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The Lion in West Africa Is Critically Endangered

The Lion in West Africa Is Critically Endangered The African lion has declined to ,35,000 individuals occupying 25% of its historic range. The situation is most critical for the geographically isolated populations in West Africa, where the species is considered regionally endangered. Elevating their conservation significance, recent molecular studies establish the genetic distinctiveness of West and Central African lions from other extant African populations. Interventions to save West African lions are urgently required. However formulating effective conservation strategies has been hampered by a lack of data on the species’ current distribution, status, and potential management deficiencies of protected areas (PAs) harboring lions. Our study synthesized available expert opinion and field data to close this knowledge gap, and formulate recommendations for the conservation of West African lions. We undertook lion surveys in 13 large (.500 km ) PAs and compiled evidence of lion presence/absence for a further eight PAs. All PAs were situated within Lion Conservation Units, geographical units designated as priority lion areas by wildlife experts at a regional lion conservation workshop in 2005. Lions were confirmed in only 4 PAs, and our results suggest that only 406 (273–605) lions remain in West Africa, representing ,250 mature individuals. Confirmed lion range is estimated at 49,000 km , or 1.1% of historical range in West Africa. PAs retaining lions were larger than PAs without lions and had significantly higher management budgets. We encourage revision of lion taxonomy, to recognize the genetic distinctiveness of West African lions and highlight their potentially unique conservation value. Further, we call for listing of the lion as critically endangered in West Africa, under criterion C2a(ii) for populations with ,250 mature individuals. Finally, considering the relative poverty of lion range states in West Africa, we call for urgent mobilization of investment from the international community to assist range states to increase management effectiveness of PAs retaining lions. Citation: Henschel P, Coad L, Burton C, Chataigner B, Dunn A, et al. (2014) The Lion in West Africa Is Critically Endangered. PLoS ONE 9(1): e83500. doi:10.1371/ journal.pone.0083500 Editor: Matt Hayward, Bangor University, United Kingdom Received September 3, 2013; Accepted November 4, 2013; Published January 8, 2014 Copyright:  2014 Henschel et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: Field survey efforts were funded by Panthera (www.panthera.org), with additional funds from National Geographic’s Big Cat Initiative (grant B1_12; http://animals.nationalgeographic.com/animals/big-cats/). LC furthermore acknowledges the support of the Oxford Martin School, and DWM the Recanati-Kaplan Foundation and Robertson Foundation. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. * E-mail: phenschel@panthera.org West African lions represent a population with unique genetic Introduction and conservation value. Recent molecular and morphological data The lion (Panthera leo) was the most successful large carnivore covering the species’ entire historical range suggests that lions in during the late Pleistocene, when the species’ range extended from Central, West and North Africa (the latter now extinct) are distinct South Africa, across Eurasia, and into the southern United States from lions in Eastern and Southern Africa and share a common [1]. Today, the lion’s range is restricted to Africa and one ancestor with lions in Asia [6–8]. These results establish that the population of the Asiatic sub-species, P. l. persica, in India [2]. principal subdivision of modern lions is within Africa [9], and While the endangered Asiatic population is currently considered question the current dichotomous division into an African and an stable, lion populations in Africa are in decline and the African Asian sub-species; a division which is still widely supported, sub-species, P. l. leo, is considered vulnerable [3]. Recent analyses including by the IUCN Red List [3]. Moreover, they demonstrate established that the African lion has lost at least 75% of its original that lions in West Africa contain mtDNA haplotypes not found in habitat, with fewer than 35,000 wild African lions remaining [4]. other lion populations, elevating the conservation significance of The main drivers of lion declines are large-scale habitat the few remaining West African populations [8,9]. conversion, prey base depletion through unsustainable hunting, Conservation interventions to save these populations are now and the retaliatory killing of lions due to perceived or real human- urgently required. However, formulating interventions is limited lion conflict [3]. The situation is most critical in West Africa, by few field data on the species’ current distribution, abundance, where lions have been considered regionally endangered since and predominant drivers of declines in West Africa. While lions 2004 [5], and where ,500 individuals may persist [4]. have been the object of extensive research effort in parts of Eastern and Southern Africa, they have been largely ignored in West PLOS ONE | www.plosone.org 1 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa Africa. Of 463 articles on African lions in the ISI Web of presence for PAs where evidence of lion occurrence was lacking, TM Science (Thompson Scientific) in 2005, not one focused and 2) establish lion population estimates for PAs where lions specifically on lions in West Africa. To address this deficit, we occurred. Although designated LCUs included both protected and first collated available data and expert opinions on lion non-protected areas, lions are largely absent outside PAs in West distribution and status in West Africa. We then undertook field Africa [11,20]. Accordingly, we restricted fieldwork to PAs (IUCN surveys in 13 large (.500 km ) protected areas (PAs) where lions PA categories I–VI) within LCUs (henceforth: LCU PAs), were reported during this process, to determine lion presence/ including two PAs awaiting formal designation (Table 1). A absence and estimate lion population size. For a further eight large primary determinant of lion extinction risk within a given PA is its PAs suspected to harbor lions, we compiled field survey data from size, and we therefore restricted our survey effort to large the literature and via interviews. Using standardized evaluation (.500 km ) LCU PAs, based on findings on critical PA size for toolkits for protected area management effectiveness [10], we lions from prior studies [21,22]. In LCUs with multiple PAs, we compared current management performance of PAs known or concentrated survey efforts in the largest PAs with the highest suspected to harbor lions and those from which lions have likely protection status (according to IUCN PA categories). been extirpated. With these data, we provide a comprehensive We also compiled data on lion presence from recent field evaluation of the tenuous status of the West African lion, and make surveys led by other institutions, from internal reports and recommendations for the taxa’s conservation. interviews with participants. We included only surveys that targeted large mammals, with survey methods and effort Study area appropriate to detect lions. Finally, we incorporated data from We restrict our analysis to West Africa, as defined by the United interviews of PA staff on lion presence/absence for several LCU Nations geoscheme (http://unstats.un.org/unsd/methods/m49/ PAs that have not been recently surveyed. We did not consider m49regin.htm), including all countries from Senegal to Nigeria reports of lion presence without physical evidence and records (Fig. 1). The same classification was used for the listing of the lion .10 years old. as regionally endangered in West Africa [5]. Historically, lions occurred in all biomes in West Africa, with the exception of the Establishing lion presence/absence coastal Upper and Lower Guinean Forests and the interior of the Survey methods commonly used for African savannah mam- Saharan Desert (Fig. 1). The collapse of lion range in West Africa mals, such as aerial surveys or line transects, typically yield few is poorly documented, but appears to be linked to large-scale observations of large terrestrial carnivores [23]. Consequently, habitat loss outside PAs through conversion to agriculture [5]. prior efforts to establish large carnivore occurrence and/or Consequently, lion range in this region is largely restricted to PAs abundance over large spatial scales relied on interviews [24], [4,11]. While several West African countries have large PAs, remote cameras [25], or track surveys [26]. We predominantly average PA performance in West Africa ranked poorest in a cross- employed track surveys, owing to their comparatively high continental comparison: large mammal populations in eleven West detection efficiency, and low effort and cost [26]. All surveys African PAs declined by an average of 85% between 1970–2005, teams included experienced observers, and we ascribed field sign compared to an average 59% decline across the continent [12]. to species based on pugmark characteristics [27]. Occasionally Population collapse within these PAs appears to be driven by teams found equivocal tracks, mainly regionally rare species such commercial bushmeat exploitation, supplying local markets in as African wild dog (Lycaon pictus) and cheetah (Acinonyx jubatus). In West Africa [13–15], and has contributed to bringing several such cases we documented pugmarks photographically, and iconic large mammal species to the brink of extinction in their presented photos to a panel of experts for species verification. West African range [16,17]. We omitted a small number of records from analysis if they could not be assigned to species unequivocally. In PAs with an intact Methods network of dirt tracks, we conducted vehicle-based track surveys, with two observers seated on the bull-bar of a vehicle driven at a Consultation of expert opinion on current lion range maximum speed of 10–20 km h [28]. In PAs without penetra- In 2005, the IUCN and the Wildlife Conservation Society ble roads, we conducted track searches on foot along roughly (WCS) organized a lion conservation workshop for wildlife predefined survey circuits, following game trails, dry riverbeds, authorities from all lion range countries within West and Central abandoned dirt tracks or other linear features commonly used as Africa [18]. The workshop consisted of a technical session to map travel routes by lions and other large carnivores. These circuits current lion range and status, followed by a strategic planning incorporated habitat features that could be expected to attract session to develop lion conservation strategies [18]. The technical larger herbivores, such as water reservoirs, floodplains, saltlicks session was modeled after the Range Wide Priority Setting process and marshes, or other sites with high herbivore abundance developed by WCS for jaguars [19]. Experts were guided to indicated by PA staff. In Mole NP and Gbele Resource Reserve in produce maps of current lion range and delimit areas harboring known or suspected populations called Lion Conservation Units Ghana, camera traps were the primary survey method. At those PAs, we deployed DeerCam DC-300 (Non Typical, Wisconsin, (LCUs) [18]. LCU delimitation relied on credible evidence of lion presence within the preceding 10 years [18], and for each LCU USA) camera units at ,1 km intervals, targeting, as with foot participants assigned lion population trends and approximate lion surveys, features expected to maximize lion capture probability population size. Although the data presented at the 2005 [29]. In Mole NP, we concentrated trapping effort in the central workshop now date from 1995 onwards, at the onset of our field and southeastern portions known to contain higher prey densities surveys in 2006 (see below), it represented the most comprehensive and key dry-season water sources. In Comoe´ National Park (NP) in and reliable dataset on lion distribution in West Africa. Coˆte d’Ivoire, West Africa’s largest NP at .10,000 km ,we conducted an aerial survey prior to our ground survey, to identify areas with important concentrations of potential lion prey. We Field surveys restricted the ensuing foot survey to those areas. Between October 2006 and May 2012, we conducted field surveys in PAs within designated LCUs, to 1) confirm lion PLOS ONE | www.plosone.org 2 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa Figure 1. Lion Conservation Units [18] and surveyed protected areas (PAs) in West Africa. Number symbols in the map correspond to PA identification numbers in Table 1 and Tables S3/S4, with numbers printed in bold representing PAs with METT assessments. doi:10.1371/journal.pone.0083500.g001 research staff. Our survey work was therefore considered an Estimating lion population size integral part of preexisting PA monitoring activities, and wildlife If lions were found in a given PA, we used systematic track authorities waived requirements for formal research clearance and counts [28] or call-ups [30] to estimate lion population size, PA entry fees. All field methods used were completely noninvasive depending on local conditions. Neither method has ever been and did not require the handling or sampling of live animals, and locally calibrated in West Africa. However, we preferred track our survey work did therefore not require approval from an ethics counts due to the generally consistent relationship between lion committee. track densities and actual lion densities observed across a wide range of different climatic and habitat-related conditions in Eastern and Southern Africa [28]. One important caveat of this Evaluating management of LCU PAs method is that the relationship between track densities and actual Prior studies highlighted the strong impact of PA characteristics densities varies with substrate type, and we therefore recorded (e.g. PA size), human population density at PA edges, and PA substrate type every 500 meters along spoor transects, to enable us management variables (e.g. PA management budgets) on lion to select the appropriate relationship from Funston et al [28]. In persistence and population status inside PAs [21,32,33]. There- two PAs with lion presence where the poor state of the road fore, we investigated correlations between lion persistence and a network precluded the use of vehicle-based track counts, we number of continuous PA variables, including annual budget, staff conducted call-ups to estimate lion population size. The call-up number, area, surrounding human population density and IUCN method requires calibration experiments to assess local response management category, using univariate analyses (Analysis of distance and response rate of lions to broadcasts [30]. Because lion Variance (ANOVA) and Mann Whitney U tests) where sample observations were extremely rare during our surveys, we obtained size allowed. We tested for univariate correlations between the PA only one response distance estimate in dense Sudano-Sahelian variables using univariate linear regression or spearman’s rank. woodland; one male lion encountered opportunistically 2.5 km We estimated human population density within a 5 km buffer from a call-up station, was observed again at the station around each PA based on human population data from the 26 minutes after the onset of the broadcast, fifty minutes after AfriPop Project (www.afripop.org), using PA outlines obtained the initial observation. As the male was traveling in the opposite from the World Database of Protected Areas (http:// direction when first seen, we consider this a conservative estimate protectedplanet.net/). We extracted continuous management for a maximum response distance in dense woodland, and used variables from WWF/World Bank Management Effectiveness this value here. To derive tentative estimates of lion population Tracking Tool (METT) assessments carried out in the region. The density based on our call-up results, we used the range of METT is one of the most widely used assessment tools for published figures on lion response rates [30,31]. Study design and Protected Area Management Effectiveness (PAME) [34], and is data analysis of our systematic track counts followed Funston et al. designed to be completed by PA managers, staff and stakeholders. [28], while our protocol for call-ups followed Ferreira and Funston Besides provisioning of continuous PA management variables such [31]. All field surveys were carried out in close collaboration with as budgets and staff numbers, the methodology encompasses a the respective national wildlife authorities, and involved senior PA rapid assessment based on a scorecard questionnaire of 30 PLOS ONE | www.plosone.org 3 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa PLOS ONE | www.plosone.org 4 January 2014 | Volume 9 | Issue 1 | e83500 Table 1. Summary of recent survey activities in LCU PAs, permitting the assessment of lion population status. Conclusion Area [km ] Year regarding lion Lion pop Map ID Name of protected area Country (IUCN PA category)surveyed Target species Type of survey occurrence size Source 1 Boe´ and Dulombi NPs Guinea-Bissau 1,943 (awaiting 2010-ongoing chimpanzee camera trap and foot considered absent n.a. J. van Schijndel gazetting) surveys (www.chimbo.org), pers. comm. 2 Niokolo-Koba NP Senegal 9,130 (II) 2011 lion track surveys (by vehicle) confirmed present 16 (0–54) this study 3 Bafing-Faleme Mali 3,326 (1094 II+672 2003–2004 chimpanzee foot surveys considered absent n.a. Granier & Martinez [54] IV+1561 none) 4 Boucle de Baoule NP Mali 5,330 (II) 2012 faunal inventory foot surveys considered absent n.a. B. Niagate, pers. comm. 5 Haut-Niger NP Guinea 1,228 (II) 2009 lion call-up and track surveys potentially present n.a. PFNH [55] (on foot) 6 Kankan Faunal Reserve Guinea 5,314 (IV) 2007 faunal inventory foot surveys potentially present n.a. Dufour [47] 7 Comoe´ NP Cote d’Ivoire 11,495 (II) 2010 lion track surveys (on foot) considered absent n.a. this study 8 Bui NP Ghana 1,897 (II) 2009 lion interview survey considered absent n.a. this study 9 Mole NP Ghana 4,522 (II) 2006–2009 lion camera trap and track surveys considered absent n.a. this study (by vehicle and on foot) 10 Gbele Resource Reserve Ghana 544 (IV) 2008 lion camera trap and track surveys considered absent n.a. this study (on foot) 11 Digya National Park Ghana 2,789 (II) 2009 lion interview survey considered absent n.a. this study 12 Nazinga Game Ranch Burkina Faso 970 (none) 1985-ongoing faunal inventory foot surveys considered absent n.a. Delvingt & Vermeulen [56] 13 Oti-Mandouri Togo 1,100 (II) 2003 faunal inventory aerial surveys considered absent n.a. Bouche´ et al. [57] National Park 14 W-Arly-Pendjari Benin/Burkina Faso/ 27,167 (14,629 2012 lion track surveys (by vehicle) confirmed present 356 this study Niger II+10,728 IV+1,809 VI) (246–466) 15 Mt Kouffe/Wari Maro Benin 3,092 (VI) 2012 lion consolidation of expert opinion considered absent n.a. CENAGREF [58] 16 Old Oyo National Park Nigeria 2,386 (II) 2009 lion interview survey considered absent n.a. this study 17 Kainji Lake National Park Nigeria 3,970 (II) 2011 lion call-up surveys confirmed present 32 (23–63) this study 18 Kamuku National Park Nigeria 1,121 (II) 2009 lion interview survey considered absent n.a. this study 19 Falgore and Lame-Burra Nigeria 2,910 (IV) 2009 lion track surveys (on foot) considered absent n.a. this study Game Reserves 20 Yankari Game Reserve Nigeria 2,244 (IV) 2011 lion call-up surveys and confirmed present 2 this study ranger-based lion monitoring 21 Gashaka-Gumti National Park Nigeria 1,900 (II) 2009 lion track surveys (on foot) considered absent n.a. this study Total 99,148 (94,378 406 suitable for lion) (250–587) For more details on the surveys, see Table S3. Aerial surveys established complete absence of large wild herbivores, strongly indicating PAs unsuitability for lions. b 2 2 Gashaka-Gumti NP encompasses 6,670 km , however, only ca 1,900 km comprise suitable lion habitat. doi:10.1371/journal.pone.0083500.t001 Lion Status in West Africa questions, with an ordinal four-point scale (0–3, with 3 population) persist in a single population in WAP, and there is representing the best management scenario). The complete strong evidence for ongoing declines in the region’s other three METT questionnaire template is provided in Supporting Infor- populations. In Nigeria, numbers dropped from an estimated 44 mation S1, and Table S1 summarizes the 13 METT scored lions in 2009 to 34 in 2011 [36]. In Senegal’s Niokolo-Koba NP, questions included in our analysis; we selected management continuing calamitous declines in prey populations (Fig. 5B) are aspects likely to influence the ability of a PA to enforce regulations almost certainly causing concomitant declines in lions. These and reduce hunting pressure, to provide insight into the managers’ trends suggest that WAP already or will soon contain .90% of perception of current PA performance). A comprehensive list of all West African lions. Given that 40–60% of a lion population PA management variables used in our analyses, variable typically consists of immature individuals [37,38], and that our track counts in WAP included large cubs and sub-adults, it is very provenance, and corresponding sample sizes can be found in likely there are ,250 adults remaining in the entire West African Table S2. All statistical analyses were performed in R [35]. region. Accordingly, our results warrant listing of the lion as critically endangered in West Africa under criterion C2a(ii), which Results applies for declining populations with ,250 mature individuals, Lion presence and absence in West African PAs where .90% of individuals persist in one subpopulation [39]. Wildlife experts attending the 2005 workshop identified 17 LCUs in West Africa (Fig. 1), totaling 254,430 km , or 5.8% of Priorities for lion conservation in West Africa historic lion range in West Africa. We identified 21 large Our surveys covered all large (.500 km ), formally designated (.500 km ) PAs within those LCUs (Fig. 1, Table 1), with a total PAs within LCUs in West Africa. While lion range in this region is area of ,95,000 km , or 37% of total LCU extent. We surveyed largely restricted to PAs [4,11], we cannot exclude the possibility thirteen of those PAs for lions, while the remaining PAs were that some lions roam outside the surveyed PAs. However, the 21 surveyed by other researchers focusing on lions (n = 2), Western LCU PAs covered in this study represent the best remaining lion chimpanzees Pan troglodytes ssp. verus (n = 2), and general faunal habitat in West Africa [4]. We deem survey effort adequate (see inventories (n = 4) (Table 1). Of the 21 LCU PAs surveyed, lions Table S3) to draw inference on the occurrence of resident lions were confirmed in only four (Table 1; Fig. 2). In two additional across sites, and we are confident that no resident lion populations PAs, both in Guinea, lions had not been observed for .10 years; were overlooked by our efforts. Further survey work may be however, credible reports of vocalizations suggest they may still be required in Haut Niger NP and Kankan FR in Guinea to assess present. Among the four PAs in which lion persistence was the possible presence of lions. However, given the lack of physical confirmed, three contain ,50 individuals, and the only large evidence for over a decade and the poor management scores of population is in the W-Arly-Pendjari (WAP), with an estimated those two PAs (see Table S4 and below), we believe any remaining 356 (range: 246–466) lions (Table 1). The total number of lions populations would be relict and close to extinction. While remaining in West Africa is estimated at 406 (range: 250–587) continued survey and monitoring work is warranted, the highest individuals, while the confirmed lion range (the total size of PAs conservation priority for lions in the region should be strength- where lions were confirmed, including potential sites in Guinea) is ening protection of the known remaining populations. estimated at 49,000 km , or 1.1% of historic lion range in West Lions are more likely to disappear from small PAs than from Africa. larger ones. Critical PA size for lions based on data from East Africa is 291 km , using an average lion density of 16.2 adults Management of LCU Protected Areas 100 km [22]. In comparison, average density across our four sites with lions was 1.0 lions 100 km , .15 times lower than in We identified METT assessments for 12 of the 21 LCU PAs East Africa. Assuming the same lower limit for a viable lion (Table S2). Details of individual assessments can be found in Table population size in West Africa, critical PA size would exceed S4. Due to the small sample sizes, we have not attempted 4,000 km at current lion densities and even that may be multivariate analyses, and have presented statistical correlations inadequate. Newmark [40] revealed that extinction rates in only where continuous data (on PA budgets, staffing, area and Ghanaian PAs were estimated to be 13–77 times higher than in human population density), as opposed to ordinal scores, were equivalent-sized PAs in Tanzania, suggesting that larger size alone available. a may provide insufficient protection against the intense hunting Protected area budget (measured as total budget (US$) and pressure impacting West African PAs. In addition to the lower budget/area (US$/km )) was positively associated with lion carrying capacity of West African savannas for large herbivores persistence (Fig. 3), and PAs with lions were, on average, more [41], higher extinction risks for West African mammals is driven than twice as large as those without, although the latter difference by intense bushmeat hunting pressure within and adjacent to PAs was not statistically significant, possibly due to the small sample [13,14], facilitated by ineffective PA management (Fig. 4) [12]. (Table 2). Total PA budget significantly increased with area Our findings highlight the urgent need for very large (spearman’s rank coefficient = 0.65 p = 0.02). (.4,000 km ) and well protected PAs to assure the survival of METT scores indicate that the majority of PAs are experiencing lions and other threatened large mammals in West Africa. Three severe management deficiencies over most facets of PA manage- of four extant lion populations in West Africa occur in PAs close to ment (Fig. 4). Scores for only two PAs, WAP (where lions are or larger than 4,000 km (Table 1), representing the best prospects present) and Mole (where lions are absent), suggest that they were for saving the taxon. being managed adequately (Fig. 4). WAP currently harbors the only population .50 animals, and is the most viable. However, lion population density is extremely low Discussion in the eastern half of WAP, i.e. the tri-national W NP (Henschel et The lion has undergone a catastrophic collapse in West Africa. al. in prep). An aerial survey covering W immediately following Our results suggest that lions have lost almost 99% of their historic our lion survey in 2012, recorded .50,000 head of cattle inside range, and that only ca. 400 individual lions persist across the the national park, underlining the weak management effectiveness region. Most of these lions (ca. 350 individuals, or 88% of the total in W NP [42]. In contrast, the western half, Arly-Pendjari, PLOS ONE | www.plosone.org 5 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa Figure 2. Lion status in West African protected areas within lion range. doi:10.1371/journal.pone.0083500.g002 supports higher lion densities, stable or increasing prey populations the WAP in aggregate, 90/km are spent in W, compared to 323/ (Fig. 5A), and incursions by livestock into the PA are rare [42].The km in Arly-Pendjari (Henschel et al. in prep). A significantly stark contrast in management effectiveness between the eastern higher operational budget is required in W to attain conservation and western halves of WAP may be due to the disparity in outcomes comparable to Arly-Pendjari. management budget allocations; of the US$197/km available for Figure 3. Boxplot illustrating median operating budgets (US$/km ) for PAs grouped by current lion population status. Hinges represent the 1st and 3rd quartiles, and whiskers represent the range of the data. Black points represent outliers. doi:10.1371/journal.pone.0083500.g003 PLOS ONE | www.plosone.org 6 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa Table 2. PA management characteristics by lion presence and absence (continuous data). Statistical test: Mann PAs where lions confirmed or Whitney U (MW); ANOVA Management characteristics All PAs potentially present PAs where lions absent (AN). Mean (SE) (n) Mean (SE) (n) Mean (SE) (n) PA area (km ) 4,721 (1,275) (21) 8,175 (3,962) (6) 3,340 (730) (15) Non-sig (AN); p = 0.13 Human population density 0.23 (0.05) (21) 0.21 (0.09) (6) 0.24 (0.06) (15) Non-sig (AN); p = 0.92 (people within a 5 km buffer) PA staff/100 km 1.45 (0.46) (12) 1.63 (0.62) (5) 1.32 (0.69) (8) Non-sig (AN): p = 0.72 PA patrol staff/100 km 0.86 (0.30) (12) 0.81 (0.26) (5) 0.91 (0.53) (8) Non-sig (AN): p = 0.76 Median (IQR) (n) Median (IQR) (n) Median (IQR) (n) 6,746 (0–117,076) (12) 185,000 (80,000–257,742) (5) 0 (0–6746) (8) Sig (MW): 29.5, p = 0.048 PA budget (total US$) 2.42 (0–30.1) (12) 35.7 (28.2–42.9) (5) 0 (0–2.41)(8) Sig (MW): W = 30.5, p = 0.03 PA budget/area (US$/km ) Percentage Percentage Percentage PA category (II/IV/VI/none) 66.3/23.7/5.2/4.7 (21) 59.0/37.3/3.7/0.0 (6) 74.2/9.1/6.8/9.9 (15) n/a doi:10.1371/journal.pone.0083500.t002 Lion populations in Niokolo-Koba and Kainji Lake NPs are lion prey species were available, WAP (represented through small and appear to be declining. While no data on management Pendjari NP, where annual wildlife counts are conducted) was also effectiveness and lion prey populations exist for Kainji Lake NP, the only site where wildlife numbers were stable or increasing management effectiveness scores are low in Niokolo-Koba NP, (Fig. 5). As a further indication of conservation success in WAP, potentially due to inadequate funding (Table S4), and prey this site harbors by far the largest remaining elephant population populations have collapsed to extremely low levels over the past 20 in West Africa [16], and one of the last remaining populations of years (Fig. 5B). Both PAs hold great potential due to their large the critically endangered Northwest African cheetah (Acinonyx size, and are surrounded by suitable lion habitat and moderate jubatus ssp. hecki) [17]. Even so, WAP’s budget is an order of human population densities (Table S3) [4]. Furthermore, Kainji magnitude below the estimated .US$2000/km budget required Lake NP is potentially still connected to WAP, through suitable to maintain lions in unfenced PAs [33], suggesting that WAPs lion habitat in Benin [4]. Besides lions, Niokolo-Koba also harbors success may not be sustainable and will require increased funding the last important population of the critically endangered Western in future. giant eland (Tragelaphus derbianus ssp. derbianus) [43], and the only Staff numbers for LCU PAs were generally low, varying confirmed population of the critically endangered West African between 0–4 staff per 100 km . Many PAs reported that staff sub-population of African wild dogs (Table S3) [44]. Both PAs will salaries were paid directly by national government and not require immediate financial and technical assistance to avert the through the individual PA management budget. Where staff are local extirpation of lions and other critically endangered taxa. paid by central government, yet few or no funds are available for At 2,244 km Nigeria’s Yankari Game Reserve is smaller than active management of PAs, the reported staff numbers likely our putative minimum and its lion population is very small and overestimate conservation effort: nine of the 12 PAs assessed using declining (Table 1). Yankari is completely surrounded by intensive METT reported having either no law enforcement activity at all cultivation [45], and the second-highest human population density (four PAs), or major deficiencies in staff capacity/resources to of all 21 LCU PAs surveyed (Table S3). As a consequence, effect patrols (five PAs). Assessors for Comoe´ NP, which has 54 Yankari’s lions and indeed all large-medium mammals are likely to patrol staff, commented that ‘‘Staffing is very low for the size of the park; be effectively isolated from neighboring populations in Nigeria only 2 of 5 sectors of the park are operational, with 1 vehicle, limited staff, no (Kainji Lake NP, ca 650 km distant) and Cameroon (Benoue´ equipment and lack of training’’. In Haut Niger, which has 15 patrol Complex, ca 260 km distant). Drastic interventions, such as staff, assessors noted that in practice, agents do no or very few fencing the reserve, may be the only solution to safeguard this patrols due to a lack of resources and motivation; as a result population [33]. Fencing Yankari could prevent inevitable poaching and illegal logging is widespread. Brugie`re [46] notes encroachment by people and livestock, reduce human-lion that existing PAs in Guinea are essentially paper-parks, i.e. they conflicts at the PA boundary and perhaps reduce penetration of have no staff, management plan or operating budget. Conse- the PA by poachers. quently, even in the largest formally gazetted PA in Guinea, the Kankan Faunal Reserve, where lions are potentially still present, The state of PAs in West Africa ca 20,000 people live within the PA, poaching pressure is high, and antelope population densities are extremely low [47]. Of 12 PAs with management assessments, six had no budget for management activities, and where budgets existed they were much WAP represents the last stronghold for lions in West Africa. lower than required to conserve lion populations effectively [33]. Conservation interventions in WAP are heavily subsidized by large PAs with confirmed or probable lion presence had larger budgets international funding bodies, such as the European Union and the than those with lions absent, and WAP had both the largest lion World Bank. However, overall investment in conservation population and the highest annual budget, at US$197/km (Table activities is extremely low in West Africa, compared to Central, S4). WAP furthermore consistently received among the highest Eastern and Southern Africa [48]. Considering that all eleven scores for management effectiveness of all PAs harboring lions former or current lion range countries in West Africa are among (Fig. 4B). For PAs where data on population trends of principal the 50 poorest countries in the world, and that six are classified as PLOS ONE | www.plosone.org 7 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa Figure 4. METT (Management Effectiveness Tracking Tool) scores for evaluated protected areas (PAs) in West Africa. (A) PAs where lions are confirmed present or are potentially still present (A); (B) PAs where lions were considered absent. The letter W represents scores for W-Arly- Pendjari while M indicates those for Mole NP (see text). Management scores range from 0–3, with 3 representing the best management scenario. For example, in the case of ‘Current Budget’ 0 = No PA budget; 1 = inadequate budget which creates serious management constraints; 2 = acceptable budget, but could be further improved to fully achieve effective management; 3 = sufficient budget which fully meets the needs of the PA. See Table S1 for full descriptions of scores. doi:10.1371/journal.pone.0083500.g004 least developed countries (data from World Bank; http:// It is imperative to address very widespread poaching of lion prey databank.worldbank.org/data/home.aspx), lion range states in species and illegal killing of lions by pastoralists within and around West Africa will be unable to mobilize the resources required to PAs [14,51,52]. We believe urgent priority must be given to a dual secure their remaining lion populations. That will rely on the strategy that focuses on 1. increasing the numbers, expertise, and provision of substantial financial and technical assistance to range operating budgets of enforcement personnel in PAs with lions, to states, principally by the international community, to increase curb the killing of lion prey and illegal incursions into PAs by management effectiveness of PAs with lions. For any such pastoralists, and 2. reducing human-lion conflict in affected investments, it will be imperative that a) conservation initiatives communities bordering PAs, by combining improved husbandry assure sound governance over the funds [49], and b) adequate practices with community sensitization, to reduce livestock losses funding levels are sustained in the long-term to achieve desired to predators and ameliorate local negative perceptions of large outcomes; a review of best practices can be found in Blom et al. carnivores [51]. Investment should also be directed toward [50]. developing and enhancing photographic tourism in politically stable countries such as Benin and Senegal. This will help to create PLOS ONE | www.plosone.org 8 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa Figure 5. Populations trends for principal lion prey species in West African protected areas. (A) Pendjari NP (which forms part of W-Arly- Pendjari); (B) Niokolo-Koba NP; (C) Mole NP. Data sources: Galat et al. [59], Sinsin et al. [60], Wildlife Division of Ghana [61], Bouche´ [62], Renaud et al. [63]. doi:10.1371/journal.pone.0083500.g005 PLOS ONE | www.plosone.org 9 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa and maintain economic incentives for lion conservation, and Table S2 PA management variables used in this study, with develop enduring revenue streams for PA management not wholly sample sizes (number of PAs). reliant on donor funding. (DOCX) Table S3 PA attributes for the 21 PAs in this study and details Conclusions & Recommendations on the field surveys, regarding survey methods and effort, and findings regarding the presence of lions and other species of The situation for the lion in West Africa is dire. We recommend conservation concern. urgent revision of lion taxonomy by the Cat Classification Task (XLSX) Force of the IUCN/SSC Cat Specialist Group [53]. Recognition of a West-Central African sub-species is supported by recent Table S4 PA METT data and scores for the 12 PAs with METT findings establishing the principal division of extant lions within data in this study (PAs labeled using PA number: see S4 for Africa, and would correctly recognize the genetic uniqueness of corresponding PA names). West African populations [8,9]. Irrespective of taxonomic status, (XLSX) we recommend listing the lion as critically endangered in West Africa. Acknowledgments Considering the relative poverty of lion range states in West Africa, we call for the mobilization of substantial and urgent We thank the respective wildlife authorities for assisting efforts to conduct lion surveys in the protected areas covered in the present study. Specific investment by the international community to assist these thanks go to M. Sidibe´ and I. Ndao in Senegal, A. Gonto, D. Azani and I. countries in improving management effectiveness of PAs contain- Herbinger from the Wild Chimpanzee Foundation in Coˆte d’Ivoire, M. ing lions. Lions persist in some of the largest and most intact Sam in Ghana, A. Tehou, M. Kiki and C. Sewade in Benin, P. Bouche´in protected landscapes in West Africa, where they co-occur with Burkina Faso, H. GARBA in Niger, as well as G. Nyanganji from WCS in some of the last remaining populations of critically endangered Nigeria. We thank L. Petracca (Panthera) for help with GIS analyses, F. mammals including Northwest African cheetahs, Western giant Leverington for the identification of additional PA management data, and elands and African wild dogs. Further deterioration of those last C. Breitenmoser, one anonymous reviewer and the editor for helpful wilderness areas in West Africa will likely cause the loss of comments on the manuscript. Supporting Information genetically distinct populations of charismatic megafauna and A template of the METT questionnaire (Supporting Information S1), a further preclude already tenuous, potential future revenue streams list of METT questions used in this study (Table S1), a complete list of PA from photographic tourism for West African nations. Without management variables used in this study (Table S2), a list of attributes for immediate action, we believe the opportunity to save both will be the 21 PAs surveyed in this study (Table S3) and a summary of lost. management data and scores for the 12 PAs with METT data (Table S4) are available online. The authors are solely responsible for the content and functionality of these materials. 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The Lion in West Africa Is Critically Endangered

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Pubmed Central
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© 2014 Henschel et al
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1932-6203
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1932-6203
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10.1371/journal.pone.0083500
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Abstract

The African lion has declined to ,35,000 individuals occupying 25% of its historic range. The situation is most critical for the geographically isolated populations in West Africa, where the species is considered regionally endangered. Elevating their conservation significance, recent molecular studies establish the genetic distinctiveness of West and Central African lions from other extant African populations. Interventions to save West African lions are urgently required. However formulating effective conservation strategies has been hampered by a lack of data on the species’ current distribution, status, and potential management deficiencies of protected areas (PAs) harboring lions. Our study synthesized available expert opinion and field data to close this knowledge gap, and formulate recommendations for the conservation of West African lions. We undertook lion surveys in 13 large (.500 km ) PAs and compiled evidence of lion presence/absence for a further eight PAs. All PAs were situated within Lion Conservation Units, geographical units designated as priority lion areas by wildlife experts at a regional lion conservation workshop in 2005. Lions were confirmed in only 4 PAs, and our results suggest that only 406 (273–605) lions remain in West Africa, representing ,250 mature individuals. Confirmed lion range is estimated at 49,000 km , or 1.1% of historical range in West Africa. PAs retaining lions were larger than PAs without lions and had significantly higher management budgets. We encourage revision of lion taxonomy, to recognize the genetic distinctiveness of West African lions and highlight their potentially unique conservation value. Further, we call for listing of the lion as critically endangered in West Africa, under criterion C2a(ii) for populations with ,250 mature individuals. Finally, considering the relative poverty of lion range states in West Africa, we call for urgent mobilization of investment from the international community to assist range states to increase management effectiveness of PAs retaining lions. Citation: Henschel P, Coad L, Burton C, Chataigner B, Dunn A, et al. (2014) The Lion in West Africa Is Critically Endangered. PLoS ONE 9(1): e83500. doi:10.1371/ journal.pone.0083500 Editor: Matt Hayward, Bangor University, United Kingdom Received September 3, 2013; Accepted November 4, 2013; Published January 8, 2014 Copyright:  2014 Henschel et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: Field survey efforts were funded by Panthera (www.panthera.org), with additional funds from National Geographic’s Big Cat Initiative (grant B1_12; http://animals.nationalgeographic.com/animals/big-cats/). LC furthermore acknowledges the support of the Oxford Martin School, and DWM the Recanati-Kaplan Foundation and Robertson Foundation. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. * E-mail: phenschel@panthera.org West African lions represent a population with unique genetic Introduction and conservation value. Recent molecular and morphological data The lion (Panthera leo) was the most successful large carnivore covering the species’ entire historical range suggests that lions in during the late Pleistocene, when the species’ range extended from Central, West and North Africa (the latter now extinct) are distinct South Africa, across Eurasia, and into the southern United States from lions in Eastern and Southern Africa and share a common [1]. Today, the lion’s range is restricted to Africa and one ancestor with lions in Asia [6–8]. These results establish that the population of the Asiatic sub-species, P. l. persica, in India [2]. principal subdivision of modern lions is within Africa [9], and While the endangered Asiatic population is currently considered question the current dichotomous division into an African and an stable, lion populations in Africa are in decline and the African Asian sub-species; a division which is still widely supported, sub-species, P. l. leo, is considered vulnerable [3]. Recent analyses including by the IUCN Red List [3]. Moreover, they demonstrate established that the African lion has lost at least 75% of its original that lions in West Africa contain mtDNA haplotypes not found in habitat, with fewer than 35,000 wild African lions remaining [4]. other lion populations, elevating the conservation significance of The main drivers of lion declines are large-scale habitat the few remaining West African populations [8,9]. conversion, prey base depletion through unsustainable hunting, Conservation interventions to save these populations are now and the retaliatory killing of lions due to perceived or real human- urgently required. However, formulating interventions is limited lion conflict [3]. The situation is most critical in West Africa, by few field data on the species’ current distribution, abundance, where lions have been considered regionally endangered since and predominant drivers of declines in West Africa. While lions 2004 [5], and where ,500 individuals may persist [4]. have been the object of extensive research effort in parts of Eastern and Southern Africa, they have been largely ignored in West PLOS ONE | www.plosone.org 1 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa Africa. Of 463 articles on African lions in the ISI Web of presence for PAs where evidence of lion occurrence was lacking, TM Science (Thompson Scientific) in 2005, not one focused and 2) establish lion population estimates for PAs where lions specifically on lions in West Africa. To address this deficit, we occurred. Although designated LCUs included both protected and first collated available data and expert opinions on lion non-protected areas, lions are largely absent outside PAs in West distribution and status in West Africa. We then undertook field Africa [11,20]. Accordingly, we restricted fieldwork to PAs (IUCN surveys in 13 large (.500 km ) protected areas (PAs) where lions PA categories I–VI) within LCUs (henceforth: LCU PAs), were reported during this process, to determine lion presence/ including two PAs awaiting formal designation (Table 1). A absence and estimate lion population size. For a further eight large primary determinant of lion extinction risk within a given PA is its PAs suspected to harbor lions, we compiled field survey data from size, and we therefore restricted our survey effort to large the literature and via interviews. Using standardized evaluation (.500 km ) LCU PAs, based on findings on critical PA size for toolkits for protected area management effectiveness [10], we lions from prior studies [21,22]. In LCUs with multiple PAs, we compared current management performance of PAs known or concentrated survey efforts in the largest PAs with the highest suspected to harbor lions and those from which lions have likely protection status (according to IUCN PA categories). been extirpated. With these data, we provide a comprehensive We also compiled data on lion presence from recent field evaluation of the tenuous status of the West African lion, and make surveys led by other institutions, from internal reports and recommendations for the taxa’s conservation. interviews with participants. We included only surveys that targeted large mammals, with survey methods and effort Study area appropriate to detect lions. Finally, we incorporated data from We restrict our analysis to West Africa, as defined by the United interviews of PA staff on lion presence/absence for several LCU Nations geoscheme (http://unstats.un.org/unsd/methods/m49/ PAs that have not been recently surveyed. We did not consider m49regin.htm), including all countries from Senegal to Nigeria reports of lion presence without physical evidence and records (Fig. 1). The same classification was used for the listing of the lion .10 years old. as regionally endangered in West Africa [5]. Historically, lions occurred in all biomes in West Africa, with the exception of the Establishing lion presence/absence coastal Upper and Lower Guinean Forests and the interior of the Survey methods commonly used for African savannah mam- Saharan Desert (Fig. 1). The collapse of lion range in West Africa mals, such as aerial surveys or line transects, typically yield few is poorly documented, but appears to be linked to large-scale observations of large terrestrial carnivores [23]. Consequently, habitat loss outside PAs through conversion to agriculture [5]. prior efforts to establish large carnivore occurrence and/or Consequently, lion range in this region is largely restricted to PAs abundance over large spatial scales relied on interviews [24], [4,11]. While several West African countries have large PAs, remote cameras [25], or track surveys [26]. We predominantly average PA performance in West Africa ranked poorest in a cross- employed track surveys, owing to their comparatively high continental comparison: large mammal populations in eleven West detection efficiency, and low effort and cost [26]. All surveys African PAs declined by an average of 85% between 1970–2005, teams included experienced observers, and we ascribed field sign compared to an average 59% decline across the continent [12]. to species based on pugmark characteristics [27]. Occasionally Population collapse within these PAs appears to be driven by teams found equivocal tracks, mainly regionally rare species such commercial bushmeat exploitation, supplying local markets in as African wild dog (Lycaon pictus) and cheetah (Acinonyx jubatus). In West Africa [13–15], and has contributed to bringing several such cases we documented pugmarks photographically, and iconic large mammal species to the brink of extinction in their presented photos to a panel of experts for species verification. West African range [16,17]. We omitted a small number of records from analysis if they could not be assigned to species unequivocally. In PAs with an intact Methods network of dirt tracks, we conducted vehicle-based track surveys, with two observers seated on the bull-bar of a vehicle driven at a Consultation of expert opinion on current lion range maximum speed of 10–20 km h [28]. In PAs without penetra- In 2005, the IUCN and the Wildlife Conservation Society ble roads, we conducted track searches on foot along roughly (WCS) organized a lion conservation workshop for wildlife predefined survey circuits, following game trails, dry riverbeds, authorities from all lion range countries within West and Central abandoned dirt tracks or other linear features commonly used as Africa [18]. The workshop consisted of a technical session to map travel routes by lions and other large carnivores. These circuits current lion range and status, followed by a strategic planning incorporated habitat features that could be expected to attract session to develop lion conservation strategies [18]. The technical larger herbivores, such as water reservoirs, floodplains, saltlicks session was modeled after the Range Wide Priority Setting process and marshes, or other sites with high herbivore abundance developed by WCS for jaguars [19]. Experts were guided to indicated by PA staff. In Mole NP and Gbele Resource Reserve in produce maps of current lion range and delimit areas harboring known or suspected populations called Lion Conservation Units Ghana, camera traps were the primary survey method. At those PAs, we deployed DeerCam DC-300 (Non Typical, Wisconsin, (LCUs) [18]. LCU delimitation relied on credible evidence of lion presence within the preceding 10 years [18], and for each LCU USA) camera units at ,1 km intervals, targeting, as with foot participants assigned lion population trends and approximate lion surveys, features expected to maximize lion capture probability population size. Although the data presented at the 2005 [29]. In Mole NP, we concentrated trapping effort in the central workshop now date from 1995 onwards, at the onset of our field and southeastern portions known to contain higher prey densities surveys in 2006 (see below), it represented the most comprehensive and key dry-season water sources. In Comoe´ National Park (NP) in and reliable dataset on lion distribution in West Africa. Coˆte d’Ivoire, West Africa’s largest NP at .10,000 km ,we conducted an aerial survey prior to our ground survey, to identify areas with important concentrations of potential lion prey. We Field surveys restricted the ensuing foot survey to those areas. Between October 2006 and May 2012, we conducted field surveys in PAs within designated LCUs, to 1) confirm lion PLOS ONE | www.plosone.org 2 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa Figure 1. Lion Conservation Units [18] and surveyed protected areas (PAs) in West Africa. Number symbols in the map correspond to PA identification numbers in Table 1 and Tables S3/S4, with numbers printed in bold representing PAs with METT assessments. doi:10.1371/journal.pone.0083500.g001 research staff. Our survey work was therefore considered an Estimating lion population size integral part of preexisting PA monitoring activities, and wildlife If lions were found in a given PA, we used systematic track authorities waived requirements for formal research clearance and counts [28] or call-ups [30] to estimate lion population size, PA entry fees. All field methods used were completely noninvasive depending on local conditions. Neither method has ever been and did not require the handling or sampling of live animals, and locally calibrated in West Africa. However, we preferred track our survey work did therefore not require approval from an ethics counts due to the generally consistent relationship between lion committee. track densities and actual lion densities observed across a wide range of different climatic and habitat-related conditions in Eastern and Southern Africa [28]. One important caveat of this Evaluating management of LCU PAs method is that the relationship between track densities and actual Prior studies highlighted the strong impact of PA characteristics densities varies with substrate type, and we therefore recorded (e.g. PA size), human population density at PA edges, and PA substrate type every 500 meters along spoor transects, to enable us management variables (e.g. PA management budgets) on lion to select the appropriate relationship from Funston et al [28]. In persistence and population status inside PAs [21,32,33]. There- two PAs with lion presence where the poor state of the road fore, we investigated correlations between lion persistence and a network precluded the use of vehicle-based track counts, we number of continuous PA variables, including annual budget, staff conducted call-ups to estimate lion population size. The call-up number, area, surrounding human population density and IUCN method requires calibration experiments to assess local response management category, using univariate analyses (Analysis of distance and response rate of lions to broadcasts [30]. Because lion Variance (ANOVA) and Mann Whitney U tests) where sample observations were extremely rare during our surveys, we obtained size allowed. We tested for univariate correlations between the PA only one response distance estimate in dense Sudano-Sahelian variables using univariate linear regression or spearman’s rank. woodland; one male lion encountered opportunistically 2.5 km We estimated human population density within a 5 km buffer from a call-up station, was observed again at the station around each PA based on human population data from the 26 minutes after the onset of the broadcast, fifty minutes after AfriPop Project (www.afripop.org), using PA outlines obtained the initial observation. As the male was traveling in the opposite from the World Database of Protected Areas (http:// direction when first seen, we consider this a conservative estimate protectedplanet.net/). We extracted continuous management for a maximum response distance in dense woodland, and used variables from WWF/World Bank Management Effectiveness this value here. To derive tentative estimates of lion population Tracking Tool (METT) assessments carried out in the region. The density based on our call-up results, we used the range of METT is one of the most widely used assessment tools for published figures on lion response rates [30,31]. Study design and Protected Area Management Effectiveness (PAME) [34], and is data analysis of our systematic track counts followed Funston et al. designed to be completed by PA managers, staff and stakeholders. [28], while our protocol for call-ups followed Ferreira and Funston Besides provisioning of continuous PA management variables such [31]. All field surveys were carried out in close collaboration with as budgets and staff numbers, the methodology encompasses a the respective national wildlife authorities, and involved senior PA rapid assessment based on a scorecard questionnaire of 30 PLOS ONE | www.plosone.org 3 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa PLOS ONE | www.plosone.org 4 January 2014 | Volume 9 | Issue 1 | e83500 Table 1. Summary of recent survey activities in LCU PAs, permitting the assessment of lion population status. Conclusion Area [km ] Year regarding lion Lion pop Map ID Name of protected area Country (IUCN PA category)surveyed Target species Type of survey occurrence size Source 1 Boe´ and Dulombi NPs Guinea-Bissau 1,943 (awaiting 2010-ongoing chimpanzee camera trap and foot considered absent n.a. J. van Schijndel gazetting) surveys (www.chimbo.org), pers. comm. 2 Niokolo-Koba NP Senegal 9,130 (II) 2011 lion track surveys (by vehicle) confirmed present 16 (0–54) this study 3 Bafing-Faleme Mali 3,326 (1094 II+672 2003–2004 chimpanzee foot surveys considered absent n.a. Granier & Martinez [54] IV+1561 none) 4 Boucle de Baoule NP Mali 5,330 (II) 2012 faunal inventory foot surveys considered absent n.a. B. Niagate, pers. comm. 5 Haut-Niger NP Guinea 1,228 (II) 2009 lion call-up and track surveys potentially present n.a. PFNH [55] (on foot) 6 Kankan Faunal Reserve Guinea 5,314 (IV) 2007 faunal inventory foot surveys potentially present n.a. Dufour [47] 7 Comoe´ NP Cote d’Ivoire 11,495 (II) 2010 lion track surveys (on foot) considered absent n.a. this study 8 Bui NP Ghana 1,897 (II) 2009 lion interview survey considered absent n.a. this study 9 Mole NP Ghana 4,522 (II) 2006–2009 lion camera trap and track surveys considered absent n.a. this study (by vehicle and on foot) 10 Gbele Resource Reserve Ghana 544 (IV) 2008 lion camera trap and track surveys considered absent n.a. this study (on foot) 11 Digya National Park Ghana 2,789 (II) 2009 lion interview survey considered absent n.a. this study 12 Nazinga Game Ranch Burkina Faso 970 (none) 1985-ongoing faunal inventory foot surveys considered absent n.a. Delvingt & Vermeulen [56] 13 Oti-Mandouri Togo 1,100 (II) 2003 faunal inventory aerial surveys considered absent n.a. Bouche´ et al. [57] National Park 14 W-Arly-Pendjari Benin/Burkina Faso/ 27,167 (14,629 2012 lion track surveys (by vehicle) confirmed present 356 this study Niger II+10,728 IV+1,809 VI) (246–466) 15 Mt Kouffe/Wari Maro Benin 3,092 (VI) 2012 lion consolidation of expert opinion considered absent n.a. CENAGREF [58] 16 Old Oyo National Park Nigeria 2,386 (II) 2009 lion interview survey considered absent n.a. this study 17 Kainji Lake National Park Nigeria 3,970 (II) 2011 lion call-up surveys confirmed present 32 (23–63) this study 18 Kamuku National Park Nigeria 1,121 (II) 2009 lion interview survey considered absent n.a. this study 19 Falgore and Lame-Burra Nigeria 2,910 (IV) 2009 lion track surveys (on foot) considered absent n.a. this study Game Reserves 20 Yankari Game Reserve Nigeria 2,244 (IV) 2011 lion call-up surveys and confirmed present 2 this study ranger-based lion monitoring 21 Gashaka-Gumti National Park Nigeria 1,900 (II) 2009 lion track surveys (on foot) considered absent n.a. this study Total 99,148 (94,378 406 suitable for lion) (250–587) For more details on the surveys, see Table S3. Aerial surveys established complete absence of large wild herbivores, strongly indicating PAs unsuitability for lions. b 2 2 Gashaka-Gumti NP encompasses 6,670 km , however, only ca 1,900 km comprise suitable lion habitat. doi:10.1371/journal.pone.0083500.t001 Lion Status in West Africa questions, with an ordinal four-point scale (0–3, with 3 population) persist in a single population in WAP, and there is representing the best management scenario). The complete strong evidence for ongoing declines in the region’s other three METT questionnaire template is provided in Supporting Infor- populations. In Nigeria, numbers dropped from an estimated 44 mation S1, and Table S1 summarizes the 13 METT scored lions in 2009 to 34 in 2011 [36]. In Senegal’s Niokolo-Koba NP, questions included in our analysis; we selected management continuing calamitous declines in prey populations (Fig. 5B) are aspects likely to influence the ability of a PA to enforce regulations almost certainly causing concomitant declines in lions. These and reduce hunting pressure, to provide insight into the managers’ trends suggest that WAP already or will soon contain .90% of perception of current PA performance). A comprehensive list of all West African lions. Given that 40–60% of a lion population PA management variables used in our analyses, variable typically consists of immature individuals [37,38], and that our track counts in WAP included large cubs and sub-adults, it is very provenance, and corresponding sample sizes can be found in likely there are ,250 adults remaining in the entire West African Table S2. All statistical analyses were performed in R [35]. region. Accordingly, our results warrant listing of the lion as critically endangered in West Africa under criterion C2a(ii), which Results applies for declining populations with ,250 mature individuals, Lion presence and absence in West African PAs where .90% of individuals persist in one subpopulation [39]. Wildlife experts attending the 2005 workshop identified 17 LCUs in West Africa (Fig. 1), totaling 254,430 km , or 5.8% of Priorities for lion conservation in West Africa historic lion range in West Africa. We identified 21 large Our surveys covered all large (.500 km ), formally designated (.500 km ) PAs within those LCUs (Fig. 1, Table 1), with a total PAs within LCUs in West Africa. While lion range in this region is area of ,95,000 km , or 37% of total LCU extent. We surveyed largely restricted to PAs [4,11], we cannot exclude the possibility thirteen of those PAs for lions, while the remaining PAs were that some lions roam outside the surveyed PAs. However, the 21 surveyed by other researchers focusing on lions (n = 2), Western LCU PAs covered in this study represent the best remaining lion chimpanzees Pan troglodytes ssp. verus (n = 2), and general faunal habitat in West Africa [4]. We deem survey effort adequate (see inventories (n = 4) (Table 1). Of the 21 LCU PAs surveyed, lions Table S3) to draw inference on the occurrence of resident lions were confirmed in only four (Table 1; Fig. 2). In two additional across sites, and we are confident that no resident lion populations PAs, both in Guinea, lions had not been observed for .10 years; were overlooked by our efforts. Further survey work may be however, credible reports of vocalizations suggest they may still be required in Haut Niger NP and Kankan FR in Guinea to assess present. Among the four PAs in which lion persistence was the possible presence of lions. However, given the lack of physical confirmed, three contain ,50 individuals, and the only large evidence for over a decade and the poor management scores of population is in the W-Arly-Pendjari (WAP), with an estimated those two PAs (see Table S4 and below), we believe any remaining 356 (range: 246–466) lions (Table 1). The total number of lions populations would be relict and close to extinction. While remaining in West Africa is estimated at 406 (range: 250–587) continued survey and monitoring work is warranted, the highest individuals, while the confirmed lion range (the total size of PAs conservation priority for lions in the region should be strength- where lions were confirmed, including potential sites in Guinea) is ening protection of the known remaining populations. estimated at 49,000 km , or 1.1% of historic lion range in West Lions are more likely to disappear from small PAs than from Africa. larger ones. Critical PA size for lions based on data from East Africa is 291 km , using an average lion density of 16.2 adults Management of LCU Protected Areas 100 km [22]. In comparison, average density across our four sites with lions was 1.0 lions 100 km , .15 times lower than in We identified METT assessments for 12 of the 21 LCU PAs East Africa. Assuming the same lower limit for a viable lion (Table S2). Details of individual assessments can be found in Table population size in West Africa, critical PA size would exceed S4. Due to the small sample sizes, we have not attempted 4,000 km at current lion densities and even that may be multivariate analyses, and have presented statistical correlations inadequate. Newmark [40] revealed that extinction rates in only where continuous data (on PA budgets, staffing, area and Ghanaian PAs were estimated to be 13–77 times higher than in human population density), as opposed to ordinal scores, were equivalent-sized PAs in Tanzania, suggesting that larger size alone available. a may provide insufficient protection against the intense hunting Protected area budget (measured as total budget (US$) and pressure impacting West African PAs. In addition to the lower budget/area (US$/km )) was positively associated with lion carrying capacity of West African savannas for large herbivores persistence (Fig. 3), and PAs with lions were, on average, more [41], higher extinction risks for West African mammals is driven than twice as large as those without, although the latter difference by intense bushmeat hunting pressure within and adjacent to PAs was not statistically significant, possibly due to the small sample [13,14], facilitated by ineffective PA management (Fig. 4) [12]. (Table 2). Total PA budget significantly increased with area Our findings highlight the urgent need for very large (spearman’s rank coefficient = 0.65 p = 0.02). (.4,000 km ) and well protected PAs to assure the survival of METT scores indicate that the majority of PAs are experiencing lions and other threatened large mammals in West Africa. Three severe management deficiencies over most facets of PA manage- of four extant lion populations in West Africa occur in PAs close to ment (Fig. 4). Scores for only two PAs, WAP (where lions are or larger than 4,000 km (Table 1), representing the best prospects present) and Mole (where lions are absent), suggest that they were for saving the taxon. being managed adequately (Fig. 4). WAP currently harbors the only population .50 animals, and is the most viable. However, lion population density is extremely low Discussion in the eastern half of WAP, i.e. the tri-national W NP (Henschel et The lion has undergone a catastrophic collapse in West Africa. al. in prep). An aerial survey covering W immediately following Our results suggest that lions have lost almost 99% of their historic our lion survey in 2012, recorded .50,000 head of cattle inside range, and that only ca. 400 individual lions persist across the the national park, underlining the weak management effectiveness region. Most of these lions (ca. 350 individuals, or 88% of the total in W NP [42]. In contrast, the western half, Arly-Pendjari, PLOS ONE | www.plosone.org 5 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa Figure 2. Lion status in West African protected areas within lion range. doi:10.1371/journal.pone.0083500.g002 supports higher lion densities, stable or increasing prey populations the WAP in aggregate, 90/km are spent in W, compared to 323/ (Fig. 5A), and incursions by livestock into the PA are rare [42].The km in Arly-Pendjari (Henschel et al. in prep). A significantly stark contrast in management effectiveness between the eastern higher operational budget is required in W to attain conservation and western halves of WAP may be due to the disparity in outcomes comparable to Arly-Pendjari. management budget allocations; of the US$197/km available for Figure 3. Boxplot illustrating median operating budgets (US$/km ) for PAs grouped by current lion population status. Hinges represent the 1st and 3rd quartiles, and whiskers represent the range of the data. Black points represent outliers. doi:10.1371/journal.pone.0083500.g003 PLOS ONE | www.plosone.org 6 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa Table 2. PA management characteristics by lion presence and absence (continuous data). Statistical test: Mann PAs where lions confirmed or Whitney U (MW); ANOVA Management characteristics All PAs potentially present PAs where lions absent (AN). Mean (SE) (n) Mean (SE) (n) Mean (SE) (n) PA area (km ) 4,721 (1,275) (21) 8,175 (3,962) (6) 3,340 (730) (15) Non-sig (AN); p = 0.13 Human population density 0.23 (0.05) (21) 0.21 (0.09) (6) 0.24 (0.06) (15) Non-sig (AN); p = 0.92 (people within a 5 km buffer) PA staff/100 km 1.45 (0.46) (12) 1.63 (0.62) (5) 1.32 (0.69) (8) Non-sig (AN): p = 0.72 PA patrol staff/100 km 0.86 (0.30) (12) 0.81 (0.26) (5) 0.91 (0.53) (8) Non-sig (AN): p = 0.76 Median (IQR) (n) Median (IQR) (n) Median (IQR) (n) 6,746 (0–117,076) (12) 185,000 (80,000–257,742) (5) 0 (0–6746) (8) Sig (MW): 29.5, p = 0.048 PA budget (total US$) 2.42 (0–30.1) (12) 35.7 (28.2–42.9) (5) 0 (0–2.41)(8) Sig (MW): W = 30.5, p = 0.03 PA budget/area (US$/km ) Percentage Percentage Percentage PA category (II/IV/VI/none) 66.3/23.7/5.2/4.7 (21) 59.0/37.3/3.7/0.0 (6) 74.2/9.1/6.8/9.9 (15) n/a doi:10.1371/journal.pone.0083500.t002 Lion populations in Niokolo-Koba and Kainji Lake NPs are lion prey species were available, WAP (represented through small and appear to be declining. While no data on management Pendjari NP, where annual wildlife counts are conducted) was also effectiveness and lion prey populations exist for Kainji Lake NP, the only site where wildlife numbers were stable or increasing management effectiveness scores are low in Niokolo-Koba NP, (Fig. 5). As a further indication of conservation success in WAP, potentially due to inadequate funding (Table S4), and prey this site harbors by far the largest remaining elephant population populations have collapsed to extremely low levels over the past 20 in West Africa [16], and one of the last remaining populations of years (Fig. 5B). Both PAs hold great potential due to their large the critically endangered Northwest African cheetah (Acinonyx size, and are surrounded by suitable lion habitat and moderate jubatus ssp. hecki) [17]. Even so, WAP’s budget is an order of human population densities (Table S3) [4]. Furthermore, Kainji magnitude below the estimated .US$2000/km budget required Lake NP is potentially still connected to WAP, through suitable to maintain lions in unfenced PAs [33], suggesting that WAPs lion habitat in Benin [4]. Besides lions, Niokolo-Koba also harbors success may not be sustainable and will require increased funding the last important population of the critically endangered Western in future. giant eland (Tragelaphus derbianus ssp. derbianus) [43], and the only Staff numbers for LCU PAs were generally low, varying confirmed population of the critically endangered West African between 0–4 staff per 100 km . Many PAs reported that staff sub-population of African wild dogs (Table S3) [44]. Both PAs will salaries were paid directly by national government and not require immediate financial and technical assistance to avert the through the individual PA management budget. Where staff are local extirpation of lions and other critically endangered taxa. paid by central government, yet few or no funds are available for At 2,244 km Nigeria’s Yankari Game Reserve is smaller than active management of PAs, the reported staff numbers likely our putative minimum and its lion population is very small and overestimate conservation effort: nine of the 12 PAs assessed using declining (Table 1). Yankari is completely surrounded by intensive METT reported having either no law enforcement activity at all cultivation [45], and the second-highest human population density (four PAs), or major deficiencies in staff capacity/resources to of all 21 LCU PAs surveyed (Table S3). As a consequence, effect patrols (five PAs). Assessors for Comoe´ NP, which has 54 Yankari’s lions and indeed all large-medium mammals are likely to patrol staff, commented that ‘‘Staffing is very low for the size of the park; be effectively isolated from neighboring populations in Nigeria only 2 of 5 sectors of the park are operational, with 1 vehicle, limited staff, no (Kainji Lake NP, ca 650 km distant) and Cameroon (Benoue´ equipment and lack of training’’. In Haut Niger, which has 15 patrol Complex, ca 260 km distant). Drastic interventions, such as staff, assessors noted that in practice, agents do no or very few fencing the reserve, may be the only solution to safeguard this patrols due to a lack of resources and motivation; as a result population [33]. Fencing Yankari could prevent inevitable poaching and illegal logging is widespread. Brugie`re [46] notes encroachment by people and livestock, reduce human-lion that existing PAs in Guinea are essentially paper-parks, i.e. they conflicts at the PA boundary and perhaps reduce penetration of have no staff, management plan or operating budget. Conse- the PA by poachers. quently, even in the largest formally gazetted PA in Guinea, the Kankan Faunal Reserve, where lions are potentially still present, The state of PAs in West Africa ca 20,000 people live within the PA, poaching pressure is high, and antelope population densities are extremely low [47]. Of 12 PAs with management assessments, six had no budget for management activities, and where budgets existed they were much WAP represents the last stronghold for lions in West Africa. lower than required to conserve lion populations effectively [33]. Conservation interventions in WAP are heavily subsidized by large PAs with confirmed or probable lion presence had larger budgets international funding bodies, such as the European Union and the than those with lions absent, and WAP had both the largest lion World Bank. However, overall investment in conservation population and the highest annual budget, at US$197/km (Table activities is extremely low in West Africa, compared to Central, S4). WAP furthermore consistently received among the highest Eastern and Southern Africa [48]. Considering that all eleven scores for management effectiveness of all PAs harboring lions former or current lion range countries in West Africa are among (Fig. 4B). For PAs where data on population trends of principal the 50 poorest countries in the world, and that six are classified as PLOS ONE | www.plosone.org 7 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa Figure 4. METT (Management Effectiveness Tracking Tool) scores for evaluated protected areas (PAs) in West Africa. (A) PAs where lions are confirmed present or are potentially still present (A); (B) PAs where lions were considered absent. The letter W represents scores for W-Arly- Pendjari while M indicates those for Mole NP (see text). Management scores range from 0–3, with 3 representing the best management scenario. For example, in the case of ‘Current Budget’ 0 = No PA budget; 1 = inadequate budget which creates serious management constraints; 2 = acceptable budget, but could be further improved to fully achieve effective management; 3 = sufficient budget which fully meets the needs of the PA. See Table S1 for full descriptions of scores. doi:10.1371/journal.pone.0083500.g004 least developed countries (data from World Bank; http:// It is imperative to address very widespread poaching of lion prey databank.worldbank.org/data/home.aspx), lion range states in species and illegal killing of lions by pastoralists within and around West Africa will be unable to mobilize the resources required to PAs [14,51,52]. We believe urgent priority must be given to a dual secure their remaining lion populations. That will rely on the strategy that focuses on 1. increasing the numbers, expertise, and provision of substantial financial and technical assistance to range operating budgets of enforcement personnel in PAs with lions, to states, principally by the international community, to increase curb the killing of lion prey and illegal incursions into PAs by management effectiveness of PAs with lions. For any such pastoralists, and 2. reducing human-lion conflict in affected investments, it will be imperative that a) conservation initiatives communities bordering PAs, by combining improved husbandry assure sound governance over the funds [49], and b) adequate practices with community sensitization, to reduce livestock losses funding levels are sustained in the long-term to achieve desired to predators and ameliorate local negative perceptions of large outcomes; a review of best practices can be found in Blom et al. carnivores [51]. Investment should also be directed toward [50]. developing and enhancing photographic tourism in politically stable countries such as Benin and Senegal. This will help to create PLOS ONE | www.plosone.org 8 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa Figure 5. Populations trends for principal lion prey species in West African protected areas. (A) Pendjari NP (which forms part of W-Arly- Pendjari); (B) Niokolo-Koba NP; (C) Mole NP. Data sources: Galat et al. [59], Sinsin et al. [60], Wildlife Division of Ghana [61], Bouche´ [62], Renaud et al. [63]. doi:10.1371/journal.pone.0083500.g005 PLOS ONE | www.plosone.org 9 January 2014 | Volume 9 | Issue 1 | e83500 Lion Status in West Africa and maintain economic incentives for lion conservation, and Table S2 PA management variables used in this study, with develop enduring revenue streams for PA management not wholly sample sizes (number of PAs). reliant on donor funding. (DOCX) Table S3 PA attributes for the 21 PAs in this study and details Conclusions & Recommendations on the field surveys, regarding survey methods and effort, and findings regarding the presence of lions and other species of The situation for the lion in West Africa is dire. We recommend conservation concern. urgent revision of lion taxonomy by the Cat Classification Task (XLSX) Force of the IUCN/SSC Cat Specialist Group [53]. Recognition of a West-Central African sub-species is supported by recent Table S4 PA METT data and scores for the 12 PAs with METT findings establishing the principal division of extant lions within data in this study (PAs labeled using PA number: see S4 for Africa, and would correctly recognize the genetic uniqueness of corresponding PA names). West African populations [8,9]. Irrespective of taxonomic status, (XLSX) we recommend listing the lion as critically endangered in West Africa. Acknowledgments Considering the relative poverty of lion range states in West Africa, we call for the mobilization of substantial and urgent We thank the respective wildlife authorities for assisting efforts to conduct lion surveys in the protected areas covered in the present study. Specific investment by the international community to assist these thanks go to M. Sidibe´ and I. Ndao in Senegal, A. Gonto, D. Azani and I. countries in improving management effectiveness of PAs contain- Herbinger from the Wild Chimpanzee Foundation in Coˆte d’Ivoire, M. ing lions. Lions persist in some of the largest and most intact Sam in Ghana, A. Tehou, M. Kiki and C. Sewade in Benin, P. Bouche´in protected landscapes in West Africa, where they co-occur with Burkina Faso, H. GARBA in Niger, as well as G. Nyanganji from WCS in some of the last remaining populations of critically endangered Nigeria. We thank L. Petracca (Panthera) for help with GIS analyses, F. mammals including Northwest African cheetahs, Western giant Leverington for the identification of additional PA management data, and elands and African wild dogs. Further deterioration of those last C. Breitenmoser, one anonymous reviewer and the editor for helpful wilderness areas in West Africa will likely cause the loss of comments on the manuscript. Supporting Information genetically distinct populations of charismatic megafauna and A template of the METT questionnaire (Supporting Information S1), a further preclude already tenuous, potential future revenue streams list of METT questions used in this study (Table S1), a complete list of PA from photographic tourism for West African nations. Without management variables used in this study (Table S2), a list of attributes for immediate action, we believe the opportunity to save both will be the 21 PAs surveyed in this study (Table S3) and a summary of lost. management data and scores for the 12 PAs with METT data (Table S4) are available online. The authors are solely responsible for the content and functionality of these materials. 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