The development of genetic studies on the African buffalo helped: to delineate subspecies number based on restricted gene flow criteria to either two or maximally three; to define three Conservation Units requiring separate management efforts, namely: (1) Eastern–Southern Africa, (2) the West–Central African forests and (3) the West–Central African savannas; to uncover major evolutionary demographic events, with the earliest identified expansion occurring 500–1000 kya; to evidence a strong population decline in Eastern–Southern Africa starting around 5 kya, and proposed to result from both climatic factors and explosive growth of human populations and their cattle. However, buffalo populations still display high genetic diversity and low genetic differentiation, and show primary sex-ratio distortion and high-frequency deleterious alleles in the buffalo genome and their potential effect on population demography and viability. Future management efforts are necessary to maintain gene flow, with the challenge that populations become more fragmented, distributed into a mosaic of conserved areas.

Much of the narrative for land clearing of wildlife is historic and frequently blames buffalo for livestock diseases, a dogma perpetrated throughout colonial history and inherited by emerging African states after decolonization. A review of this dogma indicates that the many significant problems for wildlife and cattle are related to introduced exotic livestock breeds that brought their diseases into Africa and the production and trade models that came with them. Reproducing European economic agricultural systems in Africa has failed in most African countries so far, challenging us to reconsider current agricultural economic development models in the context of human-induced global ecological changes, human relations to nature and our planetary limits. The next generation of African farmers, wildlife managers and policymakers have the opportunity to frame new coexistence and productive models between wildlife, including African buffalo, and livestock-based agriculture in the ecosystems in which they have coevolved.

Buffalo are ranched primarily to produce trophy animals, but inferior animals represent a good potential for meat production. Estimating and applying the well-reasoned stocking rates and feed supplementation during periods of nutrient shortages are essential to ensure the maintenance of good body condition. Protein supply, and the intake thereof, is the main constraining factor with regards to production performance in buffalo during the dry season on rangeland. Through optimized nutrition, year-round breeding may be supported, earlier attainment of sexual maturity can be realized, and more cows can be served by one bull (1 bull:30 cows sex ratio), thereby improving the productivity of buffalo herds in captivity. Buffalo have carcass yields similar to those of domestic beef cattle, and by applying the suggested grading system presented in this chapter, meat processing of buffalo carcasses may be optimized regarding allocation of carcasses and cuts according to their expected quality. Primal cuts should be vacuum-packed and matured for a minimum of 25 days under refrigerated conditions before being sold, with the sirloin and topside muscles being valuable cuts.

There are only a few infections that cause disease and mortality in African buffalo. These include rinderpest (historically – now eradicated), anthrax, bovine tuberculosis, bovine brucellosis, bovine papillomatosis and possibly Rift Valley fever. Historically, buffalo were probably the original source and maintenance hosts of several important cattle diseases, namely the SAT strains of FMD and theileriosis caused by Theileria parva. More recently, cattle-adapted strains of these two diseases have evolved, and they now cycle in several areas of sub-Saharan Africa in the total absence of buffalo. Buffalo may also be important reservoirs of trypanosomes in tsetse fly-infested areas, and may be a source of heartwater rickettsias in open systems within the distribution range of the Amblyomma vector ticks. Buffalo are susceptible to a whole range of important livestock diseases, including bluetongue, epizootic haemorrhagic disease, peste des petits ruminants, bovine ephemeral fever, bovine viral diarrhoea and anaplasmosis, but large knowledge gaps remain. Although these pathogens can infect and replicate in buffalo, buffalo appear to play little or no role in the epidemiology and maintenance of these livestock diseases. Buffalo appear to be totally refractory to several important infections of livestock like lumpy skin disease, contagious bovine pleuropneumonia and dermatophilosis. Buffalo, like most wildlife species, naturally carry a complement of ecto- and endo- macroparasites. With few exceptions, most of these parasites do not appear to deleteriously affect the health of free-ranging buffalo in any way. They may, however, become a problem in intensive ranching systems.

The current tribe Bovini may very well be polyphyletic. African buffalo might be descended from African Boselaphini, but the African fossil record before 8 Myr is quite poor. Palaeontological data tally well with nuclear DNA data showing that African buffalo and Asian buffalo separated some 8 Myr ago and are very distantly related. Cross-fertilization experiments and (failed) implantation tests of embryos of Asian Bovini into African buffalo wombs underscore the fact that these species are evolutionarily very distantly related. Karyotypic evolution of African buffalo is also very different from these Asian Bovini. This may warrant the establishment of a separate tribe for the African buffalo and its ancestors, namely, the Syncerini. Until recently there were two species of buffalo in Africa, Syncerus caffer and S. antiquus, which in some parts of their range coexisted. The ecology of the single surviving species of African buffalo may thus have been co-shaped by that recently extinct sister taxon. Because the present species is so distantly related to wild cattle and Asian buffalo, little or nothing can be learned from studying these species for the ecology or management of the African buffalo, even if much were known about these species in the wild (which is not the case).

The African buffalo has interacted with human societies for millennia across its vast African range. It is part of the bestiary of the few African imaginaries and mythologies that have managed to reach us. These representations of the species in African cultures seem to have percolated more recently into the imaginaries of European cultures, especially from the angle of hunting and photographic safaris. The buffalo is also at the centre of services and disservices to different actors, providing uses but also generating conflicts in African landscapes, the species being central in so-called Human–Wildlife Conflicts. For animal health services, the buffalo represents in some instances a public enemy, influencing meat trade policies, land uses and boundaries in many parts of the continent. The African buffalo is therefore an emblem of the coexistence between humans and nature in Africa.

Conservation, management and research require buffalo to be handled and sometimes moved from one place to another. Techniques providing more efficiency and a safer environment for buffalo capture and handling, including mass physical and individual chemical capture techniques, have been developed over the past few decades. These techniques, which are based on the experience and skills of staff, retain some room for improvement, e.g. using new drugs especially non-opioids for chemical immobilization, adapting technological advances (e.g. drone, scent technology) or new concepts (e.g. virtual boundary) to physical capture. The cardinal rule of buffalo or any wildlife capture, translocation and release is to regard all human interventions as potentially stressful to the animals, and therefore to strive to conduct them as far as possible as ‘short-term and low-stress management exercises’.

Whether practiced legally or illegally, formally or informally, hunting buffalo for meat occurs broadly across African cultures. Nearly all buffalo parts are prized in addition to the meat. Buffalo are also hunted for traditional medicine, social positioning, mystical reasons and in retaliation for causing damage to people and crops. The buffalo is a major game for the hunting industry in every country, but the reasons vary from place to place. In South Africa, buffalo is the first income-generating game despite being the least hunted of all important game. In Tanzania, despite a trophy fee that is lower than that of other species, buffalo is the top tax-earning game because it is the most hunted among the important game. As duly gazetted protected areas, hunting areas are contributing internationally to the global network of conservation areas. They more than double the land area that is used for wildlife conservation in sub-Saharan Africa. Acting as buffer zones of national parks and as corridors between national parks, hunting areas are the last frontier of the African buffalo outside national parks. In South Africa, where all buffalo are fenced and buffalo hunting occurs behind fences, the buffalo is subject to genetic manipulation to enlarge trophy horns and produce disease-free herds. While ‘clean buffalo’ widely contributed to expanding the land dedicated to wildlife conservation in a beef-exporting country, ‘augmented buffalo’ remain a matter of concern for the long-term conservation of the taxon. Several non-African countries imposed bans on importing hunting trophies of CITES-listed species from Africa, leading to a drop in the hunting market. The bans are having two impacts on buffalo: (i) although not CITES-listed, the buffalo is a collateral victim of the bans because many abandoned hunting areas are exposed to poaching and habitat conversion; and (ii) unintentionally, the bans are lifting the value of buffalo as a leading flagship game in an attempt to compensate for the loss of CITES-listed game. Hence, once a commodity game, the buffalo is turning into a high-value game.

African buffalo and cattle interfaces are expanding on the African continent due to the encroachment of human activities into savanna and forest habitats. These interfaces are especially important for managing the risk of pathogen transmission that can threaten small-scale and commercial livestock production, public health and biodiversity conservation. Recent technological advances in the field of remote sensing and telemetry provided opportunities to characterize buffalo/cattle interfaces with an accuracy allowing the estimation of potential infectious contacts and spillover of pathogens. Integrating environmental drivers, animal movement and pathogen transmission models is now possible, but this has not yet been done for buffalo/cattle interfaces. A better characterization and modelling of these interfaces could provide knowledge to design new management options for disease mitigation and control. The management of the sanitary risk at these interfaces is key to promoting healthy African landscapes in which production and conservation objectives coexist.

The African buffalo is host to numerous parasites, including microbial pathogens such as viruses, and bacteria and eukaryotic organisms such as worms and protozoa. Because buffalo are an important source of livestock diseases, understanding the ecology of the parasite community within African buffalo populations has both theoretical and applied importance. Competitive and synergetic interactions between parasites occur at both within- and between-host scales, and the African buffalo has served as a wildlife model for understanding how such interactions impact individual host health and parasite population dynamics. We describe the current understanding of the community ecology of parasites in African buffalo, identify general patterns that have emerged across parasite taxa, and describe key future research directions. Throughout the chapter we highlight important tools and techniques for studying parasite communities in wildlife populations.

Predation, poaching, disease and drought all impact African buffalo population numbers. Droughts in particular have important implications for the trajectories of animal populations, especially in tropical savannas. This is due to the pulse-like occurrence of droughts at intervals within the average lifespan of a buffalo. Consequently, populations are always in a state of transition and the proportions of the population in each age group are continually changing. We show that in these circumstances attempts to determine maximum stocking rates are prone to error. Furthermore, applying aggregated age groups to models may result in misleading forecasts of population trends. We believe this also holds for the populations of other mammalian species that live under so-called non-equilibrium conditions because their dynamics are then event-driven and not governed by factors such as density dependency.

Prior to colonization, humans always used African buffalo for meat and other products. The arrival of imperial powers marked the beginning of more extensive hunting, reducing buffalo populations in large areas of Africa. Buffalo production systems exist today along a gradient ranging from extensive (natural habitat) to semi-extensive (game ranches) to intensive (game farms) systems. These production systems rely on four sustainable uses: breeding, non-consumptive tourism, consumptive tourism and production of meat and other products. Private ownership and agro-sustainable biodiversity game ranching with buffalo has recently expanded in southern Africa, especially in South Africa, where it represents an extensive and productive land use.

This chapter presents the distribution, abundance patterns and trends of African buffalo in the 38 countries of its distribution area based on recent aerial and ground census data and feedback from field experts. For the period 2001–2021, we collected abundance data from 163 protected areas or complexes of protected areas and presence data from 711 localities. The savanna buffalo population is estimated in 2022 at over 564,000 individuals, after deduction of the 75,000 buffalo under intensive private management in South Africa. Its abundance is roughly equivalent to that estimated 25 years ago (625,000). The subspecies conservation status is highly unbalanced. The Cape buffalo is by far the most abundant, representing 90 per cent of the total estimated population (510,000 individuals). The West and Central subspecies respectively represent 4 and 6 per cent (>20,000 individuals and >34,000 individuals). The conservation status of the Central African savanna buffalo, whose abundance has been nearly halved over the last 25 years, is worrisome, with exception of the steadily increasing populations of Zakouma NP (Chad) and Garamba NP (DRC). Estimating the abundance of forest buffalo is challenging, as is establishing a trend. Our investigations showed that the forest buffalo is still well represented in Central Africa in areas with low human density. The forest buffalo’s most important stronghold in Central Africa is probably the Greater TRIDOM/TNS (Tri-National Dja-Odzala-Minkébé / Trinational Sangha), a vast contiguous block of mainly pristine moist forest covering 250,000 km2 and straddling Cameroon, Congo, Gabon and Central African Republic (11 per cent of the Central African forest block). In West Africa, we obtained very little information on the presence of the forest buffalo in the residual forest block, suggesting that the conservation status of the forest buffalo in this region is very worrisome.

The African buffalo is one of the best-researched of all ungulate species even though it must give way to some North American deer species, an elephant-seal species and the red deer. The African buffalo had some monographs dedicated to it, but much new research has been carried out on the species since that time, which is brought up to date in the present volume. This allowed us to make an inventory of what we do not know yet about this important species. For that purpose, we made an inventory of research topics, or questions tabulated under three different knowledge domains, (i) ‘known unknowns’, (ii) ‘unknown unknowns’ and (iii) ‘unknown knowns’. The ‘known unknowns’ we categorized as those research questions sitting as it were in the backs of the minds of the current suite of African buffalo specialist; our inventory yielded 37 research issues. The ‘unknown knowns’, we portrayed as evidence-based scientific knowledge on buffalo that a current generation of scientists appear to have forgotten. This proved difficult, but three topics were identified. Here we also draw attention to the fact that modern scientists appear to ignore francophone literature, which is rather unfortunate as West and Central Africa are to a large extent francophone. Not using this repository of information may lead to knowledge decay. Finally, we share thoughts on the ‘unknown unknowns’, which we described as ‘knowledge once we have it will upset our present thinking, perhaps about African buffalo, perhaps on ecology evolution, or on aspects of the veterinary sciences’. Under this category, we touched on 15 issues, but perhaps our imagination was too limited. So, we share in total some 60-odd questions and ideas, and we hope that at least some of these questions or ideas will kindle someone’s imagination and drive to bring knowledge on this great species further.

In this chapter we envision the possible futures of the African buffalo populations in Africa by reflecting on the regional and international factors and their relationships that could positively or negatively impact the healthiness of the buffalo species in the next 30 years. Using the expertise of the authors of this book, we drafted and validated a list of factors of change that could impact the futures of African buffalo populations on the continent and use a set of prospective methods, i.e. structural analysis, critical uncertainty matrix and morphological analysis to develop seven synopses which provided caricatural African contexts within which the consequences for African buffalo populations could be imagined. In 2050, the futures of the African buffalo will vary according to each country specific social, technical, economic, environmental, political and value contexts. In a context of climate change that will impact increasingly the environmental contexts in Africa, good futures for buffalo were often associated with political stability and good governance. The proportion of African living in cities will also be important. The ratio of urban versus rural African will not only determine the intensity of the agricultural pressure on land but also the African worldviews towards nature and its conservation. The influence of non-African states will also be determinant, especially in extractive industries and their request for land. A pivotal factor is the conservation models that will prevail in 2050: to what extent they are still influenced and constrained by part of the Western opinion; to what extent they are funded by them; and to what extent African worldviews push for the design of new conservation models based on different relationship between people and nature. Probably, landscapes associating land-sparing (e.g. national parks) and land-sharing management options, based on the sustainable use of natural resources will provide the best futures for buffalo to thrive on the continent.

African buffalo herd size varies across their distribution range from as few as 5–10 in the rainforests of West and Central Africa to as many as 2000 individuals in the floodplains of eastern and southern Africa. The home range size of African buffalo also varies greatly, with those of savanna buffalo herds generally ranging between 50 and 350 km2. The larger home ranges are generally observed in areas where resources are spatially segregated, and where herds are forced to undertake seasonal movements. In contrast, forest buffalo exhibit smaller home ranges (<10 km2) due to a less pronounced seasonality of the environment, and a more homogeneous spatial arrangement of resources. African buffalo are ruminants, essentially feeding on grass and roughage. This species is capable of subsisting on pastures too coarse and too tall for most other herbivores. The African buffalo occupies an important niche, opening up habitats that are preferred by short-grass grazers. Although the African buffalo primarily is a grazer, savanna buffalo can partially switch their diet to browse when grasses become tall and lignified. The ability of the African buffalo to cope with contrasting environmental conditions throughout most sub-Saharan ecosystems, by modulating a large array of biological traits, highlights a high degree of behavioural plasticity.

Based on genetics and ecology, it is best to discern three subspecies of African buffalo, namely the northern savanna buffalo, the Cape buffalo and the forest buffalo. In honour of the oldest written reference to the buffalo by the Syrian geographer Ibn Fadl Allah al-Umari in 1347 CE, we propose the name Syncerus caffer umarii for the northern savanna buffalo, and maintain S. c. caffer for the Cape buffalo and S. c. nanus for the forest buffalo. We think it likely that the forest buffalo is a recent form of buffalo (about 150 kyr), derived from the northern savanna buffalo in the eastern part of its range, which underwent dwarfing (i.e. miniaturization) in the rainforest. We propose that the northern savanna buffalo, because of the high amount of genetic exchange with the forest buffalo, has many hallmarks of a hybrid subspecies that expanded its range due to the creation of the Guinea savanna and Sudan savanna by Iron Age agriculturalists. The Cape buffalo shows the highest number of food web interactions with other large mammals, while the dwarfed forest buffalo is very lightly embedded in its trophic web.

Some aspects of the well-described planet- and satellite-like framework of, respectively, mixed herds and bachelor groups of buffalo recently were challenged. Associations of female buffalo are now considered more fluid than the initial idea of a stable mixed herd defined by a home range. Within mixed herds, adult female associations are unstable and transient despite using largely overlapping home ranges. Mixed herds in most instances do not seem to overlap much in space. Between mixed herds exchanges exist, apparently dominated by juvenile females (but almost no information exists on juvenile males). The dynamics of fusion–fission events within mixed herds are largely driven by habitat heterogeneity, the quality and quantity of grazing and surface water, and the influence of predation, parasitism and fires. The influence of the human/buffalo interface on these dynamics is not yet well understood. Future studies will benefit from more advances in telemetry and new technologies, new information sources (e.g. sound recorders) and non-invasive genetic studies to enhance our knowledge of buffalo social dynamics. Knowledge of buffalo social dynamics would also benefit from studies more representative of the African distribution of the species and across its subspecies.