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Part IV - Management

Published online by Cambridge University Press:  09 November 2023

Alexandre Caron
Affiliation:
Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD), France
Daniel Cornélis
Affiliation:
Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD) and Foundation François Sommer, France
Philippe Chardonnet
Affiliation:
International Union for Conservation of Nature (IUCN) SSC Antelope Specialist Group
Herbert H. T. Prins
Affiliation:
Wageningen Universiteit, The Netherlands

Summary

Type
Chapter
Information
Publisher: Cambridge University Press
Print publication year: 2023
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From a Commodity to a High-Value Species

The African buffalo has always been taken for granted. Due to its massive body size and vast herds, the species has constantly been perceived as an infinite source of wealth. Local hunters were fairly certain to bring home lots of valuable meat for their neighbours, friends and family. Scientists paid little attention to this abundant and rather unattractive cattle-looking beast compared to charismatic creatures threatened with extinction. Foreign hunters knew that a hunt for buffalo would not be in vain, that they would most certainly be challenged by a formidable adversary, with at least a thrilling fair chase as a result. Even conservationists were disinterested after the spectacular recovery of the species from the rinderpest onslaught. It is no surprise that such a commodity animal remained largely unnoticed and overlooked for so long.

Until things changed. With the human demographic upsurge, the escalating demand for game meat overtook the ability of wildlife – including buffalo – to match the needs. Buffalo habitats shrunk under unrestrained assaults of agro-pastoral encroachment. With the spread of modern weapons all over the continent, the fear of stalking buffalo faded. Buffalo started to struggle to cope with death tolls that were exceeding birth rates. They disappeared from large parts of their range, and were cornered in a few strongholds, mainly National Parks. Hunting Areas, another category of Protected Areas, became their last frontier outside National Parks, acting as critical buffer zones of National Parks and corridors in between.

Then the time came when visionary scientists such as Anthony Sinclair and Herbert Prins focused their minds on the species. Increasing numbers of tourists were excited to tick buffalo off their wish list as one of the Big Five. A new generation of veterinarians became conscious of the unique capacity of the indigenous African buffalo to resist or tolerate diseases that devastated exotic livestock. They also gained expertise in capturing and moving buffalo individuals and herds. Innovative cattle-ranchers initiated buffalo ranching as a new animal production system for multiple uses, both consumptive and non-consumptive, with a wider scope than dairy and beef, and as a means to rewild former cattle ranches.

Nearly restricted to very few countries in southern Africa, the private ownership of wildlife has changed the picture quite dramatically. By adapting livestock farming science and technology, the productivity of buffalo herds in captivity has improved. With optimized nutrition, especially during the dry season, buffalo ranchers get rid of the seasonality for breeding year-round and obtain earlier attainment of sexual maturity. However, the quest to produce ever larger trophy horns, driving prices up to at times unsustainable levels, is resulting in contentious breeding practices that include genetic manipulations such as outbreeding with East African buffalo and extreme inbreeding. Therefore, the large stock of captive buffalo is not considered part of the wild free-ranging populations for the long-term conservation of the taxon.

More recently, several non-African countries started to peremptorily impose bans on the importation from Africa of hunting trophies from charismatic wildlife species such as elephant and lion. This effectively posed an embargo on hunters from their jurisdictions to hunt for most trophies in range states of the African buffalo. Although buffalo (a non-controversial species) is not targeted by the bans, it is impacted by them. By downsizing the hunting market, the bans leave many Hunting Areas vacant and exposed to poaching and habitat conversion, rendering the buffalo a collateral victim of bans directed at other species. In the meantime, the bans are elevating the value of the buffalo, making it a foremost game in an attempt to compensate for the loss of other huntable game. Although less profitable than more prominent game, the buffalo finds itself in a position to financially contribute more to sustain hunting concessions so that they can continue to function as Protected Areas for preserving vast tracts of wilderness and their biodiversity. Hunting the African buffalo, if done judiciously and with restraint, could showcase the concept of sustainable use, which is one of the two pillars on which the Convention on Biological Diversity rests, the other being conservation.

When well-managed, the buffalo is a typical example of a high-value species producing high income from a low percentage of the population harvested. After being a commodity species throughout history, the African buffalo is now appearing as a promising prospect for Africa. It is time for buffalo to be considered a prominent asset for people rather than a banal species.

However, there are limitations that constrain the necessary change in paradigm. One is related to southern Africa, where the modified (“augmented”) buffalo stock in captivity makes an ex-situ population incompatible with conservation. Another is external to Africa, because foreign standards dictated by non-African countries are working hard to prevent Africa from using its own renewable natural resources such as wildlife, including buffalo (meat and trophies). This is leading the intrinsic values (existence values) advocated by many in the North to prevail over the use-values (utilitarian values) needed by the South. It is as if African wildlife only exist to make nice movies and serve as tourist attractions for temporary visitors. One more constraint is the need to make more progress in some fields, notably in veterinary science, for example how to manage diseases where cattle and buffalo cohabitate, and how to develop physical restraint technologies that rely less on chemical immobilization, especially opioids.

The information provided in this section is not only based on different forms of academic research, but also on extensive field experience, gained by hard work, successes and also failures. As such, the expertise acquired from field experience forms part of what is known as ‘experiential knowledge’, important in fields such as conservation science, biomedical research, farming and veterinary science. While often difficult to collate, this information is commonly useful in practice. Therefore, in this section, we formalize some of the experiential knowledge that we have acquired over the years.

13 African Buffalo Production Systems

D. Furstenburg , P.T. Gandiwa , Pa. Oberem and Pe. Oberem
Historical Perspective and Current Situation

Before the eighteenth century, the African buffalo Syncerus caffer was widespread and abundant in Africa (Furstenburg, Reference Furstenburg2015). Across the African continent, humans had used buffalo for millennia, well before domestic cattle were introduced, as a source of meat and co-products such as hides. The meat from buffalo and other game was the product of hunting, including trapping and even kleptoparasitism. Unfortunately, the use of buffalo has not always been sustainable, in particular since European explorers and settlers arrived with their guns (Chapter 12). In more modern times, human population growth, associated agricultural encroachment and modern weaponry has greatly impacted the conservation status of the African buffalo across its continental range, reducing its natural habitat and population size. In southern Africa especially, culling by white settlers from the 1650s to 1800s had a major impact. The great rinderpest epidemic of the 1890s spread south across the continent, further reducing the remaining buffalo population while also eradicating large numbers of other wildlife. This compounded the earlier impacts on the geographic distribution, population size, structure of herds, migration patterns, and hence production of buffalo.

Buffalo are asymptomatic carriers of SAT serotypes of foot-and-mouth disease (FMD), various species of Theileria causing East Coast fever, corridor disease and January disease, as well as tsetse-transmitted nagana (Chapter 9). To control and prevent the spread of these diseases to domestic stock, veterinary fences to control the movement of buffalo, other disease carriers and susceptible animals consequently have been used in southern Africa. This has had a further dramatic impact on the buffalo’s range and numbers (Oberem and Oberem, Reference Oberem and Oberem2016). It is only recently, with the introduction of community-based natural resource management, private ownership and game ranching, that the concept of sustainable utilization has again, this time consciously, become widely practiced in southern Africa. Regional wildlife populations have grown in southern Africa with the increase of private ownership.

Globally, from the year 1900 to 2000, domestic animal numbers increased by a multiple of 4.5 while wildlife numbers were halved (Smil, Reference Smil2011). Across African savanna areas, after evolving at varying times and speeds in different regions, the conservation status of habitat and species is today similar, with up to 80 per cent of wild animals lost and replaced in large areas by domestic stock, especially cattle. These developments across the continent have reached a point today where domestic livestock, although an exotic taxon, has virtually replaced buffalo, an indigenous taxon, and restricted the remaining buffalo populations to residual scattered wilderness.

Today, buffalo populations across Africa are broadly conserved in three major land-use systems, that is public protected areas owned by the State, communal land and private properties, the latter in only about five countries, all in southern Africa, out of the 37 African buffalo range countries. Variations in management objectives across these land-use regimes strongly influence the resultant production systems and the extent of the species’ utilization by land managers. Consequently, buffalo production systems have evolved and diversified between extensive models with free-ranging buffalo at low densities on large land areas and, at the other extreme, intensive models with enclosed buffalo at high stocking rates on small, fenced properties (Figure 13.1). The various categories of buffalo production systems are not compartmented; there is a continuum between categories.

Figure 13.1 Various categories of African buffalo production systems.

Adapted from Chardonnet, Reference Chardonnet2011; background picture: © Christophe Morio.

Buffalo farms are always fenced, most buffalo ranches are fenced, while most hunting areas with buffalo as a game animal are unfenced. In South Africa, however, all reserves, parks, ranches and farms where buffalo production occurs are enclosed by fences that restrict animal movement.

Buffalo Production Systems

Wildlife production systems can be classified on a scale of intensity of management. Here they are structured into three categories of property size and management intensity: (1) extensive production systems, (2) semi-extensive systems (game ranches) and (3) intensive systems (game farms).

In Zambia, the 200 game ranches existing there in 2012 (with a growth rate of six (6 per cent) per year over the past 32 years) were classified in three similar categories: (i) large-size game ranches of over 500 ha (75 ranches, that is 38 per cent of the national total), (ii) intermediate-size game farms of between 50 and 499 ha (27 game farms, 13.5 per cent of the national total) and (iii) small-size ornamental properties of less than 50 ha (98 ornamental properties, 49 per cent of the national total) (Chomba et al., Reference Chomba, Obias and Nyirenda2014).

This structure and these definitions are made in a quest to clarify and better understand the concepts. However, there are no strict limits between the three categories, it is rather a gradient of intensity.

Extensive Production Systems

In extensive production systems, buffalo are free-ranging and occur at natural densities, with or without the ability to migrate between natural resources and without managerial or veterinary intervention, as seen through most of the range states in Africa. Wildlife in extensive production systems is managed to be utilized for ecotourism and/or regulated hunting (Bothma and Du Toit, Reference Bothma, du and Du Toit2016).

Multispecies Bushmeat Hunting in Natural Ecosystems

Africa’s diverse ecosystems are endowed with wild large carnivores and herbivores that hold both ecological and socio-economic importance, and bushmeat hunting is probably as old as humans and still occurs today throughout Africa, both legally (hunting) and illegally (poaching).

In large areas, managers generally employ a more hands-off (extensive) management style utilizing multi-species in natural ecosystems. The smaller the area, the higher the likelihood that fewer species are more intensively managed.

The Cape buffalo Syncerus caffer caffer (hereafter, buffalo), given its large size and gregarious gathering in herds, was once one of the southern and eastern African mega-herbivores with the largest distribution (Hildebrandt, Reference Hildebrandt2014). In Africa, humans have, with some exceptions, mostly been transformed from traditional hunter-gatherers into sedentary village hunters and farmers (Wilkie et al., Reference Wilkie, Wieland and Boulet2016). Historically, subsistence hunting for consumption (bushmeat) in traditional systems was not considered to have a detrimental effect on wildlife populations, because hunting was regulated (Fa and Brown, Reference Fa and Brown2009). Traditional hunting or human predation in multi-species natural wildlife production systems for animal protein (bushmeat) and other wild animal products characterize many tropical indigenous communities (Marks, Reference Marks1977a; Manyanga and Pangeti, Reference Manyanga, Pangeti, Manyanga and Chirikure2017). For example, the African buffalo is among the important target species for the Valley Bisa community in the Luangwa Valley, Zambia. Their hunting techniques and selection of prey is related to the ecology and behaviour of the prey and influence the hunting patterns and timing of hunts (Marks, Reference Marks1977a, Reference Marks1977b). This traditionally organized form of wild animal hunting has facilitated the persistence of wild animals due to its selectivity and associated cultural conservation practices (Marks, Reference Marks1973). However, with the general decline in large wildlife populations (Craigie et al., Reference Craigie, Baillie and Balmford2010; Mabeta et al., Reference Mabeta, Mweemba and Mwitwa2018), species such as buffalo tend to be progressively substituted by medium to small-sized wild herbivores in response to the increasing demand of bushmeat consumption and trade in urban markets (Davies and Brown, Reference Davies and Brown2007).

Around the start of the twentieth century, the declines in wildlife populations prompted many African countries, most then under colonial rule, to criminalize the traditional livelihood strategy of bushmeat hunting (Child et al., Reference Child, Musengezi, Parent and Child2012). This led to negative relationships and conflict among local people, wildlife and the state as most local communities’ access to bushmeat was controlled. Any local hunting of wildlife now is labelled as poaching, and wildlife are mostly confined to protected game areas and national parks (Child et al., Reference Child, Musengezi, Parent and Child2012; Hildebrandt, Reference Hildebrandt2014; Mutanga et al., Reference Mutanga, Vengesayi, Muboko and Gandiwa2015).

Today, bushmeat hunting is generally non-selective and indiscriminate with regard to the animal’s sex and age and, when it is commercial, to the number of individuals taken. As rural populations grew, hunting methodologies became more modern, effective and less selective (firearms as opposed to the more traditional methods). As the land available for wild animal populations became limited by the expansion of farming and agriculture, bushmeat hunting concentrated in the remaining natural habitats was reported to threaten wildlife populations (Child et al., Reference Child, Musengezi, Parent and Child2012; Wilkie et al., Reference Wilkie, Wieland and Boulet2016). Literature points to hunting by humans, since the advent of modern firearms, having led to the extinction of wildlife species inclusive of large carnivores and herbivores (Martin, Reference Martin1966; Ripple et al., Reference Ripple, Wolf and Newsome2019). Today, wildlife provides many ecosystem services in the form of ecotourism, trophy hunting, meat, medicinal products, aesthetic enjoyment and inspiration (Tchakatumba et al., Reference Tchakatumba, Gandiwa and Mwakiwa2019).

Community-Based Natural Resources Management and Multi-Species Hunting

The introduction of community-based natural resources management (CBNRM) in the early 1980s was perceived as a necessary intervention to benefit wildlife and communities (Child et al., Reference Child, Musengezi, Parent and Child2012). Where CBNRM is implemented properly, wildlife can be used sustainably as an economic engine in communal lands while simultaneously encouraging conservation (Child et al., Reference Child, Musengezi, Parent and Child2012). For example, the Communal Areas Management Programme for Indigenous Resources (CAMPFIRE) is an example of a CBNRM programme that was designed and implemented by the Government of Zimbabwe in 1989 to stimulate the long-term development, management and sustainable use of natural resources in the country’s communal farming areas adjacent to state-protected areas. Thus, under CAMPFIRE, extensive natural wildlife areas were actively managed by local communities in order to reduce unsustainable exploitation of wildlife and human–wildlife conflicts, while also providing local communities with conservation benefits and incentives (Muboko and Murindagomo, Reference Muboko and Murindagomo2014). A major shift in the business model was the sharing of benefits inclusive of bushmeat derived from organized trophy hunting of multiple species of wild animals based on a participatory and sustainable quota setting system.

In 2022, a total of 58 of 60 districts in Zimbabwe were under CAMPFIRE programmes with a total area of 50,000 km2 (12 per cent of Zimbabwe’s surface area), which supports approximately 200,000 households (Machena et al., Reference Machena, Mwakiwa and Gandiwa2017; Campfire Association Zimbabwe, 2022). On average, CAMPFIRE generates about €1.85 million per year with trophy hunting, constituting the major source of revenue while other sources of revenue include ecotourism and lease fees (Machena et al., Reference Machena, Mwakiwa and Gandiwa2017). Thus, under CAMPFIRE, local communities realize both direct and indirect benefits from the sustainable management of local natural resources (Figure 13.2). The buffalo is identified as one of the ‘Big Five’ species, is valuable for both meat and trophy hunting and is a high-value species for photographic tourism. Local communities are tasked with conducting anti-poaching patrols and general resource monitoring in CAMPFIRE areas. Studies on CAMPFIRE show that wildlife habitats are being maintained well and have created conditions for increased wildlife populations outside protected areas (Gandiwa et al., Reference Gandiwa, Heitkönig and Lokhorst2013; Musiwa and Mhlanga, Reference Musiwa and Mhlanga2020). Nonetheless, there has been some criticism of the CAMPFIRE experience (e.g. Dzingirai, Reference Dzingirai2003). Elsewhere, similar CBNRM programmes (e.g. Botswana and Namibia) have led to enhanced conservation, benefits for local communities and recovery of wildlife populations (Mogomotsi et al., Reference Mogomotsi, Stone, Mogomotsi and Dube2020; Stoldt et al., Reference Stoldt, Göttert, Mann and Zeller2020).

Figure 13.2 Flow of direct and indirect benefits from CAMPFIRE programmes (Tchakatumba et al., Reference Tchakatumba, Gandiwa and Mwakiwa2019). *RDC refers to Rural District Council.

Source: with permission of Taylor & Francis.
Semi-Extensive Production Systems: Game Ranches

A semi-extensive production system is a natural area that is large enough for self-sustaining wildlife populations to be managed, that is a game ranch or a national, provincial or private park or reserve (Cloete et al., Reference Cloete, van der Merwe and Saayman2015). It can be fenced or unfenced, but humans need to intervene to provide either water, supplementary and/or complementary feeding, control of parasites, control of predation or the provision of health care (Cloete et al., Reference Cloete, van der Merwe and Saayman2015). Camp sizes (subportion of a game ranch/reserve) vary from several hundred to several thousand hectares depending on the habitat, climate, environment, other herbivore species, topography of the land and the nature and scope of the business. Every production system is unique, with specific ecological and animal management parameters addressed scientifically and professionally by experts. Game ranches may be considered as an innovative, sustainable form of agriculture or animal husbandry where an important outcome is the rewilding of an area.

Buffalo ranching often occurs in semi-extensive multi-species production systems as one element of the herbivory with or without natural predation. Stocking rates may exceed the natural carrying capacity of the rangeland; hence, in such cases, the need to supply supplemental feed during the dry season. Without careful rangeland management, there is subsequently a risk of ecological deterioration of natural habitat conditions. Buffalo ranching is often practiced on marginal agricultural land that was formerly severely degraded due to monocropping or domestic stock farming, and there is a need over time for sophisticated habitat rehabilitation programmes to be implemented (Chapter 14).

Such systems also require the management of sex and age structure by (i) limiting the number of mature breeding bulls (selective per individual animal profiling), normally 1 bull per 20–40 mature cows; (ii) removing surplus young bulls, mostly allowing only one bachelor group of <10, or complete removal of all young bulls, to limit social confrontation and fighting with the usually very valuable breeding bull; and (iii) removing and/or replacing post-age and non-productive females from the population.

Its reputation has given the buffalo the status of being recognized worldwide as one of the ‘Big Five’. The buffalo is the most dangerous of all African game species, especially if wounded or solitary. Its economic value has been further enhanced (Figure 13.3) by veterinary restrictions that prevent its translocation due to the danger of spreading disease. Consequently, the captive breeding of disease-free buffalo in semi-extensive confinement has become a lucrative business, but one which must be approached properly to ensure success. There was a boom in prices after the worldwide economic crises of 2008, reaching a record high in 2017 (Figure 13.3), followed by a fall to more normal pricing trends during 2018–2019.

Figure 13.3 Auction prices of live breeding buffalo bulls over time and illustrating the value initially placed by purchasers on buffalo of East African origin for reasons discussed in the section dealing with production of buffalo with large horn size, below. East African buffalo, formerly recognized as a subspecies, is phenotypically 12 per cent larger in body size, 10–20 cm higher shoulder height, with greater horn spread, lesser curve-drop and smaller bosses, than the southern African buffalo. East African buffalo was introduced into the South African production systems adding specific value market traits.

© Deon Furstenburg.

Figure 13.4 Horizon, the most expensive African buffalo bull ever bred so far, was sold at an auction for €10.8 million. Horizon was bred by Jacques and Caroline Malan of Lumarie. According to the SCI method (following the external curve of the horns, in inches), he measures an impressive 55 6/8".

© Nyumbu Game.
Intensive Production Systems: Game Farms

Intensive wildlife production systems occur in small fenced areas where wild animals are intensively managed for the production of meat, hides and live animals. Buffalo farming is also sometimes practiced intensively in small camps on game farms, mainly to produce highly priced animals for live sales, that is specific disease-free and specifically selected for phenotypes (e.g. body size, horn size and shape) of trophy buffalo (Figures 13.5 and 13.6; Bothma and Du Toit, Reference Tambling, Venter, Toit, Child, Child, Roxburgh, Do Linh San, Raimondo and Davies-Mostert2016). A camp is fenced off to more closely manage rare and valuable animals that cannot move freely. These small camps vary in size from 5 ha pens to 80 ha camps. As a result of the small surface areas of the camps, daily supplement feeding, or even a complete feed, is provided all year round, the ratio depending on the camp size and quantity (biomass) and quality (nutritional contents) of the grass production. The animal load in the camps exceeds natural vegetative carrying capacity generally by two- to threefold or more. One mature bull (selected by its animal and genetic profile) and 10–40 mature cows depending on the specific situation are usually kept as a herd in a camp. As a result of socio-spatial restrictions, only one bull is kept and all male progeny are removed to a different camp before reaching sexual maturity and the risk of intersocial confrontation, that is fighting. However, because of the recent dramatic decline in prices attained for buffalo and due to the cost of management, feed, veterinary services and medication in these intensive systems, the truly intensive breeding facilities have begun to turn more to semi-intensive methods of ranching.

Figure 13.5 Aerial view of a 460-ha intensified multi-camp buffalo production system in savanna habitat with centred pens for handling, supplement feeding and rotation of stocking between camps. Optimal habitat management entitles (i) a 2-camp system per buffalo herd and rotated every 8 months, or (ii) a 3-camp system per herd and rotated every 4 months.

Figure 13.6 Example outlay of a semi-extensive buffalo production camp system (2-camps, on average 230 ha each, per breeding herd, including two free-roaming areas >4,000 ha each for surplus animals) constructed per vegetation survey map in arid Kalahari savanna habitat.

Buffalo Products

All of the above production systems rely on one or more of the four sustainable use pillars of ‘game ranching’, namely (i) breeding for sale to new properties being converted to game ranches and rewilding; (ii) non-consumptive tourism, that is ecotourism; (iii) consumptive tourism, that is hunting; and (iv) production of meat and other animal products such as skins/leather, curios from horns, skins, hooves, bones, etc.

Live Sales of Breeding Animals

Many game ranchers took the opportunity to breed game animals for live sales. Most of these specialized in specific so-called rare species, for example bontebok, black wildebeest, sable antelope, roan antelope. Others specialized in specific, sought-after characteristics such as buffalo and sable with trophy-quality horns and body conformation. Yet others focused on multiplying colour variants that occur naturally but rarely in nature, such as black impala, golden wildebeest, etc. The breeding, sale and translocation of these animals resulted in the rewilding of marginal conventional agricultural land that was converted into game ranches. Today, this market segment is less lucrative than at its summit in 2017 but is still thriving.

Non-Consumptive ‘Ecotourism’/Wildlife-Viewing Tourism

Non-consumptive tourism in the form of wildlife-viewing tourism can also be regarded as a production system where the product or service is a photographic, educational or recreational safari sold to clients who buy a period of time spent in nature to watch fauna and flora including buffalo. Buffalo, as one of the ‘Big Five’ and with a reputation of being dangerous, are highly prized by wildlife-viewing tourists. With appropriate management, both consumptive and non-consumptive tourism can be conducted in the same area to increase and diversify the value of the ecosystem service.

Consumptive Tourism/Hunting Tourism = Sustainable Utilization

By definition, hunting tourism harvests a very low percentage of individuals within populations, old males or excess animals only, with the ecological and economic objectives of (i) conserving a buffalo population and its habitats through sustainable hunting and (ii) sustaining the hunting enterprise as well as the ranch. The trophy-hunting model aims to produce large trophies, while the sport-hunting model aims to offer fair chase hunts to tourist hunters who are more interested in the quest than in the trophy.

These buffalo-hunting production systems operate over large to very large areas where the buffalo densities appear at their natural levels, which are low compared to intensive systems. In all of the countries where buffalo tourism hunting occurs, hunting areas are unfenced open range extending in size from between 50,000 and 300,000 ha. South Africa, where buffalo are hunted in fenced hunting areas of smaller but still physically substantial sizes such as a few thousand hectares, is an exception. Given the demand and value realized by these forms of buffalo hunting, hunting buffalo for meat in these semi-extensive systems is rare, in contrast with the hunting of more common, less expensive game species (Chapter 16).

Animal Products

Game meat is considered a delicacy in many parts of the world where it is in demand for its rarity and its health benefits, such as high protein and low fat content. There are specialist harvesters who harvest excess animals for the purpose of supplying specialist game meat processors. The jurisprudence with respect to the South African Meat Safety Act 40, 2000 still needs (after >12 years of negotiations) to be amended to ensure that game meat can reach its true potential as a source of good, healthy, natural protein.

Many different curios are manufactured, formally and informally, from many parts of carcasses used for trophies and meat, including from skins (leather goods such as skin floor mats, shoes, handbags and belts, even furniture coverings), horns (door handles, lamps, wall decorations), bones (carved salt and pepper cellars, knife handles, lamp stands), etc.

Case Study: The Wildlife Ranching Industry in South Africa
The Buffalo in South Africa

In South Africa, the game ranching industry was born with the promulgation of the Stock Theft Act in South Africa in 1991, which confers ownership of game to the owner of the land so long as the land is adequately fenced. It got a further growth boost in 1996 when the new South African Constitution was adopted; Section 24 of this constitution recognizes the principle of the sustainable use of natural resources in South Africa.

The 2008 economic crisis played a further role with investors seeking different ways to invest their money. At its summit in 2015/2017, 8,000–10,000 game ranches covered almost 20 million ha (i.e. 14 per cent of the national estate, an area 2.2 times larger than the formally protected areas of the country). Many game ranches were established on marginal land, that is farmland with low agricultural production potential. Others were established on degraded agricultural farmland that was previously occupied by monocultures of domestic stock and/or crops such as maize (Cloete et al., Reference Cloete, van der Merwe and Saayman2015), thus rewilding and converting former farms into wildlife-based enterprises.

Sustainable use as a form of conservation was at the beginnings of a massive private and privately funded ‘rewilding’ of the country. This brought about a major turnaround in the numbers of many endangered species, as well as in the ‘ownership profile’ of animals in the country. As can be seen from Table 13.1, the numbers of species, including endangered species, are much higher on privately owned game ranches compared to state land.

Table 13.1 Percentage of various species, some endangered, on private land owned by private game ranches versus those on state reserves in South Africa (Nel, Reference Nel2021; Furstenburg et al., Reference Furstenburg, Otto, Van Niekerk and Lewitton2022).

Species% on private land% on state land
Black wildebeest8713
Blesbok9010
Bontebok8812
Buffalo6337
Oribi973
Roan antelope955
Sable antelope973
White rhinoceros65Footnote *15*

* Of the world population.

The same successful contribution has been made by private owners on private land to the survival of buffalo in South Africa. Table 13.2 indicates the number of buffalo in national and provincial parks versus game ranches as well as their disease status. There are only 645 disease-free buffalo in state parks compared with 75,000 disease-free buffalo on private ranches.

Table 13.2 Numbers and disease status of buffalo in South Africa; bTB = tuberculosis; CA = brucellosis; FMD = foot and mouth disease (personal research of P.T. Oberem).

Facility:
State Protected Areas versus private ranches
Size (Ha)Buffalo numbersSanitary status
Kruger NP2,000,000>35,000bTB, CA, FMD positive, theileriosis positive
Addo Elephant NP170,000440bTB, CA, FMD, Theileria free
Mountain Zebra NP28,40080bTB, CA, FMD, Theileria free
Hluhluwe–Imfolozi96,000>7000bTB, theileriosis positive
CA free, FMD free?
Camdeboo19,40075bTB, CA, FMD, Theileria free
Marakele61,00020 plusbTB positive?, CA, FMD, Theileria free
Mokala26,48550bTB, CA, FMD, Theileria free
Madikwe72,000800CA, Theileria, FMD free, bTb positive
Total in State Protected Areas2,401,285>43,465Only 645 disease-free
Total on private ranches>7,000,000 (available)>75,000ALL DISEASE-FREE
bTB, CA, FMD, Theileria free
Legal Status of Buffalo in South Africa

The South African Government Gazette No. 42464 dated 17 May 2019 amended table 7 of the Animal Improvement Act (Act no. 62 of 1998) and now lists 32 new wild animal species, including 24 indigenous mammals (e.g. the African buffalo), to provide for the breeding, identification and utilization of genetically superior animals to improve the production and performance of animals in the interest of the Republic. By declaring these wild animals as landrace breeds (in table 7 of the regulations), the Act typically provides for landrace breeds to be bred and ‘genetically improved’ to obtain superior domesticated animals with enhanced production and performance. Similarly, provision is made for the Breeders Association to lay claim to the breed and to establish specific breed standards for animals to be included in stud books. Animals declared as landrace breeds can also be used for genetic manipulation, embryo harvesting, in-vitro fertilization and embryo transfers.

Numerous concerns about the new legislation have been raised, including from scientists, over negative genetic consequences, ecological and economic risks, as well as direct conflict with other biodiversity laws in South Africa (e.g. IUCN SSC Antelope Specialist Group, 2015; IUCN, WCC 2016; Somers et al., Reference Somers, Walters and Measey2020). However, many if not all of these concerns could be mitigated by the Code of Conduct of the game breeder association (Wildlife Ranching South Africa), which intends to become the administrative and implementing agent under this legislation.

Macroeconomics

On the 20 million hectares occupied by game ranches, an income stream of €1.2 billion (ZAR 20 billion; €1 = ZAR 16.31) is generated annually, resulting in numerous decent jobs and outperforming the national economy (Oberem and Oberem, Reference Oberem and Oberem2016).

Surveys of game ranch usage in South Africa (Taylor et al., Reference Taylor, Child and Lindsey2020) revealed important facts about the benefits of private game ranching. Eighty per cent of private ranches utilized some form of consumptive sustainable use, with 5 per cent of the total land area covered by these private properties utilized for intensive breeding of rare species or colour variants. While profitability varied greatly between the properties, they produced an average return on investment (ROI) of 0.068 and employed more people at higher wages than equivalent domestic livestock operations. From the survey, it was concluded that the South African model could be a suitable option for other African countries seeking sustainable land-use alternatives.

A further survey (Taylor et al., Reference Taylor, Lindsey and Nicholson2021) assessed how the wildlife ranching sector (including intensive and semi-extensive) contributes to the conservation of herbivores. It concluded that individual ranches had a mean of 15.0 (±4.8) species, 1.9 (±1.5) threatened species and 3.6 (±3.1) extralimital species per property. In comparison to 54 state Protected Areas, wildlife ranches had significantly higher species richness, more threatened species but also more extralimital species, with total herbivore numbers estimated to be as many as 7.5 million. The report concluded that private game ranching in South Africa represents one of the few examples on earth where indigenous mammal populations are thriving and demonstrating how sustainable use can lead to rewilding.

Nel (Reference Nel2021) reported that 50 per cent of game ranches obtain an income from hunting, with hunting being the main income stream for 30 per cent of these ranches. Of these game ranches, 5 per cent conduct photographic tourism and 52 per cent are engaged in all four of the economic activity pillars. Table 13.3 indicates the income obtained from the economic activity pillars on game ranches.

Table 13.3 Income from various economic activity pillars on game ranches in South Africa (Nel, Reference Nel2021) (€1 = ZAR16.31).

ActivityAnnual income (€1 = ZAR16.31)
Subsistence hunting (meat)€735.9 million (ZAR 12 billion)
International hunting (sport/trophy)€122.7 million (ZAR 2 billion)
Processed products (meat/leather/curios)€306.6 million (ZAR 5 billion)
Live animal sales @ formal auctions€61. million (ZAR 1 billion)
Total€1.2 billion (ZAR 20 billion)

In South Africa, buffalo was the number one income-generating species in 2016 (North-West University, 2017; Table 13.4), although it does not appear on the list of the top ten most hunted species. This is an outstanding demonstration of a high-value species that produces high income with a small number of harvested individuals.

Table 13.4 Top 10 income generators (€1 = ZAR16.31) (North-West University, 2017).

Species201420152016% CHANGE
Buffalo€7.8 million
(ZAR127 million)
€8.9 million
(ZAR145 million)
€13.5 million
(ZAR220 million)
+73
Sable€3.5 million
(ZAR57 million)
€4.5 million
(ZAR73 million)
€7.2 million
(ZAR117 million)
+106
Lion€12 million
(ZAR195 million
€11.1 million
(ZAR181 million)
€6.8 million
(ZAR111 million)
–43
Kudu€4.8 million
(ZAR78 million
€6.4 million
(ZAR104 million)
€6.7 million
(ZAR110 million)
+40
White rhino€4.4 million
(ZAR72 million)
€4.7 million
(ZAR76 million)
€5.1 million
(ZAR83 million)
+14
Nyala€2.8 million
(ZAR45 million)
€2.8 million
(ZAR46 million)
€4.7 million
(ZAR76 million)
+71
Waterbuck€2.2 million
(ZAR36 million)
€2.5 million
(ZAR40 million)
€3.1 million
(ZAR51 million)
+39
Blue wildebeest€2.2 million
(ZAR36 million)
€2.4 million
(ZAR39 million)
€3.1 million
(ZAR50 million)
+39
Burchell’s zebra€2.4 million
(ZAR39 million)
€2.8 million
(ZAR45 million)
€3.1 million
(ZAR51 million)
+29
Oryx/gemsbuck€2.4 million
(ZAR39 million)
€3.1 million
(ZAR51 million)
€3 million
(ZAR49 million)
+27
Basics of the Game Ranching Technology

In general, smaller properties require far more management inputs than larger ones where the size, diversity and lower density levels of animals allow for less close oversight and interventions.

Infrastructure

Fences around game farms in South Africa are regulated by law. To own wild animals the property is required to have a Certificate of Adequate Enclosure (CoAE), which is issued by the Department of Environmental Affairs. The specifications (height, number of stands, etc.) are dictated by the law. In order to introduce and release African buffalo onto the property, a permit (WR number for the property) is required from Veterinary Services. Properties with buffalo also have specific minimum fencing requirements. Fences are not generally electrified, they are so usually only when very valuable animals are kept in small camps (<80 ha), and this to keep aggressive bulls in adjacent camps from fighting and to prevent predation of the calves.

In order for any buffalo to be moved from one property to another, both properties need to be approved and registered (WR numbers) by Veterinary Services, the animals have to be tested for the four controlled diseases, namely FMD, corridor disease (i.e. theileriosis), bovine brucellosis/contagious abortion (CA) and bovine tuberculosis (bTB). Permits must then be issued by the Department of Environmental Affairs in the provinces involved (two if moving the animals from one province to the other).

Bomas, or small, sturdily built camps of 1 ha or less, are not often used. When they are, it is mainly only for temporary housing, for example when holding animals while waiting for disease test results and permits (no animals may be moved without permits, see above), while in quarantine and/or for adaptation purposes to new farms in new and different geographic areas (Figures 13.7 and 13.8).

Figure 13.7 Buffalo in boma.

© Q. Strauss – MLP Media.

Figure 13.8 Buffalo in boma.

© J. Malan.

The sectoral focus that is the main economic driver and the size of the game ranch determine the need and type of water provision and/or water facilities provided. The biggest ranches would most likely rely mainly on natural water resources and sources such as rivers, dams and wetlands with perhaps (as is seen even in the 2 million ha Kruger National Park) some additional artificial drinking reservoirs and troughs to supplement the resource and ensure better utilization of the available habitat (pasture). At the other extreme, smaller farms and camps may rely entirely on such artificial sources.

Habitat and Feeding Management

Habitat management includes restoring the natural habitat and vegetation that generally has/had been damaged to varying degrees by earlier agricultural practices, including ploughing, overgrazing with a monoculture of species (e.g. cattle) and internal fencing/camping. It also includes providing artificial water sources, boreholes, reservoirs and dams to improve the utilization of the natural habitat across the property. Many of these former cattle farms also may be damaged as a result of bush encroachment (e.g. Dicrostachys sp., Stoebe vulgaris, various thorn trees of Senegalia and Vachellia sp.), which requires expensive interventions to restore the vegetative value and carrying capacity. It may also entail the removal of toxic invasive plants (often aliens) such as Lantana camara and Asclepias spp.

Especially on smaller properties, supplementary feeding needs to be practiced, in particular during the dry season (in South Africa this is mainly during the austral winter months) to ensure optimum health and reproductive rates. This would include vitamin and mineral supplements, protein supplements during the winter, and compounds to counteract the plant’s own defences, for example inclusion of polyethylene glycol (PEG) and propylene glycol (PG) to bind terpines and tannins allowing better utilization of especially browse but also lignified sour grasses during the winter (van Hoven and Oberem, Reference van Hoven and Oberem2018).

Breeding and Health Management

The first most important breeding management interventions are reducing the number of male animals that are kept for breeding to allow a higher percentage of female animals, that is altering the sex ratio from 1:1 to 1 male for 25–40 females. Males that are not selected for breeding are used either for trophy/sport hunting or for harvesting game meat. The second most important breeding management intervention is the selection of specific males for breeding to (i) maintain the natural characteristics of the species, (ii) improve adaptedness to that specific environment and (iii) improve general health by selecting against characteristics predisposing to parasites and diseases. For example, it is not recommended to breed with animals, in particular bulls, that habitually carry large numbers of ticks (‘tick taxis’). In the case of buffalo, selection is often specifically to restore the ‘lost’ horn length and character, which was selected against by heavy trophy hunting over many decades. A further important reason for selecting specific males and keeping records thereof is to prevent inbreeding.

Reproductive performance can be greatly improved by supplementary feeding in particular, and by reducing the numbers of male animals – and hence competition. Production management reduces the average intercalving rate of cows from extensive areas (as seen in the larger protected areas) from 22 to 14.5 months. This increases the maximum number of progeny per lifespan (20 years, first mating at age 5) from a natural n = 8 calves to n = 12 per lifespan (i.e. a 50 per cent increase per breeding cow). The age at first calving can also be reduced through the provision of constant quality feeding.

Wild animals have various adaptations to reduce the impact of parasites such as ticks and helminths on them. In some cases, this consists of migrating away from heavily parasitized areas, which often is not possible on fenced properties. This requires management interventions to reduce parasite numbers. Various ‘self-medication’ forms of acaracide applications have been developed. However, they all have negative aspects (e.g. not being able to control which animals are treated, frequency of dose and/or rate/size dose are difficult to control). Recently, acaricidal balls have been developed, shot by paintball sporting guns, meaning that the correct pour-on acaricide dose can be applied to the correct animal at the required time. Helminth treatment is usually only necessary on small properties with a higher numbers of animals per hectare and is most commonly applied to the supplementary feed.

Genetic Perspective of Buffalo Ranching

In South Africa, all buffalo are in fenced areas, either on private game ranches or National or Provincial Protected Areas. Similarly, veterinary fences and national boundaries in many cases prevent the migration and free movement of buffalo. This has created separate genetic pockets in regions, countries, reserves and private ranches. Given the earlier genetic bottlenecks the African buffalo has suffered, namely the great rinderpest epidemic, and hunting and veterinary controls, this further genetic isolation is of great concern. Private game ranchers, however, have by the nature of their businesses traded and moved animals, in particular bulls, from farm to farm, a practice of metapopulation species management. The purpose was and is twofold, namely to (i) mitigate against inbreeding and loss of genetic diversity and (ii) enhance the quality of the animals on a property by being healthy specimens of the typical buffalo in line with the descriptions recorded by Skinner and Chimimba (Reference Skinner and Chimimba2005).

A study of 4,000 buffalo from 26 private ranches (Greyling et al., Reference Greyling, Furstenburg and van Hooft2013) revealed that 11 ranches had a genetic diversity 3 per cent lower, and nine ranches had a genetic diversity greater than that of Kruger National Park. The latter indicates the enhancement obtained from metapopulation outbreeding because of frequent trading between private populations. In comparison, relative heterozygosity of private production populations ranges from 1.05 to 0.7 (disease-free) compared to protected conservancy-based populations of (i) Kruger National Park = 1 (diseased, meaning with the four main diseases cited above, that is bTB, CA, FMD and theleiriosis), (ii) Hluhluwe–iMfolozi = 0.85 (diseased), (iii) Addo = 0.65 (disease-free) and St Lucia Estuary = 0.62 (diseased) (Greyling, Reference Greyling2017).

Metapopulation macro-genetic management by private production systems could not only enhance, but also restore historically depleted genetic diversity of game species, with a positive contribution to the survival of the species. The combination of climate change and human industrial development poses increased risk to species adaptation and survival (Furstenburg and Scholtz, Reference Furstenburg and Scholtz2009; Scholtz et al., Reference Scholtz, Furstenburg and Maiwashe2010). Consequently, increased species and population heterozygosity (genetic integrity) has become directly essential for species survival, and sustained species marketing traits as incentive for production breeding being indirectly essential (Chapter 3).

Production of Specific Disease-Free Buffalo

After detecting bTB in Kruger National Park in 1990, a project was developed to preserve the Kruger buffalo genotype. In 1998, 11 disease-free calves were successfully bred and moved to private land outside of Kruger National Park. As a result of the subsequent successful breeding of more than 27,000 privately owned disease-free buffalo in South Africa, the project was terminated in 2011 (Bengis et al., Reference Bengis, Govender and Lane2016). In contrast to state and provincial parks, all buffalo in the private buffalo sector are thus currently disease-free (Table 13.2; Chapter 12).

Production of Large Horn Size Buffalo

Early travellers’ journal entries and many scientific studies indicate that buffalo, like many other species (e.g. sable, Hippotragus niger; greater kudu, Tragelaphus strepsiceros; eland, Tragelaphus oryx; elephant, Loxodonta africanus; and more), were exploited during the eighteenth and nineteenth centuries by continuous selective hunting. Trophy hunters in particular often first shot the largest individual in a herd, consequently possibly gradually depleting the natural genetic integrity and quality of the species. Studies of kudu populations by Furstenburg (Reference Furstenburg, du, Bothma and van Rooyen2005) in both free-roaming conservancy production and semi-extensive production systems in the Eastern Cape and in Namibia revealed genetic quality depletion in under 20 years by continuous selective harvesting/hunting.

During the wildlife price boom of the 2010s, East African buffalo had greater trade value for having a 12 per cent larger body size and a greater horn spread than the Kruger and Addo phenotypes. Kruger buffalo are known for thick bosses and a deep drop at the side of the head before curving upwards, and Addo buffalo have smaller body sizes and smaller bosses. East African buffalo were introduced and bred with the Southern African private populations during the late 1990s. Gradually, trophy quality increased, and the first 50-inch trophy bull was auctioned in September 2013 for €1.6 million (ZAR 26 million), and re-auctioned in February 2016 at an all-time record for buffalo of €10.8 million (ZAR 176 million; the animal shown in Figure 13.4). Indications from auctions are that today there are more than 50 bulls with greater than 50-inch trophies among the breeding stock in private production systems in South Africa.

The extent to which this is manipulated genetic engineering versus the restoration of historic natural genetic integrity continues to be debated at the national and international levels. At the national level, disagreements between various organizations are flaring, including between hunting organizations (Selier et al., Reference Selier, Nel and Rushworth2018). Somers et al. (Reference Somers, Walters and Measey2020) point out numerous concerns in the new legislation, including the process of consultation, and argue that the law will not improve the genetics of the species mentioned but will have considerable negative genetic consequences and pose ecological and economic risks. At the international level, there is much concern about intentional genetic manipulations of wildlife, for example (i) the World Conservation Congress at its session in Hawaii, United States of America, 1–10 September 2016, adopted the recommendation WCC-2016-Rec-100-EN on management and regulation of selective intensive breeding of large wild mammals for commercial purposes (IUCN WWC, 2016); and (ii) the Antelope Specialist Group of IUCN released in 2015 a position statement warning about intentional genetic manipulation of antelopes (IUCN SSC Antelope Specialist Group, 2015).

The twin impacts of indiscriminate hunting of the better trophy buffalo bulls in the rest of Africa and the managed breeding and sustainable use of these animals in South Africa are clearly visible in Figure 13.9 (Safari Club International, 2022). The growth of hunted buffalo’s average horn length in South Africa can be seen in the graph of records from the 1990s. In comparison, the horn lengths of hunted buffalo from the rest of eastern and southern Africa have shown a steady decline, probably as a result of indiscriminate hunting of the better horned bulls.

Figure 13.9 Average horn length in Cape buffalo (Syncerus caffer caffer) in South Africa (n = 777, 22 per cent) and in the ‘Rest of Africa’ (RoA), a variable composed of data from 11 countries from eastern and southern Africa: Angola, n = 19; Botswana, n = 35; Kenya, n = 89; Mozambique, n = 100; Namibia, n = 16; Rwanda, n = 3; Tanzania, n = 857; Uganda, n = 4; Zambia, n = 482; Zimbabwe, n = 811. All buffalo were hunted for trophy hunting in South Africa when buffalo in other countries may have been hunted for other reasons.

Graph drawn from data published by Safari Club International (2022).
Domestication

Domestication of a species is a process whereby, over time, and via genetic selection and modification of a species, it may be adapted for human association and use. Some species, through their genetic plasticity, are better suited for this process (e.g. the dog). It is important to distinguish domestication from ‘taming’ and ‘habituation’, both processes being short-term, individual- or small group-based and not involving genetic modification. Habituation can occur even in areas as large as Kruger National Park, where animals of all types become accustomed to and accepting of, for example, tourists in their vehicles on the roads and behave as if the latter were not there. Similarly, the concept of buffalo herding as practiced recently in Zimbabwe and historically in Mozambique is another example of habituation rather than domestication.

Domestication of the African/savanna buffalo, although unsuccessfully attempted on a few occasions, is not something to consider. First, the buffalo’s aggressive temperament, massive size and huge horns renders this a risky exercise. Second, its value as a tourism (both consumptive and non-consumptive) icon would be eroded. As domestication would require genetic selection for docility and other ‘agriculturally favourable’ traits it might, if not very carefully managed, lead to a weakening of the desirable survival traits/genes of the species.

Perspectives and Prospects

The hunger for land to feed the growing human population is rapidly driving the spread of agriculture into the remaining wilderness areas of Africa. With the disappearing wilderness and the loss of species, the need for formal conservation through the declaration of National Parks and the like increases. This in turn often leads to growing wildlife–human conflicts. Governments in many of the poorer developing countries just do not have the financial resources to fund, create and manage Protected Areas or compensate adequately those evicted from the declared areas, fuelling illegal bushmeat harvesting or, in other words, poaching.

Community-based resource management and private rewilding, funded through sustainable use, of the huge areas of marginal land already in use for other forms of agriculture should be considered to ensure the restoration and conservation of biodiversity, such as has been the case in South Africa. When the natural human inclination to want to determine one’s own destiny on one’s own piece of land is overcome through cooperation and the formation of a cooperative landscape on much larger areas, then it becomes so much easier to sustainably create wealth and create decent jobs for communities previously excluded from tourism (consumptive and non-consumptive), meat harvesting and processing, and the production of many other products. This is a system somewhere in-between the CAMPFIRE programme and the smaller private game ranching as currently found in some southern African states, particularly South Africa. The benefits are habitat conservation, improved biodiversity, improved production, sustainable job creation, integrated community economic development and improved food security and welfare through sustainable use. The iconic buffalo, as one of the Big Five, and valuable as a hunting trophy, for photographic tourism and for meat production, plays a pivotal role in such developments.

14 Management Aspects of the Captive-Bred African Buffalo (Syncerus caffer) in South Africa

L. C. Hoffman , C. A. Shepstone , K. Robertson and T. Needham
Introduction

The objective of most buffalo production systems is to produce offspring that meet specific requirements. In the case of captive-bred African buffalo in South Africa, the main aim is to produce trophy-quality animals for breeding and hunting. Managing nutrition and feeding is of the utmost importance when working with high-value species such as the African buffalo, as their nutritional status has a direct effect on their (re-)production and the profitability of the enterprise. In natural systems (game reserves, national parks), droughts cause buffalo numbers to decline due to animals not being able to source the necessary nutrients for reproduction in the available dry grazing (Chapter 7), thus reducing the animals’ reproductive performance and production. Within intensive systems focusing on individual and herd performance, particularly reproductive performance, there is a need to create management programmes and practices to assuage potential poor performance due to a lack of necessary nutrients at different times of the year. Although numerous factors, such as sexually transmitted diseases, libido, age of first mating, season and nutrition, influence reproduction rate, this section will discuss how nutrition may be used to support reproduction and production in intensively and semi-extensively housed/ranched African buffalo herds. In this chapter, we discuss the feeding preferences under semi-extensive systems together with the estimation of stocking rates for buffalo of differing physiological stages, as well as the nutritional requirements of buffalo, the effects of season on these, and how supplementary feeding may be used to ensure adequate nutrition – most of the knowledge/experience presented in this chapter comes from southern Africa, given their development of the private wildlife sector (see Chapter 13).

Furthermore, fundamental information regarding buffalo reproduction and their utilization for meat production is summarized using real-time ranch experience originating from the disease-free buffalo breeding ranches that flowed out into commercial buffalo ranching in South Africa. It is essential to give credit where it is due; the origin of buffalo ranching started with the cattle industry’s husbandry techniques, and as time progressed, more and more scientific data were collected. This scientific knowledge allowed the game industry to not only gain the knowledge of breeding disease-free buffalo, but also to introduce these animals back into the wild.

Nutrition

Buffalo are broadly classified as bulk grazers, spending 40–80 per cent of their time feeding and ruminating depending on the season. Rumination is the process through which selected forage, already in the rumen, is repeatedly regurgitated into the mouth and back to the rumen to decrease particle size and buffer rumen pH via saliva; these fine food particles are degraded in the rumen by microbial action and fermentation. These small particles pass through into the omasum, then the abomasum, and then the small and large intestine for further degradation, digestion and nutrient absorption. The African buffalo consumes a wide variety of grass species, with grass constituting a relatively high proportion of their diet (75–100 per cent), but utilize more browse during the dry season or in different vegetation zones (woodlands or forest) when they are forced to graze less selectively and browse on woody shrubs, increasing the browse proportion of their diet.

African buffalo are relatively unselective grazers, but prefer highly palatable nutrient-rich grass. In extensive systems of southern Africa within granite and basalt landscapes (Macandza et al., Reference Macandza, Owen-Smith and Cross2004), they depend primarily on Panicum spp. (mostly P. maximum) throughout the year, and as the dry season progresses, Digitaria eriantha and Urochloa mosambicensis (previously also known as U. usambarensis) and Cynodon dactylon are the predominant species consumed. Some Bothriochloa spp. become important contributors to the buffalo’s diet during the transition from wet to dry seasons, but mostly not B. insculpta, while Eragrostis spp. contribute towards the end of the dry season. On the other hand, important cattle forage species, like Themeda triandra, which generally hold more fibrous content during the dry season, are less favoured by African buffalo under low-input management conditions Furthermore, Cymbopogon plurinodis, Bothriochloa spp., Pogonarthria squarrosa, Aristida spp. and Setaria spp. tend to be rejected by buffalo, regardless of the season.

The quantity (amount) and quality (nutrients) of grazing is influenced by soil type, topography, rainfall, ambient temperature and animal stocking rate/density. In semi-extensive ranching systems, these factors need to be taken cognisance of, as the manager only has control over the number of animals placed in the ranch/camp/paddock (stocking rate/density), keeping in mind factors such as carrying capacity (see below).

Extensive Grazing and Stocking Rates

Safe stocking rate (referred to as ‘carrying capacity’ by many game and livestock ranchers) can be defined as the number of animals that a specific piece of land can accommodate annually without degrading the quality of the forage, and can be measured in different animal units, generally known as large stock or large animal units (LSU or LAU), grazing and browsing units (GU/BU) in southern Africa. With irregularities existing in stocking rate methodology and the interpretation of the animal units in these methods, researchers developed a model where the different methods could be interpreted on a metabolizable energy basis measured in megajoules (MJ ME), establishing a calculated large stock unit (LSUC), grazing and browsing unit (GUC/BUC) to be used in the model. One LSU is equivalent to a steer (cattle) with a body mass of 450 kg that is growing 500 g per day by feeding on grazing that has a mean digestible energy concentration of 55 per cent, thus supplying 75 MJ ME per day (Meissner, Reference Meissner1982). A grazing unit (GU) is a 180 kg blue wildebeest and a browsing unit (BU) is a 180 kg kudu (Van Rooyen and Bothma, Reference Van Rooyen, Bothma, du, Bothma and Du Toit2016) requiring 29.71 MJ ME per day (Shepstone et al., Reference Shepstone, Meissner and Van Zyl2022). These methods are conservative ways to calculate a piece of land’s safe stocking rate if its grazing and browsing capacity has been assessed. This prevents overutilization of the available forage and ensures that the quantity and quality of grazing do not deteriorate over time.

When estimating the carrying capacity and stocking rate on semi-extensive systems, the average LSUC or GUC/BUC value for the specific species should be used. Using averages for all production phases mentioned in Table 14.1 will undersupply energy to lactating females, as well as growing and adult males. Intensive systems should either use the fixed values in Table 14.1, where the number of animals is multiplied by the LSUC or GUC/BUC value for each respective physiological state in a spreadsheet, or use the lactating cow (cow with calf) LSU value of 1.32 as the baseline parameter. For example, using the LSUC value of 1.32, a herd of 20 breeding buffalo will need 264 ha if the carrying capacity of the property is 10 ha/LSU. On commercial ranches in southern Africa, stocking rates vary and range between 2.6 and 13.3 ha/LSU (Hildebrandt, Reference Hildebrandt2014). The methodology behind calculating the ME requirements, calculated large stock, grazing and browsing unit values and dry matter intake (DMI), is described by Shepstone et al. (Reference Shepstone, Meissner and Van Zyl2022).

Table 14.1 Calculated large stock unit equivalents and metabolizable energy values for different physiological stages of African buffalo.

Physiological stageMass (kg)ME (MJ/day)Footnote *LSUCFootnote #BUC/GUC#
Calf, 8 months14529.60.391.00
Heifer, dry, 4 years46078.11.042.63
Cow, dry, 10 years53072.60.972.44
Cow, with calf, 4 years46099.11.323.34
Cow, with calf, 10 years53093.21.243.14
Young bull, 4 years50080.21.072.70
Adult bull, 10 years64081.91.092.76

* Calculated metabolizable energy.

# Calculated large stock, grazing and browsing units.

Adapted from Shepstone et al. (Reference Shepstone, Meissner and Van Zyl2022).

When the stocking rate exceeds the assessed safe stocking rate, the property is overstocked, making it necessary to purchase or supply stored roughage of a suitable quality to reach the desired reproductive goals. In circumstances where the stocking rate equals, or is lower than, the assessed safe stocking rate, and the quality of the available grazing is not suitable for optimal reproductive performance, the specific nutrients that nature cannot supply must be provided so the ranch can reach the desired production goal. The production constraining nutrients normally found in dry grasses are digestible protein, minerals and vitamins. It is important to note that no specific nutrient guidelines currently exist for buffalo, so cattle data are used to extrapolate the nutrient requirements for buffalo and other similar species. When supplying animals with supplemental feed formulated to mitigate a deficiency, it is important to understand that content and quantity of a particular nutrient will differ from one production system to another, and from one ranch to another. The nutrient concentration and the amount to be fed are directly influenced by the nutrient requirements of the animals/herds at that specific time, the quantity and quality of the natural grazing (high correlation with rainfall) or supplied roughage, seasonal changes, availability of raw materials, manufacturing equipment and storage of the mixed feed and raw materials. In order to ensure optimal rangeland utilization on a piece of land, and to limit rangeland degradation by overutilization (i.e. too many animals), routine vegetation studies are necessary to calculate its respective annual safe stocking rate and to take measures to ensure a conservative stocking rate using either or both the calculated LSUC, GUC and BUC methods. Similar animal unit methods are used internationally, such as the animal unit (AU) used in North America and the tropical livestock unit (TLU) used in tropical countries. Be aware of the differences before translating values 1 to 1.

Nutrient Requirements

When considering nutrient requirements for wild animals, a similar well-studied species is used as a proxy when formulating feeds; when considering the bulk-grazing African buffalo, other bulk grazing species such as beef cattle and water buffalo (Bubalis bubalus) data can be used. In this document, we use nutrient requirements of beef cattle in the United States (National Academies of Sciences Engineering and Medicine, 2016) as the baseline comparative nutrition proxy because rangeland beef cattle in southern Africa select a similar diet, live in similar habitats and have a similar daily water requirement to the African buffalo. On the other hand, water buffalo are animals housed and raised similarly to how the dairy industry houses and raises their dairy cattle, making this species less comparable to the African buffalo from a comparative nutrition point of view. Using cattle nutrient requirement data to estimate the daily nutrient requirements of buffalo is of little value if the buffalo’s average weight, physiological state and daily DMI are unknown; thus, these are important factors to account for when formulating a supplement/feed for buffalo. Knowing the physiological state and average weight aids in calculating the animal’s nutrient requirements, and nutrient analysis of the grass or roughage supplied to the animals will aid in calculating what shortfalls exist.

In addition to providing sufficient quantities of feed, the quality (nutrients) of feed, which includes the energy, protein, fibre and trace elements (vitamins and minerals) content, is important for ensuring optimal production and reproduction based on the animals’ nutrient requirements. The calculated LSUC, GUC and BUC methods discussed above is currently the most accurate method to calculate the energy requirements for game animals (Shepstone et al., Reference Shepstone, Meissner and Van Zyl2022), where the energy used by the animal is expressed as ME, measured in megajoules (MJ). When conditions are favourable, ruminants eat to meet their energy requirements rather than to fill their rumens (intake capacity). However, during the dry season, when the available ME in the selected grazing is too low to meet maintenance/lactation requirements, the animals cannot consume more grass or browse to satisfy their requirements, the main reason being that the total amount of feed intake per unit of time is restricted by their thoracic cavity (restricted rumen capacity). Furthermore, a cow in her last trimester of pregnancy has even less space (as the calf is taking up a lot of the abdominal cavity) for food in her rumen. The average voluntary feed intake (VFI) for buffalo is calculated to be approximately 1.8–2.3 per cent of their live body mass (530 kg) for a dry cow and a lactating cow, respectively (Shepstone et al., Reference Shepstone, Meissner and Van Zyl2022), which compares well to the published value of 2.5 per cent of live body mass on a dry mass (DM) basis (Prins, Reference Prins1996).

The supply and intake of protein are the main factors controlling production performance in ruminants like cattle (Köster et al., Reference Köster, Cochran and Titgemeyer1996) and buffalo on dry rangeland. The minimum crude protein requirement of buffalo is 7–8 per cent (Prins, Reference Prins1996), which may be provided by browse when available. When considering dietary protein supply in ruminants, it can generally be broken down into rumen-degradable protein (RDP) and rumen-undegradable protein (UDP). Ruminants require protein (nitrogen and amino acids) for two important functions. First, specific amino acids are needed for their metabolic processes (UDP). Second, and more importantly, protein from grass and supplemental feed is needed to supply the necessary nitrogen (RDP) to the rumen microbes to multiply, playing a pivotal role in increasing the VFI of dry forage. A ruminant’s RDP requirement in general can be calculated using the following equation: RDP = live body weight0.75 × 4. The VFI of dry grass is directly related to the concentration of RDP in the forage and/or feed.

While buffalo can increase their dietary protein intake by increasing their VFI, they are constrained by the need to ruminate, which competes with grazing time. In addition to selecting more browse during periods when high quantities of mixed quality food are available or under food scarcity/poor quality conditions, buffalo apply bulk grazing, whereby they graze during periods of adequate or high grazing volume availability and spend equal amounts of time grazing and ruminating (Prins, Reference Prins1996). When the quantity and quality of grazing is poor, buffalo spend more time in search of food, therefore they have less time for rumination and digestion on a daily basis, and are limited by rumen fill (capacity) regardless of passage rate. Feed supplementation should be considered to meet the nutrient requirements of buffalo, especially with regards to RDP during the dry season or during periods of increased productive/reproductive performance.

The most important goal of supplementation, particularly RDP (nitrogen), is to maximize the VFI of dry roughage during the dry season to ensure optimal ruminal microbe proliferation and supply of microbial protein. With a limited supply of RDP (nitrogen), fewer microbes are available to ferment the finer particles of grass that have been masticated into small particles by rumination, thereby reducing the fermentation rate of the ingested food, causing food to stay in the rumen for longer periods. As it takes the animal a longer amount of time to degrade and digest the food, the animal starts losing condition, forcing it to mobilize stored energy (fat) and protein (muscle) to survive. When this problem can be diminished or even prevented by increasing the VFI of dry grass, the animal will have more nutrients to maintain its good body condition. This can be done by ensuring adequate amounts of RDP are available all year round. Green grazing normally has sufficient RDP to maintain the animal’s condition, but as the dry season progresses the plants dry out, causing the quality and supply of protein to become limited for the rumen bacteria first, thereby making it necessary to supply additional RDP. Supplements formulated to supply the appropriate amount and ratio of RDP and other nutrients will enable beneficial microbes to proliferate. With a larger microbe population, the animals can degrade and ferment more food, thereby increasing their VFI of dry grass, resulting in improved production, body condition, health, milk, and colostrum quality and strong calves. As buffalo are ruminants, they can utilise nitrogen from non-protein nitrogen (NPN) sources such as urea and convert this into microbial protein (McDonald et al., Reference McDonald, Edwards, Greenhalgh and Morgan2002), making it possible to design supplements that include NPN sources such as urea.

In addition to energy and protein, minerals (macro-minerals > 100 mg/kg feed and micro-/trace minerals < 100 mg/kg feed) and vitamins are nutrients important for herbivorous animals including buffalo. Macro-minerals, micro-minerals and vitamins are important components of supplementary feed if optimal reproductive performance is desired. As grass is the main component of a buffalo’s diet, the animal relies on the minerals and vitamins that grass supplies. The mineral content and the availability of grass to the animal are influenced by different factors, including grass species and stage of maturity of the grazing, the type of soil, climate and seasonal conditions, and the condition of the soil, such as pH and mineral content as well as fertilization and liming (McDonald et al., Reference McDonald, Edwards, Greenhalgh and Morgan2002). The (trace) elements that are typically deficient are phosphorous, sodium, chloride, sulphur, iron, iodine, copper, cobalt, manganese, and selenium (Schmidt and Snyman, Reference Schmidt, Snyman, Du, Bothma and Du Toit2016), although there may be specific minerals that are deficient in specific areas.

Vitamins are needed in small quantities by herbivorous animals, and ruminants require even less as they have microorganisms that synthesize some vitamins in the rumen that can be used in the buffalo’s body. Under natural extensive conditions, vitamin levels in green grazing are generally high enough to meet the animals’ requirements. The exception is vitamin A, which normally is found in low concentrations in dry mature grass and grains used in formulated feeds. As with most supplementation of shortages, the most effective starting point to control or manage a mineral or vitamin shortage is to accurately predict its extent. A possible way of predicting the mineral shortages of buffalo is to analyse the grass species selected by the animal on the ranch at different stages of growth, and then formulate a mineral supplement that makes up for the shortfalls of the grazing (Schmidt and Snyman, Reference Schmidt, Snyman, Du, Bothma and Du Toit2016). However, this method might prove impractical and/or costly. Alternatively, basic knowledge of the animal’s well-being and behaviour, regular observation and accurate record-keeping combined with basic knowledge of the environment (type of veld, general shortages in the area, weather patterns, parasites, etc.) should suffice to identify most shortages. The buffalo themselves are the best indicators of mineral shortages, and if the animals maintain good health with 13-month intercalving periods, and the calves display optimal growth and health, then any changes made are likely unwarranted. Any unnecessary changes to the ‘environment’ may have an adverse effect on production and cause monetary losses.

The planting of additional grazing (pastures) is an effective management method in areas that provide low-quality natural grazing (such as sourveld in the dry months; ‘sourveld’ is the term used in Southern Africa for nutrient-poor, dystrophic savannas and grassland types), especially for overwintering. By irrigating these planted pastures, the feed production of the pasture can be raised substantially, and the expense of feed costs can be lowered. Nonetheless, these planted pastures can be a potential reservoir for parasites, especially in cases where the grazing forms a thick matt at the base, such as kikuyu (Pennisetum clandestinum, also known as Cenchrus clandestinus) and should be managed accordingly. Additional advantages include the fact that many of these grasses are perennial grasses, and once established, all that is needed for growth is sufficient water and, depending on the soil type, at times fertilizer. The grass not used for grazing can be baled and either stored for drought years or sold as an additional source of income.

Influence of Season on Nutrition and Feeding

Three factors need to be taken into consideration to estimate what nutrient shortfalls exist, if any, namely: (i) a nutrient analysis of the grass or roughage supplied to the animals; (ii) an assessment of the physiological state of the particular class of animals, including body condition, to gauge their nutrient requirements; and (iii) for the same reason, an estimate of the typical weights of the particular class of animals for which one wants to estimate the possible nutrient shortfall. The available grass on reserves and game ranches is normally a combination of annual and perennial grass species. These grasses have a green growing phase and a dry phase (Figure 14.1). During the dry phase, perennial grasses go into dormancy and store most of their nutrients in their roots and seeds, while the annual grass species release their seeds and die. Under free-ranging conditions, many buffalo populations would migrate shorter or longer distances, grazing higher-quality food elsewhere, but within ranches they are stuck in an area (due to fencing) where the grass nearly always deteriorates to a point where the buffalo lose condition. Buffalo in a poor body condition have lower conception rates, fewer calves, less milk and fewer calves that reach adulthood (see Chapter 7); in addition, the general health of the adult and subadult deteriorates over time.

Figure 14.1 How the season affects the quality of the grazing and animals’ body condition in southern Africa. Considering the months of January through to December (a full year), each month has a shade of green (rainy season), or yellow to orange (dry season). The green blocks correspond with the rumen and plant (in green) and the green arrow below it, portraying the time of the year when the selected feed gets degraded and digested in less than 24 h. The yellow to orange blocks correspond with the rumen and plant (in yellow/orange) and the orange arrow below it, portraying the time of the year when the selected takes longer than 24 h to be degraded and digested.

Source: Craig Shepstone.

The changes in nutrient quality from a high-/higher-quality green plant to a dry, brownish, poor-quality plant have a direct effect on the buffalo’s ability to break down and digest the available grass. Protein, energy, macro-minerals, trace minerals and vitamin concentrations decrease as the plant dries out, while the fibre portion increases. The drastic decrease in nutrient concentration, particularly RDP, results in fewer microbes proliferating, slowing down the degradation and digestion of the ingested fibrous feed, and thus the production of volatile fatty acids (VFA). VFAs produced by these microbes are the main energy source for ruminants and thus buffalo. The small population of microbes present when buffalo feed on poor-quality feed takes longer to degrade the fibrous feed, resulting in the feed remaining in the rumen for a longer period, forcing the animals to mobilize stored nutrients to survive (Figure 14.1) and resulting in the animal losing body condition.

As shown in Figure 14.1, the nutrient-rich green grass available from January to April supplies enough nutrients for increased VFI. Bulk grazing ruminants like buffalo then only need between 8 and 24 h to digest the green grass, resulting in an improved body condition (poor to improved body condition). However, in the dry season (June to November), buffalo can take as long as 3.7 days to digest the nutrient-poor grazing available, resulting in poorer body condition. The lower the RDP concentration in the dry grazing, the longer it takes to be digested in the rumen. Supplying the correct amounts of the necessary nutrients in the dry season will aid in reaching the desired production/reproduction goals, reducing feed digestion time from 3.7 days (worst case scenario) to 24 h.

For optimal production, managers should pre-empt the negative effects of the upcoming dry season and supply supplemental feed in small amounts before the grass deteriorates and the animals start losing condition. As the dry season progresses, it may be necessary to supply more nutrients for optimal body condition; the reason for this is that grass nutritional quality deteriorates further as the dry season progresses, particularly the protein and energy contents. Supplementing ranched/managed animals with the feed nutrients they need during the times when nature cannot supply these nutrients will not only speed up the time taken to digest the ingested food, but will also improve body condition, conception, milk production and general health. Not unimportant, it will assist in raising healthy calves. Furthermore, when considering optimal production in times of drought where the quality and the quantity of the available feed gets poorer, it is imperative that animal numbers are reduced, or additional high-quality roughage be supplied. Droughts have deleterious effects on production, reproduction and growth, with young and weaned animals being the most vulnerable (Chapter 7).

Supplementary Feeding of Ranched Buffalo

High-quality feeds can be used to supplement buffalo during critical periods (without having to decrease the number of animals) and can be found in different forms. These include everything from pellets to home-mixed rations, which are normally supplied in an amount smaller or equal to one-third of total daily intake. Supplementary feed for buffalo on dry grazing focuses on supplying RDP, energy, minerals and vitamins. In situations where grazing is limited or not available (in a ‘boma’, also known as ‘kraal’ or ‘corral’), feed must be supplied daily with all of the above nutrients together with ample high-quality roughage. In the rainy season, when the quantity and quality of grazing are high, supplying adequate amounts of protein and energy, some minerals may nonetheless remain deficient. For example, phosphorus, copper and zinc are deficient in most parts of southern Africa, making it advisable to supply some minerals to the animals throughout the year. Mineral licks composed of salt, macro-, and trace minerals will supply the nutrients for the animals to reach their owners’ production goals.

For any rancher/manager interested in obtaining well-balanced feeds, licks in both meal and block form for supplying buffalo the nutrients nature cannot, feed companies throughout southern Africa (South Africa, Namibia, Zimbabwe, Zambia) can be contacted, who will formulate and supply custom diets for the game ranches’ specific need. Purchased or self-mixed feeds usually come in the form of a supplement (concentrated feed) providing ≤⅓ of total DMI a semi-ad-libitum feed supplying approximately ⅔ of total DMI and full feeds, otherwise known as total mixed rations (TMR). Semi-ad-libitum feeds are usually 50:50 concentrated nutrients: high-quality roughage. Some pellets on the market, known as high-fibre pellets, are designed to be fed as semi-ad-libitum feeds. A TMR for buffalo usually contains roughly one-third (33–40 per cent) of the total daily amount of feed as concentrated nutrients that supply all of the desired trace minerals and vitamins, and most of the protein and energy, with the rest of the roughage making up the difference. To ensure optimal rumination in the bulk-grazing African buffalo, fibre length should be at least 2.5–3.5 cm.

Intakes of concentrated supplemental feed, known in Africa as lick, in a meal (powder) or block form can be controlled by increasing or decreasing the concentration of salt, ammonium sulphate and monocalcium phosphate and by hardening the licks in block form by adding binders like calcium hydroxide, magnesium oxide and molasses syrup.

When pellets or fine meals are used as ⅔ of the total DMI, the concentrations of the nutrients, particularly protein, energy and copper, should be kept in mind. Do not supply a pellet or fine meal that is designed as a supplement as a semi-ad-libitum feed at approximately two-thirds of DMI or as a full feed at three-thirds. This will result in the overconsumption of some nutrients or minerals, which could cause deleterious effects. Additives like Poly Ethylene Glycol (PEG) can be added to a supplemental diet to aid the buffalo in degrading and digesting tannin-rich browse when they need to consume browse as a food source. Mycotoxin binders should be added to all self-mix recipes. If self-mix recipes are stored for periods longer than 2 or 3 days, the addition of a mould inhibitor is advised. The addition of an active yeast product will aid in the control of rumen pH (reducing the risk of acidosis) and enhance fibre digestion (Chaucheyras‐Durand et al., Reference Chaucheyras‐Durand, Ameilbonne and Bichat2016). All dry or semi-dry feed should be stored in a well-ventilated (dehumidified if necessary) storeroom or container (rodent-, insect- and bird-free), preferably on pallets, and never in direct sunlight.

Buffalo should receive good-quality grass hay as the largest portion of their diet. When hays (alfalfa, grass, oat or other cereal hays) are supplied to the animals in the camp (ad-libitum if limited or no grazing is available), place them in or near the feeding area, in a separate bowl or hay rack that keeps the hay off the ground (reducing losses and cost). All hay that falls on the ground should be removed, preventing young animals from eating wet/soiled/mouldy hay. Mouldy or dusty hay can cause pneumonia, colic and/or heaves. All roughage with any visible signs of mould should not be used, as it usually contains high levels of mycotoxins. If poor-quality grass is the only source of grass hay available to the animals, it would be advisable to mix 10–20 per cent legume hay (alfalfa) into the roughage diet to augment the amount of protein. Legume hay roughage, such as soya bean or peanut, can be used as a roughage source if the products are free of mycotoxins, as the pods often house the fungus. When in doubt, use a suitable mycotoxin binder when using any such roughage as feed. Macronutrient analysis of all hay should be done routinely. This will not only help illustrate what the buffalo are eating, but it will also assist in deciding how much legume hay is needed to be mixed into a ration if the grass hay is of poor nutrient quality. Exclusive use of small grain hay and alfalfa hay for the Cape buffalo is discouraged as it may lead to mineral imbalances, laminitis, colic and diarrhoea, and other dietary abnormalities.

When the season changes and the nutrient quality of the grass drops to levels where it is necessary to supplement the animals with a supplemental (concentrated) feed to keep them in good condition, it is critical to remember that the animals need to be slowly adapted to the feed (over 5 weeks) to avoid conditions like acidosis, rumenitis and Clostridium-related illnesses, which can be fatal. This is also important when changing concentrated feeds, where the new feed is mixed with the old in increasing increments of over the 5 weeks. Animals should be vaccinated against red gut (frequently caused by sudden feed changes to ruminant diets, thought to be caused by excess growth of Clostridium perfringens type A, which causes an enterotoxaemia or torsion of the gut). During the growing season, it is necessary to supply a well-balanced mineral lick, and a well-balanced supplement/semi-ad-libitum feed in the dry season.

When a particular spot in a camp is used continuously as a feeding site, problems could arise in the long term, namely parasite build up (wire and other roundworms and coccidia) and a breeding place for disease due to the high number of flies and mould growing in old food, leading to possible mycotoxicosis. Unfortunately, feeding sites that are used continuously are often associated with accumulated faeces and urine. To prevent the build up of the abovementioned problems, and to encourage the animals to eat grass in a different part of the reserve or game ranch, feeding sites should be moved regularly. Small enclosures are an exception, making dedicated feeding sites necessary. These feeding areas need to be designed so they can be cleaned and disinfected routinely. Feed should not be placed near water troughs, and placing feed bowls a short distance away will limit the amount of collected feed falling out of the animal’s mouth into the water. Water troughs and feed bowls must be cleaned routinely, and new feed should not be added on top of old remaining feed.

A rule of thumb suggested to ranchers who keep animals where both male and female animals have horns is to place a bowl per animal and an extra bowl for every four animals, but observation of the competition at feeding is necessary, and it may be necessary to increase the number of bowls. Place the bowls 2.5 animal lengths apart in a rectangular chessboard-like fashion and only feed the animals when they are near the feed bowls. Move one line of bowls every second or third day (Figure 14.2). This not only saves time but also gives the keeper the chance to clean all the bowls at least once a month.

Figure 14.2 Feed bowl placement in a camp and movement suggestions. The circles represent bowls, and the figures refer to a column of bowls that need to be moved together periodically, from left to right as shown in the diagram. Spaces between bowls should be 2.5 the animal’s length.

Source: Craig Shepstone.
Water Provision

The importance of water in ruminant nutrition is often overlooked; water being the basis of rumen fluid creates a suitable environment in which the beneficial anaerobic bacteria can degrade (ferment) and digest the selected food, making it imperative that animals like buffalo need a regular supply of clean water.

The water requirements of buffalo differ according to the different circumstances and environments. Factors affecting water requirements include the gestation, age and physiological condition of the buffalo, the composition of the vegetation fed on and environmental conditions. Watering points are needed at regular intervals to supply adequate water without the buffalo having to travel long distances. Buffalo can utilize most types of water sources but seem to prefer artificial water holes and dams over troughs. In the case of intensive ranching, it is advisable to use water sources that can be controlled. Controlled water points can be tested and analysed for quality and contaminants (which is advised when ranching intensively with high-value species) and filtered if harmful agents are found in the water, as opposed to open and stagnant ground-/rainwater which could contain toxin-producing bacteria or other harmful agents. Open groundwater also doubles as a potential mud hole for wallowing, which is a part of daily activity in warm months, especially by bulls. As buffalo tend to urinate or defecate in these mud holes, it is advised to limit these to only wallowing (and not drinking as well) to prevent any potential diseases emanating from these activities.

Reproduction

The reproduction efficiency of buffalo is influenced by body condition (like cattle), and thus nutrition plays a crucial role in a herd’s reproductive performance. It is influenced by the fact that buffalo show some degree of seasonality in their breeding activities under extensive conditions, with most births occurring during seasons when food quality and availability are at their highest (Bertschinger, Reference Bertschinger and Penzhorn1996). However, the degree of breeding seasonality may differ under semi-extensive/intensive conditions where nutrition is optimal and health challenges are minimized. The attainment of sexual maturity is primarily influenced by body condition and thus nutrition; sexual maturity in buffalo is estimated to occur when they attain two-thirds of their genetically determined adult body mass which, depending on their condition, is generally 4–5 years of age for wild free-ranging southern African buffalo heifers. Typically, mature males can weigh 650–850 kg while females can weigh 520–635 kg at maturity. In captive-bred/supplemented herds, buffalo heifers, due to better nutrition and consequently faster growth rates, regularly become sexually mature in the latter stages of their third year. The average lifespan of the buffalo is 11 years under wild conditions (disease and predation being mostly responsible for mortalities) and 16 years in captivity. Females become senescent at 15 years (Prins, Reference Prins1996). Maximum lifespans are 24 and 20 years on average, for males and females, respectively.

African buffalo are promiscuous, and under extensive, so-called natural conditions, adult buffalo bulls constitute 10–15 per cent of the herd and adult females 55 per cent (Sinclair, Reference Sinclair1977), and thus a 1:5 adult sex ratio typically exists. As reported in the survey of Hildebrandt (Reference Hildebrandt2014), southern African ranches tend to have a male:female ratio of 1:2 under intensive ranching conditions, with a ratio of active breeding bulls:cows being 1:27, within herds that ranged from 56 to 290 buffalo (mean = 156 animals/enterprise). Hildebrandt concluded that the optimal ratio of breeding bulls:cows would be 1:30, provided that their nutritional requirements are met and body condition is maintained.

The buffalo cow is polyoestrous, and her oestrous cycle is 23 days long, with oestrus lasting 24 h, and the ovulation of two ova is rare (the second is often resolved). The buffalo is a long-day breeder, being sensitive to photoperiod effects. The gestational period of buffalo is ~340 days, with an intercalving period varying from 13 to 29 months, depending on nutritional conditions (Prins, Reference Prins1996). Hildebrandt (Reference Hildebrandt2014) conducted surveys of a number of ranches in southern Africa and reported an average intercalving period of 443 days (14,6 months), with the optimum period being under 400 days. Typically, lactation lasts 10–15 months (Carmichael et al., Reference Carmichael, Paterson, Drager and Breto1977). Calves are on average 40 kg at birth. When the calf is removed shortly after birth (3 days), oestrus occurs within 5 weeks, but post-partum anoestrus depends on the conditions of the cow and population dynamics. Cows with good body condition at parturition, receiving a high-energy diet, resume oestrus within 90 days post-partum. On the contrary, poor nutrition of the cow may result in low fecundity for up to 2 years thereafter (Ryan et al., Reference Ryan, Knechtel and Getz2007). The body condition of the cow during gestation also affects the calf’s growth and survival.

Thus, prior to parturition and during lactation, the cow would benefit from improved nutrition and her body condition should be monitored. It is important to remember that as soon as implantation of the embryo takes place during gestation, the feed intake of the cow becomes the feed intake of the calf too. This continues after parturition, as the calf is now dependent on the milk from the cow, although less direct than when in the uterus, but the quality of the milk has a direct effect on the development of the calf and its performance as an adult (McDonald et al., Reference McDonald, Edwards, Greenhalgh and Morgan2002). The energy requirement of lactation is extremely high and can reach between 93 and 99 MJ/day at peak milk production (~5 weeks after parturition; Shepstone et al., Reference Shepstone, Meissner and Van Zyl2022). The weaning age of buffalo in southern African ranches ranges from 6, 12 to 18 months (Hildebrandt, Reference Hildebrandt2014). In the wild, it appears to tend more to 18 months than to shorter periods (Prins, Reference Prins1996).

The feeding of weaned male buffalo bulls is often neglected, as most of them are placed in a bull camp to grow out with as little expense as possible to the rancher. As some of the weaned bulls might prove to be the best sellers, especially when from a good genetic background, it may be worthwhile to also attend to their feed requirements to optimize growth and obtain the maximum expression of their genetic potential.

From a trophy-hunting perspective, the ‘solid boss’ of a bull is its most desirable characteristic. Wild, free-ranging southern African buffalo, which occur south of the Zambezi River in areas with 450–750 mm of annual rainfall, become sexually mature in their fifth year, but their bosses only become sufficiently hard enough to become a desirable ‘trophy’ by their ninth year (Pienaar, Reference Pienaar1969). This is the same in East Africa (Prins, Reference Prins1996). Better nutrition enables captive-bred bull buffalo to grow faster and attain their sexual maturity weight at a younger age, which makes their bosses develop sooner; a 6-year-old bull buffalo may have the appearance of a solid-bossed 9-year-old simply by providing better nutrition, which in turn enables them to be hunted as ‘trophies’ at a much younger age.

Meat Production

Presently, with the exception of Kruger National Park (KNP, South Africa), no other entity in southern Africa has sufficient numbers of buffalo to ensure a constant supply of buffalo carcasses. However, as commercial ranchers reach their safe stocking rates and the supply of trophy bulls surpasses the demand, more inferior animals will become available (and cheaper) for so-called ‘biltong’ (traditional dried lightly salted/spiced meat product) or meat hunters to hunt, or to start harvesting on a more commercial scale. Nonetheless, there are still some buffalo hunted for trophy purposes whose meat is available to enter the consumption market. In addition, the culling of animals for various management reasons is an essential component of wildlife management, and there is interest in economic opportunities for game ranchers and ecotourism. However, concerns have been raised over the practicality and financial benefits of using culled buffalo as a source of meat in South Africa, where buffalo are not normally utilized for their meat. Cape buffalo have carcass yields similar to those of domestic beef cattle and produce meat with favourable organoleptic properties (Van Zyl and Skead, Reference Van Zyl and Skead1964). Cape buffalo carcasses are typically from subadults (280–430 kg live weight), adult cows (450–680 kg) or adult bulls (600–850 kg), who yield carcass weights of 140–220 kg, 260–330 kg and 300–440 kg, respectively (Grobler, Reference Grobler and Penzhorn1996). Thus, buffalo typically have a dressing percentage of 48–58 per cent depending on their gut fill at point of slaughter – similar to that of Bos taurus cattle (Hoffman et al., Reference Hoffman, Van As, Gouws and Govender2020). Under extensive conditions, the mass gains (kg/year) from up until 42 months can be expected to be for different regions of Africa (Bothma and van Rooyen, Reference Bothma and van Rooyen2005):

  • Kruger National Park, South Africa: 113 kg/year for bulls and 108 kg/year for cows

  • Serengeti, Tanzania: 103 kg/year for bulls and 99 kg/year for cows

  • Ruwenzori, Uganda: 103 kg/year for bulls and 92 kg/year for cows

  • Northern Uganda: 105 kg/year for bulls and 100 kg/year for cows.

Currently, there is a lack of scientific information regarding expected growth rates for intensively farmed buffalo.

As outlined in Figure 14.3, the culling and processing of African savanna buffalo meat typically involve five stages, namely (1) field operation, (2) slaughter, (3) deboning, (4) production of products, and finally (5) marketing. Moreover, these logistical points (phases) will utilize a set of activities required to process a raw material into a value-added product.

Figure 14.3 Standard operating system for the major logistical operational points to be considered in a typical buffalo culling scheme, as demonstrated in Kruger National Park (KNP, South Africa). LTL = longissimus thoracis et lumborum; BF = biceps femoris; SM = semimembranosus muscles; WPS = Wildlife Processing Structure.

Source: Louwrens Hoffman.
Field Operation and Slaughter

Two of the challenges involved in the large-scale harvesting of buffalo in the field is to shoot a sufficient number of animals rapidly and efficiently ensuring that the procedures comply with animal welfare standards and human safety requirements, and to ensure that the carcasses all fall in an area as small as possible to facilitate the further slaughtering and loading of the carcasses for transport to the abattoir. Due to these challenges, a typical system that has been shown to work efficiently is to shoot a group of buffalo out of a helicopter with darts containing scoline (suxamethonium chloride, also known as suxamethonium or succinylcholine). This is a medication used to cause short-term paralysis; in most countries the use of scoline has to be managed by a veterinarian. The helicopter pilot can herd the darted group of buffalo (four to eight, depending on terrain, herd size and abattoir capacity) into a small compact group until all of the darted animals have dropped (±5 min from the first dart). Before the cull, careful selection of the ‘killing zone’ needs to be made to ensure that the animals are close to this area and that it is accessible to the ground team and all vehicles. After dropping, the animals are killed with a free bullet shot in the head with a heavy-calibre rifle.

Although cost-effective, safe for the operators and widely used since the 1980s, the use of scoline is prone to ethical issues. Herding, darting buffalo with scoline and resulting asphyxia was shown to generate high levels of cortisol concentrations compared to animals shot at a standstill (Hattingh et al., Reference Hattingh, Wright and De Vos1984). Such response could partly be ascribed to conscious perception of asphyxia in conscious animals with resulting fear (Button and Mülders, Reference Button and Mülders1983). Previous studies suggested that residues of scoline in meat and biltong are apparently considered acceptable by public health authorities (Button et al., Reference Button, Bertschinger and Molders1981). This statement should be re-examined in the light of recent technologies and standards, also taking into consideration that using scoline alters physical meat quality attributes (pale, soft and exudative meat; Hoffman, Reference Hoffman2001). In a context of evolving animal welfare standards, the cost–benefits associated with the use of scoline should also be reconsidered.

After shooting, the animals are bled/exsanguinated, preferably on a slope or suspended. Using a terrain vehicle that is equipped with a conveyer belt whereupon the buffalo carcasses are placed helps to speed up the time between exsanguination and the hanging process. Removal of the internal organs from the carcass takes place in the field to ensure that there is no bloating of the carcass (which increases the risk of contamination). Ideally, transportation to the abattoir should not take longer than 2 h. It is important to ensure that knives are sterilized throughout and that hand-cleaning facilities are available. The primary meat inspection is conducted by a state veterinarian in the field and includes inspecting the head, pluck (red offal), feet, abdominal and reproductive organs of a partially dressed game carcass with the pluck and carcass then being sent to a registered game abattoir. Most of the white offal is left in the field for predators or vultures. However, some of the white offal will be cleaned and taken for consumption by the staff, such as the plies (third stomach) and set of tripe (weasand, first, second and fourth stomach and rectum).

Following the field operation, buffalo arrive at the abattoir offloading section (Figure 14.3). The carcasses are hoisted up, suspended from both Achilles tendons, and weighed. The head is removed, and trophy-worthy heads are cleaned and dried in the sun to be sold as trophies to hotels and restaurants as decoration or to individuals, while the smaller heads and the condemned (i.e. rejected) carcasses go to the rendering plant to produce an end product known as bone meal that can be sold as fertilizer. The skin is then removed, and to prevent contamination, removal of the hides should be done carefully, preferably when the carcass is warm, and all of the cuts are made from the inside to the outside to prevent contamination of the meat, using the two knife principles. The hides are normally processed (salted and dried) to be sold at auctions.

Red offal is preferably removed at the abattoir and is hung on a separate line, in the same order as the carcasses, for inspection by the veterinarians so that they may be correlated with the carcass, which is also individually inspected. The carcasses should be split with a saw blade along the spinal column to promote chilling. Lastly, the carcasses are washed with potable water, to remove all blood and bone sawdust, quartered between the ninth and tenth ribs, and weighed before being placed in the cold room (<7°C).

If only the red offal is affected, and the rest of the carcass is normal, only the red offal is condemned (lungs, heart and liver). However, if the intestines are linked to general diseases such as enlargement of the lymph glandes, fever, or hepatitis, etc. the whole carcass is condemned. The condemned carcasses and condemned organs are sent to the rendering plant to create bone meal. Carcasses can be partially or totally condemned. Affected areas are condemned for various reasons, such as infections caused by systematic or generalized lesions.

Older buffalo, depending on where they have been reared and with which other species (generally wild dogs and carnivores) they have interacted, can contain intermediate stages of tapeworm parasites (cysticerci and hydatid cysts) and need to be frozen for a minimum of 72 h with an air temperature of at least –18°C before being deemed fit for human consumption. However, the freezing compromises the quality of the meat and it is no longer suitable for selling as tender prime meat and should thus be processed further.

Deboning and Products

While no official carcass grading system is currently used for buffalo, or any other game species really, Table 14.2 provides a suggested grading system that could be utilized to class buffalo carcasses according to age. The application of a grading system helps to categorize buffalo carcasses, guiding the deboning team to know which carcasses will go for prime steaks, ageing, value-added products and processed meat. Incorporating the grading of buffalo at the abattoir will help speed up the process and prevent adult/old buffalo meat from being sold as prime tender (and expensive) steaks.

Table 14.2 Suggested grading of African savanna buffalo (Syncerus caffer caffer) carcasses.

GradePermanent incisorsBuffalo ageCategory
A0<2 yearsJuvenile
AB1–22.5; 3–3.5 yearsSubadult
B3–64; 4.5; 5; 5.5 yearsAdult
C7–86; 6–10; >12 yearsOld

Grades A and AB (subadult) buffalo carcasses should be used for aged primal cuts as well as processing for value-added products. Grades B, C (adult) and detained (frozen) buffalo carcasses can be used for processed meats (mainly biltong, a traditional dried lightly salted/spiced meat product) and value-added products. However, the fillets of all the carcasses can be sold as primal steaks at the highest price.

Deboning is the process whereby the fore and hind quarters of the carcass are taken and processed into various primal cuts (similar to those from beef carcasses, and illustrated in Figure 14.4) at 24–36 h post-mortem. The primal cuts from the hind quarter are silverside (biceps femoris), fillet (psoas major and psoas minor), sirloin (longissimus thoracis et lumborum), prime rump (gluteus medius), T-bone, topside (semimembranosus), knuckle (vastus medialis and other related muscles) and hind shin (Peronaeus terius, extensor digitorum longus and extensor digiti terii proprius).

Figure 14.4 Primal cuts of African buffalo carcasses for further processing and marketing.

Source: Tersia Needham.

Block tests are a measure established for wholesalers and retailers to price a variety of cuts given a certain producer price. By using the block test, the quantity of meat that will be produced can be predicted, and the price per carcass can be calculated. Therefore, block tests (Table 14.3 on a buffalo with a carcass weight of 277 kg) should ideally be conducted regularly to help determine the number of different cuts that will be produced from a buffalo carcass.

Table 14.3 Block test conducted on an African savanna buffalo (carcass weight of 277 kg).

Retail cutsWeight (kg)% whole carcass
Topside11.44.11
Silverside15.25.49
Rump steak10.93.93
Thin flank8.53.07
Thick flank8.93.21
Short fillet1.90.68
Soft shin113.97
Shin3.31.19
Tail0.50.18
T-bone steak15.85.7
Blade steak13.74.94
Brisket17.36.24
Short rib6.32.27
Chuck/prime22.78.19
Neck113.97
Trimming217.58
Stew43.615.74
Goulash3.11.12
Hump4.51.62
Bones3412.27
Sinew and fat10.73.86
Band saw loss1.70.67
Total277100

Primal cuts should be vacuum-packed and matured for a minimum of 25 days under refrigerated conditions (and labels should clearly indicate the day in and suggested sale date) before being sold, with the sirloin and topside muscles in particular identified as valuable cuts. The fillets of all animals can be sold as soon as they are removed; due to their inherent nature/composition, they need not be aged. In contrast, the silverside is ideal for biltong production, as toughness will not decrease over a more extended ageing period, and it will thus remain a tough muscle. The offcuts and trimmings could be used for value-added meat products, including biltong, droëwors (traditional dried sausage product made from meat off-cuts, sheep/beef fat and traditional spices), patties, boerewors (traditional fresh sausage) and minced meat. Biltong can be produced from frozen-thawed (detained) carcasses using different drying methods, which creates a larger profit margin because frozen carcasses can only be used for processing biltong, stewing meat and mince.

15 Handling and Moving the African Buffalo

M. La Grange , N.J. La Grange , J. Mostert , J. Mostert-La Grange , R. Hoare , M. D. Kock , I. L. Lekolool and P. Chardonnet
Introduction

Archaeology and literature provide evidence that African wild animals have been live captured for a very long time, at least since the ancient Egyptians (e.g. Trinquier, Reference Trinquier2002; Mark, Reference Mark2016) and Romans (e.g. Bertrandy, Reference Bertrandy1987; Mackinnon, Reference Mackinnon2006; Christesen and Kyle, Reference Christesen and Kyle2014). However, modern wildlife capture methods are only a few decades old, and considerable progress has been achieved recently in innovative chemical and physical restraint techniques for all wildlife species. While these methods are now used all over the continent, southern Africa appears today as the leading region in wildlife capture. In South Africa, the Wildlife Translocation Association’s members annually capture and translocate approximately 130,000 game animals, and the game capture industry now has an annual turnover in excess of €7.4 million (Snyman et al., Reference Snyman, Sumba and Vorhies2021).

As a member of the famous ‘Big Five’, the African buffalo (Syncerus caffer) adds significantly to the value of the wildlife economy everywhere, both for consumptive and non-consumptive use. However, with its historical range severely impacted by human activity, attaining this value today is often dependent upon being able to physically ‘manage’ them. The conservation of buffalo is consequently massively enhanced and facilitated today by being able to capture, handle and move the species. All of these actions are very specialist undertakings because buffalo are large-bodied, live in sizeable herds and can become aggressive and dangerous to humans.

Reasons for Capturing and Moving Buffalo

Buffalo may need to be captured and released on-site for diverse reasons. Depending on the type of management (extensive or intensive; Chapter 13), a manager can request a licensed veterinarian to examine or treat injured or sick individuals. Similarly, buffalo populations at risk of disease outbreaks, posing a sanitary risk to other species (such as livestock or humans) or subject to sanitary regulations may be required to be captured for mass vaccinations to control the targeted diseases. Under extensive management (i.e. natural conditions), contexts necessitating the capture of buffalo may be more restricted (i.e. for a suffering individual).

Disease investigations or pre-movement health checks, notably for foot and mouth disease (FMD – including setting up FMD-free herds), bovine tuberculosis (bTB), tick-borne diseases (especially theileriosis) and other zoonotic or livestock diseases, are additional reasons to sample buffalo populations after immobilization. In addition to sanitary knowledge for production purposes, ecological or ecosystem research, for example collaring individuals with satellite tracking devices, may require captures for a few minutes in order to fit or release the devices.

Finally, buffalo may be captured for translocation for diverse reasons (Box 15.1). These include establishing founder populations, either for reintroduction or introduction purposes; numerically or genetically reinforcing depleted, isolated or small populations; moving vulnerable individuals or populations; and finally moving individuals to mitigate human–wildlife conflicts. Table 15.1 provides a sample of recent translocation events in eastern and southern Africa showcasing these diverse reasons for moving buffalo.

Table 15.1 A few examples of capture and translocation operations of African buffalo

DatePlaceNumber of buffalo involvedReasonOperationsResults
1984Zambezi valley, ZimbabweGroups of around 100 eachInvestigation of foot and mouth disease (FMD)Capture, process, and releaseEstablishing strains and level of disease
1985, 1986From Zambezi valley to central Zimbabwe3 herds of around 100 eachRemoval of female calves to establish FMD-free herdsCapture entire herds and remove calvesRelease natal herds on site and remove calves to safe haven to be quarantined
1990 to 2020Harare, ZimbabweSeveral groups of 1–10 individualsFMD-free herds and redistribution of blood linesIndividual darting, loading and transportation99% survival rate despite 5-day turn around
1995Malilangwe, ZimbabweSeveral herdsIndividual vaccination for anthraxCapture, vaccination in crush and releaseThe buffalo population was saved
1997, 1998Kariba dam, Zimbabwe400, then 400Rescue from lake KaribaCapture in Bumi Hills, transport by ferry and release in Gache GacheSuccessful relocation: 98% survival rate
2004From Lake Nakuru National Park (NP) to Il Ngwesi Conservancy, Kenya54Population management at source and improving genetic diversity at recipient siteCapture, transport and release100% survival rate
2004From Lake Nakuru National Park to Il Lewa Wildlife Conservancy, Kenya19Population management at source and improving genetic diversity at recipient siteCapture, transport and release100% survival rate
2008From Solio Ranch to Aberdare National Park, Kenya60Mitigation of habitat destructionCapture, transport and release100% survival rate
2009From Kibiku Forest, Ngong to Nairobi NP, Kenya17Mitigate human–wildlife conflict (HWC)Capture, transport and release100% survival rate
2009Hwange NP, Zimbabwe500Reintroduction in Mwenezi RanchCapture and translocation from Robins camp to Mwenezi RanchSuccessful reintroduction
2010Marromeu National Reserve (NR), Mozambique99Investigation of bovine tuberculosis (BTb)Capture, sampling and releaseStatus of bTB in Marromeu buffalo
2012From Marromeu NR and Gorongoza NP to Gilé NR, Mozambique20Reintroduction of buffalo in Gilé NR (species formerly extinct)Capture, transport, and releaseSpecies reintroduced in Gilé NR
2013From Niassa NR to Gilé NR, Mozambique47Reinforcement in Gilé NR (consolidation of the reintroduction)Capture in Niassa NR, transport over 900 km and release in Gilé NRThe reintroduction of the species is consolidated (150 buffalo in 2021)
2015 to 2017From Marromeu NR to Coutada 9, Mozambique50 in 2015 and 200 in 2017Reinforcement of the relict buffalo population in Coutada 9Capture in Marromeu NR, transport and release in Coutada 9Successful reinforcement in Coutada 9 (380 buffalo in 2020)
2017From Marromeu NR to Zinave NP, Mozambique250ReintroductionCapture, transport and releaseSuccessful reintroduction, 99% survival rate
2017, 2018From Chinhoyi, Zimbabwe, to DRC50, then 50IntroductionAirlift to Luanda and drive to Lubumbashi99% survival rate
2019From Kitengela to Nairobi NP, Kenya8Mitigation of HWCCapture, transport and release100% survival rate
Source: Author.

Box 15.1 Definitions of Translocations (IUCN/SSC, 2013)

Conservation translocation is the intentional movement and release of a living organism where the primary objective is conservation. This will usually comprise improving the conservation status of the focal species locally or globally, and/or restoring natural ecosystem functions or processes. Conservation translocations are classified according to the intended benefit of the process, entailing releases either within (population restoration) or outside (conservation introduction) the species’ indigenous range.

  • Population restoration involves:

    • Reinforcement, which is the intentional movement and release of an organism into an existing population of conspecifics, aiming to enhance population viability.

    • Reintroduction, which is the intentional movement and release of an organism inside its indigenous range from which it has disappeared, aiming to re-establish a viable population of the focal species.

  • Conservation introduction involves:

    • Assisted colonization, which is the intentional movement and release of an organism outside its indigenous range to avoid extinction of populations of the focal species.

    • Ecological replacement, which is the intentional movement and release of an organism outside its indigenous range to perform a specific ecological function. This is used to re-establish an ecological function lost through extinction and will often involve the most suitable existing subspecies, or a close relative of the extinct species within the same genus.

For whatever reason a capture operation is decided, it should be carefully thought through, and planned with strict adherence to veterinary regulations with respect to disease control, dangerous drugs, etc. The planning exercise should pay particular attention to behavioural characteristics and stress management.

Most of the knowledge and information presented in this chapter is based on massive experience and skills developed after many years of practice by the authors and other colleagues (La Grange, Reference La Grange2005, Reference La Grange, du, Bothma and Du Toit2010).

Behavioural Characteristics to Consider for Capturing Buffalo

Like most bovids, buffalo are herd-orientated animals occupying distinct home ranges (Chapter 6). Therefore, they try to stay with one another while being driven towards the boma (a large funnel-shaped enclosure to physically capture buffalo – see below). Once in the boma, they will continue to follow the lead individual, who is likely to exploit any weakness in the integrity of the boma and try to initiate an escape response. Consequently, identifying and managing the lead individual effectively manages the herd.

Surprisingly, under capture pressure, the individual taking the lead to escape or attack is more often a cow that both defends and leads the way to safety. In the wild, bulls will join and separate depending upon age; subadult males will often leave to form bachelor groups while post-breeding old bulls will separate. These older bulls, often referred to as ‘dagga’ bulls in southern Africa, frequently have impaired eyesight and hearing. They sometimes seek the safety (i.e. from natural predators) of areas surrounding human habitations and can become exceedingly dangerous if stumbled upon.

Mothers are strongly bonded to their calves, recognizing them immediately from the bawling vocalizations of their respective offspring. They will respond aggressively to investigate and retrieve them even under stampede situations. Yearlings are reliant on parent herd knowledge, become lost in unrecognizable environments and need to be close to their mothers and other herd members. The cow–calf bond lasts for longer periods than for many other species (a comparatively long time between birth and puberty).

Home range dispersal is generally forced through large predator interactions or human-induced disturbances. Individual adult animals driven out into surrounding communities because of forced dispersal tend to be harassed and therefore can become exceedingly dangerous and often attack with little provocation. These individuals are unlikely to be returned successfully to the area they originated from.

Behaving in many respects like sable, once cornered and unable to escape in a boma, the buffalo herd will form a tight circular gathering often referred to as ‘laager’ in southern Africa, providing an effective defence strategy, especially against predators such as lions. This strategy prevents losses which likely would occur if the herd panicked and ran in several directions.

However, in a herding situation during capture, depending upon the pressure exerted on them, buffalo will readily attack when deemed threatened. The correct reading of the situation is necessary and relies on experience in order to apply just enough pressure to solicit the required response. It may be necessary to back off before this pressure becomes too much and invokes an attack or unnecessary panic. In this way, the lead cow becomes both ally and foe, and given the opportunity, would prefer to find a way out of the capture boma. She and the herd may be cleverly manipulated through good boma design and management to achieve smooth capture of the herd. Experienced operators and helicopter pilots do make a difference!

Stress Management

Fifty years of capture experience by professional operators have proven that for all species, stress cannot be removed completely, but its intensity can be minimized during every stage of the capture operation. Observations indicate that the main activity compounding stress is uncontrolled panic and running with attempts to attack or to escape. Struggling and overexertion prior to restraint must be quickly brought under control. Uncontrolled running up and down a boma, for example, is sufficient to overexert animals, resulting in potential health complications. Similarly, continued stress observed during physical restraint requires quick, deft action to control and calm animals throughout the operation. Basic application, for example, of a blindfold prior to or during immobilization and tranquillization, will provide a calming effect. Fortunately, buffalo are less prone to panic than many other species, but these principles still apply.

Any capture operation must minimize mortality and stress. Currently, advances in translocation knowledge have resulted in minimal mortalities, even when moving entire herds over long distances. An acceptable mortality rate is considered to be <2 per cent. However, modern capture techniques have been so refined that almost zero or very low mortality is now achievable. A thorough understanding of the subtleties of buffalo herd behaviour is important to overall success. Such knowledge allows potential problems to be anticipated before they occur, providing for timely management decisions that can be corrected and adapted throughout the entire capture process, effectively minimizing stress at all stages with the goal of zero mortality.

Physical Capture/Mass Capture

Physical capture is the capture of wild animals without using drugs. Mass capture of herd animals using a temporarily erected boma is a revolutionary technique of physical capture that originated in the 1960s in Namibia (Oelofse, Reference Oelofse1970). In the animals’ natural habitat, opaque or ‘blind’ plastic polyweave sheeting is erected on poles and supported top and bottom by tension cables in the formation of a large funnel-shaped enclosure, usually named mass boma capture, often referred to simply as boma capture (Figure 15.1). The principle employed is that animals are herded into the large open end of the funnel, and while being forced to traverse it to the narrow end, will not challenge the tall, flimsy barrier because they cannot see an escape route to the outside. The funnel ends in a crush in which animals can be individually selected or handled and thereafter either released or loaded onto a custom-made transporter vehicle already situated on an exit road.

Figure 15.1

(a) Diagram of a temporary mass capture boma constructed out of woven plastic sheeting.

(b) In practice, the boma is camouflaged in the vegetation and the narrow ‘crush’ section curves towards a ramp into transport vehicles on an exit road (transport vehicle on the top).

Source: Author.

Using a small, manoeuvrable helicopter is the preferred method of driving animals to a boma (using horses, vehicles or people would be cheaper but is impractical for many reasons). It is absolutely essential that the pilot is not simply a commercially rated licence holder but someone who has experience in both low-level game capture flying and wild animal observation and behaviour. While flying the pilot has to be able to separate a workable number of animals from a herd, move them as calmly as possible in a downwind direction towards the boma, and avoid having them panic, separating far from each other and escaping from the group. At the same time, the pilot must communicate via radio and thus coordinate the activity of members of a ground team strategically positioned in and around the capture boma. The helicopter pilot flies alone to reduce helicopter mass enabling more power for manoeuvring.

The overhead noise and presence of a helicopter provide sufficient stimulus to move most wild animals including buffalo. The disturbance readily groups them while moving them in the required direction. The pilot must vary altitude and position judiciously, strategically moving around the herd, and applying varying levels of pressure to direct and keep the selected herd/group together. Directional pressure is gradually increased (lower altitude, closer distance) as the herd approaches the well-concealed wide boma entrance. Near the mouth of the boma, the helicopter is flying low, and finally activates a loud siren to provide the final stimulus while directing ground staff to close the main gate, with the process repeated to close secondary gates as the herd funnels down through the boma.

Ground teams stationed on the boma partition gates rapidly draw these plastic curtains across in sequence behind the buffalo as the herd progresses forward towards the narrow end of the boma. Employing this strategy, the herd is confined in a manageable sized space (a boma compartment), with individual movement restricted and less chance of escape. It is essential to limit outside stimuli while the boma is occupied to allow buffalo to regain their rest composure following the stressful chase. In large capture operations, the manageable number of animals driven in per helicopter drive is around 30–40 buffalo. This number is considered practical to handle and load, adding further subdrives to fill additional boma compartments if required or carrying out additional drives after the loading of the previous subdrive.

Placing buffalo under stress during mass capture is akin to them being hunted as prey in the wild where they display equivalent behavioural responses. Initially, they bunch into a circular defensive formation to prevent losses that would likely occur from individuals panicking and running in several directions. They often attack under extreme or persistent pressure, with a number of individuals charging in one direction behind a leader.

Pushing captured wild buffalo one by one into a narrow ‘cattle-like crush’ is obviously not easy and cannot be done by people on foot inside the boma. A modified four-wheel drive vehicle with strong front and side protection can be used to achieve this by slow ‘persuasion’ (Figure 15.2). Buffalo will attack the vehicle, especially the front and wheels, rather than target the people inside. Adding a curve to the handling facility, or hanging vegetation at the far end of the boma to camouflage the dead end, leads the herd into thinking that there is an attractive way of escape, and its leaders will eventually move on towards that (Figure 15.1).

(a) buffalo are chased into the boma by the helicopter;

(b) then pushed into desired sections of the boma or lorry using an adapted vehicle.

© Philippe Chardonnet.

Figure 15.2 Final stage of a buffalo mass capture:

All procedures in the capture of wildlife are in effect stress management exercises. Put simply, physical capture without drugs is achieved through a sequence of induced animal behaviours: naturally moving away from the disturbance source, encouraged to take the escape opportunity given, enabled by loss of geo-location and reluctance to challenge the unknown (see some tips in Box 15.2).

Box 15.2 Tips for Managing Buffalo within the Mass Boma

  • The helicopter drive for separating the required number of buffalo is important to avoid driving too many in at once. Too many animals will be difficult to manage in the boma. Instead, position the main herd nearby ±200 m and separate 30–40 animals for each subdrive into the boma, filling the respective boma compartments providing further options. Rather than filling each compartment, more often additional drives are conducted after the loading of the previous subdrive.

  • Identify and work with the lead cow as discussed. This requires lots of patience – offer a way to escape and capitalize on any advantage emerging.

  • Provide a suitable boma herding vehicle within the boma as operating on foot would be extremely dangerous. As indicated, prevent direct confrontation, applying targeted pressure, following or backing off, observing the response. Any old 4×4 vehicle will do as buffalo tend to attack the vehicle, especially the front and wheels, rather than the people inside.

  • The secret is: ‘Give n’take!’ Avoid applying too much pressure and AVOID direct contact!

  • The boma is flexible in design to provide for a wide range of capture applications, including:

    • Combining physical and chemical capture to enable the selection of specific animals for testing or capture. Basically, instead of a crush, providing a sufficiently large circular working area at the boma end to drive the herd in and dart individual animals selected from a vehicle. Individual animals can be marked and released back to the herd or following bulk knockdown, for example, veterinary testing, before releasing the herd back to the wild.

    • Calf removal for FMD-free buffalo breeding programmes is basically a variant of this technique, capturing the entire herd, then employing a vehicle to drive among the herd as they mill around, ‘fishing’ out the calves individually with a rope and pole noose. Male calves are separated from the females into different bulk crates. Finally, the males are released back to the herd before the final herd is released back to the wild.

    • Conducting large immunizing programmes against diseases such as anthrax. This is done by providing an extended crush at the boma end to hold 30–50 buffalo. These are pole syringed and, importantly, marked for identification before release.

    • The first gate (Figure 15.1) serves to first contain the driven buffalo herd, aiming to keep the area between it and the main gate pristine for subsequent drives.

Chemical Capture/Individual Capture

Chemical immobilization of buffalo is achieved by darting or injecting drugs. Darting and handling of immobilized wild animals require very comprehensive training in specialized courses that are available in a few selected countries. Even veterinarians are advised to attend these courses because this very detailed and specialist field is usually not fully covered in general veterinary training. Dangerous drugs, especially opioids, fall under very strict veterinary regulations. In cases of accidental exposure to these, such as the potent opioids used for buffalo, humans are unfortunately extremely susceptible to the same effects, which can rapidly be fatal. Safety procedures are paramount and must be thoroughly applied.

Darting on the ground requires approaching a buffalo within close range (maximum about 40 m), which requires knowledge of the behaviour of the species and judgement of each circumstance. It is safest and easier to approach by vehicle if the terrain allows. However, in West and Central Africa where many landscapes do not allow driving off track, and where helicopters are difficult to hire, buffalo darting is often carried out on foot (Figure 15.3). Compared to many other large mammals, buffalo are quite easy to approach on foot provided the stalking is done very strictly against the wind. However, approaching buffalo by foot close enough for darting becomes a difficult exercise in poorly managed areas where harassed buffalo become very shy. Operational success hence may drop from darting a few buffalo each day to just one buffalo every few days, especially if specific individuals are targeted (e.g. an adult female or collared individual for device removal).

Figure 15.3 Individual darting on foot of a West African savanna buffalo.

© Daniel Cornélis.

Lone males are the easiest to get close to, followed by male coalitions. Herds are more difficult, however, often with males following behind.

Darting from a helicopter requires a pilot with experience in both low-level flying techniques and interpreting animal behaviour (Figure 15.4). The darter and the pilot must have excellent intercom communication in the air. Aircraft reliability and aviation and veterinary safety procedures must be established and adhered to as there is very little margin for error (see tips in Box 15.3).

Figure 15.4 Chemical capture of a free-ranging herd of Cape buffalo by individual darting from a helicopter.

© Samy Julliand.

Box 15.3 Tips to Consider with Aerial Darting of Buffalo

  • Capitalize on the ‘window-of-opportunity’ offered by confusion, separating out smaller, manageable subgroups of buffalo, driving them to more accessible open ground away from the principal herd, forcing them to circle on themselves to promote confusion, and quickly darting all the individuals comprising the group. It is important to keep them together in suitable recovery terrain until they all go down. This process requires skill and experience from the helicopter pilot and darter working together with ground teams, who should be directed in to render timely assistance to potentially compromised animals. It is important to minimize the total downtime of the group.

  • Positive knockdown of the targeted animals is paramount, requiring correct dart placement and the appropriate drug combination. Generally, with free-range darting of wild buffalo, apply the high-dose opioid protocol (Table 15.2), avoiding underdosing that is more problematic with the potential for complications. Buffalo herded by a helicopter are more likely to be stressed and hyperthermia can be a problem, especially with a dark-skinned animal. Consider combining thiafentanyl with etorphine in a 50:50 combination dose, which significantly reduces the excitement phase, reducing time running and therefore distance travelled. Thiafentanyl alone in combination with azaperone provides quick knockdown, but thousands of buffalo in southern Africa have been successfully immobilized with etorphine and azaperone.

  • It is especially recommended to immediately redart if poor dart placement is suspected rather than waiting for drug sign. Be prepared to manage possible overdose using butorphanol.

Table 15.2 Drug recommendations for the African buffalo (taken with the kind permission of Kock and Burroughs, Reference Kock and Burroughs2021).

BuffaloOpioidTranquillizerOpioid antagonistα-2 agonist
High-dose opioid protocol
Free-ranging bullsEtorphine 7–8 mg orAzaperone 40–60 mgNaltrexone 140–160 mgNone
Thiafentanil 7–8 mgAzaperone 40–60 mgNaltrexone 70–80 mg
Mix etorphine 4 mg and thiafentanil 4 mgAzaperone 40–60 mgNaltrexone 120 mg
Free-ranging cowsEtorphine 4–6 mgAzaperone 40–60 mgNaltrexone 80–120 mg Diprenorphine at 12-18 mg is useful in loading
Thiafentanil 4–6 mgAzaperone 40–60 mgNaltrexone 40–60 mg
Mix etorphine 3 mg and thiafentanil 3 mgAzaperone 40–60 mgNaltrexone 90 mg
Adults in bomaEtorphine 3–5 mgAzaperone 40–60 mgDiprenorphine 9–15 mg or
naltrexone 60–100 mg
Low-dose opioid protocol
Adult bullThiafentanil 1.5–2 mgMedetomidine 4 mg plusNaltrexone 15–20 mgAtipamezole 4 mg plus yohimbine at 0.5 ml per mg of medetomidine
Azaperone 40mg
Adult cowThiafentanil 1–1,5 mgMedetomidine 3–4 mg plusNaltrexone 10–15 mgAtipamezole 4 mg plus yohimbine at 0.5 ml per mg of medetomidine
Azaperone 40 mg
0–6 monthsThiafentanil 1 mgNaltrexone 10 mg
6–12 monthsThiafentanil 1 mgMedetomidine 0.5 mg plus azaperone 15 mgNaltrexone 10 mgAtipamezole 0.5 mg plus yohimbine at 0.5 ml per mg of medetomidine
12–24 monthsThiafentanil 1 mgMedetomidine 1 mg plus azaperone 20 mgNaltrexone 10 mgAtipamezole 1 mg plus yohimbine at 0.5 ml per mg of medetomidine
24–36 monthsThiafentanil 1 mgMedetomidine 2 mg plus azaperone 30Naltrexone 10 mgAtipamezole 2 mg plus yohimbine at 0.5 ml per mg of medetomidine

Notes:

  • Azaperone is recommended in all buffalo immobilizing combinations.

  • Ketamine given intravenously (IV) is effective as a ‘top-up’ drug in animals that are not sufficiently immobilized by the opioid and sedative/tranquillizer mixture. Administer 100–200 mg IV and further doses can be given if required. Doses of 50–100 mg will often be sufficient.

  • Naltrexone for free release is preferred and for transport a mixture of diprenorphine and naltrexone is useful.

  • Diprenorphine at 12–18 mg is useful in loading.

  • In forest buffalo, the same drug combinations can be used but reduce doses accordingly due to smaller size. Carfentanil (4 mg) has been used successfully to immobilize this subspecies – reverse with 200–300 mg naltrexone. The use of carfentanil may result in knuckling problems post-recovery in savanna buffalo and is not recommended for them.

  • The addition of hyaluronidase to the mixture in the dart is advisable in buffalo, particularly with the high-dose opioid protocol.

  • Azaperone is the better drug to use. Buffalo are generally sensitive to the effects of the α-2 agonists so take due precautions in free-living animals. When complete reversal of the α-2 agonist is required, use atipamezole.

The darting of individual buffalo or small groups uses combinations of opioid drugs and tranquillizers (Table 15.2). Such combinations are designed to provide synergistic effects; the opioid is the ‘knockdown’ component, inducing a physiological state called ‘narcosis’ which is much different from the deeper unconscious state of ‘anaesthesia’, familiar to most humans. The addition of a tranquillizer reduces stress by smoothing the ‘induction period’ (the time for a drug’s full effect over several minutes), counters muscle rigidity during the immobilization phase, and improves the recovery process.

Historically, the traditional choice of drug combinations has been etorphine hydrochloride (M99, Captivon® Wildlife Pharmaceuticals): 5–8 mg etorphine combined with 40–50 mg azaperone for a free-ranging adult, reducing this marginally (by 20 per cent) for penned and tamer animals (Figure 15.2). With free-range darting of wild buffalo, apply the high-dose opioid protocol. In holding boma situations, pens or where buffalo are calm or habituated, the low-dose opioid protocol can be substituted. Either on the ground or from the air, if poor dart placement occurs or dart failure is suspected, it is generally recommended to redart immediately rather than waiting for signs of drug effect.

Following chemical capture, the recumbent buffalo is given an opioid antagonist, a rapidly acting antidote drug which allows the animal to regain full consciousness to normal mobility with all its vital functions intact. Recovery drugs (variously called ‘antidotes’, ‘reversals’ or ‘antagonists’) and their dosages vary according to what management procedure is required following capture. If a buffalo needs to be moved from an area inaccessible to transport, ‘partial antagonists’ drugs may be used. The remarkable efficacy of these drugs is to allow the buffalo to get to its feet and be slowly physically guided by well-trained handlers. Obviously, this is while it is still blindfolded and well restrained with ropes, but in a heavily tranquilized state and not sufficiently awake to injure the handlers or escape uncontrollably.

It is far easier to manage the effects of ‘overdosage’ of immobilizing drugs on a wild animal than ‘underdosage’. To most people, this would seem counterintuitive. The reason, however, is that if a recumbent animal is physiologically compromised, there are various ways of quickly improving its vital functions to keep it alive while it remains recumbent and manageable. In the worst-case scenario, should the procedure not be able to continue, an intravenous opioid antidote drug can quickly wake the animal up and bring it back to normality. By contrast, if the animal receives an insufficient dose, it will remain ‘half-immobilized’ on its feet and continually try to escape, whereupon its uncontrolled mobility and associated high stress levels can cause it to become rapidly compromised physiologically, which is often fatal (see tips in Box 15.4).

Box 15.4 Tips for Monitoring the Immobilized Buffalo

  • The immobilized animal should be approached slightly from behind. A blindfold should be gently lowered over its eyes, this will significantly reduce stress, effectively assisting in relaxing the animal and protecting the eyes. Earplugs are optional.

  • Immediately upon approach, the buffalo must be placed and maintained in a sternal recumbency position with the head lowered. Over the years the potential for regurgitation and aspiration of ruminal contents into the lungs has proven an issue, especially with drug combinations using α-2 agonists.

  • Throughout the whole procedure until the end, respiration must be monitored with a normal rate of 6–8 breaths/minute. Less than this, butorphanol can be injected in increments of 5 mg but beware of complete reversal at doses higher than 30 mg. Doxapram given at 5–15 ml IV also provides respiratory stimulation, but is short-acting and can produce some arousal, so constant monitoring is required. A rate of 10–12 means the animal is light and, depending on the time from darting, may require a top-up of 100–200 mg of ketamine. Avoid adding more opioid into the animal, ketamine is highly effective and safe.

  • A large buffalo’s limbs folded under a heavy body are very susceptible to lack of blood supply during longer procedures; thus, the body position must be regularly adjusted to maintain adequate circulation to the limbs and to avoid any nerve damage due to pressure.

  • The drugs to either terminate or prolong immobilization must be on hand and the required procedures known in detail by the operators. For reversal, a combination of naltrexone (25 to 100mg depending on opioid dose) and diprenorphine (12-18mg) can be used IV or IM, especially when more than one animal are woken up at the same time in a recovery crate – the lower dose of naltrexone helps with the recovery of buffalo. A significant cost reduction can be achieved using this in combination with the full diprenorphine dose.

The choice of a commercially available darting system is a matter of personal preference. There are two main types of ‘remote injection devices’, distinguished by the method of dart propulsion from the gun and drug injection from the dart. Powder-charged guns use small blank cartridges while gas-powered guns are fitted with small cylinders containing compressed carbon dioxide (CO2). Drugs are expelled from darts by either small powder charges or the release of compressed air. Two of the most used systems are called ‘Pneudart®’ and ‘DanInject®’. These darting systems are versatile enough that they can be used with darts from different manufacturers (11 mm and 13 mm barrel bore size), such as Palmer Cap-Chur® (0.50 calibre or 13 mm). It is important to select the correct needle size and length (2 × 50 mm) when darting buffalo (Kock and Burroughs, Reference Kock and Burroughs2021).

Transport of Buffalo

Transporting wild animals as large as buffalo by road is a very detailed and specialized undertaking, requiring large amounts of equipment, logistical support and organizational capability. Unless only a few individual buffalo are to be transported a very short distance (which could be done by tranquillizing and transporting them recumbent in a pickup truck), moving buffalo requires experienced ‘capture operators’.

In some African countries, there may be detailed veterinary and other legal requirements for moving buffalo, resulting in extensive prior paperwork. Buffalo can share several diseases and parasites with livestock, so official health requirements can be very stringent, expensive and time-consuming.

Customized crates replicating ‘shipping containers’ are the most used equipment for road transport. Obvious requirements for these containers are non-slip flooring, good ventilation, sliding doors and operator access via the roof for observation or animal behaviour intervention. Buffalo travel well in groups when well-designed crates and good management practices are used, and can travel for up to 36 hours after capture without food and water en route. Watering and feeding can be done at both ends of the journey. Tranquillization using injectable drugs should be limited to bulls and/or truculent animals only. Never tranquillize juveniles or yearlings because there is the risk that they may lie down in transit and be trampled by adults (see tips in Box 15.5).

Box 15.5 Tips for Loading Buffalo and Managing Their Transportation

  • Buffalo travel well as mixed groups.

  • Need to employ fully enabled management crates to properly distribute the captured animals – considered essential.

  • Compartments fitted with fully functional sliding doors – capable of separating buffalo, allowing the movement of animals back and forth between compartments as required.

  • Cross-loading capabilities – extra truck units may be needed to cross-load as required.

  • Crates providing full access from the top to inspect, move, sort, tranquillize and operate partition gates.

  • Watch packing density. Buffalo are prone to hyperthermia, which can be exacerbated by too many animals packed into a compartment.

  • Not too many animals at one time should be in a crush. It is a good idea to split the load into 20–25 subgroups at the crush (a suitable number for a truck and trailer load). Where large numbers are to be moved, consider cross-loading into additional truck units linked up.

  • Generally, calves at foot do not load and transport well; therefore, avoid the breeding season.

  • Bulls tend to be easier to load, but they will occupy a larger space and should be appropriately tranquillized.

  • Tranquillization of truculent individuals only.

  • Never tranquillize calves and yearlings that are then prone to be trampled upon!

  • Buffalo can travel for up to 36 hours after capture without food and water.

  • Should watering and feeding prove necessary, this may be achieved by moving animals into an adjacent compartment, for example, when undertaking time-consuming cross-border operations.

To transport live animals, drivers of heavy vehicles must be extensively trained and experienced. The best travel route must be researched and planned, and very reliable communications guaranteed between drivers and support staff. The weather en route is often overlooked: in hot daytime conditions buffalo can overheat if too tightly packed; on the other hand, when travelling during the coldest part of the night, a wind chill factor can reduce the ambient temperature by up to 8°C. On long journeys, vehicle stops in quiet locations at a maximum of every 200 km are essential. Extra requirements for unforeseen problems arising are a senior staff member in an accompanying 4WD ‘chase vehicle’ and an empty crate for cross-loading should some animals need to be removed during transit (see checklist in Box 15.6).

Box 15.6 Checklist Tips Prior to Transportation

  • Ensure animals are settled before transporting! Unload and reload should animals remain unsettled requiring group resorting.

  • Tranquillize as required.

  • Ensure correct paperwork is in hand: wildlife permits, vehicle clearance, border crossing. Experience dictates that it is best to use the services of an experienced clearing agent to facilitate border crossing.

  • Consider best travel route: terrain, condition and directions.

  • Weather conditions en route are more often overlooked. Consider potential for chill factor problems en route when travelling through the coldest part of the night. Wherever possible, avoid these situations, remembering the chill factor may often reduce ambient temperatures by a factor of 4–8°C.

  • Consider GPS track monitoring of the vehicle.

  • Cell/smartphone–satphone enabled.

  • Stop and rest frequently, every 200 km. Select quiet stopping places, not near gatherings of people, for example, a village.

  • Watering and feeding are not normally an issue if delivering within 24 hours. Empty one compartment and place flat troughs and fresh grass should this become necessary.

  • It is very important to ensure rehydration on final release.

  • Consider refuelling requirements for the trucks: identify places and the currency required. Think logically through all these requirements, addressing any that have the potential to interrupt smooth passage.

  • Drivers should be experienced and well briefed.

Long-distance considerations

  • Use alternate drivers.

  • Stop and rest frequently, every 200 km.

  • Emergency tranquillization may be needed (azaperone and diazepam) for particularly truculent individuals.

  • An accompanying ‘chase’ vehicle is important with a qualified person to assist the animals for cross-border deliveries.

  • May need to water down animals, drinking considerations become more important with long distances (avoid excessive water in crate, beware of danger of slipping).

  • Monitor weather en route, especially whether hot or cold spells.

Release at Destination

Herbivores with social behaviour like buffalo are easier to translocate than many other taxa, but a successful capture does not end upon reaching the destination, where much remains to be done. Possibly one of the greatest pitfalls to concluding a successful capture and translocation is the problem of subsequent maladaptation when wild animals are introduced to new surroundings.

If the transport arrives late at night, leave the buffalo inside the crates and unload them early the following day. Allowing buffalo immediate and full rehydration at the destination is essential. Immediate free release (also named ‘hard release’) might be practiced in a fenced environment. However, the construction of an adaptation boma is recommended where the buffalo can settle into new surroundings via an initial captive period. If the new habitat is substantially different from the source area, for example, subtle complications involving rumen microflora adaption can be very important. Dietary maladaptation can seriously impact the animals’ health and future survival.

When boma management is carried out correctly, buffalo do settle down relatively quickly compared to some other wild herbivores (see tips in Box 15.7). A few main points relating to this phase of handling are to allow an area of about 1 ha (100 × 100 m) for up to 25 buffalo and increase accordingly. As in a capture boma, the walls must be opaque or blanked off (e.g. woven plastic sheeting on fences), adequate shade for the entire group should be available, and good-quality grass and/or hay ad libitum should be provided, concentrate supplements can be added but only in small quantities daily. Human activity inside the pen (cleaning/animal sorting) should be kept to a minimum and during the cooler hours of the day and any disturbance, especially from spectators, must be strictly limited and only outside the boma. Domestic dogs should never be allowed anywhere near wildlife-holding bomas or pens. For final release back to the wild, pick a quiet day early in the morning and simply leave the enclosure gate open to allow the buffalo to find their own way out in their own time.

Box 15.7 Tips for Pen Management

  • In carrying out the daily chore management of the pens, it is important to establish a routine. Animals are naturally routine-orientated as they go about their daily business of feeding and resting. This should be maintained as much as possible during the penning process.

  • It is extremely important to respect the midday siesta routine. Naturally, animals rest in shade during the hot part of the day, so pen cleaning, sorting and animal feeding should be limited to early morning before 11h00 and late afternoon from 16h00 onwards.

  • Move animals to a new pen once a week. This allows for pen cleaning and rotation, which greatly helps when they finally need to be loaded and transported.

  • Ensure that any disturbances remain on the outside of the pens, never on the inside. The pen attendant and family should be accommodated nearby the pen complex, effectively providing disturbance to the outside but not invading the privacy within. This greatly assists with the taming process; animals realize that the pens represent safety.

  • For further reading, refer to Raath (Reference Raath and Penzhorn1996).

The Case of Virtual Boundary

The more that wildlife are managed, the more there are indications that a virtual component is playing out. This is particularly noticeable upon releasing animals into a new environment that they do not know; essentially, they are rendered lost. In the wild, animals are fully geo-located to their home ranges and always know accurately where they are. The establishment of ‘remembered’ boundary positions, risky areas and food and water locations is information imprinted virtually in their mindset, logged against time and season. If they are suddenly uplifted and moved in a dramatic fashion to where a new reality is foisted upon them, it renders them rather lost and unable to recognize much of their new surroundings. Experience over the years has demonstrated that release stress is best cushioned by placing animals in a blanked off and confined space (boma or pen) that they cannot see out of. In this, they quickly establish the basic information relating to the whereabouts of food, shelter and most importantly, a refuge. This position becomes ‘virtually logged’, so that release into a new wild area is undertaken from this remembered position as they gradually move away from it, exploring their new surroundings. The virtual boundary concept is interesting for future management in that this virtual knowledge is proving invaluable in developing innovative approaches to mitigate capture complications and human–wildlife conflict (La Grange et al., Reference La Grange, Matema, Nyamukure and Hoare2022).

Conclusion

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’. Achieving this objective involves well-coordinated teamwork with individual team members practicing an eclectic mixture of activities that add up to ‘an art as much as a science’. However, that said, there is still room for progress. The chemical capture of large mammals remains overly dependent upon opioids, which are problematic for two main reasons: (i) they are extremely dangerous for humans and animals, and (ii) strict procurement protocols have severely hampered access for most countries outside the few with substantial experience in wildlife capture. Hence, there is a real need to actively research non-opioid drugs, especially the alpha-2s (e.g. medetomidine) and combinations thereof. Physical capture methods also could be improved, and maybe even new strategies developed, including for example virtual applications such as a drone capture technique (under review), and applying scent technology through a guided, one-way camouflaged crush arrangement into a compacted mobile crate management arrangement, obviating the necessity for large plastic mass boma equipment, helicopters and expensive labour commitment.

16 Buffalo Hunting: From a Commodity to a High-Value Game Species

P. Chardonnet , R. Taylor , W. Crosmary , S.P. Tadjo , F.A. Ligate , R. Baldus , L. Siege and D. Cornelis
A Story Longer Than Thought
Early Days

January 1895. Makanga country, now in Mozambique. Edouard Foa, a French explorer, is struggling to gain an audience with the powerful and feared Chief Tchanetta Mendoza. Foa had come there on his way to cross the continent by foot from the Indian Ocean to the Atlantic Ocean. Eventually, after having threatened Foa, the Chief consented to grant him a clearance to walk across and hunt on this land. At that time, the country was rich in game and Tchanetta forbade unnecessary shooting. Because Arabs used to come there from the North once a year for times immemorial, the Chief had them hunt elephants exclusively, measuring the powder for each hunter himself. Buffalo (Syncerus caffer), antelope, and other game were reserved to indigenous hunters for feeding his people. The tribute to be paid to the Chief for hunting elephant was one tusk per elephant killed. When the beast had fallen, the tusk that was on the ground side was the property of the Chief of the territory. Locally, in Portuguese, this tax was named ‘o dente da terra’, the Earth’s tooth (Foa, Reference Foa1900).

The price to pay for the right to hunt existed long before Foa. As early as the sixteenth century, Portuguese records state that no elephant could be killed and consumed without the consent of the Chief in the lands south of the Zambezi, where the ‘dente da terra’ tax already existed by unwritten law (Manyanga and Pangeti, Reference Manyanga, Pangeti, Manyanga and Chirikure2017). Such hunting levies were not only restricted to this area. In western Tanzania, Foa had to pay the ‘hongo’, a tribute to walk and hunt on a Chief’s land (Foa, Reference Foa1900). In western Zimbabwe, Lobengula (1836–1894), Chief of the Ndebele, was issuing hunting concessions for foreign hunters as a way to protect Ndebele hunting rights (Moyo et al., Reference Moyo, O’Keefe and Sill1993).

These ancient situations reveal extremely important historical traits: systems of governance and management of wildlife were already in place in precolonial times, mainly enacted by traditional leaders and their ruling families (Sansom, Reference Sansom and Hammond Tooke1974; Campbell, Reference Campbell1995; Carruthers, Reference Carruthers1995), even endorsed by spirit mediums, at least in the Zambezi valley (Hasler, Reference Hasler1996). These systems did not disappear abruptly under colonial rule and often coexisted with new foreign regulations.

Today, the current trophy fee paid by the hunting tourist is nothing other than a modern form of the historical ‘dente da terra’. The present listing of particular species as fully protected is nothing other than ancient rules such as the prohibition by Lobengula of hunting hippopotamus, and the fee paid by the hunting operator to lease a hunting concession from the State is nothing other than the historical tribute to be paid to the landlord for being allowed to walk and hunt on his land. Today, by delegating the appropriate authority from central to local levels, the now widespread mechanism of community-based natural resources management is in a way reviving precolonial systems, but with more democratic efforts than under the past feudal regimes.

Colonial Times

With the establishment of colonies, foreign powers assumed that the traditional sanctions and precolonial institutions that regulated hunting were an inadequate means of conserving wildlife in the face of growing human populations and competition for wildlife resources (Child, Reference Child and Child2004). By transposing their foreign laws, many colonial regimes prescribed wildlife as res nullius: with wildlife now belonging to no one and managed by the State, traditional rulers were disempowered from controlling hunting. It is even assumed that some of them let poaching happen to steal State goods in revenge for having lost control.

The settlers who began arriving at the Cape of Good Hope in 1652 hunted wildlife for food and commercial gain (Booth and Chardonnet, Reference Booth and Chardonnet2015), and to open land to develop agriculture and livestock husbandry. In less than two centuries, wildlife had been deeply impacted by the introduction of millions of muzzleloaders, metal gin-traps, etc. (Richards, Reference Richards1980), the development of agriculture, and the expansion of livestock accompanied by several exotic diseases. The rinderpest outbreak in the 1880s wiped out up to 95 per cent of the buffalo populations (Robertson, Reference Robertson1996; Spinage, Reference Spinage2003; Chapters 9 and 12). Regarded as common game, buffalo did not benefit from special protection and were even destroyed in southern Africa in the attempt to eradicate tsetse flies. Most colonial regimes maintained special, relatively cheap meat hunting licences to feed populations and plantation workers (Anderson, Reference Anderson and Flack2017).

At the end of the nineteenth century, a number of hunters throughout Africa recognized the harm of uncontrolled hunting and played a key role in establishing protected areas (Kruger National Park in 1894 in South Africa, Selous Game Reserve in 1896 in Tanzania). In the meantime, they also introduced modern protective game laws. All over Africa, many if not most of the Hunting Reserves that were gazetted at that time are the ancestors of today’s National Parks. The turn of the century was the period when hunting for trading ivory and skins or for collecting specimens for museums (Roosevelt, Reference Roosevelt1910) gave birth to hunting for sport, adventure and exotic travels named safari (safari means travel in Swahili). Hunting tourism arose in East Africa with pioneer farmers and explorers guiding foreign hunters (Lindsey et al., Reference Lindsey, Roulet and Romañach2007). After the First World War, the hunting safari industry expanded, policed by law and administration. After the Second World War, sport hunting became more organized and regulated as a business (Booth and Chardonnet, Reference Booth and Chardonnet2015).

Independence

After independence, game and hunting laws were progressively modernized and the network of Protected Areas developed. Safari hunting continued except for a few countries like Kenya, where it was banned in 1977, which precipitated the steep decline of game numbers in the country (Western et al., Reference Western, Russell and Cuthill2009; Ogutu et al., Reference Ogutu, Piepho and Said2016). In contrast, neighbouring Tanzania, after a temporary hunting ban between 1973 and 1977, has maintained until today safari hunting on vast areas while also succeeding in maintaining the highest numbers on Earth of large mammals such as lion and buffalo. Unexpectedly, the bans on hunting in Kenya and temporarily in Tanzania made both safari hunting clients and professional hunters look for new hunting fields in other regions of Africa, which boomed following the bans in East Africa (Hurt and Ravn, Reference Hurt, Ravn, Prins, Grootenhuis and Dolan2000).

While buffalo remained common in some areas, more and more situations were arising, especially in West and Central Africa, where local buffalo populations were diminishing as human population growth drove demand for more land at the expense of wilderness, with agriculture and livestock encroachment, and with increasing poaching pressure for bushmeat. Gradually, hunting became controlled by sustainability norms and integrated into conservation strategies. The rationale was to create sustainable revenue streams for rural communities and State wildlife agencies, thus providing incentives to preserve Hunting Areas as duly gazetted Protected Areas, in a challenging attempt to prevent their conversion into agriculture or other environment-unfriendly land uses (Prins and de Jong, Reference Prins, de Jong, Kiffner, Bond and Lee2022). In several African countries, there was a gradual alignment of trophy-hunting industries with conservation and development policies, supported by a number of international donor agencies (Lindsey et al., Reference Lindsey, Roulet and Romañach2007).

Starting in the 1980s with the Communal Areas Management Programme for Indigenous Resources (CAMPFIRE) programme in Zimbabwe, new approaches aiming at increasing benefits from hunting and other wildlife uses for local populations led to a paradigm shift towards connecting sustainable use and hunting with rural development and livelihoods (Murphree, Reference Murphree2000; Chapters 1 and 13). This approach progressively expanded throughout Africa with the Administrative Management Design programme (ADMADE) in Zambia, the Programme de Développement des Zones de Chasse Villageoises (PDZCV) in CAR, the Zones d’Intérêt Cynégétique à Gestion Communautaire (ZICGC) programme in Cameroon, the Gestion Participative des Ressources Naturelles et de la Faune (GEPRENAF) programme in Burkina Faso, and the Ecosystèmes Protégés d’Afrique Soudano-Sahélienne (ECOPAS) programme (Lindsey et al., Reference Lindsey, Roulet and Romañach2007). The foundation of this Community-Based Natural Resources Management (CBNRM) approach is to allocate user rights to local people, thereby allowing for benefits from wildlife use and creating conservation incentives (Baldus, Reference Baldus2009). However, the implementation of this approach is not always that simple. In south-eastern Zimbabwe, for example, Poshiwa et al. (Reference Poshiwa, Groeneveld, Heitkönig, Prins and van Ierland2013) show the limitations of revenues from wildlife diversification, even though wildlife income is less volatile than income from the agro-pastoral system, and wildlife can be used as a hedge asset to offset risk from agricultural production without compromising on return.

In these utilization schemes, hunting tourism has in most cases the highest income potential (Booth, Reference Booth2010). As one of the most numerous large game animals, the buffalo is a core species for high-income hunting tourism (Lindsey et al., Reference Lindsey, Balme, Booth and Midlane2012). Buffalo hunts contribute a high share to community hunting income under CBNRM, for example in CAR (Bouché, Reference Bouché2010) and Tanzania (TAWA, 2019).

Hunting Buffalo Today
Hunting Buffalo for Meat and Other Reasons
Informal Bushmeat Hunting Throughout Africa

Hunting for food began millennia ago with the first humans. Today, many rural communities across the continent still heavily rely on bushmeat, both for food security and income (e.g. Loibooki et al., Reference Loibooki, Hofer, Campbell and East2002; van Vliet and Mbazza, Reference Van Vliet and Mbazza2011; Friant et al., Reference Friant, Ayambem and Alobi2020). Consumption of buffalo meat occurs broadly across the wide range of wild animal species consumed (Table 16.1). The pay-off for hunting a buffalo is high: a single buffalo represents one of the greatest amounts of meat that can be obtained per capita, and buffalo meat is one of the most nutritive among the wild species usually hunted (Cawthorn and Hoffman, Reference Cawthorn and Hoffman2015). Buffalo is highly prized in urban markets and restaurants. While not the case everywhere, in some places like in Bangui, Central African Republic (Fargeot et al., Reference Fargeot, Drouet-Hoguet and Le Bel2017), or Manica Province, Mozambique (Lindsey and Bento, Reference Lindsey and Bento2012), its meat is among the most expensive. This makes buffalo one of the species most targeted by poaching in several areas (Skikuku et al., Reference Skikuku, Makenzi and Muruthi2018; Gaodirelwe et al., Reference Gaodirelwe, Masunga and Motsholapheko2020). Buffalo meat may also be obtained as a by-product of conflicts between the species and the local communities; several communities hunt buffalo in retaliation after the species has raided their crops or attacked people (Long et al., Reference Long, Mojo and Fu2020).

Table 16.1 Uses (either legal or illegal) of African buffalo by local communities: examples across the species range by region, in West, Central, East and Southern Africa (based on data/sources in the table).

RegionCountryAreaBuffalo productUseDetailsReference
West AfricaBurkina FasoBobo-DioulassoMeatFoodBuffalo hunted in groups for commercial purpose, sold to restaurantsMontcho et al. (Reference Montcho, Ilboudo and Dayou2020)
TestisTraditional medicineAphrodisiac potions
TrophySocial prestige
GhanaMeatFoodSpecies previously regarded as totems, such as buffalo, started to appear openly on major bushmeat markets because of increasing poverty and the growing scarcity of preferred wildlife speciesFAO/CIG (2020)
Ivory CoastComoé National ParkMeatFoodAll the local residents surveyed feed on buffalo flesh and/or skinAtta et al. (Reference Atta, Soulemane, Kadjo and Kouadio2021)
Organs and other body partsTraditional medicineE.g. tail, heart, leg bones, horns, poop, urine fat, brain, bile to cure diarrhoea, eye aches, folie, heartache, bone weakness, sexual impotence, etc.
WitchcraftTurning away bad luck, banish fear, repulse bad spells, etc.
MeatFoodNear extinction of buffalo because of hunting for foodP. Henschel, unpublished data in Lindsey et al. (Reference Lindsey, Balme and Becker2015)
Table 16.1 (cont.) – Part A
RegionCountryAreaBuffalo productUseDetailsReference
NigeriaAbia, Bauchi, Edo, Kogi, Ondo and Zamfara StateMeatFoodAlarape et al. (Reference Alarape, Shuaibu and Yaduma2021)
PenisTraditional medicineUsed as an aphrodisiacAdeola (Reference Adeola1992)
Ibadab, Oyo StateBoneTraditional medicineUsed as anti-vomitingOduntan et al. (Reference Oduntan, Akinyemi and Ojo2012)
Cross River StateMeatFoodHunters prefer not to hunt buffalo because the traditional system demands them to share the meat with community members, sometimes in distant places. However, Fulani herdsmen reported that they responded to the conflict between buffalo and cattle by setting wire snares along trails and shooting buffaloEniang et al. (Reference Eniang, Ebin and Nchor2017)
South-western townsSkin, eyeballs, liver, tail, penis, etc.Traditional medicineElephantiasis, loss of hearing/speech/eyesight vertebral column fracture, prolonged pregnancy; extrusion of placenta after parturition, human skull fracture, fertilitySodeinde and Soewu (Reference Sodeinde and Soewu1999)
Nose, headWitchcraftInvoking witches appeasing traditional gods
Table 16.1 (cont.) – Part B
RegionCountryAreaBuffalo productUseDetailsReference
Central AfricaCameroonKimbi-Fungom National ParkMeatFoodBuffalo hunting is a source of incomeNda et al. (Reference Nda, Tsi and Fominyam2018)
Buffalo is generally avoided in the Pygmy groups because it is considered as having a potential harmful effect on humansDuda et al. (Reference Duda, Gallois and Reyes-García2018)
Central African RepublicBanguiMeatFoodBuffalo, with snakes, are the most expensive species sold in the marketsFargeot et al. (Reference Fargeot, Drouet-Hoguet and Le Bel2017)
Yangambi Landscape, Yangambi Biosphere Reserve and the Ngazi Forest ReserveMeatFoodThe local extirpation of buffalo is explained as the result of overhunting by armed groups (Armed Forces of the DRC, Congolese, Rwandans, and Ugandans from eastern DRC) during the periods of rebellionVan Vliet et al. (Reference Van Vliet, Muhindo and Kambale Nyumu2018)
Around Lomani National ParkMeatFoodBuffalo nearly disappeared because of overhuntingBatumike et al. (Reference Batumike, Imani, Urom and Cuni-Sanchez2021)
Democratic Republic of CongoGaramba National ParkMeatFoodDuring peacetime, protected species such as elephant and buffalo rarely appeared in the rural markets, but they comprised more than half of all bushmeat sales in the urban markets. During wartime, the sales of protected species in the urban markets increased fivefoldDe Merode and Cowlishaw (Reference De Merode and Cowlishaw2006)
South of the Salonga–Lukenie–Sankuru LandscapeMeatFoodBuffalo meat sold in large quantities, accounting for the highest percentage of total weight of carcasses found in the local marketSteel et al. (Reference Steel, Colom, Maisels and Shapiro2008)
Table 16.1 (cont.) – Part C
RegionCountryAreaBuffalo productUseDetailsReference
Democratic Republic of Congo and Republic of the CongoKinshasa and BrazzavilleMeatFoodBuffalo meat illegally sold in the restaurants. Buffalo is the most expensive meat among the ungulate speciesGluszek et al. (Reference Gluszek, Viollaz and Gore2021)
GabonGambia Complex of Protected AreasMeatFoodIn most locations with buffalo, signs of poaching were found as well. Buffalo meat sold at a price of 2200 CFA-FrancLitjens (Reference Litjens2017)
Republic of the CongoPointe NoireMeatFoodBuffalo is among the species most frequently bought and sold in markets and restaurantsBoratto and Gore (Reference Boratto and Gore2018)
East AfricaEthiopiaWestern EthiopiaBushmeat and illegal trophy hunting are the key causes of buffalo collapse. Bushmeat hunting is carried out by local poachers or local militias, whereas most illegal trophy hunters come from the remote parts of Limu, Gidda Ayana and Ebantu districts of the East Wollega Administrative ZoneErena et al. (Reference Erena, Jebessa and Bekele2019)
Oromia Regional StateTrophySocial prestigeHunting buffalo for trophies was frequently practised in the areaErena (Reference Erena2014)
KenyaMount Elgon Biosphere ReserveMeatFoodBuffalo is the mostly targeted species, after antelopes. Also hunted by poachers from UgandaSkikuku et al. (Reference Skikuku, Makenzi and Muruthi2018)
TailCultural ornamentation, sign of prestige
NationwideRetaliatory killingBuffalo is the second most commonly killed species in retaliation for damage causedLong et al. (Reference Long, Mojo and Fu2020)
Table 16.1 (cont.) – Part D
RegionCountryAreaBuffalo productUseDetailsReference
RwandaVolcanoes National ParkMeatFoodHunting buffalo for meat was the most common forest activity in the past. Less common now because of increased law enforcementMunanura et al. (Reference Munanura, Backman and Sabuhoro2018)
SudanDinder Biosphere ReserveMeatFoodDuring periods of famine, conflict and critical fallback of food sources (crop and domestic livestock), many Sudanese consume all types of wild fauna, including buffaloAdam (Reference Adam2019)
TanzaniaUzungwa Scarp Forest and Mwanihana ForestMeatFoodLocally extinct in the Reserve by the early 1970s as a result of intensive hunting for bushmeat tradeRovero et al. (Reference Rovero, Mtui, Kitegile and Nielsen2012); Hegerl et al. (Reference Hegerl, Burgess and Nielsen2017)
South West Rungwa Game ReserveMeatFoodCommunities get meat through resident hunting. Buffalo meat is mostly used for trade to generate incomeNachihangu et al. (Reference Nachihangu, Kiondo and Lwelamira2018)
Western SerengetiMeatFoodThe ethnic groups in Western Serengeti prefer medium–large wildlife such as buffalo for protein and incomeHolmern et al. (Reference Holmern, Mkama and Muya2006); Ndibalema and Songorwa (Reference Ndibalema and Songorwa2007); Mfunda and Røskaft (Reference Mfunda and Røskaft2010)
Table 16.1 (cont.) – Part E
RegionCountryAreaBuffalo productUseDetailsReference
Tarime DistrictMeatFoodThe harvesting rates of buffalo are alarming. Buffalo was reported to be reduced by 50–90% out of their rangeHolmern et al. (Reference Holmern, Røskaft and Mbaruka2002, Reference Holmern, Mkama and Muya2006); Kideghesgo et al. (2006)
UgandaNorthern UgandaMeatFoodBuffalo meat found in local marketsDell et al. (Reference Dell, Masembe and Gerhold2021)
Near Murchison National ParkMeatFoodBuffalo is perceived by poachers as the most dangerous wild animal to hunt and the most dangerous to trapDell et al. (Reference Dell, Souza and Willcox2020)
Southern AfricaBotswanaOkavango DeltaMeatFoodApproximately 1800 illegal hunters each harvest an average of 320 kg of bushmeat annually, although some reported harvesting =1000 kg. While impala was the most commonly hunted species, buffalo accounted for 30% of all bushmeat productionRogan et al. (Reference Rogan, Lindsey and Tambling2017)
In and outside Wildlife Management Areas around the Moremi Game ReserveMeatFoodCBNRM communities mostly target impala, followed by Cape buffaloGaodirelwe et al. (Reference Gaodirelwe, Masunga and Motsholapheko2020)
MozambiqueMeatFoodIllegal hunters commonly use gin traps, which are manufactured from steel car springs and used to kill animals as large as buffaloLindsey and Bento (Reference Lindsey and Bento2012)
Manica ProvinceMeatFoodBushmeat from large species such as buffalo is less frequently sold today than during the civil war. However, buffalo is one of the most commonly cited bushmeat species by intervieweesLindsey and Bento (Reference Lindsey and Bento2012)
Table 16.1 (cont.) – Part F
RegionCountryAreaBuffalo productUseDetailsReference
South AfricaPafuri in the Makuleke concessionMeatFoodCable from the dilapidated western boundary fence frequently stolen by illegal hunters to make snares to capture hippo and buffaloC. Roche, unpublished data in Lindsey et al. (Reference Lindsey, Balme and Becker2015)
Xhosa and Sotho communities in the Western Cape ProvinceBonesTraditional medicineBuffalo bone is one of the most expensive animal items soldNieman et al. (Reference Nieman, Leslie and Wilkinson2019)
Faraday market, JohannesburgSkull, horns, skinTraditional medicineBuffalo is one of the ungulate species most represented in the marketWhiting et al. (Reference Whiting, Williams and Hibbitts2011)
ZambiaLuangwa Valley, Upper and Lower Lupande, Lumimba and Sandwe game management areasMeatFoodDeclining population of buffalo in areas close to human settlements, close to boundary of the National ParkR. McRobb, M. Becker and D. Lewis, unpublished data in Lindsey et al. (Reference Lindsey, Balme and Becker2015)

Local communities also hunt buffalo for purposes other than meat (Table 16.1). In Ethiopia, for instance, poachers hunt buffalo as trophies to increase their social acceptance and respect in society (Erena, Reference Erena2014). For the Bisa people in Zambia, there are multiple dimensions to hunting buffalo, including social positioning and cohesion of their society (Marks, Reference Marks1976). In many areas, buffalo body parts are used for cultural ceremonies and in traditional medicine (Whiting et al., Reference Whiting, Williams and Hibbitts2011).

There are some communities that are reluctant to hunt buffalo. First, because hunting buffalo may be perceived as too dangerous by local hunters (Dell et al., Reference Dell, Souza and Willcox2020). In many traditional systems, hunters also have to share the meat from their hunts with a large number of community members. They therefore tend to avoid large species such as buffalo to limit the expense of delivering parts of the hunted animals to relatives living in distant places (Eniang et al., Reference Eniang, Ebin and Nchor2017). Finally, for some communities, the buffalo is regarded as a totem or taboo animal, and its hunt is not allowed (FAO/CIG, 2002; Duda et al., Reference Duda, Gallois and Reyes-García2018; Chapter 1).

That said, hunting for bushmeat largely contributes to local declines of buffalo populations, even sometimes to the vanishing of the species (Prins, Reference Prins1996; Batumike et al., Reference Batumike, Imani, Urom and Cuni-Sanchez2021). Basically, bushmeat hunting is unselective and unlimited; where snares and gin-traps are set for buffalo and other game, any calf, female or male can be taken, and with no limitation in numbers given that traps can be reset. Bushmeat hunting is often considered one of the greatest threats to biodiversity in African savannas and forests, often ahead of other major threats such as deforestation and habitat fragmentation (Wilkie et al., Reference Wilkie, Bennett, Peres and Cunningham2011; van Velden et al., Reference Van Velden, Wilson and Biggs2018).

Regulated Bushmeat Hunting

Some countries, such as Tanzania, allow hunting quotas for meat purposes (including buffalo), while others allow subsistence community hunting, like CAR (Snyman et al., Reference Snyman, Sumba and Vorhies2021). In most countries, trophy-hunting concessionaries are mandated by their lease agreements to provide local communities – free of charge – the meat obtained by tourist hunters. This is quite stringent in West and Central Africa, where wild meat is extremely sought after. In Zambia, 130 tons of fresh game meat – of which 24 per cent is from buffalo – are provided annually by the hunting tourism industry to rural communities at an approximate yearly value for the meat alone of over €500,000 exclusive of distribution costs (White and Belant, Reference White and Belant2015).

In some southern African countries, the production of wild meat constitutes a real industry, one that is organized and regulated. In Namibia, with an annual mean of between 60 and 75 kg of venison produced per square kilometre in 2013 on farmland, hunting for venison is an important sector which contributes more to national food security than livestock, as beef is mainly exported (Lindsey et al., Reference Lindsey, Havemann and Lines2013). However, most of the venison is from antelopes, not from buffalo, which is restricted by veterinary regulations. In South Africa, ‘biltong hunting’ is a recreational hunting by local hunters who harvest wild meat and process it into biltong (dried meat) or sausage (Taylor et al., Reference Taylor, Lindsey and Davies-Mostert2015). It is a major value chain in this country, much larger than trophy hunting; however, it mainly targets common game rather than buffalo.

Buffalo Hunting Tourism
What Are We Talking About?

This section addresses lawful and regulated hunting only, in contrast with outlawed and unregulated hunting, commonly called poaching (see Prins, Reference Prins2020). The terminology of hunting categories has been debated at length (Booth and Chardonnet, Reference Booth and Chardonnet2015). One reason is that the terms used in each language are often difficult to translate literally into other languages. Another reason is that the various categories of hunting often overlap (IUCN, 2016). For IUCN, ‘trophy hunting is hunting of animals with specific characteristics and involves the payment of a fee by a foreign or local hunter for a hunting experience, usually guided; it may be a distinct activity or overlap with recreational or meat hunting’. While trophy hunting reflects the quest for an outstanding trophy, sport hunting rather reflects the quest for a challenging fair chase of the game by tracking on foot, whatever the trophy. The trophy is a key part of a safari, but the hunting experience and adventure in the bush are also what attracts clients, and there also has to be the feeling of a fair chase to the proper hunter with no guarantee of success (Hurt and Ravn, Reference Hurt, Ravn, Prins, Grootenhuis and Dolan2000). While some authors prefer the term ‘regulated hunting’ (Dickson et al., Reference Dickson, Hutton and Adams2009; Booth and Chardonnet, Reference Booth and Chardonnet2015), many other terms are commonly used, for example safari hunting, recreational hunting, tourism hunting, hunting tourism. For Spenceley (Reference Spenceley2021), ‘hunting tourism is a consumptive mode of nature-based tourism that uses renewable natural resources in a wild or undeveloped form for the purpose of enjoying natural areas or wildlife and contribute to conserve and value wilderness areas’. It is a typical tourism value chain with (i) emitting countries, that is countries of origin of the clients (hunting tourist or tourist hunter), and (ii) receiving countries, that is countries selling operating rights to tourism operators (hunting company or hunting operator or outfitter), themselves selling tourism services (hunting safari or hunting trip or hunting party or hunt) to their clients.

Throughout Africa

To most hunters, the buffalo is a fascinating game for being (i) one of the so-called ‘dangerous game’ and (ii) one of the ‘Big Five’, the term commonly used to describe the five major big game species. Hunting accidents with buffalo are not uncommon, even with experienced professional hunters. The buffalo is widely regarded as dangerous to hunt, which certainly adds to the attractiveness of its hunt: ‘He looks as if you owe him money’ (Ruark, Reference Ruark1987, italics added for emphasis). In 2022, buffalo can be legally hunted by hunting tourists in 16 sub-Saharan African countries, that is in 43 per cent of the 37 buffalo range countries (Figure 16.1). The COVID-19 pandemic in 2020 and 2021 prevented hunting tourists from travelling, which severely impacted hunting tourism like all forms of tourism. The situation slightly returned to normal in 2022.

Figure 16.1 Buffalo range countries where hunting tourism is lawful in 2022 for the four subspecies of buffalo recognized by the IUCN Red List so far. Note: Buffalo in northern and central Angola were categorized as ‘Cape buffalo’ by IUCN (2019), but phenotypically and perhaps even genetically they are ‘forest buffalo’.

Source: Author.

Among the four subspecies thus far recognized by the IUCN Red List (Chapters 3 and 4), the Cape buffalo is by far the most hunted, being legally hunted in nine countries. This obviously reflects its much higher abundance than the other subspecies, but also other factors like a greater development of the tourism industry, a safer security situation, a larger expansion of CBNRM programmes, etc. The forest buffalo is the least hunted subspecies with only three countries where it can be hunted legally, a situation resulting from a more restricted range, landlocked hunting grounds, the difficulty of the hunt in thick habitats, and also probably a degraded conservation status. The West and Central African savanna buffalo, both subspecies being rather similar and intermixed, can be hunted in eight countries. However, the overall number of buffalo trophy-hunted annually in these two regions has always been quite low, about 300 a year. We need to mention that the hunting community recognizes a fifth subspecies, the Nile buffalo, which ranges in Ethiopia, northern and western Uganda, and appears as an intermediate form between the Central African savanna buffalo and the Cape buffalo. The reality of the transitional shape of its trophy explains that hunters specifically hunt this particular buffalo and register their trophies distinctly in the records books.

Hunting quotas (the maximum number of adult male buffalo allowed to be hunted per year per Hunting Area) and offtakes (number of buffalo effectively harvested per year per Hunting Area) vary greatly between regions, with the highest figures in Tanzania and Southern Africa and the lowest in West and Central Africa (Table 16.2). The national offtake rate (ratio of offtake to quota) is not only the result of the number of buffalo taken per Hunting Area, but also of the percentage of Hunting Areas being leased and operational, which is a sign of the functionality of the industry in the country. In nearly all of the hunting countries, the hunt concerns free-ranging buffalo in unfenced Hunting Areas. South Africa, where buffalo hunting happens behind fences, is a major exception. Another peculiar feature of South Africa is that hunting quotas are set by the landowner, while they are generally set by government authorities quasi-everywhere else.

Table 16.2 Buffalo hunting quotas and offtakes in selected countries throughout Africa.

2011/20122012/20132013/20142014/20152015/20162016/20172017/20182018/20192019/2020Average
West AfricaBenin (PNP, 2018, 2019; PNW, 2018, 2019)Buffalo quota (N buffalo)n/an/an/an/an/an/a7575n/a75
Buffalo offtake (N buffalo)n/an/an/an/an/an/a5759n/a58
Buffalo offtake rate (%)n/an/an/an/an/an/a7679n/a77
Burkina Faso (DFRC, 2018)Buffalo quota (N buffalo)147163153153183183181n/an/a166
Buffalo offtake (N buffalo)115136129821158181n/an/a106
Buffalo offtake rate (%)78838454634445n/an/a64
Central AfricaCameroon (MINFOF, 2020)Buffalo quota (N buffalo)n/an/an/an/an/a352341356381358
Buffalo offtake (N buffalo)n/an/an/an/an/a15618412599141
Buffalo offtake rate (%)n/an/an/an/an/a4454352639
East AfricaTanzania (Wildlife Division, personal communication, 2021)Buffalo quota (N buffalo)2130213021301948145614561456145614561735
Buffalo offtake (N buffalo)1129901889940828672655625737820
Buffalo offtake rate (%)53424248574645435147
Southern AfricaZimbabwe (ZPWMA, personal communication, 2022)Buffalo quota (N buffalo)n/an/a17941751120513081343125212891420
Buffalo offtake (N buffalo)n/an/717699593642592585200575
Buffalo offtake rate (%)n/n/4040494944471641
Namibia (only for Communal Conservancies) (MEFT and NACSO, personal communication, 2022)Buffalo quota (N buffalo)n/an/a106106108122122122132117
Buffalo offtake (N buffalo)n/an/a889393110991146194
Buffalo offtake rate (%)n/an/a8385869081934680
West Africa

Three countries of West Africa allow legal hunting of buffalo. In Senegal, with a relict population of West African savanna buffalo in the far south-eastern corner of the country, buffalo trophy hunting is anecdotal. In contrast, Benin and Burkina Faso have developed a well-organized and regulated big game hunting tourism industry with the West African savanna buffalo as the main attraction together with the roan antelope (Hippotragus equinus). Buffalo hunting there is renowned for being a challenging, fair chase by stalking on foot with excellent local trackers.

In Burkina Faso, in 2017, 303 hunting tourists (9 per cent of all tourists) harvested 424 mammals for a production of 86 tons of meat and a direct revenue of about €827,000 (Ouedraogo, Reference Ouedraogo2018). Over seven years between 2012 and 2018, the average national annual quota was 166 buffalo/year (147–183), and the average national annual offtake was 106 buffalo hunted/year (81–136) for a national annual offtake rate of 64 per cent (44–84) (DFRC, 2018; Table 16.2).

In Benin, over the two hunting seasons 2017–2018 and 2018–2019, the five existing Hunting Areas (only four of which were operational) harvested an annual average of 58 buffalo out of an average annual quota of 75 for an average annual offtake rate of 77.3 per cent (PNP, 2018, 2019; PNW, 2018, 2019; Table 16.2). In 2018, the W National Park ecosystem earned 76 per cent of its revenue from 19 hunting tourists visiting the Mekrou Hunting Area and 2 per cent from 476 photographic tourists visiting the W National Park (PNP, 2018, 2019; PNW, 2018, 2019; Table 16.2).

Since 2019, the severe degradation of the security situation in the region (with terrorism taking over vast wilderness areas) has prevented many National Parks and Hunting Areas from operating in West Africa.

Central Africa

Central Africa is the region where buffalo are the most diverse, with three subspecies occurring out of four. Buffalo there is not the first game of appeal for tourist hunters, who mainly look for the Eastern giant eland (Tragelaphus derbianus gigas) and the Western or lowland bongo (Tragelaphus eurycerus eurycerus). However, buffalo is part of the hunting package and is sought after for providing serious stalking by foot with outstandingly skilful trackers from local communities.

Cameroon is the country with the highest number of legal big game hunters in all of West and Central Africa in recent years. In 2018, 285 tourists came to Cameroon for hunting (MINFOF, 2019). In this country, Hunting Areas are a major component of the national network of Protected Areas: 71 gazetted Hunting Areas (Zones d’Intérêt Cynégétique) cover 57,000 km² (11.9 per cent of the country), that is 1.5 times the size of National Parks and Reserves (39,000 km², 8.2 per cent of the country; MINFOF, 2019). Over four hunting seasons between 2016 and 2020, the average annual quota was 358 buffalo (341–381) with 69 per cent savanna buffalo and 31 per cent forest buffalo. During this period, an annual average of 141 buffalo (99–184) were hunted for an average annual offtake rate of 39 per cent (MINFOF, 2020; Table 16.2). Such a low offtake rate reflects an important proportion of unleased Hunting Areas, as a number of them are no longer operational due to degradation by all sorts of activities which are illegal in protected areas: poaching, livestock invasion (Figure 16.2), cotton encroachment, gold mining, logging, and the charcoal trade.

Figure 16.2 Livestock sightings in the BSB landscape covering the transboundary national parks of Bouba Ndjidda (Cameroon) and Sena Oura (Chad) as well as the seven neighbouring Hunting Areas (Cameroon), during the aerial wildlife survey in 2018 (total surface of about 10,500 km²). The estimated livestock population (117,134 heads) was six times higher than the estimated population of the 11 largest wild mammals (20,136 individuals), and located mostly within the Hunting Areas surrounding the National Parks.

(data and illustration reproduced from WCS and MINFOF, 2018, with permission)

The Central African Republic (CAR) could be named the ‘buffalo country’, as it is the only one on the continent where three subspecies of buffalo occur and can be legally hunted, although the forest buffalo is rarely hunted there. In this country, 89 gazetted Hunting Areas cover 220,000 km² (35 per cent of the country), that is 3.6 times the size of the National Parks and Reserves (61,000 km², 10 per cent of the country). Before the political unrest initiated in 2012, CAR was a prime destination for big game hunting. It is still practiced in 2022, but so far remains marginal. Before the collapse of tourism, the buffalo was the second most abundant large game species after the giant eland in the Zones cynégétiques villageoises (ZCV, Village Hunting Zones) of northern CAR, with a density of 1.1 buffalo per km² (Bouché, Reference Bouché2010). In these ZCV only, the buffalo was the most hunted game species: in the 2008–2009 hunting season, 44 buffalo were harvested by hunting tourists, ahead of 26 giant eland (Bouché, Reference Bouché2010).

Chad is renowned for hosting the typical form of Syncerus caffer aequinoctialis with its wide, flattened horn shape. The country used to be famous for big game hunting until the contemporary civil turmoil. Despite these constraints, hunting tourism continued to be practiced in 2022, but at a lower scale.

In the Republic of Congo, hunting tourism has recently resumed with only a few forest buffalo harvested per year.

The security situation in Central Africa has been deteriorating for a longer time than in West Africa, and this has undermined the hunting industry as well as conservation. The region is experiencing what Scholte et al. (Reference Scholte, Pays, Adam and Chardonnet2021) call a conservation overstretch: with increasing insecurity and declining revenues, governments find themselves confronted with too few resources to protect vast areas.

East Africa

In East Africa, three countries have developed a well-structured hunting tourism industry. In Ethiopia, few buffalo are hunted for the simple reason that the Hunting Areas are not exactly located within the buffalo range in this country. The buffalo is not the game of appeal for tourist hunters coming to this country. In Uganda, the hunting industry has developed over the last 20 years to a point where it is now a real alternative to the other East and Central African hunting destinations. A special attraction is the so-called Nile buffalo, and Uganda is the place to find it (Siege and Siege, Reference Siege and Siege2020).

Tanzania, which hosts the largest number of African buffalo on Earth, unsurprisingly comes first among all African countries for regulated hunting of free-ranging buffalo. Tanzanian buffalo are famous for their large herds and their magnificent wide horns.

The hunting domain is an essential pillar of the national network of Protected Areas in this country. In 2004, proclaimed Protected Areas gazetted as Hunting Areas covered over 250,000 km² (26.4 per cent of the country), nearly twice the size of the National Parks (134,881 km², 14.1 per cent; Baldus and Cauldwell, Reference Baldus and Cauldwell2004). The number of Hunting Areas was progressively reduced from 164 to 113 in 2020 with the gazetting of several Game Reserves as National Parks. However, Hunting Areas still cover nearly a quarter of Tanzania’s surface. Many if not most Hunting Areas are not viable for other conservation options such as ecotourism due notably to remoteness, lack of scenery and poor visibility of wildlife compared to the top National Parks.

Hunting tourism is an important and organized sector in Tanzania. For the 2013–2018 period, out of 164 Hunting Areas, 149 were awarded to 60 hunting companies. However, there was considerable financial pressure during this period due to adverse publicity regarding sport hunting, and the impact of hunting bans on elephant and lion trophy imports to the USA, Europe and Australia (TAWA, 2019). With fewer hunting clients visiting Tanzania than in previous years, hunting tourism revenues dropped from €44 million in 2008 with 1673 hunters (Booth, Reference Booth2010) to €28.3 million in 2014 with 708 hunters (Booth, Reference Booth2017). When the cost of maintaining Hunting Areas became higher than the income, many hunting companies returned their Hunting Areas to the wildlife authorities. By the end of 2018, 81 Hunting Areas were handed back, representing slightly less than 130,000 km² (approximately 52 per cent) of the area set aside for hunting (TAWA, 2019).

This downtrend also impacted buffalo conservation in two contrasting ways. First, when the Hunting Areas formerly leased for hunting were abandoned, these 13 million ha of wilderness became vacant, and hence were exposed to poaching and encroachment by other land uses detrimental to the environment. Highly susceptible to these threats, buffalo became a collateral victim of the bans on the importation of hunting trophies directed at elephants and lions, two species listed on CITES Appendices. Second, as the buffalo is not a CITES-listed species, the bans turned the buffalo, once considered a secondary game species, into a first-choice species for hunters travelling to Tanzania. The character of the Tanzania hunting industry has changed over the last 10 years from being a ‘big four’ game hunting destination to one that is now heavily dependent on leopard and buffalo (TAWA, 2019).

However, although the trophy fee for buffalo is cheaper than that of the flagship game species, buffalo remains the first tax-earning species in this country due to the larger number harvested: in 2019/2020, the trophy fees (€2080 per buffalo) of 737 buffalo hunted in 77 Hunting Areas earned €1.53 million, to which all other revenue sources should be added, that is hunting block fees, licences, daily fees (Wildlife Division, personal communication).

Over eight years between 2012 and 2020, the average national annual quota was 1681 buffalo/year (1456–2130), and the average national annual offtake was 781 buffalo hunted/year (625–940) for a national annual offtake rate of 46.3 per cent (41.7–56.8; Wildlife Division, personal communication). A yearly offtake of 781 individuals represents an annual taking of about 0.3 per cent of the roughly evaluated 250,000 buffalo population in Tanzania (see Chapter 4 for actual best estimates).

Southern Africa

In Southern Africa, there are six countries with legal hunting tourism, and the Cape buffalo is a major game. In Botswana, buffalo hunting was resumed in 2020. Hunting is organized in registered Hunting Areas covering 75,000 km² (13 per cent of the country) for an annual revenue of €40 million in 2012 (Di Minin et al., Reference Di Minin, Leader-Williams and Bradshaw2016). In Zambia, buffalo is a major game species for 36 hunting concessions within Game Management Areas covering 170,000 km² (23.6 per cent of Zambia; Snyman et al., Reference Snyman, Sumba and Vorhies2021). In Zimbabwe, hunting is undertaken in 78,000 km² (20 per cent of the country) and generated €24.4 million revenue in 2015 (Chitauro, 2016 in Snyman et al., Reference Snyman, Sumba and Vorhies2021). Buffalo is an important game outside the central plateau in both State land and in the 10 CAMPFIRE communal areas. In Mozambique, buffalo is also a major game species for the various categories of Hunting Areas (Coutadas, Fazendas do bravio, etc.) covering 135,000 km² (17 per cent of the country) (Di Minin et al., Reference Di Minin, Leader-Williams and Bradshaw2016). In Namibia, buffalo hunting is restricted to the Caprivi strip because existing veterinary policies prevent the reintroduction of buffalo, although it is a key species for tourism and safari hunting (Lindsey et al., Reference Lindsey, Havemann and Lines2013). Hunting is a major driver of the wildlife-based tourism in Namibia, with €26.6 million direct revenue in 2016 (Snyman et al., Reference Snyman, Sumba and Vorhies2021) over 287,000 km² (Lindsey et al., Reference Lindsey, Havemann and Lines2013). Hunting is undertaken in two land categories: (i) communal conservancies (86 of them in 2021 cover 166,000 km², i.e. 20.2 per cent of Namibia), which collect 100 per cent of the hunting fees (€2.3 million in 2018) in their 48 hunting concessions (Snyman et al., Reference Snyman, Sumba and Vorhies2021); and (ii) private game ranches (so-called ‘freehold lands’), which contain 21–33 times more wildlife than Protected Areas (Snyman et al., Reference Snyman, Sumba and Vorhies2021).

South Africa has the largest African hunting industry in terms of numbers of operators, visiting hunters, animal collected, and revenues generated (Lindsey et al., Reference Lindsey, Roulet and Romañach2007). South Africa also hosts the highest number of buffalo in southern Africa, yet with a peculiar situation that contrasts sharply with the rest of the continent: there are no free-ranging buffalo in this country, all of them being enclosed, so that buffalo are always hunted behind fences (Chapter 13). Hunting Areas there are hence considerably smaller in size than anywhere else in Africa, largely due to the requirement for fencing (Taylor et al., Reference Taylor, Lindsey and Nicholson2020). The average size of a game ranch is slightly less than 3000 ha (Cloete et al., Reference Cloete, Van der Merwe and Saayman2015), that is in the order of between 10 and 100 times smaller than Hunting Areas in the rest of Africa (e.g. the average size of the 17 Hunting Areas of Niassa Special Reserve in Mozambique is 2486 km²). Overall, Hunting Areas cover 150,000 km² in South Africa, that is 12 per cent of the country (Snyman et al., Reference Snyman, Sumba and Vorhies2021). Since the Game Theft Act of 1991, properly fenced wildlife in South Africa is the property of the landowner, a situation almost non-existent in most other African countries. This ownership of wildlife allowed the private sector to develop a dynamic wildlife industry providing substantial benefits to local and national economies (Snyman et al., Reference Snyman, Sumba and Vorhies2021). For half of the nearly 10,000 game ranches, hunting is a source of income, and for 30 per cent of them hunting is the main source of income (Nel, Reference Nel2021).

Buffalo in South Africa is a typical example of a high-value species producing high income from a very low percentage of the population harvested. It does not appear on the list of the 10 most hunted game species in South Africa (NWU, 2017 in Snyman et al., Reference Snyman, Sumba and Vorhies2021), yet it is the top income-earning species with €13.2 million generated in 2016 and €9.2 million in 2019 (South African Professional Hunters statistics, 2019), well ahead of the second high-value game species, sable (Hippotragus niger).

Since the amendment in 2019 of the Animal Improvement Act of 1998, buffalo are legally subject to selection programmes for enlarging and reshaping their horns in order to raise their commercial value for live sales and hunting trophies (e.g. the first 50-inch-wide trophy live bull in South Africa was auctioned at an all-time record for buffalo of €10.5 million). The selection methods combine (i) extreme inbreeding among the most desired individuals and (ii) outbreeding with East African buffalo, which have greater horn spread than South African buffalo. Whether this development is a matter of manipulated genetic engineering or the restoration of historic natural genetic integrity is an issue of tense debate, including in the international arena (IUCN SSC Antelope Specialist Group, 2015; IUCN WCC, 2016). There is considerable concern about the negative genetic consequences of intensive selective breeding of wildlife, as well as about the image and tourism economy of South Africa (e.g. Selier et al., Reference Selier, Nel and Rushworth2018; Russo et al., Reference Russo, Hoban and Bloomer2019; Somers et al., Reference Somers, Walters and Measey2020). Game ranching in South Africa is certainly a success story in many ways (socioeconomic, rewilding, recovery of endangered species, etc.; Chapter 13), for example there are roughly three times more wildlife in private game ranches than in the National Parks (Kitshoff-Botha, Reference Kitshoff-Botha2020). The sustainable-use approach of wildlife ranching has furthermore proved to be a legitimate way to conserve biodiversity, and one that may even be advisable for other African countries to be considered (Taylor et al., Reference Taylor, Lindsey and Nicholson2020). However, a great many stakeholders and observers disapprove of the creation of so-called ‘superior’ bigger trophy animals, as well as of introducing exotic taxa and canned or put-and-take hunting (Snyman et al., Reference Snyman, Sumba and Vorhies2021).

Administration and Management of Buffalo Hunting
Legal Framework at a Glance
International Scene

The Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) does not list the African buffalo in any of its Appendices of protected animals (CITES, 2022). No CITES Party has passed stricter domestic measures for the African buffalo to date. For example, the European Union does not list this species in the Annexes of the EU Wildlife Trade Regulations (European Commission, 2010), and the USA do not include this species in the list of foreign species of its Endangered Species Act (ESA) (US Fish and Wildlife Service, 2022). Therefore, international trade of buffalo and their parts including trophies is not subject to specific controls beyond general custom, wildlife and veterinary regulations. In 2022, the African buffalo is listed in the ‘Near Threatened’ Category of the IUCN Red List, the second lowest category on the risk scale (IUCN SSC Antelope Specialist Group, 2019). Thus far, the Red List does not distinguish between buffalo subspecies, a matter for discussion as the conservation status of each subspecies is evolving differently (Chapter 4).

National Settings

Each buffalo range country has established its own environmental legislation with an array of laws and regulations to protect and manage biodiversity. All of the countries that allow the legal hunting of buffalo have set their respective permit systems with precise rules, so that hunting buffalo without the proper licences is taken as poaching and subject to penalties. In most countries, the rules, taxes and fees are different between citizens, resident expatriates and foreigners. The cost to hunt a buffalo is much higher for foreign hunting tourists than for citizens. Costs consist of government levies, payments for the services of safari operators and royalties or retention schemes for local communities and landowners (Hurt and Ravn, Reference Hurt, Ravn, Prins, Grootenhuis and Dolan2000). The revenues generated by buffalo hunting provide incentives for (i) the State to preserve the national network of Protected Areas, and (ii) communities and landowners to keep game on their lands and avoid landscape conversion into alternative land uses that are environmentally unfriendly.

Monitoring Buffalo Hunting

Monitoring is an essential process for the assessment of population trends in evaluating the conservation status of species at multiple scales over time. For management purposes, monitoring helps determine whether an intervention like hunting is on track to meet its objective and, if not, when, where and how changes may need to be made (Bell, Reference Bell and Owen-Smith1983, Reference Bell, Bell and McShane-Caluzi1984; CSIR, Reference Ferrar1983; Martin, Reference Martin, Bell and McShane-Caluzi1984).

Monitoring Buffalo Populations

Knowing how many animals there are in a given area at different times helps to measure the population trend. However, this is not simple, and a selection of appropriate methods and techniques (Collinson, Reference Collinson1985) is crucial, underpinned by clear objectives and a decision-making process (Caughley, Reference Caughley1977). While the aerial survey is often the method of choice in open savanna landscapes (Norton-Griffiths, Reference Norton-Griffiths and Grimsdell1978), it is not appropriate for forest or savanna–forest mosaics. However, as a herding species, buffalo are usually non-randomly distributed in clusters, which makes the count less reliable than for more evenly distributed species (Norton-Griffiths, Reference Norton-Griffiths and Grimsdell1978; Taylor and Mackie, Reference Taylor and Mackie1997). Nevertheless, the aerial survey (with photography) remains the most cost-effective approach in large savanna landscapes (1000–10,000 km² and above). Ground counts using distance sampling methods (Buckland et al., Reference Buckland, Anderson and Burnham2001), also referred to as road strip or line transect counts, are also used either on foot or in vehicles, including for community-based game counts (NACSO, 2021).

Counting buffalo in forest landscapes is much more tedious and time- and money-consuming, using either transect surveys (line, recce or strip transects) or point sampling in, for example, forest clearings. More recently, camera traps have been utilized to assess densities by using distance sampling methods (Hofmeester et al., Reference Hofmeester, Rowcliffe and Jansen2017; Howe et al., Reference Howe, Buckland, Després‐Einspenner and Kühl2017). Another method, the Pooled Local Expert Opinion (PLEO) method, is based on traditional knowledge. A number of local hunters are asked to estimate wildlife abundance in a specified area, after which densities are calculated per species, and the estimates are pooled and extrapolated for the whole area (Van der Hoeven et al., Reference Van der Hoeven, de Boer and Prins2004). Using citizen science and local communities as resource managers contribute to improving conservation monitoring (Rigava et al., Reference Rigava, Taylor and Goredema2006; Keeping et al., Reference Keeping, Burger and Keitsile2018).

Quota Setting for Hunting

The primary objective of monitoring a hunted population is to assess the demographic trend in that population in order to set hunting quotas that allow sustainable hunting. However, detecting trends on a regular basis is often fraught with the difficulty of making decisions based on inadequate and/or imprecise data (Taylor, Reference Taylor, Hulme and Murphree2001). Consequently, it is important to consider multiple sources or lines of evidence that can provide more robust data or information on the species being hunted. In addition to survey data, other indices of abundance should be used as well as the local knowledge of multiple stakeholders ultimately involved in the management and use of the species. Fortunately, the buffalo lends itself comparatively easily to this approach.

In a number of African countries, annual trophy hunting quotas are still set by the wildlife management authorities as a percentage of the total population size of the given species, for example 1–2 per cent of the buffalo population size. However, such a method appears impractical in most African conditions where population sizes are usually either unknown or imprecisely known or not updated on a yearly basis (Bell, Reference Bell, Bell and McShane-Caluzi1984). Quota-setting methods relying on wildlife censuses face serious limitations because estimating the density or population size of large herbivores with high precision and accuracy is difficult, especially over large areas, and requires considerable investment of time, people and money (Morellet et al., Reference Morellet, Gaillard and Hewison2007). In these situations, it is meaningless to attempt to set quotas on a percentage basis, and it is preferable to set quotas either (i) by specifying biological rules such as minimum trophy size or age of individuals to be taken (Morellet et al., Reference Morellet, Gaillard and Hewison2007) and/or (ii) by adjusting quotas according to participative assessments of population trends (WWF, 1997, 2000) as has been done with success for decades in several southern African countries as well as in North America and Europe.

The quota-setting method based upon trends requires the involvement of an ‘extended peer community’ consisting of those with a stake in the issue of concern (Funtowicz and Ravetz, Reference Funtowicz and Ravetz1993). This is counter to conventional wisdom, which seeks to maintain centralized control (Bell, Reference Bell, Anderson and Grove1987). Failure to integrate knowledge held among all stakeholders undermines effective resource management (Hulme and Taylor, Reference Hulme, Taylor, Lee and Kirkpatrick2000). Participatory quota setting for the harvesting of wildlife species reflects a relatively recent departure from the conventional norm, whereby local resource managers become active participants in an adaptively managed process with greater devolution of responsibility and accountability (Taylor, Reference Taylor, Hulme and Murphree2001; Rigava et al., Reference Rigava, Taylor and Goredema2006).

The Participatory Quota-Setting Process

The process should ideally bring together all of the parties involved in establishing a quota and its subsequent use. Typically, this would include wildlife authority managers and ecologists, land occupiers (farmers or resource managers), safari operators and hunters, local communities and even hunting trackers as applicable, regardless of background, education or training. Each stakeholder brings different sets of information, recognizing the importance and value of the information and its source. The use of a facilitator provides greater understanding and demystifies the process of establishing and using a quota. This information provides a set of matrices that can be triangulated. Triangulation comprises an iterative process of examining, assessing and sense-making of information, which results in a reliably informed decision being made (Greyson, Reference Greyson2018). Trend data are assembled by participants and graphically represented for each species and entered into the matrix. The current quota is assessed against the available data and information, and the proposed quota adaptively determined using the full set of indices (Table 16.3). The proposed quota can be submitted to the regulatory wildlife authority for review and approval with or without adjustment, and subsequently used by the safari operator in the coming hunting season.

Table 16.3 An example of the participatory triangulation matrix summarizing the trends in key indicators for individual species in view of proposing new hunting quota.

Species (males only)Current quota (Year N)Aerial survey groundsGround count trendsTrends from other monitoring methodsTrophy quality trendsCatch-effortIllegal activityHWC and PACOther type of infoNew quota (Year N + 1)
Greater kudu8⇐⇑⇐⇑UNK⇐⇑X6
Buffalo10⇐⇑UNK⇐⇑⇐⇑X8
Impala20UNK⇐⇑⇐⇑XX20
Other species

X, information not available or irrelevant; UNK, information unknown; HWC, human–wildlife conflict; PAC, problem animal control.

⇑, indicator shows a population increasing trend; ⇓, indicator shows a population decreasing trend; ⇐⇑, indicator shows a stable population.

Monitoring Buffalo Hunts

Hunting during the season is monitored by representatives of the stakeholders and in compliance with applicable laws and regulations as required by specific countries. Regardless of such requirements, completion of a ‘Hunt Return Form’ (HRF) is essential. This is a crucially important monitoring tool that captures key biological and economic variables associated with every individual hunt. At the end of the hunting season, the set of HRFs collected per hunting area is analysed and used at both national and hunting area levels for the establishment of further sustainable hunting quotas. Subsequent data analysis provides insights into trends in quotas, offtakes, trophy quality and assessments of ‘catch-effort’ (Grobbelaar and Muselani, Reference Grobbelaar and Masulani2003).

Using Quotas for Buffalo

Hunting quotas for buffalo are only set for adult males, ideally old ones. Neither females nor subadult males are hunted by trophy hunters. However, hunting buffalo for trophies is challenged by the fact that the size of the trophy does not well reflect the age of the individual because the horns of old buffalo tend to wear down (Grobbelaar and Muselani, Reference Grobbelaar and Masulani2003). The largest trophies are thus obtained from animals at or just above middle age, which coincides with the age at which males are breeding bulls. Males aged 5–10 years constitute the breeding cohort, a period when they wear their largest horns. Moreover, trophy males have to be replaced by maturing younger males in order to have trophies available in the next seasons. Trophy hunting will be unsustainable if inappropriate hunting practices take place that remove these younger males in their prime instead of harvesting the oldest bulls. For this reason, trends in trophy quality and age should be carefully monitored (Crosmary et al., Reference Crosmary, Loveridge and Ndaimani2013).

Trophy Quality

For most species, trophies only represent a small fraction of the older adult males in the population, mainly after their breeding time, and therefore a very small proportion of the total population. Removing this segment of the population does not impact the survival of the population because no females are hunted and only a tiny proportion of the old males are harvested as trophies. However, selection pressure on bulls actively breeding can impact on characters in a population such as horn length. Removing breeding animals with superior horns can possibly result in a decrease in such specimens in the population, and increase specimens with inferior horns (Crosmary et al., Reference Crosmary, Loveridge and Ndaimani2013). Therefore, trophy quality should be monitored per hunting area per hunting season. The trophy quality is indexed by the trophy size of hunted individuals.

The Rowland Ward (RW) system of measurement, founded in 1870, has been the traditional method for measuring hunting trophies, for example 30th Edition for Africa in 2020 of Rowland Ward’s Records of Big Game (Rowland Ward, 2020). In 1977, North American trophy hunters introduced the Safari Club International (SCI) Record Book of Trophy Animals (SCI, 2022) with a measurement system built upon the original RW system, but nonetheless quite different. For buffalo, the RW system measures the greatest outside spread of the horns, which is not affected by the wear of the horns (RW method 12-a for Cape, Central African and Nile buffalo, rowlandward.org; Figure 16.3). Note that RW uses a different method (12-b) for West African and Dwarf buffalo. The SCI system measures the so-called ‘tip to tip length of the horns’ following the curves all along both horns, which is obviously much affected by the horns’ wear (SCI method 4 for African buffalo, safariclub.org). Thus, by penalizing worn horns, the SCI system encourages hunters to hunt younger breeding bulls with longer tip-to-tip lengths (Grobbelaar and Muselani, Reference Grobbelaar and Masulani2003; Taylor, Reference Taylor2005). Using Taylor’s (Reference Taylor1988) predictive tooth wear and age relationship, and relating this to trophy score with both RW and SCI systems (Taylor, Reference Taylor2005), it is clear that the SCI scoring system favours the attributes of younger individuals and leads to rates of offtake that are too high for sustaining trophy quality. The Namibian Professional Hunters Association is considering adopting an Age-Related Measuring System that scores according to age in addition to other criteria, and where immature animals are disqualified (NAPHA, 2021).

Figure 16.3 Method for measuring the trophies of Cape, Central African and Nile buffalo according to Rowland Ward’s Records of Big Game, Rowland-Ward-Method-12-a-Cape.pdf (rowlandward.org).

Illustration reproduced from © RowlandWard.org with permission.

While determining the age of individual hunted animals provides an additional refinement to monitoring, it can also be considered as a further imposition on safari operators, professional hunters and their hunters. However, where there may be concern over sustainability and possible diminishing trophy size, the measurement of the first molar tooth for age determination of hunted buffalo (Taylor, Reference Taylor1988) should be implemented as part of good adaptive management. This will necessitate the proper collection, labelling and storing of lower jaws (mandible).

Overall and simply, when hunting a male trophy buffalo, ideally:

  1. (i) do not hunt buffalo males in herds; rather, hunt males in bachelor groups or individually,

  2. (ii) think RW not SCI when selecting the individual to hunt,

  3. (iii) select the oldest of the old males; however, if none of the bulls is old enough refrain from hunting,

  4. (iv) post-hunt measure trophy using RW should be mandatory and SCI optional,

  5. (v) hunter/hunting guide/hunting operator must determine age of hunted buffalo by extracting the first permanent molar and measuring tooth cusp height.

Strengths and Weaknesses of Buffalo Trophy Hunting
Buffalo Hunting, Conservation and Livelihood

According to IUCN (2016), legal, well-regulated trophy hunting can, and does, play an important role in delivering benefits for both wildlife conservation and for the livelihoods and wellbeing of indigenous and local communities living with wildlife.

Hunting Areas More Than Double the Land Area Dedicated to Wildlife Conservation

Buffalo hunting tourism is conducted in officially gazetted Hunting Areas proclaimed as such by the law of each country. Hunting Areas are recognized by IUCN as Protected Areas under both IUCN Categories IV and VI. They contribute to the national networks of Protected Areas covering the percentage of a country’s surface internationally declared as set aside by the country as Protected Areas. In sub-Saharan Africa, Hunting Areas cover a minimum area of 1,394,000 km², exceeding the area encompassed by National Parks (Lindsey et al., Reference Lindsey, Roulet and Romañach2007). This means that financial incentives from trophy hunting effectively more than double the land area that is used for wildlife conservation, relative to the area that would be conserved by national parks alone (Lindsey et al., Reference Lindsey, Roulet and Romañach2007). Hence, trophy hunting sustains these immense wilderness areas acting as biodiversity reservoirs, carbon sinks and ecosystem service providers.

The large proportion of Hunting Areas that neighbour National Parks act as buffer zones amortizing the human pressure from outside. Many Hunting Areas are also the last ecological corridors linking National Parks that otherwise would become conservation islands in a human landscape devoid of wildlife. In the final analysis, Hunting Areas are the ‘last frontier’ of buffalo and large wildlife outside National Parks. Typical examples are two buffalo strongholds: the three National Parks (W, Arly, Pendjari) of the transboundary WAP complex (Benin, Burkina Faso, Niger) in West Africa, and the three National Parks (Faro, Bénoué, Bouba Ndjidda) of northern Cameroon in Central Africa. These National Parks are all embedded in Hunting Areas that also link the parks together with no discontinuity.

In South Africa and Zimbabwe, trophy hunting has been the entry point for the conversion of thousands of livestock ranches to wildlife ranches with the reintroduction of locally extinct species like buffalo and the subsequent multiplication of wildlife populations (Bond et al., Reference Bond, Child, de la Harpe and Child2004; Leader-Williams et al., Reference Leader-Williams, Hutton, Woodroffe, Thirgood and 478Rabinowitz2005). Similarly, trophy hunting was the initial driver for local communities to establish the CAMPFIRE programme in Zimbabwe, Community Conservancies in Namibia, Wildlife Management Areas in Tanzania, and Village Hunting Zones in CAR, etc. where wildlife often are more abundant than in neighbouring National Parks. In Mozambique, trophy hunting played an important role in facilitating the recovery of wildlife populations in Hunting Areas after the war (Lindsey et al., Reference Lindsey, Alexander and Franket2006) by permitting income generation from wildlife without jeopardizing wildlife population growth (Bond et al., Reference Bond, Child, de la Harpe and Child2004). Buffalo in particular is making a remarkable comeback in this country, with Hunting Areas within Niassa Special Reserve and Marromeu complex as sources of founders for reintroducing locally extinct or depleted buffalo populations in National Parks like Gilé and Zinave (Chardonnet et al., Reference Chardonnet, Fusari and Dias2017; Fusari et al., Reference Fusari, Lopes Pereira and Dias2017; Macandza et al., Reference Macandza, Bento and Roberto2017). Trophy hunting may be a viable alternative for Protected Area-based wildlife conservation in countries or areas where National Parks fail to protect their wildlife (e.g. Western et al., Reference Western, Russell and Cuthill2009), in regions of political instability, in remote wilderness areas, or where wildlife densities are low (Lindsey et al., Reference Lindsey, Alexander and Franket2006).

Conservation Funding from Buffalo Hunting

Not only are Hunting Areas the only Protected Areas that cost nothing to the State, they also provide funds to the State through leasing taxes, hunting taxes, income taxes, etc. that sustain wildlife administrations and, in several countries, even represent the main source of income for the wildlife administration. In Tanzania, while the funding of TANAPA (Tanzania National Parks, in charge of wildlife within National Parks) mainly comes from park entry fees, 80 per cent of the funding of TAWA (Tanzania Wildlife Management Authority, responsible for wildlife all over the country outside the jurisdiction of TANAPA) comes from hunting tourism (TAWA, 2019). Buffalo is the top tax-earning game in this country (TAWA, 2019), making it crucial for TAWA to maintain all of the Protected Areas other than National Parks in a country where 68 per cent of the Protected Areas rely on income from trophy hunting (Lindsey et al., Reference Lindsey, Allan and Brehony2020). In South Africa, becaue buffalo is the top income-earning game species for the hunting tourism sector (DEA, 2016; South African Professional Hunters statistics, 2019), it is a pillar sustaining the privately owned wildlife conservation areas. In this country, trophy hunting contributed more than €341 million and supported more than 17,000 employment opportunities in 2015/2016 (Saayman et al., Reference Saayman, van der Merwe and Saayman2018). In Zimbabwe, 80 per cent of the budget of the Zimbabwe Parks and Wildlife Management Authority comes from tourism, including trophy hunting (Lindsey et al., Reference Lindsey, Allan and Brehony2020). In Benin, in 2018, the W National Park ecosystem earned 33 times more money from hunting tourism in the neighbouring Hunting Areas (which provide income to the State) than from photographic tourism within the National Park (which costs the State) with 25 times fewer hunting tourists (19) than photographic tourists (476) (PNW, 2019). In South Africa, in 2013, each foreign leisure hunter spent about €8250, that is about 14 times more than that spent by the average foreign tourist arriving by plane (Oberem and Oberem, Reference Oberem and Oberem2016). According to Hurt and Ravn (Reference Hurt, Ravn, Prins, Grootenhuis and Dolan2000), safari hunting produces an income per hectare some seven times higher than that from cattle or game ranching and from far fewer animals harvested. They also reckon that wildlife-viewing tourism can generate even higher returns, but only in areas that are scenic and have very high concentrations of wildlife, and from massive numbers of tourists (Earnshaw and Emerton, Reference Earnshaw, Emerton, Prins, Grootenhuis and Dolan2000). Lindsey et al. (Reference Lindsey, Balme, Booth and Midlane2012) hold a different view, observing that net returns from livestock in semi-arid African rangelands ($10–30/km2/year in areas with 400–800 ml of annual rainfall according to Norton-Griffiths Reference Norton-Griffiths2008) are similar to those from trophy hunting in some areas ($24–164/km2). However, they conclude that maximizing returns from hunting is key to ensuring the competitiveness of wildlife-based land uses.

Some critiques of the socioeconomic effects of trophy hunting suggest that its contributions to country-level gross domestic product (GDP) are small relative to non-hunting wildlife tourism (Ghasemi, Reference Ghasemi2021). ’t Sas-Rolfes et al. (Reference ’t Sas-Rolfes, Emslie, Adcock and Knight2022) disagree, arguing that the claim is misleading because national GDP contributions are a poor indicator in terms of both broader socioeconomic relevance and appropriate scale of analysis: (i) GDP metrics fail to consider essential ecosystems services and natural capital (Costanza et al., Reference Costanza, d’Arge and de Groo1997) and (ii) nation states are an arbitrary level at which to make such assessments. More relevant are the global benefits of effective species conservation and ecosystem services provided by intact habitats, functionally populated with large game, and the more localized benefits that flow to specific rural landowners and communities, who are thereby incentivized to actively support conservation (’t Sas-Rolfes et al., Reference ’t Sas-Rolfes, Emslie, Adcock and Knight2022).

Overall, hunting tourism drives a virtuous chain with financial flows of hard currency originating from developed countries (tourist-emitting countries) and directed to developing countries (tourist-receiving countries), from wealthy individuals to poorer people, and supporting vast conservation areas and local communities, as well as providing States of the South with revenues from their renewable natural resources.

Buffalo Hunting Sustaining Livelihood

When sustainable, consumptive utilization of wildlife can promote conservation beyond the borders of National Parks while at the same time generating revenue for local communities (Crosmary et al., Reference Crosmary, Côté and Fritz2015a). Where properly managed, trophy hunting can provide income for impoverished and often landlocked rural communities (IUCN, 2016), that is royalties, employment, venison, community infrastructures, social services, etc. Namibia is one of the best examples in this regard, well ahead of many other countries. Trophy hunting finances the budgets of 82 communal conservancies, which cover ~20 per cent of the country (162,000 km²) and encompass ~189,000 community members, or 9 per cent of Namibia’s population (Naidoo et al., Reference Naidoo, Weaver and Diggle2016). However, a number of other countries or areas are not as successful for various reasons, for example when the benefits from hunting are captured by local elites (Leader-Williams et al., Reference Leader-Williams, Baldus, Smith, Dickson, Hutton and Adams2009) or when the benefits are substantial at the community level but too small at the household level. In northern Cameroon, Mayaka et al. (Reference Mayaka, Hendricks, Wesseler and Prins2005) proposed a series of recommendations to improve the benefits of wildlife harvesting, notably by increasing the return to local communities for resource custodianship. In the same area, Akito Yasuda (Reference Yasuda2011) pointed out that while sport hunting certainly generates tax revenues and provides profit sharing and employment opportunities to local communities, the latter two are too limited and inequitably distributed in the community. Similarly, in south-eastern Zimbabwe, Poshiwa et al. (Reference Poshiwa, Groeneveld, Heitkönig, Prins and van Ierland2013) described the benefits of wildlife tourism but emphasized their limited magnitude. Because high levels of poverty (Matseketsa et al., Reference Matseketsa, Krüger and Gandiwa2022) and poor governance, such as the leakage of hunting revenues for communities (Burn et al., Reference Burn, Underwood and Blanc2011), are powerful drivers to poaching by local communities, the allocation of sufficient benefits of Hunting Areas to communities is an absolute critical factor for a successful deal between the local community (living on the land), the State (owning the land) and the hunting operator (protecting and valorizing the land).

Access to natural resources is also important for the livelihoods of local communities. In northern Cameroon, populations complain that locals’ rights over natural resource use are regulated (Akito Yasuda, Reference Yasuda2011). However, while National Parks are strict exclusion areas for local communities, most Hunting Areas are less stringent and allow for some activities by local communities, such as harvesting firewood and non-timber forest products.

Finally, concerns about the negative cultural and environmental impacts of tourism are growing with mass wildlife tourism in Africa (Spenceley, Reference Spenceley2005; Lindsey et al., Reference Lindsey, Roulet and Romañach2007), for example in the Okavango Delta, Botswana (Mbaiwa, Reference Mbaiwa2003). However, due to their very small number, the impacts of hunting clients, such as habitat conversion for infrastructure development and all sorts of pollution, are considerably lower compared to mass tourism.

Threats to Buffalo Hunting
Ill-Managed Hunting Undermines Well-Managed Hunting

There have been and there are cases of hunting poorly conducted by some hunting operators and of hunting sectors poorly regulated by some wildlife administrations (IUCN, 2016). A variety of problems may hamper the proper functioning of the hunting tourism industry and undermine the conservation role of sustainable wildlife utilization, for example depending on countries, poor governance of the hunting sector (Burn et al., Reference Burn, Underwood and Blanc2011), lack of professionalism in the administration and control of the hunting activity (Booth and Chardonnet, Reference Booth and Chardonnet2015) and risk of corruption (Leader-Williams et al., Reference Leader-Williams, Baldus, Smith, Dickson, Hutton and Adams2009). We concur with Lindsey et al. (Reference Lindsey, Roulet and Romañach2007) that the inequitable distribution of hunting revenues represents the most serious threat to the long-term sustainability of the industry. In some countries, there is insufficient sharing of hunting taxes by government administrations reluctant to decentralize and empower communities. Too often, benefits are centralized into the hands of elites or captured by local rulers so that promises from trophy hunting fail to materialize at the grassroots level (Nelson et al., Reference Nelson, Nshala and Rodgers2007). In a number of situations, the management of Hunting Areas certainly needs to be improved. One failure, for example, is the reduction of anti-poaching activity outside the hunting season. Another is the lack of proper monitoring by hunting operators, which weakens their credibility and constraints the sustainability of the activity (Selier and Di Minin, Reference Selier and Di Minin2015). Nevertheless, all of these problems are far from being specific to the hunting industry, they are also fully shared by other industries, including photographic ecotourism (Christie and Crompton, Reference Christie and Crompton2001; Walpole and Thouless, Reference Walpole, Thouless, Woodroffe, Thirgood and Rabinowitz2005). Finally, poorly managed trophy hunting can cause local wild population declines (Packer et al., Reference Packer, Brink and Kissui2011). However, in the case of buffalo, no example is known of a buffalo population driven to extinction by hunting tourism, while poaching is well recognized as being responsible for many local extinctions across the buffalo’s range.

Hunters Their Worst Enemies?

While the hunting community is certainly skilled, with a great deal of field experience and knowledge of the bush, members rarely produce or publish peer-reviewed scientific articles which nevertheless largely make the basis of conservation politics. Moreover, a number of hunting professionals tend to be reluctant to seek the collaboration of scientists. As a result, reliable standardized data on the hunting sector are certainly missing (Lindsey et al., Reference Lindsey, Roulet and Romañach2007; Snyman et al., Reference Snyman, Sumba and Vorhies2021). This situation appears detrimental to the hunting industry at a time it badly needs more science in all sorts of domains, for example biological, socioeconomic, management. In Western Zimbabwe, Crosmary et al. (Reference Crosmary, Côté and Fritz2015b) showed that harvested populations of large herbivores in trophy Hunting Areas may perform as well, and sometimes even better, than in National Parks where trophy hunting is not authorized. However, Buckley and Mossaz (Reference Buckley and Mossaz2015) pointed out that this study represented only one example, concluding that more studies are needed to understand the benefits of hunting tourism to wildlife conservation. Crosmary et al. agree and concur with Selier and Di Minin (Reference Selier and Di Minin2015) that scientists are needed to establish long-term wildlife monitoring systems that also integrate the social and financial benefits of trophy hunting for local communities.

There is probably some kind of misunderstanding on the part of hunting stakeholders, who find it difficult to accept critics in a polemic context. However, and counterintuitively, the hunting activity holds a broad set of very strong assets in favour of conservation, not only of the hunted game, but also of non-game species and their habitats, of the entire biodiversity in fact (fauna and flora), of all ecosystem services, without even talking about the livelihoods of local communities. In other words, hunters are poor advocates of their achievements. This said, some poorly performing individuals, companies and administrations certainly jeopardize the profession, like in any profession, whether because they lack training, professionalism, ethics or something else. While these kinds of internalities probably affect all sectors, they cannot be hidden in the hunting industry.

Beyond these internalities, there are also powerful externalities that fall beyond the responsibility of the hunting community and severely affect Hunting Areas and the hunting activity. The current hunting industry inherited ancient situations that are no longer suitable today, for example Hunting Areas that are very (too?) large to take care of in view of the fast-growing human population, and which require much more funding than before for their proper management (Scholte et al., Reference Scholte, Pays, Adam and Chardonnet2021). The profession is also facing newly arising tricky situations such as increasing numbers of all sorts of new arrivals claiming to be local communities despite not being indigenous people, more pastoralists with ever larger herds of livestock replacing wildlife in Hunting Areas (e.g. Figure 16.2 in Cameroon; Bouché et al., 2012 and Aebischer et al., Reference Aebischer, Ibrahim and Hickisch2020 in CAR; Musika et al., Reference Musika, Wakibara, Ndakidemi and Treydte2021 and Musika et al., Reference Musika, Wakibara, Ndakidemi and Treydte2022 in Tanzania), illegal goldminers, wild loggers, without mentioning bandits and even terrorists. Other contemporary constraints are the intense pressure of lobbies promoting commercial crops at all costs, especially the cotton value chain, which are heavily supported by national and international agencies with hardly any exception. Overall, many externalities have appeared on the scene and reshuffle the game, making hunting work more difficult, less viable and threatening ever more the conservation of natural resources. There is definitely a need to reform the governance and administration of hunting tourism (Booth and Chardonnet, Reference Booth and Chardonnet2015), but given the above-mentioned externalities, the reform should not be considered in isolation (Leader-Williams et al., Reference Leader-Williams, Baldus, Smith, Dickson, Hutton and Adams2009).

Poaching Versus Hunting

The African buffalo does not give the impression of being a fragile animal. However, it is indeed extremely sensitive to poaching, notably because it is quite easy to stalk on foot provided you strictly approach against the wind. The buffalo shows little resilience under poaching pressure. Poaching means limitless and indiscriminate offtake of any kind of buffalo, whatever sex and age, whereas tourism hunting harvests a tiny percentage of only old bulls (Table 16.4). Legal and illegal hunting are mutually exclusive: where poaching flourishes, hunting tourism deteriorates and even fails. Just like National Parks, Hunting Areas require anti-poaching engagement to be protected and avoid wildlife depletion.

Table 16.4 Comparison between poaching and tourism hunting.

Illegal unregulated hunting (poaching)Legal regulated tourism hunting
Wildlife offtakeSpeciesAll wildlife species with value as food or trophy, e.g. ivory, claws, etc.Only a few selected game species
Number of individualsUnlimitedSmall % (approx. 1%) of the population
Sex of individualsMales and femalesOnly males (exceptions in South Africa)
Age of individualsAny age including calvesMostly old individuals, often beyond reproductive age
Impacts of the wildlife offtakeFor local communitiesMeat and other wildlife productsMeat
Livelihood but with limited income from trade of meat and trophiesLivelihood and formal employment by hunting companies
Negative impact due to overexploitation leading to depletion of the wildlife resourceAccording to countries: share of the taxes (% of leasing tax, trophy fees, etc.), royalties
Conflict with law enforcement leading to fines and prison sentencesCommunity infrastructures and services (schools, dispensaries, wells, etc.)
For illegal wildlife tradersHigh financial gainsNo business
For the Government financesNegative impact due to absence of revenue from the activityTaxes (income tax, etc.)
For the wildlife administrationNegative impact due to the cost of anti-poachingTaxes (leasing fees for hunting areas, trophy fees, hunting permits, operating licenses, etc.)
For the private sectorNegative impact due to the cost of anti-poaching and the depletion of the wildlife resourceReturn from daily fees, paid hunting services
For the national network of Protected AreasNegative impact due to the degradation of the Protected AreasHunting Areas as Protected Areas of the IUCN Cat. VI are maintained by the income of hunting tourism (ecotourism rarely viable in these areas)
For animal welfareLong death and suffering for animals caught by snares, gin traps, pits or other trapping devicesInstant death in most cases
For the conservation of biodiversityNegative impact due to the degradation of the wildlife conservation status leading to loss of biodiversityImproved conservation status of (i) the few income-generating game species, and (ii) all the non-game species of fauna and flora
Hunting Bans and the Future of Buffalo

One of the biggest challenges facing the hunting industry is the prescriptive unilateral decision by Western countries to ban imports of hunting trophies from Africa (Ares, Reference Ares2019), which could have a long-lasting negative impact on many economies, and in turn on conservation, in Africa (Snyman et al., Reference Snyman, Sumba and Vorhies2021). For local communities in northern Botswana, the safari hunting ban of 2014 led to a reduction of tourism benefits to local communities, for example income, employment opportunities, social services and scholarships. This led to the development of negative attitudes by community-based organizations of rural residents towards wildlife conservation and to an increase in incidents of poaching (Mbaiwa, Reference Mbaiwa2018; Blackie, Reference Blackie2019; Strong and Silva, Reference Strong and Silva2020). For game ranchers and other owners of private conservation areas in South Africa, most believe that the economic viability of their enterprises, biodiversity conservation and the livelihoods of owners and employees would be lost following a hunting ban (Parker et al., Reference Parker, De Vos and Clements2020). Without hunting activity, most Hunting Areas would no longer protect buffalo, which means that the persistence of buffalo outside of National Parks would be short-lived, as experienced in CAR after the 2012 political events when buffalo was one of the first large mammals to disappear from the Hunting Areas (Matthieu Laboureur, personal communication). With the authoritarian restrictions by Western countries on imports of elephant and lion hunting trophies from Africa, many Hunting Areas were returned to the governments in Tanzania and Zambia. Without funding or surveillance, these areas are left to poaching, greatly impacting the fate of buffalo.

Hunting trophies import bans dictated by some northern countries without an alternative global conservation framework providing conservation incentives will likely reverse the gains in wildlife conservation and rural development in some southern countries where sustainable utilization is an integral part of the wildlife conservation practice (e.g. Di Minin et al., Reference Di Minin, Leader-Williams and Bradshaw2016; Dickman et al., Reference Dickman, Cooney and Johnson2019; Nyamayedenga et al., Reference Nyamayedenga, Mashapa, Chateya and Gandiwa2021). Where trophy hunting is planned to end, alternatives should be implemented to avoid land conversion and biodiversity loss in Hunting Areas (Di Minin et al., Reference Di Minin, MacMillan and Goodman2013). However, most of these areas appear unsuitable for alternative wildlife-based land uses such as photographic ecotourism because of, for example, difficult and expensive access, absence of infrastructure, lack of attractive scenery and of high densities of viewable wildlife (Wilkie and Carpenter, Reference Wilkie and Carpenter1999; Lindsey et al., Reference Lindsey, Alexander and Franket2006; Winterbach et al., Reference Winterbach, Whitesell and Somers2015). IUCN (2016) states that unless better land-use alternatives exist, hunting reforms should be prioritized over bans, while such reforms have proved effective (Booth and Chardonnet, Reference Booth and Chardonnet2015; Begg et al., Reference Begg, Miller and Begg2018).

Surprisingly, bans and restrictions on importing hunting trophies of game species listed on CITES Appendices diverted the attention of the hunting industry to buffalo, a non-CITES-listed species. While becoming a new focus, the buffalo has either reinforced or taken the lead as a flagship game in an attempt to compensate the loss of CITES-listed game, even though it does not attract as much income. Buffalo hunting does not draw much public awareness, in contrast with the hunting of the four other representatives of the Big Five, a bit like the wild boar in Europe compared to red deer or chamois. Therefore, the less-charismatic member of the Big Five is now gaining more importance for sustaining Hunting Areas and for wildlife conservation outside National Parks. In other words, from a commodity game, buffalo is turning out to be a high-value game species.

In 2021, Van Houdt et al. surveyed international networks to investigate the divergent views on trophy hunting in Africa. Unlike European respondents, African respondents showed significantly more support for trophy hunting and, unlike North Americans, African respondents supported external subsidies of wildlife areas presently funded by hunting. Oddly, while Europeans and North Americans carry out trophy hunting in their own countries, they tend to oppose it in African countries. The inquiry concluded that policies on African hunting should better integrate African perspectives, in particular those of rural communities (Van Houdt et al., Reference Van Houdt, Brown and Wanger2021). While opponents to hunting tourism in Africa often qualify this activity as a colonial relic, it cannot be denied that most Protected Areas have deep roots in the colonial period, either National Parks for wildlife viewing tourism or Hunting Areas for hunting tourism, ‘but that makes it even more important that today, the decisions-making and rights of African countries and communities are respected; Westerners must not continue to externally impose their own ideals upon Africans, such as pushing trophy hunting bans and restrictions’ (Dickman et al., Reference Dickman, Child, Hart and Semcer2021). A group of African countries called for a ‘New Deal’ for rural communities (Southern Africa Trust, 2019) that allows them to achieve the self-determination to sustainably manage wildlife and reduce poverty. Dickman et al. (Reference Dickman, Cooney and Johnson2019) stated that it is incumbent on the international community not to undermine that. More recently, in response to the call of a UK parliamentary committee in 2022 for ending trophy hunting in Africa (but not in the UK), the Community Leaders Network of Southern Africa responded: ‘It’s a form of colonialism to tell us Africans what to do with our wildlife’ (Louis, Reference Louis2022).

Footnotes

13 African Buffalo Production Systems

* Of the world population.

14 Management Aspects of the Captive-Bred African Buffalo (Syncerus caffer) in South Africa

* Calculated metabolizable energy.

# Calculated large stock, grazing and browsing units.

15 Handling and Moving the African Buffalo

16 Buffalo Hunting: From a Commodity to a High-Value Game Species

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Figure 0

Figure 13.1 Various categories of African buffalo production systems.

Adapted from Chardonnet, 2011; background picture: © Christophe Morio.
Figure 1

Figure 13.2 Flow of direct and indirect benefits from CAMPFIRE programmes (Tchakatumba et al., 2019). *RDC refers to Rural District Council.

Source: with permission of Taylor & Francis.
Figure 2

Figure 13.3 Auction prices of live breeding buffalo bulls over time and illustrating the value initially placed by purchasers on buffalo of East African origin for reasons discussed in the section dealing with production of buffalo with large horn size, below. East African buffalo, formerly recognized as a subspecies, is phenotypically 12 per cent larger in body size, 10–20 cm higher shoulder height, with greater horn spread, lesser curve-drop and smaller bosses, than the southern African buffalo. East African buffalo was introduced into the South African production systems adding specific value market traits.

© Deon Furstenburg.
Figure 3

Figure 13.4 Horizon, the most expensive African buffalo bull ever bred so far, was sold at an auction for €10.8 million. Horizon was bred by Jacques and Caroline Malan of Lumarie. According to the SCI method (following the external curve of the horns, in inches), he measures an impressive 55 6/8".

© Nyumbu Game.
Figure 4

Figure 13.5 Aerial view of a 460-ha intensified multi-camp buffalo production system in savanna habitat with centred pens for handling, supplement feeding and rotation of stocking between camps. Optimal habitat management entitles (i) a 2-camp system per buffalo herd and rotated every 8 months, or (ii) a 3-camp system per herd and rotated every 4 months.

(Furstenburg, 2017a)
Figure 5

Figure 13.6 Example outlay of a semi-extensive buffalo production camp system (2-camps, on average 230 ha each, per breeding herd, including two free-roaming areas >4,000 ha each for surplus animals) constructed per vegetation survey map in arid Kalahari savanna habitat.

(Furstenburg, 2017b)
Figure 6

Table 13.1 Percentage of various species, some endangered, on private land owned by private game ranches versus those on state reserves in South Africa (Nel, 2021; Furstenburg et al., 2022).

Figure 7

Table 13.2 Numbers and disease status of buffalo in South Africa; bTB = tuberculosis; CA = brucellosis; FMD = foot and mouth disease (personal research of P.T. Oberem).

Figure 8

Table 13.3 Income from various economic activity pillars on game ranches in South Africa (Nel, 2021) (€1 = ZAR16.31).

Figure 9

Table 13.4 Top 10 income generators (€1 = ZAR16.31) (North-West University, 2017).

Figure 10

Figure 13.7 Buffalo in boma.

© Q. Strauss – MLP Media.
Figure 11

Figure 13.8 Buffalo in boma.

© J. Malan.
Figure 12

Figure 13.9 Average horn length in Cape buffalo (Syncerus caffer caffer) in South Africa (n = 777, 22 per cent) and in the ‘Rest of Africa’ (RoA), a variable composed of data from 11 countries from eastern and southern Africa: Angola, n = 19; Botswana, n = 35; Kenya, n = 89; Mozambique, n = 100; Namibia, n = 16; Rwanda, n = 3; Tanzania, n = 857; Uganda, n = 4; Zambia, n = 482; Zimbabwe, n = 811. All buffalo were hunted for trophy hunting in South Africa when buffalo in other countries may have been hunted for other reasons.

Graph drawn from data published by Safari Club International (2022).
Figure 13

Table 14.1 Calculated large stock unit equivalents and metabolizable energy values for different physiological stages of African buffalo.

Adapted from Shepstone et al. (2022).
Figure 14

Figure 14.1 How the season affects the quality of the grazing and animals’ body condition in southern Africa. Considering the months of January through to December (a full year), each month has a shade of green (rainy season), or yellow to orange (dry season). The green blocks correspond with the rumen and plant (in green) and the green arrow below it, portraying the time of the year when the selected feed gets degraded and digested in less than 24 h. The yellow to orange blocks correspond with the rumen and plant (in yellow/orange) and the orange arrow below it, portraying the time of the year when the selected takes longer than 24 h to be degraded and digested.

Source: Craig Shepstone.
Figure 15

Figure 14.2 Feed bowl placement in a camp and movement suggestions. The circles represent bowls, and the figures refer to a column of bowls that need to be moved together periodically, from left to right as shown in the diagram. Spaces between bowls should be 2.5 the animal’s length.

Source: Craig Shepstone.
Figure 16

Figure 14.3 Standard operating system for the major logistical operational points to be considered in a typical buffalo culling scheme, as demonstrated in Kruger National Park (KNP, South Africa). LTL = longissimus thoracis et lumborum; BF = biceps femoris; SM = semimembranosus muscles; WPS = Wildlife Processing Structure.

Source: Louwrens Hoffman.
Figure 17

Table 14.2 Suggested grading of African savanna buffalo (Syncerus caffer caffer) carcasses.

Figure 18

Figure 14.4 Primal cuts of African buffalo carcasses for further processing and marketing.

Source: Tersia Needham.
Figure 19

Table 14.3 Block test conducted on an African savanna buffalo (carcass weight of 277 kg).

Figure 20

Table 15.1 A few examples of capture and translocation operations of African buffalo

Source: Author.
Figure 21

Figure 15.1(a) Diagram of a temporary mass capture boma constructed out of woven plastic sheeting.

Figure 22

Figure 15.1(b) In practice, the boma is camouflaged in the vegetation and the narrow ‘crush’ section curves towards a ramp into transport vehicles on an exit road (transport vehicle on the top).

Source: Author.
Figure 23

Figure 15.2(a) buffalo are chased into the boma by the helicopter;

Figure 24

Figure 15.2(b) then pushed into desired sections of the boma or lorry using an adapted vehicle.

© Philippe Chardonnet.
Figure 25

Figure 15.3 Individual darting on foot of a West African savanna buffalo.

© Daniel Cornélis.
Figure 26

Figure 15.4 Chemical capture of a free-ranging herd of Cape buffalo by individual darting from a helicopter.

© Samy Julliand.
Figure 27

Table 15.2 Drug recommendations for the African buffalo (taken with the kind permission of Kock and Burroughs, 2021).

Figure 28

Table 16.1 Uses (either legal or illegal) of African buffalo by local communities: examples across the species range by region, in West, Central, East and Southern Africa (based on data/sources in the table).

Figure 29

Table 16.1 (cont.) – Part A

Figure 30

Table 16.1 (cont.) – Part B

Figure 31

Table 16.1 (cont.) – Part C

Figure 32

Table 16.1 (cont.) – Part D

Figure 33

Table 16.1 (cont.) – Part E

Figure 34

Table 16.1 (cont.) – Part F

Figure 35

Figure 16.1 Buffalo range countries where hunting tourism is lawful in 2022 for the four subspecies of buffalo recognized by the IUCN Red List so far. Note: Buffalo in northern and central Angola were categorized as ‘Cape buffalo’ by IUCN (2019), but phenotypically and perhaps even genetically they are ‘forest buffalo’.

Source: Author.
Figure 36

Table 16.2 Buffalo hunting quotas and offtakes in selected countries throughout Africa.

Figure 37

Figure 16.2 Livestock sightings in the BSB landscape covering the transboundary national parks of Bouba Ndjidda (Cameroon) and Sena Oura (Chad) as well as the seven neighbouring Hunting Areas (Cameroon), during the aerial wildlife survey in 2018 (total surface of about 10,500 km²). The estimated livestock population (117,134 heads) was six times higher than the estimated population of the 11 largest wild mammals (20,136 individuals), and located mostly within the Hunting Areas surrounding the National Parks.

(data and illustration reproduced from WCS and MINFOF, 2018, with permission)
Figure 38

Table 16.3 An example of the participatory triangulation matrix summarizing the trends in key indicators for individual species in view of proposing new hunting quota.

Figure 39

Figure 16.3 Method for measuring the trophies of Cape, Central African and Nile buffalo according to Rowland Ward’s Records of Big Game, Rowland-Ward-Method-12-a-Cape.pdf (rowlandward.org).

Illustration reproduced from © RowlandWard.org with permission.
Figure 40

Table 16.4 Comparison between poaching and tourism hunting.

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  • Management
  • Edited by Alexandre Caron, Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD), France, Daniel Cornélis, Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD) and Foundation François Sommer, France, Philippe Chardonnet, International Union for Conservation of Nature (IUCN) SSC Antelope Specialist Group, Herbert H. T. Prins, Wageningen Universiteit, The Netherlands
  • Book: Ecology and Management of the African Buffalo
  • Online publication: 09 November 2023
  • Chapter DOI: https://doi.org/10.1017/9781009006828.018
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  • Management
  • Edited by Alexandre Caron, Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD), France, Daniel Cornélis, Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD) and Foundation François Sommer, France, Philippe Chardonnet, International Union for Conservation of Nature (IUCN) SSC Antelope Specialist Group, Herbert H. T. Prins, Wageningen Universiteit, The Netherlands
  • Book: Ecology and Management of the African Buffalo
  • Online publication: 09 November 2023
  • Chapter DOI: https://doi.org/10.1017/9781009006828.018
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  • Management
  • Edited by Alexandre Caron, Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD), France, Daniel Cornélis, Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD) and Foundation François Sommer, France, Philippe Chardonnet, International Union for Conservation of Nature (IUCN) SSC Antelope Specialist Group, Herbert H. T. Prins, Wageningen Universiteit, The Netherlands
  • Book: Ecology and Management of the African Buffalo
  • Online publication: 09 November 2023
  • Chapter DOI: https://doi.org/10.1017/9781009006828.018
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