Habitat preferences of the secretive forest buffalo (Syncerus caffer nanus) in Central Africa

The forest buffalo Syncerus caffer nanus is one of the three subspecies of African buffalo inhabiting the rainforests of Western and Central Africa. Because of its secretive behaviour and main habitat (dense rainforests), there is little quantitative information on the habitat preferences of this buffalo. We present here the first data on the frequencies of this species along a habitat gradient ranging from clearings and rivers to forests, as well as the characteristics of the buffalo's resting places. We recorded information from a buffalo herd during the period January 2002–January 2004 in the Bai-Hokou area (Dzanga-Ndoki National Park, Central African Republic). Resting places were firstly compared with available habitat (i.e. resting vs. random sites) and, successively, comparisons were made between diurnal versus nocturnal and wet versus dry season resting places. Forest buffalos were found to be highly dependent on clearings, as well as on the more open forest stands, characterized by large trees and open canopy. Such preferences could be due to the tendency of the buffalos to rest all together; open patches are likely to facilitate social interactions between the members of the herd.     

Trade-offs of predation and foraging explain sexual segregation in African buffalo

1. Many studies have investigated why males and females segregate spatially in sexually dimorphic species. These studies have focused primarily on temperate zone ungulates in areas lacking intact predator communities, and few have directly assessed predation rates in different social environments. 2. Data on the movement, social affiliation, mortality and foraging of radio-collared African buffalo (Syncerus caffer) were collected from 2001-06 in the Kruger National Park, South Africa. 3. The vast majority of mortality events were due to lion (Panthera leo) predation, and the mortality hazard associated with being an adult male buffalo in a male-only 'bachelor' group was almost four times higher than for adult females in mixed herds. The mortality rates of adult males and females within mixed herds were not statistically different. Mortality sites of male and female buffalo were in areas of low visibility similar to those used by bachelor groups, while mixed herds tended to use more open habitats. 4. Males in bachelor groups ate similar or higher quality food (as indexed by percentage faecal nitrogen), and moved almost a third less distance per day compared with mixed herds. As a result, males in bachelor groups gained more body condition than did males in breeding herds. 5. Recent comparative analyses suggest the activity-budget hypothesis as a common underlying cause of social segregation. However, our intensive study, in an area with an intact predator community showed that male and female buffalo segregated by habitat and supported the predation-risk hypothesis. Male African buffalo appear to trade increased predation risk for additional energy gains in bachelor groups, which presumably leads to increased reproductive success.     

Can small wildlife conservancies maintain genetically stable populations of large mammals? Evidence for increased genetic drift in geographically restricted populations of Cape buffalo in East 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.     

Phylogeography of the African buffalo based on mitochondrial and Y-chromosomal loci: Pleistocene origin and population expansion of the Cape buffalo subspecies

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.     

Basic haematological values in the African buffalo (Syncerus caffer caffer) and in the red buffalo (Syncerus caffer nanus)

Basic haematological values in 13 African buffaloes (Syncerus caffer caffer), 11 cows and 2 bulls, and in 2 red buffaloes (Syncerus caffer nanus), 1 bull and 1 cow, are reported. In the African buffalo, the erythrocyte count--mean 9.76 X 10(12) 1(-1)--ranged from 6.72 to 12.42 X 10(12) 1(-1), the haematocrit values--mean 0.391--varied from 0.32 to 0.47 and the haemoglobin content--mean 148.17--was in the range from 122.1 to 172.5 g 1(-1). For the red buffalo only individual values are given. In the African buffaloes, the leukocyte counts--mean 7.39 X 10(9) 1(-1)--ranging from 5.1 to 12.4 X 10(9) 1(-1) were slightly increased compared with man. For the red buffalo only individual values are reported. These values are compared with data abstracted from the literature for related buffalo species. The age-dependent decrease of erythrocyte counts and the age-dependent rise of the eosinophile proportion in the differential count are discussed.     

The numbers and distribution of buffalo in the Ruwenzori National Park*, Uganda

The buffalo (Syncerus coffer (Sparrman)) in the Ruwenzori National Park, Uganda were censused by means of aerial survey. Details of earlier censuses of buffalo from the air are given. Herds were located by flying along flight lines 1–6 km apart at a height of 300 m above ground. Buffalo in small herds and in bachelor groups containing less than thirty animals were counted individually but most herds were photographed and the number of buffalo counted from the prints. Two full photographic counts were made in November 1968 and December 1969 and subsequently, counts of herds only were made four times a year during each wet and dry season. The total numbers of buffalo were estimated from the herd counts on the basis of the mean herd size and percentage of bachelors recorded earlier. A total of 17 835 buffalo, comprising 16 749 in 165 herds and 1086 bachelor bulls, was recorded in 1968 and 18 040 buffalo made up of 17 141 in 162 herds and 899 bachelors were counted in 1969. The mean herd size was 101-5 in 1968 and 105-8 in 1969 with bachelors representing 6-1% and 5-0% of the totals in 1968 and 1969 respectively. The mean size of the bachelor groups was 4–7 and 3-3 in 1968 and 1969 respectively. The density of the buffalo was the same each year at twelve animals per sq. km. If only the preferred habitat is considered, the density becomes thirty-eight buffalo per sq. km. The mean areas occupied by a single herd were 9-4 and 9-6 sq. km in 1968 and 1969 respectively. The mean area of preferred habitat used by one herd was 2–9 sq. km each year. It is concluded that herd counts are not a satisfactory method for assessing the total buffalo population although they have value as indices, provided the same observer is used. However, they do suggest in the present work that there is a slight but definite seasonal movement of buffalo into forested areas or towards permanent water during the dry season.     

Potential effects of prescribed savannah burning on the diet selection of forest buffalo (Syncerus caffer nanus) in Lopé National Park,Gabon

Seasonality and management are factors that may affect the diet selection of the forest buffalo (Syncerus caffer nanus). Fire is considered a major driving force in savannah systems and prescribed burning is a commonly applied conservation tool in protected areas such as Lopé National Park, Gabon. Prescribed annual fires contribute to the maintenance of open areas and provide high‐quality forage for forest buffalo, a major herbivore in the park. We used microhistological faecal analysis to determine the diet selection of forest buffalo and measured the extent of variation between a dry season, preburn and a wet season, postburn sampling period. The buffalo diet comprised mainly of monocotyledons, primarily grasses (Poaceae) and sedges (Cyperaceae). Intake of open‐area‐associated plant species was higher in the wet season, postburn treatment sampling period (97%) than the dry season, preburn sampling period (87%), which corresponded conversely to a reduction in forest‐associated Marantaceae plants (10% versus 1%). High proportions of grasses and sedges in the diet signify the importance of open areas for forest buffalo. Controlled burning as tool for maintenance of open areas may play a key role in the meta‐population management of the forest buffalo.     

Buffalo Herd Structure and its Repercussions for Condition of Individual African Buffalo Cows

The African buffalo Syncerus caffer was studied in Lake Manyara National Park, Tanzania. Emphasis was placed on the study of (individual) buffalo cows, which live in mixed herds. Buffalo herds are discrete social units and females were never observed in another herd than their own. The herd showed a structure with respect to the distribution of sex-age classes. Individual cows generally kept the same location within the herd. The location in the herd appeared to be coupled to food intake and was strongly related to physical condition. The best location (highest intake and best condition) was between the front and the centre of the herd, the worst location was the rear of the herd (when moving or grazing). Females with calves appeared to have the highest position in the hierarchy as determined from the rate of displacement over food; adult bulls did not interact with cows. Few births were observed during the late dry season and it appears that there is a calving peak at the end of the long rains. Conception rate increased when cows increased in condition and dropped when cows lost condition. Cows showed a strong seasonality in condition but bulls on average hardly changed in condition, except for a loss in condition during the inferred conception peak. Buffalo herds in Manyara showed a fusion-fission pattern independent of season but strongly influenced by the size of the herd: large herds split more often than smaller ones. In large herds, buffalo grazed closer together than in small herds and it appeared likely that competition was more severe in large herds. Animals in the rear of a large herd lost condition faster during the dry season than animals in the best location in the herd, and especially cows in the rear split off most frequently from the herd to graze in a smaller fragment. From the literature on cattle, it is inferred that the reproductive success of cows in the rear of the herd will be lower than of cows in the best location, and this differential is confirmed by the behaviour of adult bulls. It is as yet unclear what the advantage is for adult cows in the rear of a large herd to stay in that herd but the sharing of information with more successful individuals seems a good candidate.     

Group size, density and biomass of large mammals in the Réserve de Faune du Petit Loango, Gabon

Group size, density and biomass of large‐bodied diurnal mammal species in the Réserve de Faune du Petit Loango, Gabon (now Parc National de Loango) was determined over a 12‐month period using standard line‐transect methods. Petit Loango encompasses a range of distinct habitat types, including coastal scrub, savanna, swamps and disturbed and mature forest. Such intact coastal habitats are increasingly rare on the Central‐West African coastline. Faecal censusing indicated highest forest elephant (Loxodonta africana cyclotis) and buffalo (Syncerus caffer nanus) ecological densities at the extreme coast (2.48 and 1.29 km⁻² respectively), probably reflecting high intensity of use of this habitat. Ape density was comparable with that at other Central African study sites at 1.01 individuals km⁻². Mean total biomass of diurnal primates, elephants and other ungulates over the 20 km² site was 3290 kg km⁻². Forest elephants and red river hogs (Potamochoerus porcus) constituted the bulk of the biomass, at 67% and 14% respectively. Primates made up 5% of the biomass. This is the first estimation of mammal density and biomass over an annual cycle at a Central African coastal site, and provides baseline data for long‐term studies in such habitats and to aid habitat and wildlife management decisions.     

Home on the range: factors explaining partial migration of African buffalo in a tropical environment

Partial migration (when only some individuals in a population undertake seasonal migrations) is common in many species and geographical contexts. Despite the development of modern statistical methods for analyzing partial migration, there have been no studies on what influences partial migration in tropical environments. We present research on factors affecting partial migration in African buffalo (Syncerus caffer) in northeastern Namibia. Our dataset is derived from 32 satellite tracking collars, spans 4 years and contains over 35,000 locations. We used remotely sensed data to quantify various factors that buffalo experience in the dry season when making decisions on whether and how far to migrate, including potential man-made and natural barriers, as well as spatial and temporal heterogeneity in environmental conditions. Using an information-theoretic, non-linear regression approach, our analyses showed that buffalo in this area can be divided into 4 migratory classes: migrants, non-migrants, dispersers, and a new class that we call "expanders". Multimodel inference from least-squares regressions of wet season movements showed that environmental conditions (rainfall, fires, woodland cover, vegetation biomass), distance to the nearest barrier (river, fence, cultivated area) and social factors (age, size of herd at capture) were all important in explaining variation in migratory behaviour. The relative contributions of these variables to partial migration have not previously been assessed for ungulates in the tropics. Understanding the factors driving migratory decisions of wildlife will lead to better-informed conservation and land-use decisions in this area.     

Ecological determinants of herd size in the Thornicroft’s giraffe of Zambia

Ecological factors have a pervasive impact on animal population sizes and the structure of their social systems. In a number of ungulate species, predator pressure exerts a major influence on group size. Given that giraffe (Giraffa camelopardalis) live in an extremely flexible social system, and that breeding is nonseasonal, they are an ideal species for examining how ecological variables contribute to fluctuations in herd size. We present an analysis of 34 years of data on a population of Thornicroft’s giraffe (G. c. thornicrofti Lydekker 1911) that reveal how herd size changes with season and habitat. Sex differences in herd size were apparent, with bulls often travelling as singletons, whereas cows were generally observed with conspecifics. Herds were larger during the wet than dry season, but herd size changed in a parallel fashion across habitats. Giraffe herds were smaller in woodland and thicket areas than in open habitats, regardless of season. We suggest that the regular fluctuations in herd size among giraffe indicate a fission/fusion social system embedded within a larger social community. We conclude that changes in herd size among giraffe reflect a dynamic process regulated by individuals adjusting the number of associates based upon an interaction of foraging, reproductive, social and antipredator strategies.     

Current population estimate and distribution of the African buffalo in Chebera Churchura National Park,Ethiopia

An investigation into the population status and distribution of the African buffalo (Syncerus caffer Sparrman, 1779) in Chebera Churchura National Park, Ethiopia, was carried out during the wet and dry seasons of 2012–2015. This study tested the hypothesis that buffalo would demonstrate seasonal habitat preferences and changes in population density. Sample counts were carried out in an area of 1215 km². The estimated buffalo population was 5193 individuals, with the population density of 4.3/km². The population showed an increase from 2617 to 5194 individuals during 2006–2015. Males comprised 42.6%, while females 46.7% of the population. Age structure was dominated by adults, which constituted 52.5% of the total population. Subadults comprised 24.3% and young 12.4% of the population. Larger herds of up to 30 individuals were observed during the wet season, and smaller herds of a minimum of four individuals were seen during the dry season. The mean herd sizes during the wet and dry seasons were 29.59 and 16.95, respectively. They were observed more in the riverine vegetation types during the dry season. Of the total, 57.6% utilized riverine habitat during the dry season, whereas 39.8% used this habitat during the wet season. Relative abundance of food sources, green vegetation cover and availability of water were the major factors governing their distribution in the present study area.     

Inter-Group Social Behavior,Contact Patterns and Risk for Pathogen Transmission in Cape Buffalo Populations

In 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.     

Age determination of the West African buffalo Syncerus caffer brachyceros and the constancy of tooth wear

Maxillae of 34 West African buffalo Syncerus caffer brachyceros were collected from three sites in Burkina Faso, West Africa, and age determined from tooth appearance and the enamel height of M1 according to the criteria of Grimsdell (1973) for Ugandan buffalo Syncerus caffer caffer. Good agreement between the two methods was found. Ground and decalcified sections were prepared from M1 and the interradicular cementum lines counted. Although this gave inconsistent results in terms of assumed age compared with the other two methods, it is concluded that agreement was sufficiently close to suggest that the rate of attrition with age was similar in both populations.     

Spatial properties of a forest buffalo herd and individual positioning as a response to environmental cues and social behaviour

Many animals aggregate into organized temporary or stable groups under the influence of biotic and abiotic factors, and some studies have shown the influence of habitat features on animal aggregation. This study, conducted from 2002 to 2004 in the Dzanga-Ndoki National Park, Central African Republic, studied a herd of forest buffaloes (Syncerus caffer nanus) to determine whether spatial aggregation patterns varied by season and habitat. Our results show that both habitat structure and season influenced spatial aggregation patterns. In particular, in open habitats such as clearings, the group covered a larger area when resting and was more rounded in shape compared to group properties noted in forest during the wet season. Moreover, forest buffaloes had a more aggregated spatial distribution when resting in clearings than when in the forest, and individual positions within the herd in the clearing habitat varied with age and sex. In the clearings, the adult male (n = 24) was generally, on most occasions, located in the centre of the herd (n = 20), and he was observed at the border only four times. In contrast, females (n = 80) occupied intermediate (n = 57), peripheral (n = 14) and central positions (n = 9) within the group. Juveniles (n = 77) also occurred in intermediate (n = 64) and peripheral positions (n = 13). Based on these results, we concluded that habitat characteristics and social behaviour can have relevant effects on the spatial distribution of animals within a group.     

Antibodies to Brucella spp. among blue wildebeest and African buffalo in Kenya

A serologic survey of blue wildebeest (Connochaetes taurinus Burchell) and African buffalo (Syncerus caffer Sparrman) in the Masai Mara area was conducted. Antibodies to Brucella spp. were found in 18% of the blue wildebeest and 30% of the African buffalo examined. There were titers in all age groups and in both sexes. Hygromata were seen in both species. The increase in numbers of blue wildebeest and African buffalo which share grazing and watering areas with cattle of the Masai people, makes the presence of infections by Brucella spp. in wildlife an important consideration in any program for control of brucellosis.     

Epididymal sperm from the African buffalo (Syncerus caffer) can be frozen successfully with AndroMed and with Triladyl but the addition of bovine seminal plasma is detrimental

Numerous diseases are carried and can be transmitted from the African buffalo (Syncerus caffer) to livestock. Buffaloes free of specific diseases (BFSD) are thus in demand amongst game farmers. Current BFSD derive from a small genetic pool and hence there is a special interest in bringing new genetic material into such herds. In this study epididymal sperm from 16 mature African buffalo bulls was frozen with Triladyl and AndroMed extender (Minitüb, Tiefenbach, Germany) with and without addition of bovine seminal plasma. Post-thaw motility, longevity and acrosomal integrity were compared. In all but one animal, post-thaw motility was higher, although not always significant, if sperm was frozen with Triladyl than with AndroMed. Seminal plasma was detrimental to the post-thaw motility. Neither semen extender nor seminal plasma had an influence on post-thaw acrosomal integrity. It can be concluded that bovine seminal plasma at a concentration of 10% is detrimental rather than beneficial for the post-thaw motility of African buffalo sperm. Even though being inferior AndroMed does, however, have the advantage that it is a defined semen extender and therefore clearly has a lower risk of contamination.     

Bayesian inference of population expansions in domestic bovines

The past population dynamics of four domestic and one wild species of bovine were estimated using Bayesian skyline plots, a coalescent Markov chain Monte Carlo method that does not require an assumed parametric model of demographic history. Four domestic species share a recent rapid population expansion not visible in the wild African buffalo (Syncerus caffer). The estimated timings of the expansions are consistent with the archaeological records of domestication.     

Population structure and ecology of the African buffalo (Syncerus caffer Sparrman, 1779) in Chebera Churchura National Park,Ethiopia

An investigation of population structure and ecology of the African buffalo (Syncerus caffer Sparrman, 1779) was carried out in the Chebera Churchura National Park, Ethiopia during the wet and dry seasons of 2005–2006. Sample counts of African buffaloes were carried out in an area of 1,215 km². The estimated population of buffalo was 2,617 individuals. Males comprised 35.0%, while females 52.8% of the population. The remaining 12% of the population was young of both sexes and of all ages of unknown sex. It was difficult to categorize the young into male and female in the field, as their primary sexual characteristics were not easily visible. Male to female sex ratio was 1.00 : 1.51. Age structure was dominated by adults, which constituted 72.06% of the total population. Subadults comprised 22.02%, and young accounted for 5.9% of the population. Larger herds of up to 27 individuals were observed during the wet season and smaller herds of eight individuals during the dry season. The mean herd size during wet and dry seasons was 24.81 and 7.77, respectively. The African buffaloes were distributed in four habitat types such as grasslands, woodlands, montane forests and riverine habitat in the study area. They were observed more in the riverine vegetation types during the dry season. Relative abundance of food resources, green vegetation cover and water availability in the area were the major factors governing their distribution in the present study area. They spent a greater proportion of the time in feeding and resting/ruminating activities. On the average, 49.7% of the daytime was spent in feeding, and 38.75% of the day on resting (lying down and standing). Morning and the late afternoon activity peaks were more pronounced during the dry season than the wet season.     

Cape buffalo mitogenomics reveals a Holocene shift in the African human-megafauna dynamics

Africa is unique among the continents in having maintained an extraordinarily diverse and prolific megafauna spanning the Pleistocene-Holocene epochs. Little is known about the historical dynamics of this community and even less about the reasons for its unique persistence to modern times. We sequenced complete mitochondrial genomes from 43 Cape buffalo (Syncerus caffer caffer) to infer the demographic history of this large mammal. A combination of Bayesian skyline plots, simulations and Approximate Bayesian Computation (ABC) were used to distinguish population size dynamics from the confounding effect of population structure and identify the most probable demographic scenario. Our analyses revealed a late Pleistocene expansion phase concurrent with the human expansion between 80 000 and 10 000 years ago, refuting an adverse ecological effect of Palaeolithic humans on this quarry species, but also showed that the buffalo subsequently declined during the Holocene. The distinct two-phased dynamic inferred here suggests that a major ecological transition occurred in the Holocene. The timing of this transition coincides with the onset of drier conditions throughout tropical Africa following the Holocene Optimum (～9000-5000 years ago), but also with the explosive growth in human population size associated with the transition from the Palaeolithic to the Neolithic cultural stage. We evaluate each of these possible causal factors and their potential impact on the African megafauna, providing the first systematic assessment of megafauna dynamics on the only continent where large mammals remain abundant.