African Buffalo Social Dynamics: What Is a Buffalo Herd?

Documenting within species group size variation is important to completely understand social organization within species and to interpret variation among species. Here, I investigated group size of African buffalo Syncerus caffer over 2 years in a heterogeneous landscape. African buffalo use closed continuous forest and vast open savannas, and anecdotal observations suggest that habitat type influences their social structure. While the Cape buffalo Syncerus caffer caffer is well studied, few data exist for the forest‐dwelling Syncerus caffer nanus. I observed forest buffalo at Lopé National Park, Gabon, and examined variation of group size. Eighteen forest buffalo herds used the study area with an estimated population of 342 individuals (∼5 buffalo km−2). The mean group size for the 18 herds was 12 ± 2 (range of means=3–24), considerably smaller than Cape buffalo herds. For eight radiocollared forest buffalo, the mean group size was stable, varying little with time of day, across seasons, or between savanna and marsh habitat. However, herd size varied widely across herds, from fewer than 10 individuals in the smallest herds to more than 20 buffalo in the largest. Large herd size is associated with home ranges that contain substantial areas of open habitat, and thus more food resources than forested habitats.

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.

Foot-and-mouth disease (FMD) inflicts severe economic losses within infected countries and is arguably the most important trade-restricting livestock disease in the world. In southern Africa, infected African buffaloes (Syncerus caffer) are the major reservoir of the South African Territories (SAT) types of the virus. With the progressive expansion of transfrontier conservation areas (TFCAs), the risk of FMD outbreaks is expected to increase due to a higher probability of buffalo/livestock contacts. To investigate the dynamics of FMD within and around the Great Limpopo TFCA (GLTFCA), 5 herds of buffaloes were sampled in June 2010 to characterize circulating viruses in South Africa and Zimbabwe. Three SAT-2 and three SAT-3 viral strains were isolated in both countries, including one that was genetically linked with a recent SAT-2 outbreak in Mozambique in 2011. In addition, two groups of unvaccinated cattle (n = 192) were serologically monitored for 1 year at the wildlife/livestock interface of Gonarezhou National Park (GNP) in Zimbabwe between April 2009 and January 2010, using the liquid-phase blocking ELISA (LPBE) and a test for antibodies directed against non-structural proteins (NSP). Neither clinical signs nor vaccination of cattle were reported during the study, yet a high proportion of the monitored cattle showed antibody responses against SAT-3 and SAT-1. Antibodies against NSP were also detected in 10% of the monitored cattle. The results of this study suggest that cattle grazing in areas adjacent to the GLTFCA can be infected by buffalo or other infected livestock and that cattle trade movements can act as efficient disseminators of FMD viruses to areas several hundred kilometres from the virus source. Current methods of surveillance of FMD at the GLTFCA interface seem insufficient to control for FMD emergence and dissemination and require urgent reassessment and regional coordination.

Population genetic structure is often used to infer population connectivity, but genetic structure may largely reflect historical rather than recent processes. We contrasted genetic structure with recent gene-flow estimates among 6 herds of African buffalo (Syncerus caffer) in the Caprivi Strip, Namibia, using 134 individuals genotyped at 10 microsatellite loci. We tested whether historical and recent gene flows were influenced by distance, potential barriers (rivers), or landscape resistance (distance from water). We also tested at what scales individuals were more related than expected by chance. Genetic structure across the Caprivi Strip was weak, indicating that historically, gene flow was strong and was not affected by distance, barriers, or landscape resistance. Our analysis of simulated data suggested that genetic structure would be unlikely to reflect human disturbances in the last 10-20 generations (75-150 years) because of slow predicted rates of genetic drift, but recent gene-flow estimates would be affected. Recent gene-flow estimates were not consistently affected by rivers or distance to water but showed that isolation by distance appears to be developing. Average relatedness estimates among individuals exceeded random expectations only within herds. We conclude that historically, African buffalo moved freely throughout the Caprivi Strip, whereas recent gene flow has been more restricted. Our findings support efforts to maintain the connectivity of buffalo herds across this region and demonstrate the utility of contrasting genetic inferences from different time scales.

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.

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.

Bulk feeder or selective grazer: African buffalo space use patterns based on fine-scale remotely sensed data on forage quality and quantity

Using age-related infection rates derived from serological data in available deterministic and specially developed stochastic simulation models, it has been possible to establish that the basic reproductive rates for South African Territory (SAT) type foot and mouth disease virus in buffalo (Syncerus caffer) are high. The models predict that there is a periodicity of infection within herds and possibly the population as a whole. Thus, buffalo herds are likely to be more infectious at some times than at others. However, because most infections in buffalo are inapparent, such episodes are difficult to identify. There is wide intratypic variation within the SAT type virus populations circulating in buffalo. This was determined by sequencing part of the 1 D gene of buffalo isolates and establishing antigenic profiles with neutralising monoclonal antibodies and conventional antisera.

Spatial and temporal changes in group dynamics and range use enable anti‐predator responses in African buffalo

Ante-mortem bovine tuberculosis (bTB) tests for buffaloes include the single comparative intradermal tuberculin test (SCITT), interferon-gamma (IFN-γ) release assay (IGRA) and IFN-γ-inducible protein 10 release assay (IPRA). Although parallel test interpretation increases the detection of Mycobacterium bovis (M. bovis)-infected buffaloes, these algorithms may not be suitable for screening buffaloes in historically bTB-free herds. In this study, the specificities of three assays were determined using M. bovis-unexposed herds, historically negative, and a high-specificity diagnostic algorithm was developed. Serial test interpretation (positive on both) using the IGRA and IPRA showed significantly greater specificity (98.3%) than individual (90.4% and 80.9%, respectively) tests or parallel testing (73%). When the SCITT was added, the algorithm had 100% specificity. Since the cytokine assays had imperfect specificity, potential cross-reactivity with nontuberculous mycobacteria (NTM) was investigated. No association was found between NTM presence (in oronasal swab cultures) and positive cytokine assay results. As a proof-of-principle, serial testing was applied to buffaloes (n = 153) in a historically bTB-free herd. Buffaloes positive on a single test (n = 28) were regarded as test-negative. Four buffaloes were positive on IGRA and IPRA, and M. bovis infection was confirmed by culture. These results demonstrate the value of using IGRA and IPRA in series to screen buffalo herds with no previous history of M. bovis infection.

African buffalo were introduced into a wildlife conservancy in the southeast of Zimbabwe in an effortto increase the conservancy's economic viability, which is primarily based on eco-tourism. The buffalo were infected with SAT serotypes (SAT-1, SAT-2 and SAT-3) of foot and mouth disease (FMD) virus, and in order to isolate the conservancy and prevent the transmission of FMD to adjacent populations of domestic livestock, the conservancy was surrounded by a double-fence system, 1.8 m in height. The intention was to prevent the movement of both wildlife and domestic animals across the perimeter. However, two years after the buffalo were introduced, FMD occurred in cattle farmed just outside of the conservancy. Using serological and molecular diagnostic tests, epidemiological investigations showed that it was most likely that antelope (impala or kudu), infected through contact with the buffalo herd within the conservancy, had jumped over the fence and transmitted the virus to the cattle.

Following the recent invasion of bovine tuberculosis (BTB) into the Kruger National Park, South Africa, we conducted a study on the maintenance host, African buffalo, to investigate associations between BTB prevalence and calf:cow ratio, age structure, body condition, and endoparasite load. Statistical analyses compared herds of zero, medium (1–40%), and high (>40%) BTB prevalence. To control for ecological variation across the park we collected data in northern, central, and southern regions and restricted some analyses to particular regions of the park. Body condition declined over the course of the 2001 dry season, and buffaloes in the southern region of the park, with the highest BTB prevalence, were in worse condition than buffaloes in the northern region (which receives less annual rainfall but is still virtually BTB‐free). Herd‐level analyses of the entire park, the south and central regions, and just the southern region all indicated that herds of higher BTB prevalence were in worse condition and lost condition faster through the dry season than herds of lower BTB prevalence. Fecal endoparasite egg counts increased during the dry season and were associated with both decreased body condition and increased BTB prevalence. Although we did not detect any obvious effect of BTB on the age structure of the buffalo population, our findings indicate early symptoms of wider scale BTB‐related ecological disturbances: buffalo herds with high BTB prevalence appear more vulnerable to drought (because of a decrease in body condition and an increase in endoparasite load), and because lions selectively kill weak buffaloes their prey base is accumulating a disproportionately high prevalence of BTB, to which lions are susceptible.

BackgroundIn Zambia, translocation of wildlife from National Parks to private owned game ranches demands that only animals free of infectious diseases that could adversely affect the expansion of the wildlife industry should be translocated to game ranches. Sarcoptes mange (Sarcoptes scarbiei) has been involved in the reduction of wildlife populations in some species.ResultsSarcoptes mange (Sarcoptes scarbiei) was detected and eradicated from two herds of African buffalo (Syncerus caffer) calves captured in the Kafue GMA in July 2004 and August 2005. The overall prevalence was estimated at 89.5% (77/86). Sex had no influence on the occurrence and severity of the disease. Of the 86 calves used in the study, 72.1% had good body condition scores, 20.9% were fair and 7.0% were poor. Of the 77 infected calves, 53.2% were mildly infected, 28.6% were moderately and 18.2% were severely infected. Body condition score was correlated to the severity of the infection (r = 0.72, p < 0.000, n = 86) at capture. Eradication of Sarcoptes mites from the entire herd using ivermetcin was dependant on the severity of the infection. The overall ability of ivermectin to clear the infection after the first treatment was estimated at 81.8% (n = 77). It increased to 94.8% and 100% after the second and third treatments respectively.ConclusionThis is the first report on the epidemiology and treatment of Sarcoptes mange in African buffaloes in Zambia. This study improves our understanding about Sarcoptes scabiei epidemiology and treatment which will have further applications for the safe animal translocation.

The ubiquity and importance of parasite co-infections in populations of free-living animals is beginning to be recognized, but few studies have demonstrated differential fitness effects of single infection versus co-infection in free-living populations. We investigated interactions between the emerging bacterial disease bovine tuberculosis (BTB) and the previously existing viral disease Rift Valley fever (RVF) in a competent reservoir host, African buffalo, combining data from a natural outbreak of RVF in captive buffalo at a buffalo breeding facility in 2008 with data collected from a neighbouring free-living herd of African buffalo in Kruger National Park. RVF infection was twice as likely in individual BTB+ buffalo as in BTB- buffalo, which, according to a mathematical model, may increase RVF outbreak size at the population level. In addition, co-infection was associated with a far higher rate of fetal abortion than other infection states. Immune interactions between BTB and RVF may underlie both of these interactions, since animals with BTB had decreased innate immunity and increased pro-inflammatory immune responses. This study is one of the first to demonstrate how the consequences of emerging infections extend beyond direct effects on host health, potentially altering the dynamics and fitness effects of infectious diseases that had previously existed in the ecosystem on free-ranging wildlife populations.

Newborn mammals have an immature immune system that cannot sufficiently protect them against infectious diseases. However, variation in the effectiveness of maternal immunity against different parasites may couple with temporal trends in parasite exposure to influence disparities in the timing of infection risk. Determining the relationship between age and infection risk is critical in identifying the portion of a host population that contributes to parasite dynamics, as well as the parasites that regulate host recruitment. However, there are no data directly identifying timing of first infection among parasites in wildlife. Here, we took advantage of a longitudinal dataset, tracking infection status by viruses, bacteria, protists and gastro-intestinal worms in a herd of African buffalo (Syncerus caffer) to ask: how does age of first infection differ among parasite taxa? We found distinct differences in the age of first infection among parasites that aligned with the mode of transmission and parasite taxonomy. Specifically, we found that tick-borne and environmentally transmitted protists were acquired earlier than directly transmitted bacteria and viruses. These results emphasize the importance of understanding infection risk in juveniles, especially in host species where juveniles are purported to sustain parasite persistence and/or where mortality rates of juveniles influence population dynamics.