13652028, 2022, 3, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/aje.12946 by Readcube (Labtiva Inc.), Wiley Online Library on [15/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License Received: 15 February 2021 | Revised: 18 August 2021 | Accepted: 25 November 2021 DOI: 10.1111/aje.12946 S H O R T C O M M U N I C AT I O N Persistence of wild felids after a protracted civil war in Quiçama National Park and Quiçama Game Reserve, Angola Franciany Braga-­Pereira1,2,3 | Carmen Van-­Dúnem Santos3 | Rômulo Romeu Nóbrega Alves2,4 | Luke Hunter5,6 1 Department de Sanitat i Anatomia Animals, Universitat Autònoma de Barcelona, Barcelona, Catalunya, Spain 2 Department of Ecology and Systematics, Universidade Federal da Paraíba, João Pessoa, Brazil 3 Department of Biology, Universidade Agostinho Neto, Luanda, Angola 4 Laboratory of Ethnobiology and Ethnoecology, Universidade Estadual da Paraíba, Campina Grande, Brazil 5 Wildlife Conservation Society, Bronx, New York, USA 6 Centre for Functional Biodiversity, School of Life Sciences, University of KwaZulu-­Natal, Pietermaritzburg, South Africa Correspondence Franciany Braga-­Pereira, Department de Sanitat i Anatomia Animals, Universitat Autònoma de Barcelona, Barcelona, Catalunya, Spain. Email: franbraga83@yahoo.com.br Funding information Rufford Foundation, Grant/Award Number: 20153-­1; Coordenação de Aperfeiçoamento de Pessoal de Nível Superior, Grant/Award Number: 20151025938; Faculdade de ciências-­Universidade de Agostinho Neto; Universitat Autonoma De Barcelona 1 | I NTRO D U C TI O N but species still believed to occur included the medium-­large species lion Panthera leo, leopard P. pardus, serval Leptailurus ser- Wild felids are globally vulnerable to increasing anthropogenic val, cheetah Acinonyx jubatus, spotted hyaena Crocuta crocuta pressures, with some populations severely declining in many and African wild dog Lycaon pictus (Huntley, 1973; Teixeira et al., areas of their distribution range (Bauer et al., 2015; Durant et al., 1967). However, the intermittent 27-­year Angolan civil war (1975–­ 2017; Terraube et al., 2020). Many felids, especially larger species 2002) combined with very low management presence even during are mostly restricted to protected areas, and their persistence is peaceful periods facilitated extensive overexploitation of wildlife, strongly dependent on adequate protection. However, profound inducing population collapse of most mammals in the park area limitations in operating funds and management capacity create a sit- (Braga-­Pereira, Peres, et al., 2020). Post-­war recovery of wildlife uation in which felid populations are strikingly depleted inside many has been severely limited by poor governance, endemic poverty protected areas, especially in Africa (Lindsey et al., 2018) or their and very widespread availability of military firearms used for hunt- current status is too poorly known to assess the prospects for long-­ ing (Braga-­Pereira et al. 2020). term persistence. Until recently, these same constraints have precluded undertak- The Quiçama National Park and adjacent Quiçama Game ing research in Quiçama. Aside from anecdotal reports, there are Reserve, Angola, are a striking example of these challenges. One of few published data on the presence or status of most large mam- the most important protected areas of Angola, it hosted abundant mals including felids since the beginning of the Angolan civil war. and widespread wildlife including five felid species until the 1960s–­ Accordingly, we undertook the first rigorous attempt to collect 1970s (Huntley, 1973; Teixeira et al., 1967). Carnivores were once data on felids since 1975. Literature increasingly suggests the use abundant but were reduced in the 1960s by a predator removal cam- of local ecological knowledge (LEK)-­based methods for carnivore paign that employed strychnine-­laced carcasses (Huntley, 2017). inventories, as these methodologies are accurate, cost-­ and time-­ By the onset of the civil war, carnivores were significantly depleted effective and can rapidly inventory large areas (Braga-­Pereira et al., This is an open access article under the terms of the Creative Commons Attribution-­NonCommercial-­NoDerivs License, which permits use and distribution in any medium, provided the original work is properly cited, the use is non-­commercial and no modifications or adaptations are made. © 2021 The Authors. African Journal of Ecology published by John Wiley & Sons Ltd. 800 |  wileyonlinelibrary.com/journal/aje Afr J Ecol. 2022;60:800–806. | 13652028, 2022, 3, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/aje.12946 by Readcube (Labtiva Inc.), Wiley Online Library on [15/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License BRAGA-­PEREIRA et al. 801 2021; Madsen et al., 2020; Mbise et al., 2020; Petracca et al., 2018; observation location on a map (Ernoul et al., 2018). We asked inter- Rafiq et al., 2019). Thus, using LEK-­based methods combined with viewees their opinion of felid abundance before, during and after the opportunistic first-­hand observations, we provide here an update on civil war. the presence and distribution of lion, leopard and serval in Quiçama Interviewees who perceived that felids had declined were asked National Park and Quiçama Game Reserve. We also provide infor- to name the drivers of declines. For hunters, we additionally asked mation on the level of human hunting of felids and their conserva- i) their motivation for hunting felids; ii) their revenue from selling tion prospects in the protected areas. skins (the primary product sold); iii) the date of the most recent felid killing; and iv) how often each species was killed, either the number killed per year or the total number killed since he/she started hunt- 2 | M ATE R I A L A N D M E TH O DS ing. We calculated an annual kill rate of each species for each inter- viewee. We opportunistically recorded physical evidence of felids 2.1 | Study area including body parts (skins, skulls in villages, etc.), tracks and direct observations. Quiçama National Park and Quiçama Game Reserve (hereafter col- The capture frequency was calculated annually, so if the capture lectively referred to as Quiçama) consist of two contiguous protected frequency occurs at intervals greater than 1 year, the frequency areas encompassing 9960 Km² with a fenced ‘Special Conservation value will be less than 1. Area’ of 99.6 km² established for reintroduction efforts of herbi- vores during the 2000s. Currently, Quiçama is occupied by almost 9000 people. Although 2.3 | Ethics approval this local population is not legally permitted to occupy the two pro- tected areas, they have lived in the area prior to park delimitation. This research was approved by the i) Brazilian National Health Subsistence hunting of non-­endangered species is permitted in Council (Resolution 466/12), through the Research Ethics Angola (Law No. 6/17 of January 24) although it is not legal inside Committee of the Universidade Federal da Paraíba (licence num- protected areas. Nonetheless, hunting by residents in Quiçama is ber 59846816.3.0000.5188); ii) Environmental Ministry of Angola very prevalent. (148INBAC.MINAMB/2016); iii) municipal administration of Quiçama (licence number 017/GAB.ADM.MQ/2017); and by the iv) leaderships of all sampled villages. An informed consent form was 2.2 | Data acquisition and compilation given to the interviewee, stating the purpose of the interview and ensuring the interviewee's anonymity, so after having the informed Data were collected from January to April 2017. Interviewees were consent from signed, the interview was started. selected using the snowball sampling technique (Bailey, 1994), in which interviewed experts indicate additional candidates, repeated until the candidate pool is exhausted. Previous personal contact 3 | R E S U LT S A N D D I S CU S S I O N between FB-­P and key interviewees (from January to July 2014) established a relationship of trust that reduced potential problems, Of 123 interviewed people, 94.3%, 83.7% and 59.3% reported see- for example the researcher being perceived as law enforcement. ing (alive or dead) at least one serval, leopard and lion, respectively, We selected 123 interviewees (120 men and 3 women), compris- in her/his life. No one definitively reported seeing cheetah, other ing 116 local hunters and 7 park rangers, ranging in age from 20 to than general references to the species being present before the 80 years old. All interviewees had lived or worked in the park for at war. The most recent observation of serval and leopard occurred in least 15 years, ensuring they had been living in the study area since the preceding 5 years for most people (85% and 76% respectively). the civil war period. Given our data present only the most recent year in which felids We conducted structured interviews with an illustrated were seen or killed, they are not indicative of population trends checklist, which provided visual stimulation with drawings of spe- of cats. From these data, we can only conclude that more people cies potentially present in the study area. We also included images reported seeing servals and leopards in recent years than lions of species that have never occurred to verify interviewees' com- which reflects the relative scarcity of the latter but tells us little petence at identifying species. Although African wildcat Felis ly- about changes over time. For lion, the most recent observation by bica is definitely present (Groom et al., 2018), our questionnaire most (56%) interviewees occurred 20–­49 years preceding the in- focused on medium-­large mammals (including non-­c arnivores, terviews. Eight people reported seeing a lion in six locations in the Braga-­Pereira et al., 2020) and we did not collect location data preceding 5 years, which represents 13% of the total reported lion on this species. observations (Table 1). We asked each interviewee the date of the most recent felid Regarding the distribution of records, for the period 2012–­ observation and, if it occurred in the last 5 years, to indicate the 2017, interviewees from all regions of Quiçama reported seeing | 13652028, 2022, 3, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/aje.12946 by Readcube (Labtiva Inc.), Wiley Online Library on [15/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License 802 BRAGA-­PEREIRA et al. serval and we also collected opportunistic observations for ser- Overhunting was the main reason associated with felid declines val throughout Quicama, which agrees with Groom et al. (2018). and most (n = 86) hunters reported killing a wild felid at least once. Leopard reports for this period (sighting and killing by the inter- For servals and leopards, 60% and 31% of respondents, respec- viewees) were restricted to the south of the park, except for two tively, reported cats being captured unintentionally in snares and recent sighting reports near the Special Conservation Area. Our leg-­hold (‘gin’) traps set for herbivores. However, interviewees opportunistic observations follow a similar pattern, with the only also reported intentionally killing felids related to i) depredation physical evidence (tracks, sightings of live individuals, skin and of domestic goats and chickens, as well as pilfering of fish from skulls) of leopards occurring in southern Quiçama (Figures 1 and drying racks in the case of servals; ii) because felids are seen as 2). All lion reports from 2012 to 2017 occurred in the south of competitors for wild prey species sought by hunters; and iii) to sell Quiçama (Figure 2). We did not record lions through opportunistic their parts, mainly the skin (Table 2), and sometimes bones, ca- observation. nines, claws and meat in the case of leopard. Regardless of whether TA B L E 1 Record of the most recent Leptailurus serval Panthera pardus Panthera leo year that each interviewee reported Sightings Sightings (live Sightings (live seeing or killing Leptailurus serval, Year (live animals) Killing animals) Killing animals) Killing Panthera pardus and Panthera leo 2017 41 4 19 5 2 1 2016 16 12 17 7 1 0 2015 15 2 20 5 0 0 2014 7 1 11 4 1 1 2013 6 2 9 1 2 2 2012 4 0 1 3 2 0 2007–­2011 16 5 10 4 8 3 2002–­2006 6 3 5 4 8 2 1997–­2001 2 4 3 1 8 2 1992–­1996 10 2 2 6 1 1987–­1991 12 1 1 8 4 1982–­1986 1 10 2 3 6 5 1977–­1981 7 1 1 10 6 1972–­1976 1 7 2 10 7 1967–­1971 6 1 4 Note: Data are presented in 5-­year increments for 1967–­2011 and annually thereafter. (a) (b) (c) (d) F I G U R E 1 Physical evidence of felids in Quicama National Park and Quiçama Game Reserve in Angola. (a) Leopard track. (b) Leopard skin in possession of local hunter. (c) Road-­killed serval. (d) Adult serval (‘servaline’ form) in northern Quicama NP. The individual was observed resting in a tree prior to being approached; it had not been chased there by dogs or people (photographs FB-­P) | 13652028, 2022, 3, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/aje.12946 by Readcube (Labtiva Inc.), Wiley Online Library on [15/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License BRAGA-­PEREIRA et al. 803 capture was accidental or intentional, felids were always killed is reflected in data on other species where, except for the fenced when found in traps and the parts sometimes sold. Special Conservation Area, most medium-­large mammals now exist The average of annual catch frequency per hunter between in the more remote and less populated south-­east (Braga-­Pereira, 2007 and 2017 was higher for serval and leopard, than for lion Peres, et al., 2020). The north region is open savannah with a more (Table 2). Most hunters reported that the peak in killing servals and developed road network where use of rifles and vehicles makes un- leopards for skins occurred between the 1970s and 1990s when gulate hunting and transport to markets particularly efficient (Braga-­ demand was high. According to interviewees, demand waned as Pereira et al., 2020). Its greater proximity to Angola's capital Luanda the fur industry shifted to imitation furs (international bans on intensifies the problems of access and demand for wild meat from spotted cat skins implemented during the 1970s were the actual residents outside the area. underlying factor, see Nowell & Jackson, 1996) and prices plum- Bushmeat hunting is pervasive among the human population of meted. Hunters reported significant declines in killing cats for their Quiçama, is inter-­generational and is driven by local needs as well skins as a result. Nonetheless, there is still a clear present market as demand from Luanda. It has severely depleted available prey for for felid skins which are still sold whenever one is killed. Lion killing large felids and is undoubtedly the key driver in population declines. peaked during the same period of high hunting for spotted cat furs This is compounded by targeted and opportunistic hunting of cats. although mainly for conflict retaliation, as lions were commonly Although few hunters view felids as the major motivation for hunt- perceived as a threat to livestock, as well as a competitor for wild ing in Quiçama, they are always killed when opportunity presents prey species sought by hunters. The use and local manufacture of and the very widespread prevalence of hunting across the system large leg-­hold traps is a relatively recent development and were as- translates to steady pressure from direct anthropogenic mortalities. sociated with a recent records in killing leopards (Table 1) accord- Rapidly and drastically curtailing this hunting is essential if extant ing to interviewees. These traps are now prevalent in the south of felids are to persist and for any future prospect of recolonisation or Quicama, specifically targeting leopards for their skins as well as restoration of lions and cheetahs. being deployed to capture other wildlife. Despite the high mon- In the immediate term, we recommend a significant increase of etary value attributed to lion skin, it was rarely captured recently ranger presence and effort in the south of Quiçama. Its greater con- (Table 2), presumably because the population has been very low servation value arises mostly from its remoteness, closed-­canopy for a long time. forest habitat which is not favoured by local hunters and lower Large felids have undergone a significant decline in Quiçama. human residency rather than a systematic anti-­poaching presence Both the cheetah and lion are likely extinct or possibly reduced which is currently extremely limited. Focusing on securing the south to a few individuals in the case of the latter. Neither species was would elevate protections for Quicama's remaining leopards while recorded during a camera-­trap survey of Quiçama following our also protecting relict populations of other large mammals includ- fieldwork, in 2017 (Groom et al., 2018). All recent (2012–­2017) ing buffaloes and elephants. Concurrently, it is urgent to identify interviewee reports of lions were of solitary individuals and we pathways that promote long-­term food security and employment found no evidence for breeding. Transients from eastern Angola opportunities among local people. Quicama's proximity to Luanda may explain our records, although the presence of lions is uncer- creates significant potential for tourism to become a major industry, tain everywhere in the country except for a small population in with concomitant potential benefits in job creation directly linked to the Luengwe-­Luiana complex, approximately 1000 km away in the wildlife conservation. However, that is a long-­term prospect even for south-­east of the country (Beja et al., 2019). Most recent reports relatively intact ecosystems and it will require many years of such of lions come from a mostly roadless section in southern Quiçama, investment to take hold in Quiçama. In the meantime, a collabora- particularly Omba, Morro Liso and Caleba (Figure 2). Further sur- tive management agreement with a private partner focused on im- veys for lions focusing on these areas would be valuable to estab- proving management and anti-­poaching capacity may be desirable lish whether the species is extant. (Baghai et al., 2017). Such partnerships are increasingly common and Leopards have fared somewhat better. They appear to be largely lend themselves to rapid implementation compared to large-­scale absent from the northern half of Quiçama, but there are indica- commercial investments in large and depleted ecosystems such as tions of a relatively healthy population in the park's south-­east. This Quiçama. Whatever the collaborative management formula, we F I G U R E 2 Map of Quiçama National Park (demarcated by the green line) and Quiçama Game Reserve (demarcated by the orange line) in Angola. The blue and red polygons indicate locations where leopards and lions, respectively, were reported in the past 5 years. One lion sighting was reported by more than one interviewee. Locations from where lions were reported are numbered: 1. Omba, 2. Morro Liso, 3. Baixa do Dondo, 4. Caleba, 5–­6. Quilonga. Serval locations occurred throughout the park in a polygon analogous to the park border and are not shown. Solid yellow triangles indicate the surveyed townships, which encompassed more than one human settlement. Triangle sizes are proportional to the population size of the townships. Green and greyish—­orange background areas represent forest and savannah environments, respectively. Map generated using ArcGIS 10.3.1; Datum: WGS84. Source: ESRI, Edited in Adobe Photoshop and Elaborated by Ana Caroline Imbelloni and Franciany Braga-­Pereira in July/2021 13652028, 2022, 3, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/aje.12946 by Readcube (Labtiva Inc.), Wiley Online Library on [15/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License BRAGA-­PEREIRA et al. 5 6 4 3 2 1 1.93 0.36 2.5 , |804 | 13652028, 2022, 3, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/aje.12946 by Readcube (Labtiva Inc.), Wiley Online Library on [15/02/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License BRAGA-­PEREIRA et al. 805 conclude by emphasising that the window for securing Quicama's Sightings (live (skin or (number of records) collected in skulls Body parts First hand physical evidence wildlife and the well-­being of its human inhabitants is closing very 4 4 0 rapidly. individuals) AC K N OW L E D G E M E N T S We sincerely thank the people from local communities who shared 2 1 0 their knowledge on wildlife and supported us in data collection Tracks through opportunist observations. Without them, this work would 2017 9 11 0 not be possible. We would like to thank the Environmental Ministry of Angola, Quiçama municipal administration, Quiçama Park ad- 0.19 max Annual catch frequency per hunter, 2007–­2017 7 1 ministration and all village coordinators for authorising and support this research. We thank The Rufford Foundation, the Faculdade de 0.03 0.03 0.02 min Ciências of Universidade de Agostinho Neto and the Universitat Autonoma de Barcelona for the financial support. FB-­P thanks the Average 0.54 0.07 Brazilian Ministry of Education-­ CAPES, for the master scholar- 0.2 ship. RRNA thanks National Council for Scientific and Technological 2007–­2017 killing cats Development by a productivity research grant. of hunters Number 26 29 9 C O N FL I C T O F I N T E R E S T TA B L E 2 Interviewees' information and opportunistic observations of Leptailurus serval, Panthera pardus and Panthera leo We declare no conflict of interest. Response rate calculated from all responses (hunters and rangers, N = 123) who reported perceiving decline in cats' abundance. (USD)c value Skin 20 400 400 DATA AVA I L A B I L I T Y S TAT E M E N T Commercial The data that support the findings of this study are available on re- quest from the corresponding author. The data are not publicly avail- Motivation for killing trade 53.5 68.3 28.9 able due to privacy or ethical restrictions. (response rate)b retaliation ORCID Conflict 81.6 100.0 34.1 Franciany Braga-­Pereira https://orcid.org/0000-0002-7848-2850 Rômulo Romeu Nóbrega Alves https://orcid. org/0000-0001-6824-0797 killing cats of hunters reporting Number Luke Hunter https://orcid.org/0000-0001-9653-6544 86 41 38 REFERENCES Perceived drivers of depletion (response Overhunting Baghai, M., Miller, J., Blanken, L., Dublin, H., Fitzgerald, K., Gandiwa, P., Laurenson, K., Milanzi, J., Nelson, A., & Lindsey, P. (2017). 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