Kormos, R., Boesch, C., Bakarr, M., and primate body parts by traditional medicine using more in- Butynski, T. (Eds), IUCN, Gland, Switzerland. Pp.55–61. depth techniques, such as participant observation or long- Groves, C., (2001). Primate Taxonomy. Smithsonian Institution term observation, will allow the authors to draw up Press, Washington, DC. possible differences in the use of distinct animal parts and IUCN (2010). IUCN Red List of Threatened Species. Version determine how such practices are disseminated. 2010.4. www.iucnredlist.org. Viewed on 21 April 2011. Lee, P.C., Thornback, J. and Bennett, E.L. (1988). Threatened Primates of Africa. Gland, Switzerland. ACKNOWLEDGEMENTS Leypey F., and Fomine M. (2010). Edible Wild Animals as Source of Medication in Cameroon. Petits Propos The authors are enormously thankful to the Institute of Culinaires. 90:99–117. Biodiversity and Protected Areas (IBAP) in Guinea-Bissau for Mallart Guimera, L. (1981). Ni dos ni ventre. Religion, magie facilitating their research in Guinea-Bissau, and for providing et sorcellerie Evuzok. Paris: Société d’ethnographie. logistical support. Particular thanks go to Milza Nanqui whose Marshall, N.T. (1998). Searching for a Cure: Conservation of assistance facilitated the gathering of information at the market Medicinal Wildlife Resources in East and southern Africa. in Bissau and to all the informants for agreeing to and trusting TRAFFIC International. 112 pp + xiv. the authors to talk about sensitive issues. The authors are also Minhos, T., Sá, R., Ferreira da Silva, M., Wallace, E., Barata, very grateful to M.W. Bruford, C. Sousa and C. Casanova for A., Carmo, M. and Bruford, M.W. (in prep.). Characteri- their helpful discussion and comments that improved the quality zation of the urban primate bushmeat trade in Guinea Bissau of the manuscript. The authors would also like to thank David using molecular tools for species identification. Greer, WWF’s African Great Apes Programme Co-ordinator; Rubin, H.J. and Rubin, I.S. (1995). Qualitative Interview: The Marine Robillard, expert in environmental anthropology, Art of Hearing Data. 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Microbiol., 0(0), 2011 pp. 1–3 © 2011 The Author(s) Journal of Eukaryotic Microbiology © 2011 International Society of Protistologists DOI: 10.1111/j.1550-7408.2011.00586.x Distribution of the Entodiniomorphid Ciliate Troglocorys cava Tokiwa, Modry´ , Ito, Pomajbı´ kova´ , Petrzˇ elkova´ , & Imai, 2010, (Entodiniomorphida: Blepharocorythidae) in Wild and Captive Chimpanzees KATERˇ INA POMAJBI´ KOVA´ ,a KLA´ RA J. PETRZˇ ELKOVA´ ,b,c JANA PETRA´ Sˇ OVA´ ,a ILONA PROFOUSOVA´ ,a,b BARBORA KALOUSOVA´ ,b MILOSLAV JIRKU˚ ,d RUI M. SA´ e,f and DAVID MODRY´ d,g aDepartment of Pathology and Parasitology, Faculty of Veterinary Medicine, University of Veterinary and Pharmaceutical Sciences, Palacke´ho 1-3, 612 42, Brno, Czech Republic, and bInstitute of Vertebrate Biology, Academy of Sciences of the Czech Republic, Kveˇtna´ 8, 60365, Brno, Czech Republic, and cLiberec Zoo, Masarykova 1347/31, 460 01, Liberec, Czech Republic, and dBiology Centre, Institute of Parasitology, Academy of Sciences of the Czech Republic, Branisˇovska´ 31, 370 05, Cˇeske´ Budeˇjovice, Czech Republic, and eBiodiversity and Ecological Processes Group, School of Biosciences, Cardiff University, Cardiff, United Kingdom, and fAnthropology Department, Human and Social Sciences Faculty, Universidade Nova de Lisboa, Portugese, and gCEITEC – Central European Institute of Technology, University of Veterinary and Pharmaceutical Sciences, Brno, Czech Republic ABSTRACT. Trophozoites of Troglocorys cava were detected in all but one of the wild chimpanzee populations from Rubondo Island (Tanzania), with a prevalence ranging between 20% and 78%. However, the ciliate was absent in all captive groups. Preva- lence appeared to increase with the number of sequential samples taken from a particular individual and reached 95.5% in wild individuals sampled at least 4 times. Key Words. Chimpanzees, Entodiniomorphida, intestinal ciliates, prevalence, Troglocorys cava. HE intestine of apes is colonized by a spectrum of pro- Chimpanzee Sanctuary, Kenya (n = 42). In addition, the T tists that cause no or little harm to their host and can be numbers of samples from Ugalla and Rubondo Island NP were classified as mutualists or commensals. Among them, ciliates extended to 119 and 206, respectively. Samples were preserved are the most prominent group, as they apparently participate by 10% formalin and examined by merthiolate-iodine-formal- in the hindgut fermentation (e.g. Profousova´ et al. 2010). dehyde sedimentation (MIFC), following Pomajbı´ kova´ et al. Reports about the presence of intestinal ciliates in wild apes (2010). were mostly a by-product of parasitological examinations We calculated the prevalence of T. cava differently for (Ashford, Reid, and Wrangham 2000; Murray et al. 2000). habituated (chimpanzees are used to being around people) and However, the proper classification of ciliates requires special- unhabituated (the opposite to previous) populations. For a ized techniques other than those used for the routine copro- habituated population, the prevalence was expressed as the scopic detection of parasites, thus representing a serious percentage of individuals infected with T. cava; for unhabitu- drawback in their research (Imai et al. 1991). ated populations, as the percentage of samples positive for Most intestinal ciliates of great apes belong to the order T. cava. To determine the effect of sequential sampling on the Entodiniomorphida and are classified in the genera Troglody- prevalence of T. cava, we calculated the cumulative prevalence tella, Gorillophilus and Prototapirella. In addition, a species for selected animals from Kalinzu Forest Reserve. We chose that had been referred to as a “small” or “unidentified ciliate” samples only from the wet season (n = 195) and calculated the (Ashford et al. 2000; Bakuza and Nkwengulila 2009; Murray cumulative prevalence based on number of samples per indi- et al. 2000) was named Troglocorys cava and assigned to the vidual: (i) one sample (n = 36 animals); (ii) two sequential family Blepharocorythidae (Tokiwa et al. 2010); the study samples (n = 29); (ii) three sequential samples (n = 25); and aimed to investigate the occurrence of T. cava in both wild (iv) four or more sequential samples (n = 22). and captive chimpanzees. Herein, we report on the prevalence, For scanning electron microscopy (SEM), trophozoites of diagnostics, and geographical distribution of T. cava in popu- T. cava from a chimpanzee from Kalinzu FR were obtained lations of wild chimpanzees and discuss the absence of T. cava from a filtrated fecal sediment and were preserved with 10% in captive chimpanzees in European facilities, African sanctu- formalin, following the protocol of Jirku˚ et al. (2009). aries, and in an introduced chimpanzee population in Tanza- nia. RESULTS AND DISCUSSION MATERIAL AND METHODS The general trophozoite morphology, determined by light microscopy and confirmed by NIC, was uniform throughout The set of fecal samples of wild (n = 511) and captive chimpan- all localities and consistent with the original description of zees (n = 203) largely corresponds with those discussed by T. cava (Tokiwa et al. 2010) (Fig. 1). Pomajbı´ kova´ et al. (2010). It also includes two additional wild The geographical distribution of T. cava includes localities chimpanzee populations and one captive facility: Goualougo in Tanzania, Uganda, and Gabon (e.g. Krief et al. 2005; Triangle, Republic of Congo (DRC) (n = 20); Cantanhez Landsoud-Soukate, Tutin, and Fernandez 1995; Murray et al. National Park (NP), Guinea Bissau (n = 102); and Sweetwaters 2000). We detected T. cava in chimpanzee fecal samples from seven of the eight studied wild populations and extended its known distribution in Uganda and Tanzania, and added local- Corresponding Author: Katerˇ ina Pomajbı´ kova´ , Department of ities in Nigeria, DRC, and Guinea Bissau. Its prevalence in Pathology and Parasitology, Faculty of Veterinary Medicine, Uni- versity of Veterinary and Pharmaceutical Sciences, Palacke´ ho 1-3, wild populations ranged from 5% to 98% (Ashford et al. 612 42 Brno, Czech Republic—Telephone number: +420541562269; 2000; File, McGrew, and Tutin 1976; Krief et al. 2005; FAX number: +420541562266; e-mail: [email protected] Landsoud-Soukate et al. 1995; Murray et al. 2000). Our 1 2 J. EUKARYOT. MICROBIOL., 0, NO. 0, SEPTEMBER 2011 Fig. 1–7. Diagnostic features of Troglocorys cava revealed by light microscopy (1–4) and scanning electron microscopy (5–7) from a formalin- fixed fecal sample from Pan troglodytes schweinfurthii, Kalinzu Forest Reserve, Uganda. 1. Right side of the trophozoite showing frontal lobe (arrowhead) surrounded by adoral ciliae. 2. Trophozoite showing vestibulum (asterisk). 3. Left concavity (arrow) in dorso-ventral view. 4. Left side of the trophozoite showing the concavity clearly demarcated by a hexagonal line formed by bases of lips (white arrow). 5. Right side of two trophozoites showing typical surface longitudinal striation and both adoral (bottom) and left cilia (above). Arrowhead indicates position of frontal lobe hidden by adoral ciliae. 6. Dorso-ventral view showing asymmetrical shape of the trophozoite. The concavity, invisible in this view, is located on the flat left side. 7. Dorso-lateral view of the trophozoite showing the prominent left concavity possessing small round projections on its surface. Scale bars = 20 lm. Fig. 1–4 and 5–7, respectively, are in the same scale. results fell within this range: Budongo FR, Ugalla – 20%; tive prevalence. Prevalence of T. cava reached 55.5% in indi- Cantanhez NP – 25%; Gashaka Gumti NP – 33%; Kyambura viduals sampled only once, 79.3% in individuals sampled George – 40%; Goualougo Triangle – 65%; and Kalinzu FR twice, 92% in individuals sampled 3 times, and 95.5% if four – 78%. In contrast, we did not find T. cava in any captive and more samples were collected. These results conform to chimpanzees. Troglocorys cava was absent also in the wild other studies suggesting that ideally three or four samples per population of chimpanzees on Rubondo Island, having been animal are necessary to assess the real prevalence of a particu- introduced onto the island in the 1960s using founders that lar parasite/symbiont (Huffman et al. 1997; Muehlenbein had spent some period in captivity (Petrzˇelkova´ et al. 2010). 2005). Therefore, we predict that T. cava is present in all indi- Thus, T. cava apparently disappears during the captivity; this viduals in wild chimpanzee populations, a pattern documented is confirmed by its total absence in zoos and sanctuaries. previously for T. abrassarti (Pomajbı´ kova´ et al. 2010). Despite These findings are in contrast to the common occurrence of the fact that none of previous studies included either morpho- Troglodytella abrassarti in captive chimpanzees (Pomajbı´kova´ logical descriptions or microphotographs, we assume that also et al. 2010). Since nothing is known about the biology of other findings of the “small ciliate” in chimpanzees by previ- T. cava in hindgut ecosystem, its absence cannot be unambig- ous researchers refer to T. cava. uously explained. Generally, the populations of commensal ciliates fluctuate ACKNOWLEDGMENTS greatly, which was demonstrated both in rumen ciliates and T. abrassarti (Pomajbı´ kova´ 2008; Williams and Coleman This work could not have been done without the willingness 1991). As a result of this fluctuation, the numbers of T. cava and close collaboration with many zoos, localities in Africa, and in some fecal samples can be reduced below the threshold level local authorities. We are grateful to all keepers, curators, and of detection. To address this problem we examined serial fecal veterinarians, namely Liberec Zoo, CZ (David Nejedlo); samples from the same individual and calculated their cumula- Plzenˇ Zoo, CZ (Ivo Tetˇ a´ l); Ostrava Zoo, CZ (Jana Ka´ lnova´ ); POMAJBI´ KOVA´ ET AL.—DISTRIBUTION OF TROGLOCORYS CAVA IN CHIMPANZEES 3 Hodonı´ n Zoo, CZ (Jaroslav Hyja´ nek); Dvu˚r Kra´ love´ Zoo, CZ LITERATURE CITED (Zdena Jerˇ a´ bkova´ ); Brno Zoo, CZ (Stanislav Maza´ nek); Bratislava Zoo, SK (Martin Krug); Spisˇ ska´ Nova´ Ves, SK Ashford, R. W., Reid, G. D. F. & Wrangham, R. W. 2000. Intestinal (Karol Dzurik); Zoologischer Garten Augsburg, Germany parasites of the chimpanzee (Pan troglodytes) in Kibale Forest, (Wilhelm Moeller); Burger‘s Zoo En Safari Arnhem, Netherlands Uganda. Ann. Trop. Med. Parasitol., 94:173–179. (Wineke Schoo); Antwerp Zoo, Belgium (Francis Vercammen); Bakuza, J. S. & Nkwengulila, G. 2009. Variation over time in parasite Chester Zoo, UK (Stewe Unwin); Twycross Zoo, UK (Nie prevalence among free-ranging chimpanzees at Gombe National Park, Tanzania. Int. J. Primatol., 30:43–53. Masters); Zoo Aquarium Madrid, Spain (Eva Martinez File, S. K., McGrew, W. C. & Tutin, C. E. G. 1976. The intestinal Nevado); Parc Zoologic De Barcelona, Spain (Teresa Abello´ ); parasites of a community of feral chimpanzees, Pan troglodytes Dierenpark Amersfoort, Netherlands (Adrien van Zanten); schweinfurthii. J. Parasitol., 62:259–261. Zu¨ rich Zoo, Switzerland (Hanspeter W. Steinmetz); La Valle´ e Huffman, M. A., Gotoh, S., Turner, L. A., Hamai, M. & Yoshida, K. des Singes Zoo, Fr (Jan Vermeer); Zoologischer Garten Leip- 1997. Seasonal trends in intestinal nematode infection and medical zig, Germany (Klaus Eulenberger); La Palmyre Zoo, Fr (Be- plant use among chimpanzees in the Mahale Mountains, Tanzania. noit Quintard and Thierry Petit); Biomedical Primate Primates, 38:111–125. Research Centre, Netherland (Peter J. Heidt); Phoenix Park Imai, S., Ikeda, S. I., Collet, J. Y. & Bohnome, A. 1991. Entodinio- Dublin, Ireland (Sandra Devaney); Limbe Wildlife Centre, morphid ciliates from the wild lowland gorilla with description of a new genus and three new species. Europ. J. Protist., 26:270–278. Cameroon (Felix Lankester) and Sweetwaters Chimpanzee Jirku˚, M., Jirku˚, M., Obornı´ k, M., Lukesˇ , J. & Modry´, D. 2009. Sanctuary, Ol Pejeta, Kenya (Martin Mulama), who provided Goussia Labbe´ , 1986 (Apicomplexa, Eimeriorina) in Amphibia: us with samples of captive apes. We express our sincere thanks diversity, biology, molecular phylogeny and comments on the sta- to Nicol Simmons (Kyambura Gorge, Queen Elizabeth tus of the genus. Protist, 160:123–136. National Park, Uganda), Klaus Zuberbuhler, Zarin Machanda Krief, S., Huffman, M. A., Se´ venet, T., Guillot, J., Bories, C., Hladik, and Fred Babweteera (Budongo Conservation Field Station, C. M. & Wrangham, R. W. 2005. Non-invasive monitoring of the Uganda and Royal Zoological Society of Scotland), Alex Piel health of Pan troglodytes schweinfurthii in the Kibale National (Ugalla Primate Project, Tanzania), Crickette Sanz and David Park, Uganda. Int. J. Primatol., 26:467–490. Morgan (Goualougo Triangle Chimpanzee Project, Republic Landsoud-Soukate, J., Tutin, C. E. G. & Fernandez, M. 1995. Intesti- nal parasites of sympatric gorillas and chimpanzees in the Lope´ of Congo), and Volker Sommer (Gashaka Primate Project, Reserve, Gabon. Ann. Trop. Med. Parasitol., 89:73–79. Nigeria) for the collection of samples from wild apes. Our Muehlenbein, M. P. 2005. Parasitological analyses of the male chim- research in Kalinzu FR (Uganda) was approved by the panzees (Pan troglodytes schweinfurthii) at Ngogo, Kibale National Uganda National Council for Science and Technology, Park, Uganda. Am. J. Primatol., 65:167–179. Uganda Wildlife Authority and Uganda National Forest Murray, S., Stem, C., Boudreau, B. & Goodall, J. 2000. Intestinal Authority. We would like to acknowledge the help of Chie parasites of baboons (Papio cynocephalus) and chimpanzees (Pan Hashimoto, Takeshi Furuichi (PRI, Kyoto University, Japan; troglodytes) in Gombe National Park. J. Zoo. Wildl. Med., 31:176– supported by the Japanese Ministry of Environment Global 178. Environment Research Fund #F- 061). Our research on the Petrzˇelkova´ , K. J., Hasegawa, H., Appleton, C. C., Huffman, M. A., Archer, C. E., Moscovice, L. R., Mapua, M. I., Singh, J. & Kaur, T. Rubondo Island NP (Tanzania) was approved by the Tanza- 2010. Gastrointestinal parasites of the chimpanzee population nian Wildlife Institute, Tanzanian Commission for Science introduced into Rubondo Island National Park, Tanzania. Am. J. and Technology and Tanzania National Parks. We express Primatol., 72:307–316. our special thanks to the Rubondo Island NP management, Pomajbı´ kova´ , K. 2008. Entodiniomorph Ciliates of Gastrointestinal Mike A. Huffman (PRI, Kyoto University, Japan), Taranjit Tract of Great Apes [Diploma Thesis]. University of Veterinary Kaur and Jatinder Singh (Virginia Polytechnic Institute and and Pharmaceutical Sciences, Brno, Czech Republic. State University).). Our research in Cantanhez National Park Pomajbı´ kova´ , K., Petrzˇelkova´ , K. J., Profousova´ , I., Petra´ sˇ ova´ , J., in Guinea-Bissau was approved by IBAP (Biodiversity and Kisˇ idayova´ , S., Vara´ dyova´ , Z. & Modry´, D. 2010. A survey of Protected Areas Institute), whom we would like to acknowl- entodiniomorphid ciliates in chimpanzees and bonobos. Am. J. Phys. Anthropol., 142:42–48. edge as well as the CNP guards. This work was supported by Tokiwa, T., Modry´, D., Ito, A., Pomajbı´ kova´ , K., Petrzˇelkova´ , K. J. the project “CEITEC – Central European Institute of Tech- & Imai, S. 2010. A new entodiniomorphid ciliate, Troglocorys cava nology” (CZ.1.05/1.1.00/02.0068) from the European Regional n. g., n. sp., from the wild eastern chimpanzee (Pan troglodytes Development Fund, by the grants from the Grant Agency of schweinfurthii) from Uganda. J. Eukaryot. Microbiol., 57:115–120. the Czech Republic (524/06/0264 and 206/09/0927), the Williams, A. G. & Coleman, G. S. 1991. The Rumen Protozoa. Leakey’s Foundation, the Grant Agency of the Academy of Springer-Verlag New York Inc., New York. Sciences of the Czech Republic (KJB600930615), and Hlavka’s Foundation. Received: 08/02/2011; accepted: 08/31/2011