Introduction

The degradation of biodiversity is one of the most important problems facing humanity today. Every day, species are disappearing due to a combination of factors, mainly linked to anthropogenic activities. Fragmentation and degradation of habitats and illegal wildlife trade are among the most important factors that threaten wildlife (Wilkie and Carpenter, 1999; Ceballos and Ehrlich, 2002). In this context, protected areas are emerging in many places around the world as the last refuges for a large number of species and are therefore critical for safeguarding biodiversity (Geldmann et al., 2013). The larger they are, the more they present a guarantee for effective and sustainable conservation of animal resources (Newmark, 1996). This is often a characteristic of transboundary protected areas. Do these transnational areas manage to guarantee the safeguarding of biodiversity better than national protected areas? Moreover, a transboundary national area implies concerted management between several countries and several regions, which is neither obvious nor easy.

Among the wildlife species that are highly threatened by human activities are the large carnivores. Because of their important space requirements, their position in the trophic chain and their economic importance, large carnivores are for the most part threatened with extinction (Woodroffe, 2000; Ripple et al., 2014), particularly on the African continent. In West Africa, the situation is particularly alarming, with these species being reduced, more so than in other parts of the continent, to protected areas. The lion is an iconic large carnivore species whose range includes West Africa.

This synthesis seeks to address, through a brief literature review and an analysis of actions on the conservation of the lion and other large carnivores in West Africa and in the W (Benin-Burkina Faso-Niger)-Arly (Burkina Faso)-Pendjari (Benin) transboundary complex, also known as WAP, the challenges of transnational management for the successful conservation of threatened species.

Conservation status of large carnivores in West Africa

Large carnivores are considered valuable indicators of the sustainability of protected area systems and ecosystems because they occupy a high position in the food chain, require large territories and are highly susceptible to human activities (Woodroffe & Ginsberg, 1998). Their conservation is therefore crucial for the preservation of biodiversity in general.

The term ‘large carnivores’ refers to the lion, hyena, cheetah, leopard and wild dog species. The most abundant of these species is the spotted hyena, classified as Least Concern on the IUCN Red List (Bohm and Höner, 2015). Nevertheless, very few studies have been conducted on the population dynamics and ecology of the spotted hyena in West Africa. The leopard, on the other hand, is considered Vulnerable in Africa and Asia (Stein et al., 2016). Present in just over a dozen countries, it remains little studied in the region. As for the cheetah, it is considered Vulnerable in Africa and Asia (Durant et al., 2015) and Critically Endangered in West and North Africa (Belbachir, 2008). Four populations occur in West Africa in five countries, the largest of which is the one in the transboundary WAP complex. The wild dog is considered Endangered in Africa and Critically Endangered in West Africa (Woodroffe and Sillero-Zubiri, 2012). It is the most threatened large carnivore in West Africa. The Red List reports the species in four West African countries, in the WAP complex and in Senegal. However, the only recently confirmed population is in the Niokolo Koba Park in Senegal, which does not appear to be sustainable. The lion, considered Vulnerable in Africa, is Critically Endangered in West Africa (Henschel et al., 2015). The WAP complex supports approximately 90% of the West African lion population (Henschel et al., 2016).

Due mainly to anthropogenic activities and their corollaries, including habitat degradation and fragmentation, direct persecution of species, conflicts with humans, poaching, trade in live animals and by-products, populations of these species are all decreasing in Africa, and particularly in West Africa. Insecurity and armed conflict in the subregion pose an increasing threat to these species (Bauer et al., 2020). The status of the species assessed in the subregion (lion, cheetah, wild dog) shows that they are more threatened than in the rest of the continent. While in Southern and Eastern Africa, several large populations of large carnivores are found outside protected areas, these species are confined to protected areas in Western Africa. In this part of Africa, conservation is often not a priority for governments, and protected areas are under great pressure from people (Bauer et al., 2020) and do not guarantee the survival of the species.

Wildlife management in the WAP complex

The WAP complex is composed of the W transboundary biosphere reserve extending over Benin, Burkina Faso and Niger, the Arly Park and its associated areas in Burkina Faso, and the Pendjari Biosphere Reserve in Benin (Figure 1). It covers more than 30,000 km2 in the three countries concerned. The protected areas that make up the reserve were created in the mid-1950s and have seen their status evolve, as well as their surface area for some. Each country has its own rules, and the protected areas making up the complex have different conservation statuses (biosphere reserve, national park, hunting area, wildlife reserve, World Heritage property), so the management methods differ. As in several protected areas in the subregion, management is highly dependent on projects, often financed by international donors. This operation favours, between projects, the illegal occupation of areas, often reducing to nothing the efforts made. This problem is aggravated by the connectivity of the areas, which can encourage the spread of threats. The Protected Ecosystems in Sudan-Sahelian Africa (ECOPAS) programme, established in early 2001, has made it possible to move towards concerted management, which has been strengthened with the various regional projects that have followed, such as the Programme d’Appui aux Parcs de l’Entente (PAPE) and the current GIZ-RBT-WAP project. An analysis at the beginning of this process had highlighted, among other things, the limited and unequal competences of the managers, the differences between the policies and strategies and the weak involvement of the riparian populations in the management (UNDP, 2014). More than 15 years later, the situation has positively evolved, thanks to efforts to harmonize and strengthen actions (Amahowé et al., 2013). Regional anti-poaching initiatives have been implemented; regular exchanges between the different teams allow knowledge and know-how to be shared. Even if differences remain between countries and efforts still need to be made, the WAP complex represents a privileged area for wildlife conservation in West Africa. It is home to the largest population of elephants in the region, important populations of species typical of the African savannah, such as the buffalo, and rare species, such as the manatee and the cheetah.

Figure 1.
figure 1

Location of the WAP complex (The names and boundaries shown and the designations used on this map do not imply official endorsement or acceptance by the United Nations)

Source: Sogbohossou, 2011.

Full size image

The active consultation framework within this complex has probably favoured the recent inscription of the Benin and Burkina Faso parts onto the World Heritage List, after the Niger part of the W Park, which has been a World Heritage property since 1996.

Lion conservation in the WAP complex: Research status and threats

The lion, like several species of large carnivores, is an iconic wildlife species. The species’ range, reduced by 75% in Africa, has been reduced by 99% in West Africa (Riggio et al., 2013). The lion is now found in five countries in the region, but only the transnational population in the WAP complex in Benin, Burkina Faso and Niger is the most viable (Henschel et al., 2014).

For a long time, the lion and other large carnivore species were neglected in West Africa. Awareness began in the early 2000s with the Limbe workshop on the status and conservation needs of the lion in West and Central Africa in Cameroon in 2001 (Bauer et al., 2001). The West and Central African Lion Conservation Network (ROCAL) was formed in 2002 following several meetings and a few initiated projects. Regular monitoring of the lion population thus began in the Pendjari Biosphere Reserve, and then throughout the WAP complex as of 2002 (Di Silvestre, 2002; Sogbohossou and Tehou, 2007; Sogbohossou, 2009). Counts on a regional scale (Henschel et al., 2014, 2016) have highlighted the threatened nature of the species and the crucial importance of the WAP Complex population for lion conservation in West Africa. Indeed, the WAP complex population was estimated in 2012 to be 350 lions corresponding to 90% of the lion population in West Africa (Henschel et al., 2014). The species is more abundant and better distributed in the Arly-Pendjari block than in the W block and appears to be at least stable based on the results of the 2014 count. Collaring individuals between 2007 and 2010 (Sogbohossou, 2011), and again from 2018 (African Parks Network, personal communication) showed movement of individuals between Pendjari and Arly and between Pendjari and W. The average territory size of a lioness which was 256 ± 154 km2 (100% MCP) (Sogbohossou, 2011) confirms the high space requirement of the species.

Other aspects were discussed, such as human-lion conflicts, genetics, ecology of the species, threats such as sport hunting and trade in by-products.

Genetic studies, including samples from the WAP complex, have confirmed the need to focus on the lion in West Africa. Indeed, the West and Central African lion is more closely related to the Asian lion than to the South and East African lion and is therefore quite different genetically from its continental brethren (Bertola et al. 2011, 2015). Although this difference is not sufficient to classify the West and Central African lion into a specific subspecies, it is significant enough to justify the need to conserve this diversity at the species level.

On the Pendjari side, Sogbohossou et al. (2011) and Efio et al. (2018) showed that hyena, lion and baboon were the main perpetrators of livestock depredation, while crops were mainly attacked by baboon, warthog and elephant. Fishermen complained of crocodile and hippo attacks. The costs of conflicts with large carnivores have been estimated at US$138 per household around the W Niger and US$196–350 around the Pendjari (Bauer et al., 2010). These conflicts have decreased in the Pendjari, with costs ranging from US$77 to US$207.2 (Efio et al., 2018). Of the methods used, guarding and fires were the most effective, even if in general, conflict reduction methods remain inefficient. Improved enclosures have proven to be effective (Bauer et al., 2010) but have not been promoted because of the relatively high cost to herders. While on the Pendjari side, lion revenge killings were not known, cases have been observed around the W. In 2018, three lions were poisoned by local people in W Benin.

Apart from human-wildlife conflicts, the trade in by-products, which encourages lion poaching, is a major threat to the conservation of the species in the subregion. Investigations, concentrated on the Benin side (Sogbohossou, 2006; Ipavec/ZSL, personal communication), have shown that lion by-products are everywhere on the markets and come from both the countries of the complex and from Central African countries.

Sport hunting, although controlled, appears to have drawbacks for lion survival in the WAP complex (Sogbohossou et al., 2014). It is one of the factors whose management varies very significantly between the countries of the complex. Indeed, this hunting is absent on the Niger side. In Benin, which hosts about 43% of the complex, the annual quota for all hunting areas, halved since 2002 after the first monitoring results, is five lions. Hunting reports indicate that on average one lion is killed for sport hunting each year. In Burkina Faso, where 36% of the complex is located, the quota was around 20 lions with an average of 11.9 lions killed per year (IUCN/PACO, 2009). It has been significantly reduced to six lions per year since 2015, after the development of the large carnivore conservation strategy in the WAP. This is an important achievement in collaborative management in the complex.

General discussion and outlook

The WAP complex in West Africa represents the last hope for saving the lion and many other species in the region. Biodiversity conservation in this ecosystem is complicated by its transnational nature. Fortunately, for nearly two decades, collaborative efforts have been made to harmonize management and research actions in the complex. Such collaborative initiatives are not legion around the world (Linnell and Boitani, 2012) and the example of WAP should be encouraged. With the various projects, the level of management has improved in all countries. The existence of a regional conservation plan for large carnivores in the WAP complex is evidence of this collaboration, although some resolutions, such as periodic regional counts of lions and other carnivores, are not automatic and the reduction or even abolition of the lion hunting quota is not supported by all parties. Research, which should guide management efforts and ensure that management is effective, needs to be improved. Studies are more national than regional, and research efforts are unevenly distributed. Several authors have shown in the case of carnivores that transnational-scale studies generate more accurate results, density and survival estimates than national-scale studies (Bischof et al., 2016; Gervasi et al., 2016; Vitkalova et al., 2018). National-level studies often tend to overestimate abundance and do not fully appreciate the consequences of differential management between protected areas. Regional counts within the WAP should therefore be continued, and this regionalization should be extended to other types of studies. The large carnivore count planned for 2019 is part of this framework, and the initiative should be continued.

Because of the lion’s large space requirements, the WAP complex offers the best habitat in the West African subregion for the species to thrive. This space must therefore be safeguarded from all sides. Even the involvement of surrounding communities in conservation must be harmonized between countries, so that conservation efforts in one part are not wiped out in another.

Actions are favoured by the fact that all the countries of the WAP complex have ratified international agreements (Convention on Biological Diversity, CITES, CMS, World Heritage Convention, etc.) and even regional agreements aimed at safeguarding wildlife, including the lion. The problem of harmonization of laws therefore does not arise, except perhaps with regard to sport hunting.

Because of the lion’s space requirements and position in the trophic chain, the WAP complex represents an important ecosystem for safeguarding the species in West Africa. Harmonization of management and research actions and sharing of results are essential, and should continue to be promoted.