Temporal Evolution of Bushmeat Traded in High Niger National Park, Guinea, West Africa

Temporal evolution of bushmeat traded in High Niger National Park, Guinea, West Africa

L UCIE D UONAMOU,ALEXANDRE K ONATE,JILIANG X U and T ATYANA H UMLE

Abstract The High Niger National Park is one of the most Introduction important protected areas for biodiversity conservation in Guinea. This study examined the temporal evolution of the unting, which was once considered a subsistence and bushmeat trade in three rural markets in the Park and in Htraditional activity supplying protein resources for the nearest urban centre, Faranah. We collected data in mar- rural populations in West Africa, has become an important  kets during August–November  in three villages around commercial activity (Bowen-Jones, ). Bushmeat hunting the Mafou core area of the Park and in Faranah, and com- is pervasive across West Africa and is one of the main threats   pared these data with equivalent published data from the to mammal populations in the region (Fa et al., , ;   same rural areas in  and  and from Faranah in Lindsey et al., ; Humle & Konate, ; Ripple et al.,   , ,  and . Across all study periods, mammals ). Unsustainable hunting practices are affecting c. % predominated in the bushmeat trade. In rural markets we of wildlife species in West and Central Africa, and are direct-   noted a marked increase in the number of carcasses and bio- ly affecting c. mammalian species (Bowen-Jones, ). ’ mass offered for sale from  onwards, whereas in Faranah Depletion of wildlife is increasingly affecting people s well- there were no differences over time other than a peak in . being and access to sources of protein and revenue, especially  Overall, there was an increase in the sale of smaller sized in rural areas (Nasi et al., ). To date most bushmeat stud- species (,  kg), and a marked increase in the sale of species ies have been carried out in forested areas (van Velden et al.,  that forage on crops, including the green monkey Chloroce- ); few have focused on savannah-dominated ecosystems ’ bus sabaeus and warthog Phacochoerus africanus,inspiteof in West Africa, especially within Guinea s dry forest systems  religious taboos against the consumption of primates and (Lindsey et al., ). It is critical to gather knowledge from Suidae. Green monkeys were not sold in markets during the such landscapes, to better understand the impact of hunting s but were the dominant species in Faranah in  and on wildlife and wildlife depletion on ecosystem integrity, as ’ . Our findings suggest a marked shift in traded species, well as on people s livelihoods and well-being in less forested associated with crop protection by farmers and economic in- and more arid regions of sub-Saharan Africa. centives to kill and trade crop-foraging species. This study Studies in Africa generally suggest that urban demand is  highlights the value of a longitudinal perspective for investi- a major driver of the bushmeat trade (Brashares et al., ;  gating the dynamic relationship between local livelihoods Fa et al., ). However, other factors such as poverty and and biodiversity conservation. human population growth also influence this trade (Lindsey et al., ;Pimmetal.,; Ripple et al., ). Farmers in Keywords Bushmeat trade, crop-protection, Guinea, High many regions also practice lethal wildlife control using snares Niger National Park, human–wildlife conflict, hunting, and, increasingly, shotguns, to protect their crops, although it market surveys, primate conservation remains unclear to what extent such mitigation strategies fuel  Supplementary material for this article is available at the bushmeat trade (Alexander et al., ;Humle&Konate,  doi.org/./S ). Such practices often target rodent, primate and Suidae species, which are amongst the most common crop-foraging species in Africa (Brooks et al., ;McNamaraetal.,; Garriga et al., ). In West Africa, primates account for –% of carcasses recorded in bushmeat markets (Davies LUCIE DUONAMOU ( orcid.org/0000-0002-6912-5923) School of Nature &Brown,). Wildlife authorities in Uganda have declared Conservation, Beijing Forestry University, Beijing, China baboons Papio sp., the vervet monkey Chlorocebus pygery- ALEXANDRE KONATE ( orcid.org/0000-0003-4896-8929) Institute Superior of Agronomy and Veterinary of Faranah, Faranah, Guinea thrus and red river hog Potamochoerus porcus to be vermin and farmers can therefore hunt these species to reduce dam- JILIANG XU (Corresponding author, orcid.org/0000-0002-2542-6441) School  of Nature Conservation, Beijing Forestry University, Beijing 100083, 35 ages and losses to their crops (Naughton-Treves et al., ). Qinghua East Road, Haidian District, China. E-mail [email protected] Bushmeat consumption studies in Lalehum near the Gola TATYANA HUMLE (Corresponding author, orcid.org/0000-0002-1919-631X) North Forest Reserve in Sierra Leone showed that the greater Durrell Institute of Conservation and Ecology School of Anthropology and cane rat Thryonomys swinderianus was more frequently killed Conservation, Marlowe Building, University of Kent, Canterbury CT2 7NR, UK E-mail [email protected] and consumed during the so-called hungry season, when rice Received  May . Revision requested  September . is growing, whereas duikers were hunted and consumed Accepted  December . First published online  September . more during the dry season (Davies & Brown, ). Although

This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, Downloadeddistribution, from https://www.cambridge.org/core and reproduction in any medium,. IP address: provided 170.106.33.22 the original work, onis 02 properly Oct 2021 cited. at 17:10:01, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/termsOryx, 2021, 55(5), 717–724 ©. https://doi.org/10.1017/S0030605319001443The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319001443 718 L. Duonamou et al.

hunting is an important source of income for many rural Kouroussa, Dabola and Faranah (Brugière et al., ). populations in Africa (Wilkie et al., ), some species are Circa , people live around the Park, with , of buffered from such practices as they are not hunted because these in the Mafou area near the urban centre of Faranah, of their cultural value, or because of religious beliefs (Reuter dominated by Malinke people, who are Muslim (Brugière et al., ; MacCarthy, ). For example, in the sin et al., ). Islam as practised in the region traditionally Lalehum in Sierra Leone, the western chimpanzee Pan troglo- prohibits the consumption of Suidae and primate species dytes verus was the species most commonly cited as taboo to (Mbotiji, ). Mean annual rainfall is ,–, mm eat and was not hunted for meat (Davies & Brown, ). and mean monthly temperature – °C, with a maxi- High Niger National Park is one of the last remaining dry mum of  °C at the end of the dry season in April–May forest–savannah mosaics in West Africa, and is a priority site (Fleury-Brugière & Brugière, ; Ziegler et al., ). for the conservation of ungulates and the western chimpanzee Agricultural activities in the region are concentrated within (Fleury-Brugière & Brugière, ;Kormosetal.,). In the Park’s buffer zones. Rice, maize, peanuts and cassava are Guinea, subsistence hunting of wildlife requires a permit the dominant crops and most people are engaged in farm- and so, in theory, offtake could be regulated and sustainable ing. The Park has two core areas, where human settlement (Saïdou & Djellouli, ). Nevertheless, many protected or activity are strictly prohibited, and two buffer zones  areas across West and Central Africa, including in Guinea, (c. , km ) that vary in the degree to which local people are affected by human activities such as hunting, agricul- are permitted to harvest and utilize resources. The two  ture and mining, and these activities remain poorly regulated strictly protected core areas are Mafou ( km ) and  (Brugière & Magassouba, ). National statistics on the Kouya ( km ), separated by  km (Brugière et al., bushmeat trade are not available for Guinea, but local-scale ). We focused our study on the Mafou sector, which studies have indicated that bushmeat is the main source of has benefitted from most protection efforts since . animal protein for rural people, as in many other African A collaboration between the Chimpanzee Conservation countries (Humle & Konate, ). Commercial hunting for Center, the Park authorities and the United Nations bushmeat is a serious threat to animal species across many Office for Project Services has facilitated the deployment protected areas as a result of a lack of law enforcement and of trained ecoguards in and around the Mafou core area unsustainable hunting practices. The failure to comply with (Fig. ). The Park harbours c. % of all known mammal hunting periods recommended by law and poor respect for species in Guinea (Ziegler et al., ; Brugière & the prohibition of hunting in strictly protected core areas Magassouba, ), including the Vulnerable lion Pan- therefore threaten the sustainability of wildlife resources thera leo and Critically Endangered western chimpanzee (Dia, ). (Kamyk, ; Humle et al., ). There were studies of bushmeat use in the Park during – (Ziegler, )andin (Fleury-Brugière &  Brugière, ). Both these studies focused on market surveys, Methods revealing that the Park harbours a diverse mammalian fauna that is threatened by hunting, but did not seek to understand Data on bushmeat hunting were collected during August– the drivers of the bushmeat trade or resource utilization. The November  in three rural villages (Sidakoro, Mansir- existing regulations for the control of bushmeat trade in this amoribaya and Koumandi Koura) in the Mafou sector of area remain poorly enforced and protection efforts are incon- the Park and in the urban centre of Faranah. The three vil- sistent across the Park (L. Duonamou & A. Konate, pers. obs., lages each have a small market in which bushmeat is sold. ). Here we explore how the bushmeat trade has evolved Data collection was based on market surveys, for comparison since these earlier studies, using data gathered in  (Humle and consistency with previous studies (Ziegler, ;Brugière & Konate, )and. We focus on the species hunted, & Magassouba, ). For each bushmeat item, we counted the number of carcasses recorded, biomass harvested, and the number of carcasses and identified the species. For esti- the body size of hunted species and whether they are crop- mating biomass, we used the mean body weight of each spe- foragers. We aim to provide critical information for the over- cies (Kingdon, ). This was also used to categorize species due update of the Park’smanagementplan(–)and by weight (, , – and .  kg). Based on local knowl- help to understand local drivers of the bushmeat trade and edge, species were categorized as non-pest (not reported to alleviate current pressures on wildlife within the Park. to forage on crops), pest (forages on crops) or carnivore (reported to depredate livestock; Supplementary Table ). To evaluate the temporal evolution of the bushmeat Study area trade, we compared our data with those collected in previous studies. For the rural area, we compared our data to  The , km High Niger National Park lies in the north- those of Humle & Konate (), gathered in , and east Republic of Guinea across the Prefectures of Kankan, Brugière & Magassouba (), collected in  across

Oryx, 2021, 55(5), 717–724 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319001443 Downloaded from https://www.cambridge.org/core. IP address: 170.106.33.22, on 02 Oct 2021 at 17:10:01, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605319001443 Wildmeat sales 719

FIG. 1 The Mafou area of High Niger National Park indicating the location of the main urban centre, Faranah, and villages located within the Park, including the study villages Mansiramoribaya, Sidakoro and Koumandi Koura (adapted from Brugière & Magassouba, ).

the same three villages and during the same months Suidae, Numididae), and Bovidae, Hystricidae, Cercopithe- (August–November). Our data from the urban centre of cidae, Thryonomyidae and Suidae were the most commonly Faranah were compared to those of Humle & Konate recorded taxa in both areas. The Suidae included two spe- (), collected in , and Ziegler (), collected in cies: the red river hog Potamochoerus porcus and the wart- ,  and , during the same months (September– hog Phacochoerus africanus. The consumption of Suidae is November). prohibited by Islam and, although the red river hog was All statistical analyses were performed SPSS . (IBM already hunted and traded in –, the presence of Corp., Armonk, USA). We used a one-way ANOVA fol- the warthog in the markets was first recorded in ,and lowed by a least significance difference post hoc test to assess in rural areas the number of carcasses doubled from  to differences in mean monthly number of carcasses recorded  (Supplementary Table ). and biomass of hunted species in urban and rural areas For Faranah, there was a significant difference in the mean  across years. A χ test was used to explore associations monthly number of carcasses recorded across years (one- between frequencies of carcasses by weight and wildlife way ANOVA F(,)=.,P=.), and the least signifi-  categories across years. For χ tests with more than a  ×  cant difference post hoc test showed that the mean monthly contingency design, z-scores based on the adjusted stan- number of carcasses recorded in  was significantly great- dardized residuals were used to assess the cells’ contribution er than in ,  and , but not than in  (Fig. a). to any significant results, with values .|.| indicating sig- For the three villages, the mean monthly number of carcasses nificance at P , .. Statistical significance for all other in  was significantly greater than in  and  (one- inferential statistics was at P , .. way ANOVA F(,)=.,P=.) but there were no significant differences between  and  (Fig. b). The monthly mean biomass exhibited the same patterns and sta- Results tistical differences across study periods in Faranah (except that a significantly greater biomass was recorded in both The market surveys revealed that birds, mammals and rep-  and  than in other years) and the villages (one- tiles were traded. We recorded a total of , carcasses way ANOVA, Faranah: F(,)=.,P=.;one-way of  wild animal species in  families, corresponding to ANOVA, villages: F(,)=.,P=.; Fig. ). , kg of biomass in the three villages and Faranah Overall, seven species were the most traded, comprising (Supplementary Tables  & ). Taxa of eight families were . % of all carcasses recorded in both rural or urban settings. recorded in both urban and rural areas (Bovidae, Hystrici- Six of these species were traded in both urban and rural dae, Canidae, Cercopithecidae, Nesomyidae, Thryonomyidae, areas: the greater cane rat, red-flanked duiker Cephalophus

FIG. 2 Monthly mean number of carcasses recorded in (a) urban and (b) rural areas (Fig. )over and  years, respectively. Bars marked with the same letter are not significantly different (P . .).

FIG. 3 Monthly mean biomass of carcasses recorded in (a) urban and (b) rural markets (Fig. )over and  years, respectively. Bars marked with the same letter are not significantly different (P . .).

FIG. 4 The most common species (as per cent of total carcasses) recorded in (a) urban and (b) rural markets (Fig. )over and  years, respectively.

rufilatus, blue duiker Cephalophus maxwelli,bushbuck The occurrence of the baboon decreased markedly in Faranah Tragelaphus scriptus, baboon Papio sp. and green monkey after  (Fig. a), and was only recorded in rural markets in Chlorocebus sabaeus (Fig. ). The patas monkey Erythrocebus . In Faranah (Fig. a), the greater cane rat and the red- patas never exceeded % of all carcasses recorded in rural flanked duiker were the most abundant species in  and areas, but comprised % of all carcasses recorded in Faranah , whereas the green monkey was the most abundant spe- in . The green monkey was never reported in Faranah dur- cies in  and . In the rural areas (Fig. b), the red- ing –, but it was the most abundant species in  and flanked and blue duikers were the most abundant species in . Although its presence in rural markets was not as prom-  but the greater cane rat and the bushbuck were the inent, its relative abundance increased there from  to . most abundant in  and .

FIG. 5 Number of carcasses by species weight category in (a) urban and (b) rural markets (Fig. ) over  and  years, respectively. The signs indicate where the observed frequency is significantly more (+) or less (-) than the expected frequency, based on standardized residuals and z-score values.

FIG. 6 Per cent of carcasses in three species categories (pest, non-pest and carnivore) in (a) urban and (b) rural markets (Fig. )over and  years, respectively. The signs indicate where the observed frequency is significantly more (+) or less (-) than the expected frequency, based on standardized residuals and z-score values.

The number of carcasses in each species weight cate- significantly more than expected in  but significantly gory varied significantly between study years in both Fa- less in . The carcasses of pest species comprised .%  ranah (χ ()=.;P, .) and the three villages of all species sold in Faranah in , significantly more than  (χ ()=.;P, .; Fig. ). Although all weight cat- expected (Fig. a). Carnivore species were traded in Faranah egories of species were traded as expected in  in Faranah, as expected in ,  and  but significantly less than in  fewer species ,  kg and more species .  kg were expected in  and  (Fig. a). In rural areas, non-pest traded than expected. In  similarly significantly fewer species were traded significantly more than expected in species ,  kg and more species weighing – kg were , and pest species significantly less. However, in  traded than expected (Fig. a). In  and , however, pest species were traded significantly more than expected there was a shift, with significantly more species ,  kg, in villages, and there was no significant difference between fewer species weighing – kg, and species .  kg traded observed and expected values in , although these species as expected (Fig. a). In rural areas in  species ,  kg comprised .% of all recorded carcasses in  (Fig. b). and .  kg were traded less than expected, and species No significant differences were noted for carnivore species weighing – kg were traded significantly more than ex- in rural areas. pected (Fig. b). In  and  small and large species were traded as expected and medium sized species were  traded significantly less than expected (Fig. b). However, Discussion overall, the majority of species were small in  and  in both Faranah and the villages (Fig. ). Our findings reveal that mammalian species comprise the There was a significant association in the frequency of greatest proportion of wildlife traded in the study region, carcasses recorded across years and wildlife categories corroborating studies carried out elsewhere in West and  (non-pest, pest and carnivore) in Faranah (χ ()=., Central Africa (Fa et al., ). In the urban centre of  P , .) and the rural areas (χ ()=.,P, .). Faranah, the number of carcasses and offtake of biomass In Faranah, the carcasses of non-pest species were traded were similar in  and , despite peaks in  and

. However, bushmeat trade in the three rural areas in- religious taboos against the consumption of these taxa. Tra- creased from  to  whether based on carcass num- ditionally, a religious taboo prohibiting the consumption bers or biomass. This finding is at odds with previous of Suidae species amongst Muslims in the region mainly studies indicating that bushmeat demand is driven by in- focused on the warthog rather the red river hog. These spe- creased demand from urban markets (Fa et al., ). cies are, however, increasingly been killed to protect crops T. Humle & F. St John (unpubl. data) uncovered evidence and, whereas in the past the meat was either donated to that many urban residents in Faranah preferred meat of do- non-Muslims or given to dogs, farmers now gain econom- mestic animals to bushmeat. However, demand from other ically from their sale, providing an additional incentive for urban centres in the forest region of Guinea and the role of killing (L. Duonamou et al., unpubl. data). Warthog meat in intermediaries connecting villages with urban traders fur- particular is also now been termed the ‘meat of the youth’ ther afield may still be important drivers of the bushmeat in rural areas, where the younger Muslim generation is dis- trade and wider regional and transborder trade patterns; creetly preparing and consuming this meat (L. Duanomou such influences need to be better understood. The absence & A. Konate, pers. obs., ). In addition, primate meat is of an increase in the bushmeat trade in Faranah since  also often supplied to local restaurants in Faranah, where could potentially reflect a depletion of wildlife species, al- people tend not to question the type of meat that is being though sales in the villages increased between  and served and where taboos are not respected as fully as in . This could also indicate that recent law enforcement the household (A. Konate, unpubl. data). initiatives, with road blocks at critical exit points from the Our findings also indicated a significant shift in the dom- Park, are successfully discouraging transport of bushmeat inant species for sale in  compared with earlier years. to the city. However, the increase in the bushmeat trade in Although the red-flanked and blue duikers were amongst rural areas could also be linked to an increase in the human the most traded species during – in both rural population combined with limited access to alternative and urban areas, they were markedly less evident in  sources of protein such as beef and chicken, which are sel- and . In addition, although the cane rat remained the dom sold in villages (T. Humle & F. St John, unpubl. data). most abundantly traded species in rural areas, its trade de- Although we noted a decrease in carnivore and non-pest creased markedly in  and  in the urban centre of species being hunted over time, our findings indicated a Faranah. This shift may be related to increased demand significant increase in the trading of species that forage on in rural areas for the greater cane rat, the meat of which crops, in both rural and urban areas. Crop-damaging species is highly appreciated and readily available (A. Konate & are increasingly being traded in bushmeat markets across L. Duonamou, pers. obs., ), and a shift in consumption rural West Africa (e.g. in Ghana: Alexander et al., ). of bushmeat species such as primates, warthog and red river Farmers also use such bushmeat to pay off loans and/or se- hog, which are not traditionally consumed in villages, and cure additional income for agricultural activities, including hence preferentially transported to and traded in urban the purchase of herbicides and fertilizers, which are now areas. The decline in duiker species may indicate these spe- widely used locally (A. Konate, pers. obs., ). The major- cies have declined as a result of unsustainable hunting. In ity of hunters in the three study villages practice farming,  and , these species were replaced by the green and social science surveys have indicated that farm pro- monkey in the urban centre and by species such as the tection is a prime driver for hunting (L. Duonamou et al., Gambian pouched rat Cricetomys gambianus in the rural unpubl. data). Primate species, such as baboons, green areas (Supplementary Table ). The bushbuck was the and patas monkeys, and Suidae species, are often cited as only species whose relative abundance remained relatively being the most common crop-foraging species in rural consistent across years in both urban and rural settings, areas (L. Duonamou et al., unpubl. data). Our findings re- suggesting that the population of this species has remained vealed a general increase in the diversity of primate species stable despite hunting. hunted, with the exception of the chimpanzee, which was Although large-bodied species such as the bushbuck only recorded once, in , and a substantial intensification were relatively constant in their presence in the markets in the sale of green monkeys in the urban centre of Faranah. across years, our study revealed a significant shift in more Our market surveys also indicated a marked increase in the recent years from medium-sized species towards small spe- sale of red river hogs and warthogs, especially in villages, cies (,  kg) in both rural and urban settings. Market sur- since . Even if some people in Faranah consume mon- veys in some West African countries have similarly revealed key and wild pig meat, sale and consumption are more a marked shift towards smaller species such as rodents, common and widespread in south-eastern Guinea where generally reflecting a depletion in larger mammals linked people are predominantly animists; Faranah and surround- to unsustainable levels of hunting (Cowlishaw et al., ). ing villages have become a source of supply of these spe- Our study highlights temporal changes in bushmeat cies for this region of Guinea (T. Humle, pers. obs., ). available for sale across rural and urban settings, sheds However, such hunting practices are evidently challenging light on the dynamics of the bushmeat trade in and around

the Mafou area of the High Niger National Park, and pro- References vides some insights into the drivers of this trade. We focused, however, on only one area of the Park and it remains ALEXANDER, J.S., MCNAMARA, J., ROWCLIFFE, J.M., OPPONG,J.& MILNER-GULLAND, E.J. () The role of bushmeat in a West unclear whether the patterns we observed are the same African agricultural landscape. Oryx, , –. in other prefectures, such as Kouroussa. Nevertheless, our BOWEN-JONES,E.() Bushmeat: traditional regulation or findings suggest that, contrary to patterns noted elsewhere, adaptation to market forces. In Trade in Wildlife: Regulation for especially in Central Africa, urban demand does not appear Conservation (ed. S. Oldfield), pp. –. Earthscan, London, UK. to be driving the bushmeat trade in this region, especially as BRASHARES, J.S., GOLDEN, C.D., WEINBAUM, K.Z., BARRETT, C.B. &  supply in rural areas has increased steadily, whereas trade in OKELLO, G.V. ( ) Economic and geographic drivers of wildlife consumption in rural Africa. Proceedings of the National Academy the local urban center of Faranah remained the same as it of Sciences of the United States of America, , –.  was in . Our findings also suggest that agricultural ex- BROOKS, J.E., BRUGGERS, R.L. & HARRIS, M.A. () Vertebrate pansion, crop protection practices and the economic status Damage Control Research in Agriculture, Fiscal Years  and . of farmers require greater consideration when addressing Agency for International Development, Washington, DC, USA.  the bushmeat trade in and around the High Niger Na- BRUGIÈRE,D.&MAGASSOUBA,B.( ) Mammalian diversity in the National Park of Upper Niger, Republic of Guinea—an update. tional Park. In addition, there appears to be an erosion in Oryx, , –. religious taboos regarding the killing and consumption of BRUGIÈRE,D.&MAGASSOUBA,B.() Pattern and sustainability primates and Suidae species. 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