Globally, wildlife is threatened by many factors including climate change [1], poaching, and agricultural expansion that leads to habitat loss [2][3]. Pastoralism, which is noted for generating human-wildlife conflict, is one of the main factors threatening the survival of wildlife, especially large carnivores. In Africa, the increase in human population has led to more land being grabbed for economic development and also for a variety of anthropogenic activities that promote subsistence [4]. Consequently, wildlife is now mainly restricted to protected areas that serve as tools for conservation. The lack of zoning for different land use forms in most Central African countries, including Cameroon, creates the scenario where communities are located in the vicinity of most protected areas. Therefore, interactions between humans and wildlife for space and other resources are frequent in these communities, leading to various human-wildlife conflicts [5]. Predation by carnivores on livestock is one of the most damaging forms of human-wildlife conflict, especially in the savannahs where pastoralism remains the main occupation of the inhabitants [6][7].

In Cameroon, there are many threats to wildlife, especially large carnivores. The main threats to the lion population include natural prey depletion, transhumance pastoralism and human-lion conflict [8]-[11]. Conflict between humans and lions due to livestock depredation is the driving factor behind the decline in the lion population in Waza National Park (Waza NP). Pastoralists (in this study includes agro-pastoralists, who are usually resident, and nomadic herders on transhumance) suffer from high levels of predation by carnivores; lions are responsible for losses of 65,597,000 FCFA ($111,952 USD) per year [12] in Waza NP. In the W National Park in Niger, 3271 domestic animals were killed between 2000 and 2006 by large carnivores (lion, hyena and jackal) representing an economic loss of 103,247,750 FCFA ($176,221 USD) [13]. The species responsible are most often killed by local communities in retaliation to attacks on livestock or in an attempt to prevent them [9][14]. Therefore, the number of large carnivores is rapidly decreasing as a direct result of conflict with pastoralists over livestock depredation [12][15]. Also, a decline in the numbers of lions, leopards, wild dogs, and hyenas has been linked to poisoning by pastoralists around Benoue National Park [8]. Livestock losses due to carnivore attacks in and around the Waza NP range from the loss of a head of livestock, about 170549.6 FCFA ($291 USD) for resident pastoralists to the loss of two heads of livestock, about 341099.2 FCFA ($582 USD) for nomadic pastoralists annually [12].

The Far North region of Cameroon is one of the most populated regions in the country and is home to many protected areas including Waza NP [16]. The region has been experiencing an influx of refugees as a result of the Boko Haram armed conflict since 2014, which is impacting natural resources in and around protected areas. The Boko Haram armed conflict has resulted in 100,000 to 150,000 internally displaced people and nearly 200,000 Nigerian refugees in the Far North Region [17]. The Waza NP was created in 1968 with biodiversity conservation as its main objective [10]. Its north-eastern periphery and part of the park is a floodplain zone suitable for livestock pastoralism, especially for transhumant and resident pastoralists in search of water and grazing during periods of drought, such as the dry season [18]. The concentration of pastoralists and their livestock in the periphery of Waza NP usually leads to incursions into the park, generating conflict over livestock depredation. The lion population in Waza NP appears to be in an alarming decline from 40 - 60 lions in 2002 to 14 - 21 lions in 2009 [9][19] mainly as a consequence of retaliatory killing of lions by pastoralists for predating on livestock.

With the Boko Haram insurgency, the number of pastoralists seeking refuge around Waza NP has increased exponentially, and the number of cattle around the park has doubled [20]. This high prevalence of pastoralists and their livestock within the home range of carnivores can significantly increase human-carnivore conflict, especially with lions. The presence of livestock in and around protected areas represents competition between wildlife and livestock for water and pasture. It also represents a threat, as pastoralists engage in killing carnivores to protect their livestock from predators [21]. Human-carnivore conflict is a global issue with significant implications for both carnivore conservation and human livelihoods. It is a major factor responsible for causing hardship in local communities and reducing human tolerance for carnivore conservation. Though there is scanty literature suggesting human-carnivore conflict in the North-eastern periphery of Waza NP [12], there is a dearth of information on the nature and magnitude of human-carnivore conflict in the park since the Boko Haram armed conflict. However, dynamics in pastoralist numbers and movements due to the Boko Haram insurgency have witnessed an increase in carnivore attacks on livestock in the North-eastern periphery. Lack of understanding of the scope and severity of human-carnivore conflict limits or actually impedes the development of evidence-based policy responses tailored in the park and its periphery. A comprehension of the nature of the conflict is vital to guide policy that will protect human livelihoods, carnivore populations, and promote coexistence of carnivores and humans. Therefore, this study was carried out to assess human-carnivore conflict and to provide knowledge for the conservation of carnivores and protection of human livelihoods in the North-eastern periphery of Waza NP.

A total of 4209 heads of livestock were recorded in all villages surveyed during the study period. All villages experienced carnivore attacks, and the frequency of attacks differed between the villages.

Predation on livestock is one of the threats to human livelihood around Waza NP, leading to the retaliatory killing of lions [12]. Villages northeast of the park and beyond now experience attacks from carnivores comparable to those in the southwest [15]. This may be explained by changes in pastoral dynamics, with movements to some villages in the northeast and even into the park for safety from armed conflicts. Actors involved in the human-carnivore conflict were predominantly resident pastoralists, with very few nomadic pastoralists. Even though this corresponds with previous findings in the park [24] and in the Pendjari Biosphere Reserve [25], the very low proportion of nomadic pastoralists may be attributed to the Boko Haram insurgency and ethnic conflicts that caused modification of their movements in the area. During the period of December 2021, the park became a refuge to an estimated 2500 pastoralists and 35,000 cattle fleeing from ethnic war. These pastoralists incurred considerable losses of livestock to carnivores during their stay in the park [11][26]. Three main carnivores are responsible for predation on livestock northeast of Waza NP: lion (Pantheraleoleo), jackal (Canis aureus), and hyena (Crocutacrocuta). These are reported to be the three main large carnivores remaining in Waza NP after the extirpation of the leopard around the early 2000s [27].

Water and pasture continue to attract pastoralists into the park, where they suffer loss of livestock to carnivores. Thus, it is not uncommon that the majority of attacks occur during the day when livestock are grazing and at waterholes. The findings of this study are in conformity with those in the periphery of the W National Park in Niger, where most attacks (56.66%) occurred during grazing [13]. Most attacks occurred during the dry season, confirming earlier authors [12][19] who reported that carnivores in this park attacked 65.20% of livestock in the dry season compared to 34.91% in the wet season. This, however, differs from findings by Garba and Di Silvestre [13] and Croes et al. [8] in the W National Park and the Benue complex. High temperatures during the dry season cause many pastoralists to enter the park in search of water, thereby increasing the availability of domestic prey for carnivores. Lougouma, Mahe, and Mbele were presented to be predation hotspots in the study, comparable to villages southwest of the park. This expansion of predation into villages not known to experience such intensity of predation in the past is a result of pastoral dynamics caused by the Boko Haram armed conflict.

Goats were the most vulnerable livestock, while the jackal killed more livestock than any other carnivore. Economically, the lion caused the highest economic loss to livestock keepers in the North Eastern periphery of Waza NP, due to the high economic value of cattle. This finding agrees with previous reports in the southern part of the park [8][12][28] and around the Benoue complex. However, they differ from those in W-National Park, where the jackal is responsible for causing the highest economic loss [13]. Although the lions killed fewer livestock compared to jackals in this study, they killed mostly cattle with a higher price value compared to goats and sheep. The loss of $390 USD on average per year per pastoralist calculated in the study induces financial stress on pastoralists and deprives them of an important economic and socio-cultural inheritance. Lions killed more small ruminants than cattle, contrary to the findings of Bauer [28] and Tumenta et al. [15] in the same study area, which demonstrated that lions preferred larger prey. This contradiction is justified by the fact that resident pastoralists represented the majority of respondents and they are usually the owners of large herds of small ruminants.

Predation as the main cause of livestock loss compared to disease and theft, accounted for the highest economic loss. These results differ from those of Bauer et al. [29] in Pendjari NP, who reported disease as the main cause of livestock loss, followed by theft and predation. As the main source of income and having great cultural importance, the loss of livestock has serious implications for pastoralists in this area.

Herders use the construction of night enclosures for their livestock, and guarding during grazing as methods to reduce predation on their livestock by carnivores. Chasing predators with noise was commonly used by pastoralists when livestock was attacked by carnivores. These results corroborate those of Garba and Di Silvestre, [13] in W National Park, who reported that 46.75% of the herders chased predators away by shouting. However, Bauer [28] recorded that hyenas were chased by herders at night by lighting torches south of Waza NP. The use of these methods does not appear to be general; just like the park authorities, some respondents reported that they had no concrete solutions to carnivore attacks.

Predation on livestock remains one of the threats to human livelihood around Waza NP, engendering the retaliatory killing of lions. Villages in the north-eastern periphery of the park now experience attacks from carnivores comparable to those in the south-west. This may be explained by changes in pastoral dynamics, with movements of resident pastoralists to some villages in the north-east and even into the park for safety from armed conflicts. The Boko Haram insurgency has influenced the movement of nomadic pastoralists; few of them use the north-east periphery of the park. Approximately 1531 livestock were predated by carnivores in the north-eastern periphery of Waza NP within three years in retrospect, resulting in an estimated total economic loss of 68,650,000 XAF ($117176.11 USD). This loss is significant for pastoralists who depend on revenue from livestock to meet basic needs such as education, health, and certain socio-cultural obligations. Although night enclosures and guarding of livestock were reported as strategies used to protect livestock from predation, these remain to be proven effective. The general consensus is that there are no concrete solutions to carnivore predation on livestock around Waza NP. Community engagement through education, awareness raising, and empowerment remains necessary. Furthermore, practical measures are necessary to mitigate the losses generated by human-carnivore conflict, to preserve human livelihoods and save populations of large carnivores in Waza NP. These measures could include targeted support for building enforced enclosures as well as community-managed waterholes away from the park.

The data used in this study are available from the corresponding author upon reasonable request.

Partial funding was received from the Leo Foundation for this study.

Pricelia N. Tumenta: Conception, design, finances, supervision, data validation, analysis and interpretation, writing and editing.

Parfait Nkontcheu Kamta: Fieldwork, data validation, drafting of the initial manuscript.

Daniel Brice Kenko Nkontcheu: Data analysis and interpretation, writing and editing.

All authors participated in preparation and critical review of the manuscript. All authors have read and approved the final version of the manuscript.

The authors sincerely thank the Lion Project Coordinator of Waza for having financially supported this work. They are thankful to the staff of the CEDC in Maroua, Cameroon, and the team of the Conservation Department of the Waza National Park (MINFOF) for their generous collaboration.