Knowns and Unknowns in African Buffalo Ecology and Management

Taxonomy The name Bos caffer was attributed by Sparrman in 1779. Since then, 92 species names have been given to the African buffalo. Taxonomists initially thought that each buffalo form represented a distinct species. Brooke (1873, 1875), who established the first classification of the African buffalo, reduced the number to three. Later, Blancou (1935) described up to 12 subspecies of buffalo. Haltenorth (1963), Ansell (1972) and Grubb (1972) summarized the first classifications of Christy (1929), Schouteden (1945) and Blancou (1935, 1954), concluding that all forms should be considered as monospecific. Although there are considerable morphological variations in body size, fur colour, horn shape and size throughout the range of distribution, the African buffalo is currently considered as a single species by various authorities (IUCN 2013; Prins & Sinclair 2013), with a subdivision into four subspecies: Cape buffalo ( S. c. caffer ), forest buffalo ( S. c. nanus ), West African savanna buffalo ( S. c. brachyceros ), and Central African savanna buffalo ( S. c. aequinoctialis ). Additionally to those four subspecies, a mountain form( S. c. mathewsi ) was also described in East Africa and may be distinct (Kingdon 1982).

Introduction: Rift Valley fever (RVF) is an arthropod-borne disease that affects both animals and humans. RVF phlebovirus (RVFPV) is widespread in Africa and Arabian Peninsula. In Mozambique, outbreaks were reported in South; seroprevalence studies performed in livestock and water buffaloes were limited to central and south regions. We evaluated the seroprevalence of RVFPV among domestic ruminants and African buffaloes from 7 of 10 provinces of Mozambique, to understand the distribution of RVFPV and provide data for further RVF control programs. Materials and methods: A total of 1581 blood samples were collected in cattle, 1117 in goats, 85 in sheep and 69 in African buffaloes, between 2013 and 2014, and the obtained sera were analyzed by ELISA. Results and discussion: The overall seroprevalence of RVFPV domestic ruminants and African buffaloes was 25.6%. The highest was observed in cattle (37.3%) and African buffaloes (30.4%), which were higher than in previous studies within Mozambique. In south and central regions, the overall seroprevalences were higher (14.9%–62.4%) than in the north. Conclusion: This study showed the presence of anti-RVFPV antibodies in animals from all sampled provinces, suggesting that RVFPV is actively circulating among domestic ruminants and African buffaloes in Mozambique, therefore surveillance should be intensified.

BackgroundBovine tuberculosis (BTB) is a zoonotic disease of global importance endemic in African buffalo (Syncerus caffer) in sub-Saharan Africa. Zoonotic tuberculosis is a disease of global importance, accounting for over 12,000 deaths annually. Cattle affected with BTB have been proposed as a model for the study of human tuberculosis, more closely resembling the localization and progression of lesions in controlled studies than murine models. If disease in African buffalo progresses similarly to experimentally infected cattle, they may serve as a model, both for human tuberculosis and cattle BTB, in a natural environment.Methodology/Principal findingsWe utilized a herd of African buffalo that were captured, fitted with radio collars, and tested for BTB twice annually during a 4-year-cohort study. At the end of the project, BTB positive buffalo were culled, and necropsies performed. Here we describe the pathologic progression of BTB over time in African buffalo, utilizing gross and histological methods. We found that BTB in buffalo follows a pattern of infection like that seen in experimental studies of cattle. BTB localizes to the lymph nodes of the respiratory tract first, beginning with the retropharyngeal and tracheobronchial lymph nodes, gradually increasing in lymph nodes affected over time. At 36 months, rate of spread to additional lymph nodes sharply increases. The lung lesions follow a similar pattern, progressing slowly, then accelerating their progression at 36 months post infection. Lastly, a genetic marker that correlated to risk of M. bovis infection in previous studies was marginally associated with BTB progression. Buffalo with at least one risk allele at this locus tended to progress faster, with more lung necrosis.Conclusions/SignificanceThe progression of disease in the African buffalo mirrors the progression found in experimental cattle models, offering insight into BTB and the interaction with its host in the context of naturally varying environments, host, and pathogen populations.

This chapter presents the buffalo genetic resources in Africa. The first section (Sect. 12.1) discusses the origin and distribution of African buffalo with emphasis on the domestic water buffalo types (swamp and river buffalo). The African water buffalo phylogeny is presented in Sect. 12.2, while the various approaches to mapping the genes of buffalo and genetic diversitycharacterization are presented in Sects. 12.3 and 12.4, respectively. The production and husbandry systems of African buffalo and genetic resources conservation are discussed in Sects. 12.5 and 12.6. This chapter concludes with a presentation of the challenges facing buffalo production and future perspectives on the development of sustainable genetic resources management for the African buffalo.

BackgroundTick-borne diseases (TBDs) are very important in relation to domestic ruminants, but their occurrence among wild ruminants, mainly in the African buffalo Syncerus caffer, remains little known.MethodsMolecular diagnostic methods were applied to detect Anaplasma marginale, Anaplasma centrale, Anaplasma phagocytophilum, Ehrlichia ruminantium and Ehrlichia chaffeensis in 97 blood samples of African buffalo captured at the Marromeu Reserve in Mozambique. Molecular detection of agents belonging to the family Anaplasmataceae were based on conventional and qPCR assays based on msp5, groEL, 16S rRNA, msp2, pCS20 and vlpt genes. Phylogenetic reconstruction of new Anaplasma isolates detected in African buffalo was evaluated based on msp5, groEL and 16S rRNA genes.ResultsAll the animals evaluated were negative for specific PCR assays for A. phagocytophilum, E. ruminantium and E. chaffeensis, but 70 animals were positive for A. marginale, showing 2.69 × 100 up to 2.00 × 105msp1β copies/μl. This result overcomes the conventional PCR for A. marginale based on msp5 gene that detected only 65 positive samples. Sequencing and phylogenetic analyses were performed for selected positive samples based on the genes msp5, groEL and 16S rRNA. Trees inferred using different methods separated the 29 msp5 sequences from buffalo in two distinct groups, assigned to A. centrale and A. marginale. The groEL sequences determined for African buffalo samples revealed to be more heterogeneous and inferred trees could not assign them to any species of Anaplasma despite being more related to A. marginale and A. centrale. The highly conserved 16S rRNA gene sequences suggested a close relationship of the new 16 sequences with A. centrale/A. marginale, A. platys and A. phagocytophilum.ConclusionsOur analysis suggests that different species of Anaplasma are simultaneously present in the African buffalo. To the best of our knowledge, this is the first study that diagnosed Anaplasma spp. in the African buffalo and inferred the taxonomic status of new isolates with different gene sequences. The small fragment of msp5 sequences revealed to be a good target for phylogenetic positioning of new Anaplasma spp. isolates.

Bovine tuberculosis (BTB) is a zoonotic disease of global importance endemic in African buffalo (Syncerus caffer) in sub-Saharan Africa. Zoonotic tuberculosis is a disease of global importance, accounting for over 12,000 deaths annually. Cattle affected with BTB have been proposed as a model for the study of human tuberculosis, more closely resembling the localization and progression of lesions in controlled studies than murine models. If disease in African buffalo progresses similarly to experimentally infected cattle, they may serve as a model, both for human tuberculosis and cattle BTB, in a natural environment. We utilized a herd of African buffalo that were captured, fitted with radio collars, and tested for BTB twice annually during a 4-year-cohort study. At the end of the project, BTB positive buffalo were culled, and necropsies performed. Here we describe the pathologic progression of BTB over time in African buffalo, utilizing gross and histological methods. We found that BTB in buffalo follows a pattern of infection like that seen in experimental studies of cattle. BTB localizes to the lymph nodes of the respiratory tract first, beginning with the retropharyngeal and tracheobronchial lymph nodes, gradually increasing in lymph nodes affected over time. At 36 months, rate of spread to additional lymph nodes sharply increases. The lung lesions follow a similar pattern, progressing slowly, then accelerating their progression at 36 months post infection. Lastly, a genetic marker that correlated to risk of M. bovis infection in previous studies was marginally associated with BTB progression. Buffalo with at least one risk allele at this locus tended to progress faster, with more lung necrosis. The progression of disease in the African buffalo mirrors the progression found in experimental cattle models, offering insight into BTB and the interaction with its host in the context of naturally varying environments, host, and pathogen populations.

The Long-Horned African Buffalo ( Pelorovis antiquus) is an Extinct Species

Context An understanding of large herbivore habitat choices in heterogeneous African protected areas is important for the better management of these key ecosystems. Aims To determine habitat use of African buffalo (Syncerus caffer) and plains zebra (Equus quagga) in a heterogeneous protected area. Methods Zambezi National Park (ZNP), Zimbabwe, was divided into five vegetation types using an unsupervised classification on a Landsat satellite image that was classified into five land cover classes, using the K-means classification algorithm. African buffalo and plains zebra densities were then determined in each vegetation type using road transect surveys monthly between January 2013 and December 2015. Normalised difference vegetation index (NDVI), grass biomass, grass height and grass quality (nitrogen, calcium, phosphorus and acid detergent fibre content) were determined in each vegetation type during the wet (November to April) and dry (August to October) seasons to establish their quality as habitats for African buffalo and plains zebra. Key results Both African buffalo and plains zebra mostly foraged in mixed and grassland areas, and avoided Zambezi teak vegetation type. Zambezi teak vegetation type had high NDVI due to the dense tree cover. Both African buffalo and plains zebra preferred vegetation types with intermediate grass biomass (approximately 300 g m−2) and grass height (approximately 16 cm). Grass nutritive value (in terms of nitrogen, phosphorus and acid detergent fibre) declined from wet to dry season in all vegetation types. Conclusions African buffalo and plains zebra in the ZNP confined their habitat use mostly to two vegetation types (mixed and grassland), which together covered 25% of the protected area. Implications Teak (Baikiaea plurijuga) vegetation, which accounted for about 60% of the ZNP, was avoided by both African buffalo and plains zebra, suggesting that a significant part of the protected area was not used by the two herbivores.

Molecular characterisation of Anaplasma species from African buffalo (Syncerus caffer) in Kruger National Park, South Africa

The Cape buffalo (Syncerus caffer caffer) is one of the dominant and most widespread herbivores in sub-Saharan Africa. High levels of genetic diversity and exceptionally low levels of population differentiation have been found in the Cape buffalo compared to other African savannah ungulates. Patterns of genetic variation reveal large effective population sizes and indicate that Cape buffalos have historically been interbreeding across considerable distances. Throughout much of its range, the Cape buffalo is now largely confined to protected areas due to habitat fragmentation and increasing human population densities, possibly resulting in genetic erosion. Ten buffalo populations in Kenya and Uganda were examined using seventeen microsatellite markers to assess the regional genetic structure and the effect of protected area size on measures of genetic diversity. Two nested levels of genetic structure were identified: a higher level partitioning populations into two clusters separated by the Victoria Nile and a lower level distinguishing seven genetic clusters, each defined by one or two study populations. Although relatively small geographic distances separate most of the study populations, the level of genetic differentiation found here is comparable to that among pan-African populations. Overall, correlations between conservancy area and indices of genetic diversity suggest buffalo populations inhabiting small parks are showing signs of genetic erosion, stressing the need for more active management of such populations. Our findings raise concerns about the future of other African savannah ungulates with lower population sizes and inferior dispersal capabilities compared with the buffalo.

Population genetics and phylogeography of the African buffalo (Syncerus caffer) are inferred from genetic diversity at mitochondrial D-loop hypervariable region I sequences and a Y-chromosomal microsatellite. Three buffalo subspecies from different parts of Africa are included. Nucleotide diversity of the subspecies Cape buffalo at hypervariable region I is high, with little differentiation between populations. A mutation rate of 13-18% substitutions/million years is estimated for hypervariable region I. The nucleotide diversity indicates an estimated female effective population size of 17 000-32 000 individuals. Both mitochondrial and Y-chromosomal diversity are considerably higher in buffalo from central and southwestern Africa than in Cape buffalo, for which several explanations are hypothesized. There are several indications that there was a late middle to late Pleistocene population expansion in Cape buffalo. This also seems to be the period in which Cape buffalo evolved as a separate subspecies, according to the net sequence divergence with the other subspecies. These two observations are in agreement with the hypothesis of a rapid evolution of Cape buffalo based on fossil data. Additionally, there appears to have been a population expansion from eastern to southern Africa, which may be related to vegetation changes. However, as alternative explanations are also possible, further analyses with autosomal loci are needed.

Fission–fusion dynamics allow animals to manage costs and benefits of group living by adjusting group size. The degree of intraspecific variation in fission–fusion dynamics across the geographical range is poorly known. During 2008–2016, 38 adult female Cape buffalo were equipped with GPS collars in three populations located in different protected areas (Gonarezhou National Park and Hwange National Park, Zimbabwe; Kruger National Park, South Africa) to investigate the patterns and environmental drivers of fission–fusion dynamics among populations. We estimated home range overlap and fission and fusion events between Cape buffalo dyads. We investigated the temporal dynamics of both events at daily and seasonal scales and examined the influence of habitat and distance to water on event location. Fission–fusion dynamics were generally consistent across populations: Fission and fusion periods lasted on average between less than one day and three days. However, we found seasonal differences in the underlying patterns of fission and fusion, which point out the likely influence of resource availability and distribution in time on group dynamics: During the wet season, Cape buffalo split and associated more frequently and were in the same or in a different subgroup for shorter periods. Cape buffalo subgroups were more likely to merge than to split in open areas located near water, but overall vegetation and distance to water were very poor predictors of where fission and fusion events occurred. This study is one of the first to quantify fission–fusion dynamics in a single species across several populations with a common methodology, thus robustly questioning the behavioral flexibility of fission–fusion dynamics among environments.

ABSTRACTIn social species, the transmission and maintenance of infectious diseases depends on the contact patterns between individuals within groups and on the interactions between groups. In southern Africa, the Cape buffalo (Syncerus caffer caffer) is a vector for many pathogens that can infect sympatric livestock. Although intra‐group contact patterns of Cape buffalo have been relatively well described, how groups interact with each other and risks for pathogen transmission remain poorly understood. We identified and compared spatial behavior and contact patterns between neighboring groups of Cape buffalo under contrasting environments: within the seasonally flooded environment of the Okavango Delta in Botswana and the semi‐arid environment of northern Kruger National Park in South Africa. We used telemetry data collected between 2007 and 2015 from 10 distinct groups. We estimated seasonal overlap and proximity between home ranges of pairwise neighboring groups, and we quantified seasonal contact patterns between these groups. We defined contact patterns within variable spatiotemporal windows compatible with the transmission of diseases carried by the Cape buffalo: bovine tuberculosis, brucellosis, and Rift Valley fever (mosquito‐borne transmission). We examined the effects of habitat and distance to water on contact location. In both study populations, neighboring buffalo groups were highly spatially segregated in the dry and rainy seasons. Inter‐group contact patterns were characterized by very few direct and short‐term indirect (within 0–2 days) contacts, lasting on average 1 hour and 2 hours, respectively. Contact patterns were generally consistent across populations and seasons, suggesting species‐specific behavior. In the drier study site, the probability of indirect and vector‐borne contacts generally decreased during the dry season with increasing distance to water. In the seasonally flooded area, only the probability of vector‐borne contact decreased with increasing distance to water. Our results highlight the importance of dry season water availability in influencing the dynamics of indirectly transmitted Cape buffalo pathogens but only in areas with low water availability. The results from this study have important implications for future modeling of pathogen dynamics in a single host, and the ecology and management of Cape buffalo at the landscape level. © 2021 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.

The African buffalo (Syncerus caffer) is an iconic species of South African megafauna. As the farmed buffalo population expands, the potential impacts on population health and disease transmission warrant investigation. A retrospective study of skin biopsy and necropsy samples from 429 animals was performed to assess the spectrum of conditions seen in buffaloes in South Africa. Determination of the cause of death (or euthanasia) could not be made in 33.1% (136/411) of the necropsy cases submitted due to autolysis or the absence of significant lesions in the samples submitted. Infectious and parasitic diseases accounted for 53.5% (147/275) of adult fatal cases and non-infectious conditions accounted for 34.9% (96/275). Abortions and neonatal deaths made up 11.6% (32/275) of necropsy cases. Rift Valley fever, bovine viral diarrhoea, malignant catarrhal fever, tuberculosis, bacterial pneumonia, anaesthetic deaths, cachexia and hepatotoxic lesions were the most common causes of death. The range of infectious, parasitic and non-infectious diseases to which African buffaloes were susceptible was largely similar to diseases in domestic cattle which supports concerns regarding disease transmission between the two species. The similarity between diseases experienced in both species will assist wildlife veterinarians in the diagnosis and treatment of diseases in captive African buffaloes. The present study likely does not represent accurate disease prevalence data within the source population of buffaloes, and diseases such as anthrax, brucellosis and foot and mouth disease are under-represented in this study. Hepatic ductal plate abnormalities and haemorrhagic septicaemia have not, to our knowledge, been previously reported in African buffaloes.

Sarcocystis infections have been reported from the African buffalo ( Syncerus caffer ), but the species have not been named. Here we propose a new name Sarcocystis cafferi from the African buffalo. Histological examination of heart (92), skeletal muscle (36), and tongue (2) sections from 94 buffalos from the Greater Kruger National Park, South Africa, and a review of the literature revealed only 1 species of Sarcocystis in the African buffalo. Macrocysts were up to 12 mm long and 6 mm wide and were located in the neck muscles and overlying connective tissue. They were pale yellow; shaped like a lychee fruit stone or cashew nut; turgid or flaccid and oval to round (not fusiform). By light microscopy (LM) the sarcocyst wall was relatively thin. By scanning electron microscopy (SEM), the sarcocyst wall had a mesh-like structure with irregularly shaped villar protrusions (vp) that were of different sizes and folded over the sarcocyst wall. The entire surfaces of vp were covered with papillomatous structures. By transmission electron microscopy (TEM), the sarcocyst wall was up to 3.6 μm thick and had highly branched villar protrusions that were up to 3 μm long. The villar projections contained filamentous tubular structures, most of which were parallel to the long axis of the projections, but some tubules criss-crossed, especially at the base. Granules were absent from these tubules. Longitudinally cut bradyzoites were 12.1 × 2.7 μm in size, had a long convoluted mitochondrion, and only 2 rhoptries. Phylogenetic analysis of 18S rRNA and cytochrome C oxidase subunit 1 (cox1) gene sequences indicated that this Sarcocystis species is very closely related to, but distinct from, Sarcocystis fusiformis and Sarcocystis hirsuta. Thus, morphological findings by LM, SEM, and TEM together with molecular phylogenetic data (from 18S rRNA and cox1) confirm that the Sarcocystis species in the African buffalo is distinct from S. fusiformis and has therefore been named Sarcocystis cafferi.