Introduction

The African buffalo (Syncerus caffer) has historically been maligned in African colonial and post-colonial veterinary and livestock communities because of its reputation for being maintenance hosts for several infectious diseases that can impact the viability of the commercial livestock industry (Michel and Bengis, Reference Michel and Bengis2012). We provide here some historical context that justified this position, but will argue that this is an unfortunate and perhaps misguided and out-of-date narrative. The dogma perpetrated throughout colonial times that African livestock systems would naturally follow northern hemisphere production systems and disease control models has been, through retrospective social and economic analysis, challenged: ‘(…) the assumptions that the high-value/high cost option [in terms of livestock industry] is necessarily the best [in Africa] – and the one that should be striven for – and the low value/low cost [of extensive livestock] is automatically bad news are not upheld’ (Scoones et al., Reference Sintayehu, Heitkönig and Prins2010). The mandates of veterinary services are almost exclusively set up to protect intensive animal-based agriculture investments and trade. This paradigm is promoted by high-income industrialized countries, and most international trade ignores the impacts on wildlife economic opportunities and the realities of domestic livestock limitations in African countries that face restricted international trade opportunities. There is also a vested interest in maintaining this situation of perceived risk from disease, especially for the expansion of European breeds and modern intensive systems of livestock agriculture, which are often described as ‘improved, productive and efficient’ in contrast to African extensive and pastoral systems, reflecting a kind of neo-colonialism.

We argue that as the ‘new deal’ required by recognition of the Anthropocene becomes accepted, the dogma will change. Future economic, agricultural and overall development models will need to fit into the finite environmental envelop that will constrain human activities by choice or by force. There is a greater appreciation of the negative environmental, climate, health and socioeconomic externalities of intensive livestock systems, and of the need for more inclusive landscapes where biodiversity conservation and local citizens’ well-being meet and more value is put on indigenous knowledge and value systems (cf. Gordon et al., Reference Gradwell, Schutte, Van Niekerk and Roux2016). African continental ownership of future policy has been controversial for decades and may well be the final arbiter on this controversy (Artz et al., Reference Artz, Motsamai, Zacharias and Tueller1991; Prins, Reference Prins and Verwey1989).

The Development of Coexistence of African Pastoralism with Wildlife

Contemporary views on buffalo reach back to the introduction of Eurasian cattle along with the colonization of the African continent. In the following sections, we focus on eastern and southern regions of Africa where most of the ‘conflict’ is expressed, and mostly exclude western and northern regions of the continent, where cattle herds have deep histories, including truly African breeds (Prins, Reference Prins, Prins, Grootenhuis and Dolan2000). Issues between cattle and buffalo in relation to disease are less pronounced in these regions, if only because buffalo are absent in the north and there are only small, scattered buffalo populations in the west where pastoral systems dominate (Kock et al., Reference Kock, Kock, de Garine-Wichatitksy, Chardonnet, Caron, Melletti and Burton2014; Chapter 4).

Before colonization, in the seventeenth century, large areas of the southern and western regions of southern Africa were sparsely populated by the nomadic hunter-gatherer San people and nomadic farmers (Khoikhoi/Khoisan) who were probably the first livestock owners in these southern lands. The more eastern and northern areas of southern Africa were inhabited by primarily Bantu groups who were also pastoralists, owning Sanga (Nguni) cattle, sheep and goats, and also growing edible crops (Maggs, Reference Maggs, Cameron and Spies1986).

The situation in East Africa in the pre-colonial context was composed of extensive grasslands and traditional cattle owning coexisting with abundant wildlife. Wildlife behaviour and traditional livestock practices such as pastoralism and transhumance coevolved in East Africa, sharing space and participating in the engineering of open grassland savannas (Chapter 2). Bos taurus africanus or Sanga cattle is generally considered the indigenous African cattle from eastern African origins. African cattle were likely derived from complex introductions over centuries, and even from aurochs in Egyptian times from the Near East (Prins, Reference Prins, Prins, Grootenhuis and Dolan2000). Some species from North Africa are recognized to have existed beyond Egypt but are now extinct, for example Bos primigenius mauretanicus (Tikhonov, Reference Tjaronda2008). Along with hybridization between B. taurus and B. indicus or zebu cattle, a variety of breeds now constitute cattle populations in sub-Saharan Africa. These breeds coexisted with wildlife for over a thousand years and developed some resilience to infection, parasites and drought. Both pastoralism and wildlife may have benefited from each other, co-creating integrated landscapes. Wild ungulates used human-occupied areas to avoid predation by wild carnivores or, in the case of smaller antelopes, to benefit from areas grazed by cattle, feeding off early shoots after heavy cattle grazing (Augustine et al., Reference Asibey2011; Georgiadis et al., Reference Georgiadis, Ihwagi, Olwero and Romanach2007; Odadi et al., Reference Odadi, Karachi, Abdulrazak and Young2011). In the case of the hirola antelope (Beatragus hunteri), the most endangered artiodactyl in Africa today, its survival in a narrow range in Kenya was closely linked to local pastoralists. Hirola, having become extinct elsewhere apart from a small part of Kenya, often concentrate, feeding around nutrient-rich old boma sites and short grasses established by livestock grazing pressure where predators are persecuted. Hence, they benefited inadvertently from traditional pastoralism while indigenous people considered the presence of hirola as a good sign for their cattle (Andanje, Reference Andanje2002). The relationship with wildlife was used to predict resource availability (tracking movements) and as a source of culture and food when necessary (Lankester and Davis, Reference Lankester and Davis2016). Presently, with the wide availability of guns, rifles and other weaponry in the hands of pastoralists, this has changed, and the hirola is now next-to-extinct and features on the IUCN Red List.

Archaeological data indicate that diseases at the wildlife/livestock interface may also have been an important component of this interaction. Even with indigenous breeds of cattle, the establishment of pastoralism in some African ecosystems was constrained by wildlife and vector-mediated diseases, such as tick-borne diseases (e.g. East Coast fever, ECF), trypanosomiasis and malignant catarrhal fever (MCF). Practices such as fire, bush-clearing and contact avoidance, respectively, have taken time to evolve to counter sanitary threats and allow pastoralism to colonize new landscapes (Gifford-Gonzalez, Reference Glover, Macfarlane and Bengis2000). Most of Africa’s livestock farmers practiced (and still do) unfenced extensive systems with indigenous breeds of cattle (historical hybrid B. indicus and B. taurus africanus). These breeds and systems have proven to be more sustainable and resilient to infectious diseases, parasites, heat and droughts when compared to most B. taurus breeds and livestock systems imported from Europe (Mattioli et al., Reference Mattioli, Pandey, Murray and Fitzpatrick2000; Morris, Reference Morris2007). Under conditions of widespread vectors and pathogens in Africa, acquired resistance enables greater sustainability of traditional livestock keeping and resilience in the face of epidemic and endemic diseases. Little control is practiced, and fencing is very limited in its use in African rangeland, mostly to separate a few historical ranches from open range. Most of the separation between buffalo and African cattle herds derives from behavioural determinants, mostly human aggression and use of dogs for reasons other than disease, with an avoidance response from buffalo herds leading to their predominance in protected areas where humans and domestic animals are mostly excluded.

Re-Drawing of the African Landscape – Colonization and De-Colonization – Emergence of Production-Oriented Livestock Systems in Africa

Southern Africa was one of the earliest subregions to be colonized by European settlers, which took place in the seventeenth and eighteenth centuries. The settlers landing in the southern Cape were European with a drive towards settled agriculture and land tenure, which was not part of indigenous tribal cultures. These moves led to major conflicts not only between different European cultures but with several tribes over land and resources. This territorial expansionism by the settlers accelerated in the nineteenth century and culminated in the forming of several nation-states. These included the Boer republics in South Africa and periods of British sovereignty over large areas of southern Africa up to the miombo belts and forests of East and Central Africa, as well as the German colonization of South West Africa (Namibia) and East Africa (Tanganyika, Ruanda-Urundi). In addition, around the end of the nineteenth century, serious colonization of East Africa by the Germans and British happened with Tanganyika ultimately falling under British rule on behalf of the League of Nations.

During these territorial expansion periods, settlers, European hunters, war and disease all eventually had major impacts on wildlife populations. The settlers were heavily reliant on hunting to supply their daily protein needs, and there are many historical accounts of wild herds stretching from ‘horizon to horizon’ (e.g. Beard, Reference Beard1977; Prins and De Jong, Reference Prins, de Jong, Bond, Kiffner and Lee2022). There was also a large amount of commercial hunting for dried meat and hides as well as sport hunting for trophies, the slaughter continuing unabated as though the resource was limitless and infinite. These practices were also used to clear land to make space for grazing domestic stock, with wildlife numbers suffering through depredation and competition. In South Africa, the kwagga (Equus quagga), Cape lion (Panthera leo melanochaita) and bluebuck (Hippotragus leucophaeus) were driven to extinction, and the bontebok (Damaliscus dorcas dorcas), white rhinoceros (Ceratotherium simum) and Cape mountain zebra (Equus zebra) were pushed to the brink. Countless animals were killed to feed trading safaris, and colonial armies also were fed meat from game. Lands were cleared of wildlife species to create space for white settlers and local farmers alike, as in Kenya (Adamson, Reference Adamson1968, pp. 97 ff, 120) and many other places. The total number of African elephants (Loxodonta africana) killed by elephant control officers in East and southern Africa may perhaps be equal to the number that was poached for their ivory. The onslaught on Kenyan wildlife during World War II is jaw-dropping, where herds of game were used as targets to represent enemy troops and elephant herds were even bombed (Prins and De Jong, Reference Prins, de Jong, Bond, Kiffner and Lee2022).

In southern Africa, the power lay with cattle keeping and agriculture communities during colonization, while National Parks (NPs) were seen as the only way to address perceptions of irrational pastoralism (Lankester and Davis, Reference Lankester and Davis2016). Ironically, the strong militarization of protected areas, including the post-independence exclusion of people from traditional lands and even the banning of hunting (both sport and traditional), generated animosity which may even have laid the foundations for the poaching of elephant, rhinoceros and other species after the colonies collapsed. In each of these arenas, there was inevitably conflict. For example, in southern Rhodesia, wildlife were culled extensively to create buffer zones between wildlife-rich areas and colonial farmer production areas for various reasons (Mutwira, Reference Mutwira1989), and to target the preferred hosts of tsetse flies in order to reduce tsetse-infested areas and the occurrence of trypanosomiasis (e.g. Zululand; Andersson and Cumming, Reference Anderson, Doughty, Anderson and Paling2014).

With colonization, pastoralism was also catastrophically reduced through the introduction of contagious bovine pleuropneumonia, followed by the great rinderpest pandemic (the Masai–Maa cultural group was reduced by two-thirds: Prins and De Jong, Reference Prins, de Jong, Bond, Kiffner and Lee2022; Box 12.1). Some wildlife species also suffered huge losses in East and southern Africa, deregulating African savannas’ trajectories in the following decades with alteration of the habitat (e.g. bush encroachment; Holdo et al., Reference Holdo, Sinclair and Dobson2009) and wildlife diversity and abundance (reduction then increase following the space liberated by human populations). The phylogenetic relationship between African buffalo and cattle, although quite distant (Chapter 2), leads to shared pathogens and led to the frequent accusations of the buffalo being a reservoir of key livestock diseases during colonial and post-colonial eras. The buffalo became synonymous with the early concept of the so-called wildlife–livestock interface in Africa (Kock, Reference Kock, Barrett, Pastoret and Taylor2006), showing the buffalo’s prominent role in several diseases (Chapter 9). During colonial times, the epidemiology was poorly understood and quantified, and most of the evidence was derived from a few human-altered ecosystems, mainly in southern Africa. At least in this region, the negative perception of wildlife and diseases led to clear segregation of land uses dedicated to livestock production and the separation of domestic and wild ecosystems. New diseases and pressure from white colonialists forced traditional pastoralists and their livestock into newly emerging tsetse fly (Glossina spp.) belts (Prins and De Jong, Reference Prins, de Jong, Bond, Kiffner and Lee2022). In East Africa, this was less of an issue because local livestock and forestry, for instance, could be combined (Brasnett et al., Reference Brasnett, Richmond and Dimbleby1948). This divergence between concepts of ‘modern’ livestock agriculture and traditional pastoralism, between the south and the north, took on sociocultural and political dimensions. During colonial or European rule, a narrative against traditional local livestock keeping in Africa developed, and this has persisted to some extent, frequently justified by the disease paradigm and from where the power in control policies lies, which is mostly within industrialized western nations. This was understandable given that epidemic and vector-borne livestock diseases were a major constraint to the expansion of European livestock systems in southern Africa (e.g. Gunn, Reference Ferguson and Hanks1932), while in the east and west of the continent, the pastoral systems thrived. Attempts at ranching cattle gradually declined over the twentieth century in East Africa, with large landholdings reverting successfully to wildlife–cattle integrated management with meat and tourism activities and, sometimes, communal ownership (NRT, 2020).

Positively, the colonial era inspired protection of game and areas of land in law. However, as royalty had given hunting rights exclusively to the wealthy in Europe, colonialists discriminated against local people and limited people of pastoral communities’ access, making wildlife a preserve of the rich behind a conservation banner. People and animals were negotiated or shifted away and excluded from extensive productive rangelands, and this has persisted to this day.

The pattern of wildlife decline was repeated in eastern and Central Africa during the period of decolonization from the mid-twentieth century. Indigenous communities and rebel armies slaughtered game in the transition periods, partly during conflict for food, and to push back against colonial masters, conducting revenge killings after years of exclusion and to avoid future restrictions by eliminating game. However, some positive post-colonial developments based on the colonial systems should be noted. For example, in Kenya, these included much-debated bans on hunting (Anon., 1977) in attempts to slow the decline in wildlife; improved protection agencies; and eradication of rinderpest, a big killer of buffalo (Kock, Reference Kock, Barrett, Pastoret and Taylor2006). These measures appear to have stabilized buffalo numbers, at least in Kenya, over the last three decades (Grunblatt et al., Reference Guerrini, Pfukenyi and Etter1996; KWS, 2021). Ironically, after the end of the colonial era, the same colonial model of disease management was adopted by emerging states with identical results. For the natural ecologies in these areas to recover, old paradigms had to be overturned or remain to be challenged. Interestingly, in the fields of human health and global health, this theme has also been gathering momentum as shackles remain on poor countries and a few high-income countries dictate the human health and health industry agenda (Büyüm et al., Reference Büyüm, Kenney and Koris2020).

Current Situation at The Interface, The Burden of History and the Weight of Changes

Today, many new pressures, including climate change, human population growth, associated buffalo–cattle–human conflicts (Matseketsa et al., Reference Martínez-Avilés, Iglesias and De La Torre2019) and agriculture (Prins and De Jong, Reference Prins, de Jong, Bond, Kiffner and Lee2022) are reducing the viability of buffalo populations, now highly dependent on protection. In much of sub-Saharan Africa, with under-investment from abroad, land-use pressure is mounting with heavy investment in extractive industries, including forestry, and mining, while the human population is growing at a particularly high rate. This human population impact on land is, to some extent, compensated by urbanization, but demand for food continues to grow. As a result, the lands that were designated for wildlife and pastoralism are being encroached on and put under increasing pressure. Increasing populations of livestock, more or less replacing wild ungulates, have been documented in some countries and regions such as Ethiopia (Gebretsadik, Reference Gifford-Gonzalez2016), and countries with strong wildlife economies like Kenya have also been affected, but at a slower rate (Ogutu et al., Reference Ogutu, Piepho and Said2016). Political power lies in the hands of urban communities and agriculturalists, while pastoralists are weakly represented in government. These communities have historically been persecuted and discriminated against through land-use policy that removes key resource areas from their control. Some of these conflicts, especially in West and Central Africa, remain serious and violent, with recent examples within the Fulani (a.k.a. Peul) pastoralists community around the Lake Chad basin. Today, there is an East and Central African model, where open rangeland remains available and true pastoralism and transhumance are still practiced, and a southern African model, where land tenure has dominated the scene for the past 200 years, and little ‘open’ rangeland still exists.

The ‘trade sensitivity’ around the main diseases impacting livestock production that were prominent in Europe at the time of colonization was simply transplanted into African colonial systems (see, for instance, Empire Marketing Board, 1930). In some regions of sub-Saharan Africa, disease in wildlife is still seen by animal health agencies as a significant barrier to agricultural development. However, we posit, that is mostly a result of residual dogma and supporting narrow policies that benefit the main agribusinesses trading in a globalized world. Veterinary fencing, with a primary role to separate buffalo from cattle populations, has been used extensively across southern Africa (Chigwenhese et al., Reference Channumsin, Ciosi and Masiga2016; Ferguson and Hanks, Reference Ferguson, Cleaveland and Haydon2010), but is cropping up in Europe too (like the wild boar fence separating Denmark from Germany) and has been in place in Australia since the end of the 1800s. Even until the present day, the government veterinary services of South Africa, Zimbabwe, Botswana and Namibia all practice so-called ‘hard edge’ disease control measures, which include barrier fences such as the Kruger NP (KNP) western and southern boundary fence (restricting contact with livestock in South Africa – while the eastern Mozambique fence is only partially closed), the Ngamiland buffalo fence (partly separating the Okavango Delta from livestock areas in Botswana) and the Namibian veterinary cordon fence to prevent contact between commercial livestock and wildlife, particularly buffalo. In Zimbabwe, 55 per cent of the buffalo population is fenced today, and this policy is sometimes supported by conservation non-governmental organizations (NGOs) sharing an interest in preventing cattle and people from entering the protected areas and wildlife from getting out. A similar cordon line exists between Western Zambia and Angola. These veterinary measures clearly separate commercial livestock production that is protected for their markets versus wildlife systems and small-scale livestock production imprisoned in the fenced areas and unable to be marketed. Indeed, Botswana’s fences protect blocks that are designated for commercial trade with a foreign region (European Union) and certainly constrain local production and compromise local food security, with devastating consequences on wildlife communities and ecosystems. To show the tenacity of such beliefs and practices, the Veterinary Cordon Fence in Namibia was enacted by the German Reichstag in 1905, and still discriminates between livestock producers from beyond the fence and those ‘on the right side’ of it (e.g. Miescher, Reference Miescher2012; Tjaronda, Reference Thomson, Bastos, Coetzer and Tustin2008). In addition, certain designated geographical control zones have been declared for diseases such as foot and mouth disease (FMD), African swine fever and corridor disease (buffalo-associated theileriosis) in South Africa. As in Botswana, these are primarily intended to protect designated production zones exporting to high-end meat markets. Its mirror side, however, is that by framing cattle meat as coming from potentially ‘dangerous’ areas where veterinary control may be wanting, farmers in these importing countries are enabled to prevent competing meat from coming to ‘their’ market (Robinson, Reference Robinson2017; Whittington et al., Reference Whittington, Donat and Weber2019). This fencing policy, mostly imposed by the state on local stakeholders is costly, opposed by pastoralists and accepted by commercial livestock and crop farmers for obvious reasons. In all but a few cases (UNEP, 2011), fences are detrimental to wildlife movements and conservation, and they require significant maintenance (De Jong et al., Reference de Garine-Wichatitsky, Caron and Gomo2020; Gadd, Reference Gebretsadik2012). In the absence of good maintenance, fencing deteriorates and becomes porous (Chigwenhese et al., Reference Channumsin, Ciosi and Masiga2016).

However, to some extent fencing for disease control and for reducing wildlife conflict with people and agriculture has taken hold in much of the continent. In East Africa, arguments against this approach seem to have helped to slow any progression down this path for reasons of disease control (Kock, Reference Kock, Ferguson and Hanks2010). In many situations, with open interfaces between protected areas and communal land, buffalo’s natural and adaptive avoidance of cattle reduces the risk of disease spillover, and these systems show high tolerance to infection without many diseases expressed (Caron et al., Reference Caron, Cornélis and Foggin2016; Meunier et al., Reference Meunier, Sebulime, White and Kock2017; Valls-Fox et al., Reference Valls-Fox, Chamaillé-Jammes and de Garine-Wichatitsky2018). In times of drought, contact rates can change dramatically and disease epidemics are more likely to occur (Bengis et al., Reference Bengis, Kock and Fischer2002; Kock, Reference Kock, Osofsky, Cleaveland and Karesh2005). A severe drought in Kenya from 1993 to 1994 probably precipitated the large rinderpest outbreak in the 1990s in East Africa (Kock et al., Reference Kock, Wambua, Mwanzia and Wamwayi1999, Reference Kock, Wamwayi and Rossiter2006), killing 60 per cent of buffalo in the Tsavo National Park ecosystem. However, this drought killed some 70 per cent of the buffalo in the Masai Mara even before the disease arrived (Dublin and Ogutu, Reference Dublin and Ogutu2015). In East Africa, livestock remained in open pastoral systems and coexisted with wildlife, which thrived (Homewood et al., Reference Homewood, Trench and Brockington2012), but mostly with buffalo only surviving in protected areas and buffer zones. Agropastoral, farm or ranching communities and pastoralists are known to use self-managed movement control to avoid epidemics. However, increasing densities of people and their livestock ultimately can continue to lead, in the absence of new policy, to the demise of wildlife (Prins, Reference Prins1992; Prins and de Jong, Reference Prins, de Jong, Bond, Kiffner and Lee2022). This is not inevitable; for example, more integrated developments in pastoral land use, such as in Kenya, have led to remarkable overall stability in wildlife populations over decades, despite rapid development, human population expansion and declines of wildlife on state lands (KWS, 2021).

When there is insistence on disease elimination rather than control in livestock, the interface becomes more threatening. De Vos et al. (Reference De Vos and van Niekerk2016) clearly described, for example, the challenge in South Africa of perceptions around certain species and diseases stating,

In more extensive pastoral systems, wildlife and livestock remain integrated to some degree, with designated protected areas allowing the survival of core buffalo populations. The protected area models adopted in West and Central Africa, with core protected areas surrounded by buffer zones with limited human activities (e.g. game hunting, some pastoral activities) offered management of the buffalo/cattle interface that has allowed the survival of core buffalo populations, even if isolated (Bauer et al., Reference Bauer, Chardonnet and Scholte2020), as long as there was no security crisis. That system has subsequently collapsed in many places in West and Central Africa (Scholte et al., Reference Scholte, Pays and Adam2021), and perhaps only timber concessions and privately managed reserves appear to maintain buffalo (Chapter 4). Even though open systems have allowed wildlife to thrive, buffalo are not tolerated by pastoral livestock owners due to the aggression sometimes shown by buffalo to pastoralists and direct competition for water and grazing. This has led to the virtual extirpation of buffalo from some communal lands (Metzger et al., Reference Metzger, Sinclair, Hilborn, Hopcraft and Mduma2010). There are a few exceptions with forest buffalo (Syncerus caffer nanus) and some savanna buffalo in forested areas such as Boni Dodori in Kenya, where large populations >10,000 share habitat with hunter-gatherer/small-scale cropping communities (Chapter 4). A similar peaceful coexistence can be seen in along the Kazinga Channel between Lakes Edward and George in Uganda (Kock, personal observation).

Where a wildlife economy dominates as a source of foreign exchange revenue over agriculture, such as in Kenya and Tanzania, the political establishment lends a more sympathetic ear. In addition, and perhaps as a result, all attempts at draconian veterinary measures detrimental to the wildlife and pastoral economy have never been applied successfully, even if policies exist on paper. A sustainable balance is often achieved, allowing for livestock keeping and healthy wildlife populations to be conserved and contributing to tourism and the economy. Increasingly this tourism industry is locally owned and beneficial to indigenous communities (Mureithi et al., Reference Mureithi, Verdoodt, Njoka, Olesarioyo, Van Ranstithi, Kideghesho and Rija2019; Tyrrell et al., Reference Tyrrell, Russell and Western2017; Western et al., Reference Western, Tyrrell and Brehony2020). While in the south of the continent, where livestock owners were well connected politically and largely dominated the land-use arguments in favour of agriculture for over a century (Munangándu et al., Reference Munangándu, Siamudaala and Mambota2006), this has been reversed to some extent more recently, with an expansion of wildlife ranching and conservancies (Chapter 13). In many of these ranches and conservancies in South Africa, integrated farming with livestock and wildlife now takes place, but with the exclusion of buffalo and large predators. Legislation dictates that buffalo and cattle may not be kept on the same property.

In addition to these influences, a failure to invest in local communities around wildlife protected areas brings more pressure. Estimates of locally shared revenue from conservation areas like the Serengeti are only 5 per cent of total annual income and only go to a few households, with the majority of beneficiaries being a distant private sector and government exchequers (Homewood et al., Reference Homewood, Trench and Brockington2012; Lankester and Davis, Reference Lankester and Davis2016). In Zimbabwe, where Operation CAMPFIRE first resulted in much higher revenue sharing with local communities, this community benefit fell to only a few euros per year after the CAMPFIRE strategy was ‘invaded’ by local politicians and bureaucrats (Poshiwa et al., Reference Poshiwa, Groeneveld and Heitkönig2013a, Reference Poshiwa, Groeneveld and Heitkonig2013b). We are not judging what was or is right or wrong in this debate on ‘human versus biodiversity rights’, but are trying to present the different perspectives and historical precedence around disease which may explain past and current actions.

Recovery of African Pastoralism and Wildlife in Africa – Is This Possible?

With ongoing climate change and continuing human population growth, as resources decline and drylands increase, the use of available water from rivers and wetlands will probably increase the likelihood of buffalo and cattle meeting. Increased grazing pressure may result in ecological disturbance and degradation of natural resources. If this progresses, the natural disease regulation benefits of the ecosystems may begin to decline and vector–host–pathogen dynamics may be disturbed, which will impact livestock more. Eventually, wildlife may also suffer as malnutrition and stress erode even their resilience to disease, and whole ecosystems may begin to decline with population crashes. As buffalo are removed from pastoral or agriculturally designated lands, they are still frequently blamed as a source of diseases for livestock kept by communities surrounding the conserved areas where buffalo are mostly found. In addition, the ecological consequences of agriculture, ranching and overall degradation of fenced land, especially with high stocking rates, create poor conditions for ungulates and increased vulnerabilities to disease irrespective of the presence of carrier animals peripherally or in the parks (Glover et al., Reference Gordon, Prins and Squire2020; Kinne et al., Reference Kinne, Kreutzer, Kreutzer, Wernery and Wohlsein2010). However, the conflict remains high in livestock keepers’ minds. As populations of cattle grow, the domestic animals themselves become more epidemiologically significant, through a mere numerical relationship, and become the main carriers of diseases and a preferred food source for disease vectors (Channumsin et al., Reference Condy and Vickers2019; Clausen et al., Reference Chigwenhese, Murwira and Zengeya1998). This can change the epidemiological dynamics of pathogens locally, shifting the role of wild and domestic species, and can drive the further spread of disease within the domestic population, the community of wild and domestic ungulates, which may include spillback to wildlife.

With today’s improved epidemiological knowledge, better diagnostic tests and better livestock vaccines, it is hoped that African endemic disease control can become less conflictual and more environmentally friendly. The movement of diseases between wildlife and livestock is in fact bi-directional. With dwindling wildlife numbers in many countries (especially in West and Central Africa: Chapter 4), wild animals can also be threatened by persistent livestock disease spillover (e.g. bovine tuberculosis, peste des petits ruminants and brucellosis) to naive and sometimes critically endangered wildlife (Pruvot et al., Reference Pruvot, Fine and Hollinger2020; Shury et al., Reference Shiferaw, Abditcho, Gopilo and Laurenson2015; Viggers et al., Reference Viggers, Lindenmayer and Spratt1993; White et al., Reference White, Wallen and Geremia2011; Chapter 9).

More recently, changing views on livestock management and values of wildlife have resulted in the fading away of earlier red lines on diseases such as FMD (Ferguson et al., Reference Gadd, Somers and Hayward2013; Weaver et al., Reference Weaver, Domenech, Thiermann and Karesh2013). Strengthening wildlife-based economies in Africa, and innovative thinking around integrative management of wildlife and livestock and the rangelands in which they coexist, are increasing environmentally friendly land uses (Ferguson et al., Reference Gadd, Somers and Hayward2013). Development of softer disease policies such as the use of commodity-based trade (CBT) to circumvent FMD trade restrictions renders FMD elimination an obsolete goal (Thomson et al., Reference Tikhonov2013) even if currently its acceptance remains slow. Nevertheless, there are now increasing opportunities for trade without disease control burdens. The emphasis on intensive husbandry of livestock with production and profit as the main goal is shifting towards more sustainability in food systems. Other benefits of mixed rangeland management include climate change mitigation and reduction in disease control costs. Perhaps the ultimate arbiter of future livestock systems will be the concerns over their role in biodiversity loss, as competitors for food crops which might otherwise be used for humans and climate change ramifications. A shift from animal- to plant-based diets is gaining momentum in many countries in Africa where meat consumption remains low per capita compared to other continents while biodiversity remains high (Figure 12.1).

Figure 12.1 Meat supply per person in 2017.

(Source: FAO, see interactive map at https://ourworldindata.org/meat-production)

This illustrates how the narratives around Africa and meat consumption are largely distorted. Africa has much more sustainable low-impact animal-based agriculture and wild meat consumption in terms of environment, biodiversity and climate change. A wildlife economy continuing and developing alongside an agricultural economy in Africa could address and prevent many of the challenges currently facing advanced economies.

In South Africa, a buffalo production model emerged in the 1990s and has evolved to this today with controversial outcomes (Box 12.2). The more visionary wildlife ranching and community-based natural resource management (CBNRM) practiced in Namibia and Zimbabwe since the 1970s, utilizing mainly the extensive and relatively free-range systems and conservancies, has been important in bringing communities on board. Kenya and Tanzania have taken strides in recent decades, with integrated pastoral livestock and wildlife ecosystems such as the Northern Rangeland Trust and Ngorongoro Conservation Area, respectively. However, upheavals and violence in the latter area in 2022 may throw another light on the success of this narrative (Kihwele et al., Reference Kihwele, Veldhuis and Loishooki2021; ROAPE, 2022). In regard to such initiatives, South Africa is lagging behind, with Uganda and Ethiopia and countries in West Africa (with large and small buffalo populations) even more so.

Under several proposed future alternative development scenarios, if land is released from animal production for rewilding, disease epidemics are likely to decrease without abundant domestic animal host populations. Historic concerns about wildlife as disease reservoirs will dissipate, resource competition will decline and wildlife-based economic opportunities will arise.

Are Buffalo and the Diseases They Carry Still a Concern in the Modern African Landscape?

As observed, the diseases which have been much thought of in the context of buffalo and cattle are African strains of FMD, corridor disease (theileriosis), bTB, brucellosis, trypanosomiasis, heartwater and Rift Valley fever (Michel and Bengis, Reference Michel and Bengis2012; Chapter 9). African buffalo are believed to serve as maintenance or incidental hosts and amplifiers of these cattle diseases with potential spillback (Musoke et al., Reference Musoke, Hlokwe, Marcotty, du Plessis and Michel2015). However, there is relatively little supportive evidence for spillback happening, and new evidence is challenging long-held assumptions. In truth, cross-species infections are rarely documented or confirmed, but epidemics can occur, especially where there is a policy on disease elimination in livestock, and a hard boundary between disease-free and infected populations created by fencing. The following paragraphs provide examples to illustrate the trends.

There is no doubt that FMD SAT strains are maintained by buffalo, and they may represent the original coevolved host (Anderson et al., Reference Andersson, Cumming, Andersson, de Garine-Wichatitsky, Cumming, Dzingirai and Giller1979; Thomson and Bastos, Reference Thomson, Penrith and Atkinson2004). However, as with many ‘emerging’ infectious diseases, confirming origins retrospectively is nearly impossible. Cattle that are FMD-free can be at risk of a breakdown in status at any interface with buffalo and other carriers of the virus (Guerrini et al., Reference Gunn2019; Hargreaves et al., Reference Hargreaves, Foggin and Anderson2004; Miguel et al., Reference Miguel, Grosbois and Caron2013). However, even this strong narrative of buffalo being the original sole ‘source’ of SAT strains of FMD in Africa is now in question. Buffalo may perhaps have been so historically, but more recent evidence in certain regions of the continent shows that cattle can also maintain cattle-adapted SAT strains for extended periods (Omondi et al., Reference Omondi, Gakuya and Arzt2020; Wekesa et al., Reference Wekesa, Sangula and Belsham2015). Not all cattle outbreaks with SAT viruses might have been from buffalo, but in several southern African countries, cattle outbreaks have been shown to be caused mainly by buffalo isolates. Many of these buffalo isolates that are regularly mutating have subsequently been incorporated into cattle vaccines.

Some vector-borne diseases – for example, the tick-borne disease theileriosis and heartwater – can cause very high mortality (up to 100 per cent) in naive African cattle (Lawrence, Reference Lawrence, Norval, Perry and Young1992; Neitz et al., Reference Neitz, Canham and Kluge1955), while certain exposed cattle populations living at extensive buffalo/cattle interfaces suffer fewer losses (Young, Reference Young, Irvin, Cunningham and Young1981). As with FMD, cattle-adapted Theileria strains have evolved and emerged, causing the diseases known as East Coast fever (ECF) and Zimbabwe theileriosis (January disease), and these diseases can circulate independently in cattle without any buffalo presence. In addition, with regards to heartwater, the development of premunity and endemic stability has resulted in fewer losses. Other vector-borne diseases can have multiple reservoir hosts. With trypanosomiasis, wildlife certainly provides reservoir hosts, some preferred by tsetse flies, such as buffalo, wild porcines, spiral-horned antelopes and elephants, but from a risk perspective, these populations can also dampen environmental infection loads away from livestock and humans, reducing disease risk and impacts (Channumsin et al., Reference Condy and Vickers2019; Clausen et al., Reference Chigwenhese, Murwira and Zengeya1998). As with the Rift Valley fever virus, buffalo are susceptible but are only one among a myriad of susceptible wild and domesticated ruminants (Swanepoel, Reference Swanepoel1976).

On the bacterial side, even if livestock-origin brucellosis and bTB have crossed the ‘species barrier’ many times, their impact on free-ranging populations of buffalo appears to be ecologically insignificant. For brucellosis, this is true even in populations where brucella spp. antibody prevalence is quite high, as in KNP, South Africa (De Vos and van Niekerk, Reference De Vos, Cumming and Cumming1969; Herr and Marshall, Reference Herr and Marshall1981; Ndengu et al., Reference Ndengu, Matope and de Garine-Wichatitsky2017) with occasional reports of disease in buffalo and spillover to other species (Condy and Vickers, Reference Clausen, Adeyemi and Bauer1972; Gradwell et al., Reference Grunblatt, Said and Wargute1977). Certainly, the observation of hygromas in older buffalo is not uncommon throughout the buffalo range. Much more has been published on bTB in buffalo, but almost exclusively focused on South Africa (but see Sintayehu et al., Reference Sintayehu, Prins, Heitkönig and de Boer2017a), where it is thought to have been introduced with cattle imports in the eighteenth and nineteenth centuries (Paine and Martinaglia, Reference Paine and Martinaglia1928). The most attention is given to the ‘compressed’ fenced or semi-fenced protected areas such as KNP and Hluluwe/Umfolozi NP (Bengis et al., Reference Bengis, Kriek and Keet1996; de Garine-Wichatitsky et al., Reference Dasmann2010), with high buffalo densities. Here, there are concerning trends with buffalo apparently suffering some disease and mortality. There is evidence that this species is also driving infection in other wildlife species such as lions, Panthera leo (Keet et al., Reference Keet, Kriek, Bengis, Grobler and Michel1996), greater kudu, Tragelaphus strepsiceros (Keet et al., Reference Keet, Kriek, Penrith, Michel and Huchzermeyer2001) and chacma baboons, Papio ursinus (Keet et al., Reference Keet, Kriek, Bengis and Michel2000). This has raised conservation concerns regarding less-populous species and predator/scavenger impacts. In more open unfenced systems in Uganda, bTB was confirmed around the 1960s and seroprevalence for bTB has been consistently high in buffalo over the intervening years, yet the disease is rarely reported (Guilbride et al., Reference Guilbride, Rollison and McAnulty1963; Meunier et al., Reference Meunier, Sebulime, White and Kock2017; Woodford, Reference Woodford1982a, Reference Woodford1982b). In Ethiopian pastoral systems, and likely in other pastoral systems too, the patterns of bTB are closely linked to human social networks (Sintayehu et al., Reference Sintayehu, Prins, Heitkönig and de Boer2017a). There are also efforts in Ethiopia to establish risk factors for cattle TB associated with wildlife (Sintayehu et al., Reference Swanepoel, Crafford and Quan2017b). These are multi-species and slowly developing diseases with a long time course before clinical expression, and they are both zoonoses, so need to be monitored.

Anthrax is another multi-species disease that occurs on most continents, and in Africa epidemics occur in wildlife and these are sometimes associated with epidemics in livestock (Mukarati et al., Reference Mukarati, Matope and de Garine-Wichatitsky2020). Epidemiologically, this spillover may well be mostly driven by insect mechanical vectors such as blowflies and biting flies and through contamination of browse, pasture or water (Bengis, Reference Bengis, Miller and Fowler2012; De Vos and Turnbull, Reference De Vos, Turnbull, Coetzer and Tustin2004; Ebedes, Reference Ebedes1976; Hugh-Jones and De Vos, Reference Hugh-Jones and De Vos2002; Prins and Weyerhauser, Reference Prins and Weyerhaeuser1987). Evidence from Ethiopia suggests some significant recorded events in wildlife occurred after a series of livestock outbreaks (Shiferaw et al., Reference Scoones, Bishi and Mapitse2002), and in the Serengeti, wildlife anthrax epidemics tend to occur during droughts, clustered around contaminated sites such as water holes/salt licks and similar locations where aggregation and mixing of species can occur (Hampson et al., Reference Hampson, Lembo and Bessell2011).

It certainly can be argued that introduced cattle diseases on the African continent have had impacts on both the cattle industry and wildlife. Indirectly, the impact on wildlife has been seen through the implementation of control measures. Fortunately, in open mixed rangelands systems, these introduced diseases and the buffalo-endemic diseases are of little consequence to wildlife populations, other than rinderpest. There is a need to re-evaluate historic and modern disease dialogues, rather than perpetuating the old narratives and the prejudice against wildlife as disease reservoirs.

Buffalo have attracted significant research, often becoming the centre of investigations, with this focus perhaps reinforcing preconceptions of their relative disease role and significance. Of 79 publications recorded in a scoping review on viruses in ungulates (Swanepoel et al., Reference Shury, Nishi, Elkin and Wobeser2021), 41 were on FMD in buffalo. This high number is most probably due to the funding available and international interest among researchers for this disease. Buffalo are overstudied without considering other species found in the same environment or the role of cattle themselves in the persistence and spread of infection. As a consequence, the roles of these other wild ungulates (e.g. greater kudu, Thompson’s gazelle [Gazella thomsonii], impala [Aepyceros melampus] or blue wildebeest [Connochaetes taurinus]) are relatively unknown despite evidence of their role in a few specific breakdowns and cattle epidemics historically (Weaver et al., Reference Weaver, Domenech, Thiermann and Karesh2013). In KNP, which has an endemically infected buffalo population, clinical spillover FMD has also been confirmed in impala, greater kudu, bushbuck (Tragelaphus scriptus), common warthog (Phacochoerus africanus) and giraffe (Giraffa cameleopardalis), but with relatively mild symptoms observed rarely (e.g. impala; Vosloo et al., Reference Vosloo, Thompson and Botha2009).

Towards a New Vision

In the context of disease transfer between buffalo and cattle, who is to blame? The most significant problems for wildlife and cattle stem from the introduction of exotic breeds and their diseases into Africa and the production and trade model that came with them. Models of coexistence between buffalo and African cattle breeds existed during the pre-colonial era only to be disturbed by the introduction of susceptible European breeds and their northern hemisphere pathogens during the colonial period. In addition, the endemic disease risks to introduced cattle, especially for the so-called improved breeds, are very high. The full benefits of buffalo (as a comparable bovid) in the context of the animal and land use are clear but are not being realized, except in some specific land use examples. Most buffalo populations have been reduced to small relict herds, especially in West and Central Africa. If buffalo go extinct, cattle will provide poor compensation for this highly adapted species.

On the other hand, from a disease risk perspective, the management of wildlife species such as buffalo along similar lines to livestock production makes little sense. Extensive free-ranging unfenced systems are already of proven value for harvesting, sport hunting and tourism, and as bulk grazers in maintaining ecosystem’s integrity and function. The history of the development of both cattle and wildlife managed systems in Africa and trends in associated disease problems provide abundant evidence for this conclusion. The resilience and health of species are highly connected to the ecological resilience of the systems in which they have coevolved. Therefore, it is likely that community-based (pastoral) systems that can be mixed, rather than agribusiness-driven fenced monoculture (wild or domestic), are a route to sustainable tourism and animal-based food production systems and economies in Africa. Much of African livestock and wildlife will remain more or less in open conditions for the foreseeable future, and thus these systems should be reinforced by appropriate policy and investment rather than discriminated against. This approach has the added value of ecological recovery of highly degraded landscapes from over-intensive livestock agriculture, reinforcing biodiversity conservation, supporting the delivery of natural ecosystem goods and services as well as related income streams. It could innovatively contribute to the target of 30 per cent of land protected by 2030 recently set by the High Ambition Coalition adopted by 69 countries (HAC, 2021). The ambition is high indeed, but realities on the ground suggest it will simply be too late in many countries to progress before major land-use changes and settlements have degraded habitats, especially in Central African savannas (Scholte et al., Reference Scholte, Pays and Adam2021). Climate change mitigation benefits will also accrue, while still contributing high-quality protein and food security, in a continent that has the lowest meat consumption per capita on Earth. However, to achieve this means shedding colonial legacies around land use and tenure, livestock development and animal health production systems imposed on Africa. It may require reversion to more traditional views on extensive animal use, harvesting and integrated low-cost–low-risk systems of management that are not new and were widely discussed in the twentieth century (Asibey,Reference Asibey1974; Dasmann, Reference De Jong, van Hooft and Megens1964; Ledger, Reference Ledger1964).

As a new community of scientists and veterinarians emerges across Africa with a novel vision, knowledge from the past and ideas about future agro-ecologies and mixed land use will likely come into play. What may help is that there is now a major shift in perception of animal-based food systems in the very countries that promoted intensive livestock production in Africa. The question that really matters is what do Africans think of alternate futures? Will they remain embedded in old development dogma, or will they surf on the economic and cultural opportunities offered by wildlife? Could mixed land use become a dominant policy? Already, new land management, with mixed livestock and conservation initiatives, has shown considerable success in Namibia and Kenya, building on earlier innovation in Zimbabwe under the CAMPFIRE project, which under difficult political and economic circumstances remains nascent.

New ideas and opportunities beyond conventional systems of agriculture and wildlife protected areas will undoubtedly emerge, becoming hopefully more conducive to both local economic growth, ecosystem stability, resilience and biodiversity conservation. Planetary health demands it. In this chapter, we have shown the need for a reappraisal of history and the risk in the context of buffalo and diseases of concern to the livestock industry. Much of the narrative for ‘land clearing’ of wildlife is historic and ‘blames’ buffalo for diseases such as FMD, tick-borne infections, brucellosis and bTB, which early on justified fencing and the compartmentalization of land. In many cases their roles in the epidemiology of these pathogens are tangential and no longer highly significant in evolving contexts in which cattle populations are exploding and buffalo populations are maintained or decreasing in many ecosystems.

The genetic modification of farmed animals for objectives of higher production creates breeds more susceptible to pathogens when buffalo are disease-resistant in African contexts. Putting them together does not make the buffalo the culprit. The current industrial approach may work in highly transformed temperate systems but not in Africa’s landscapes, where there is high microbial and vector diversity and a multitude of hosts. This failure of livestock intensification to develop in Africa without subsidization has led to the narrative that it is only through promoting greater separation and higher biosecurity that animal-based food economies can develop and reduce the risk of catastrophic disease epidemics in heavily invested livestock industries. This is basically underpinning policy on disease control and has been supported by unrealistic standards in disease management generated by the World Animal Health Organization (WOAH). These industries are effectively highly subsidized through international funding, and agencies are dedicated to forcing through these agricultural development agendas. Ironically, most of the benefits from these policies accrue to high-income countries that are not African, for instance through blocking access to their lucrative markets. Globally, these policies have come at a major cost to biodiversity and ecosystems in return for little more than cheap protein and high profits for agribusinesses. Africa now stands to gain substantially from shifts in diet, with reduced meat consumption and an increasing acknowledgement of the value of natural land and biodiversity. None of the externalities from, for example, climate change impacts to the loss of habitats or biodiversity are currently born by the livestock industry, although the internalization of these externalities has been called for by the Ecosystem Approach (Principle 4) under the Convention of Biological Diversity (CBD Decision COP V/6) and again under the Principles for Sustainable Use by the same (binding) Convention (CBD Decision COP VII/12). This situation will change as there is increasing pressure for accountability and determination of externalities of various industries for future sustainable development. Buffalo may well still have a bright future.