Population trends of large non-migratory wild herbivores and livestock in the Masai Mara ecosystem, Kenya, between 1977 and 1997

The total of all non-migratory wildlife species in the Masai Mara ecosystem has declined by 58% in the last 20 years. This decline ranges from 49% in small brown antelopes to 72% in medium brown antelopes. In individual wildlife species, the decline ranges from 52% in Grant's gazelle to 88% in the warthog. Declines of over 70% have been recorded in buffalo, giraffe, eland and waterbuck. Only elephant, impala and ostrich have not shown any significant decline or increase. Overall, there has not been any significant difference in decline of all wildlife population sizes inside and outside the reserve, except for Thomson's gazelle and warthog. Livestock have not significantly declined over the entire analysis period. However, livestock and cattle populations significantly declined during the 1983–88 period. Donkey declined by 67%, while shoats (goats and sheep) remained stable. In the case of wildlife, land use and vegetation changes, drought effects and poaching are considered to be among the potential factors that may have been responsible for the decline; the decline in livestock during the 1983–84 period was probably due to drought effects.     

Rainfall influences on ungulate population abundance in the Mara-Serengeti ecosystem

1. Rainfall is the prime climatic factor underpinning the dynamics of African savanna ungulates, but no study has analysed its influence on the abundance of these ungulates at monthly to multiannual time scales. 2. We report relationships between rainfall and changes in age- and sex-structured abundances of seven ungulate species monitored monthly for 15 years using vehicle ground counts in the Maasai Mara National Reserve, Kenya. 3. Abundance showed strong and curvilinear relationships with current and cumulative rainfall, with older topi, Damaliscus korrigum (Ogilby); warthog, Phacochoerus aethiopicus (Pallas); waterbuck, Kobus ellipsyprimnus (Ogilby); and impala, Aepyceros melampus (Lichtenstein) responding to longer lags than younger animals, portraying carryover effects of prior habitat conditions. 4. The abundances of newborn calves were best correlated with monthly rainfall averaged over the preceding 5-6 months for topi, waterbuck, warthog, and 2 months for the migratory zebra Equus burchelli (Gray), but with seasonal rainfall averaged over 2-5 years for giraffe, Giraffa camelopardalis (L.); impala; and kongoni, Alcelaphus busephalus (Pallas). The cumulative late wet-season rainfall was the best predictor of abundance for quarter- to full-grown animals for most species. Monthly rainfall exerted both negative and positive effects on the abundances of zebra, impala and waterbuck. Ignoring age, both sexes responded similarly to rainfall. 5. Births were strongly seasonal only for warthog and topi, but peaked between August and December for most species. Hence abundance was strongly seasonal for young topi and warthog and the migratory zebra. Pronounced seasonality in births for warthog and topi obliterated otherwise strong relationships between abundance and rainfall when both month and rainfall were included in the same model. Aggregated density produced relationships with rainfall similar to those for fully grown animals, emphasizing the necessity of demographic monitoring to reliably reveal rainfall influences on ungulate abundance in the Mara. 6. Strong relationships between abundance and rainfall suggest that rainfall underpins the dynamics of African savanna ungulates, and that changes in rainfall due to global warming may markedly alter the abundance and diversity of these mammals. Ungulates respond to rainfall fluctuations through movements, reproduction or survival, and the responses appear independent of breeding phenology and synchrony, dietary guild, or degree of water dependence. Newborns and adults have contrasting responses to rainfall. Males and females respond similarly to rainfall when age is ignored.     

Community‐based wildlife management area supports similar mammal species richness and densities compared to a national park

Community‐based conservation models have been widely implemented across Africa to improve wildlife conservation and livelihoods of rural communities. In Tanzania, communities can set aside land and formally register it as Wildlife Management Area (WMA), which allows them to generate revenue via consumptive or nonconsumptive utilization of wildlife. The key, yet often untested, assumption of this model is that economic benefits accrued from wildlife motivate sustainable management of wildlife. To test the ecological effectiveness (here defined as persistence of wildlife populations) of Burunge Wildlife Management Area (BWMA), we employed a participatory monitoring approach involving WMA personnel. At intermittent intervals between 2011 and 2018, we estimated mammal species richness and population densities of ten mammal species (African elephant, giraffe, buffalo, zebra, wildebeest, waterbuck, warthog, impala, Kirk's dik‐dik, and vervet monkey) along line transects. We compared mammal species accumulation curves and density estimates with those of time‐matched road transect surveys conducted in adjacent Tarangire National Park (TNP). Mammal species richness estimates were similar in both areas, yet observed species richness per transect was greater in TNP compared to BWMA. Species‐specific density estimates of time‐matched surveys were mostly not significantly different between BWMA and TNP, but elephants occasionally reached greater densities in TNP compared to BWMA. In BWMA, elephant, wildebeest, and impala populations showed significant increases from 2011 to 2018. These results suggest that community‐based conservation models can support mammal communities and densities that are similar to national park baselines. In light of the ecological success of this case study, we emphasize the need for continued efforts to ensure that the BWMA is effective. This will require adaptive management to counteract potential negative repercussions of wildlife populations on peoples' livelihoods. This study can be used as a model to evaluate the effectiveness of wildlife management areas across Tanzania.     

The biomass of game animals in Nairobi National Park, 1960–66

The large animals of Nairobi National Park have been censused intermittently once a month from mid-1960 until the end of 1966. Of the 23 species discussed, the density per kilometre has been determined accurately for the following: Grant's gazelle, Thomson's gazelle, water-buck, impala, wildebeest, hartebeest, eland, buffalo, warthog, giraffe, zebra and ostrich. From these figures the biomass has been calculated for each species each year. After a peak biomass of 12,775 kg/sq. km during the drought of 1960–61 the population declined to and remained fairly level at about 5690 kg/sq. km from 1962 to 1966. This figure includes about 1050 kg/sq. km of cattle and sheep. The large predators account for only about 1.4 % of the total biomass of wild animals while removing about 15.5% from this group annually.     

Antipredator behaviour of African ungulates around human settlements

As human populations grow and come into more frequent contact with wildlife, it is important to understand how anthropogenic disturbance alters wildlife behaviour. Using fine‐scale spatial analyses, we examined how proximity to human settlements affects antipredator responses of ungulates. We studied seven common ungulate species (Kirk's dik‐dik, Thomson's gazelle, impala, common warthog, common wildebeest, common zebra and Masai giraffe) in the Tarangire–Manyara ecosystem in northern Tanzania. In zebra and giraffe, flight responses to humans were significantly more likely when closer to settlements; however, there was a weak relationship between flight responses and distance to settlement in all other species. While there was largely a weak relationship between proximity to human settlements, the distribution of settlements in the landscape appears to affect wildlife behaviour, suggesting that animals perceive and respond to spatial variation in risk exerted by humans.     

Long‐term persistence of wildlife populations in a pastoral area

Facilitating coexistence between people and wildlife is a major conservation challenge in East Africa. Some conservation models aim to balance the needs of people and wildlife, but the effectiveness of these models is rarely assessed. Using a case‐study approach, we assessed the ecological performance of a pastoral area in northern Tanzania (Manyara Ranch) and established a long‐term wildlife population monitoring program (carried out intermittently from 2003 to 2008 and regularly from 2011 to 2019) embedded in a distance sampling framework. By comparing density estimates of the road transect‐based long‐term monitoring to estimates derived from systematically distributed transects, we found that the bias associated with nonrandom placement of transects was nonsignificant. Overall, cattle and sheep and goat reached the greatest densities and several wildlife species occurred at densities similar (zebra, wildebeest, waterbuck, Kirk's dik‐dik) or possibly even greater (giraffe, eland, lesser kudu, Grant's gazelle, Thomson's gazelle) than in adjacent national parks in the same ecosystem. Generalized linear mixed models suggested that most wildlife species (8 out of 14) reached greatest densities during the dry season, that wildlife population densities either remained constant or increased over the 17‐year period, and that herbivorous livestock species remained constant, while domestic dog population decreased over time. Cross‐species correlations did not provide evidence for interference competition between grazing or mixed livestock species and wildlife species but indicate possible negative relationships between domestic dog and warthog populations. Overall, wildlife and livestock populations in Manyara Ranch appear to coexist over the 17‐year span. Most likely, this is facilitated by existing connectivity to adjacent protected areas, effective anti‐poaching efforts, spatio‐temporal grazing restrictions, favorable environmental conditions of the ranch, and spatial heterogeneity of surface water and habitats. This long‐term case study illustrates the potential of rangelands to simultaneously support wildlife conservation and human livelihood goals if livestock grazing is restricted in space, time, and numbers.     

The effect of drought on the large mammal populations of Zambezi riverine woodlands

Large mammals were counted in Zambezi alluvial woodlands and on the adjacent ecotone in the north of Mana Pools National Park, Zimbabwe, from April 1981 until October 1989. Densities of all large herbivores on the alluvium in the daytime increased during the dry season. Year-to-year variation was studied by comparing end-of-dry-season densities. Flooding of the Zambezi River, controlled by Kariba Dam 100 km upstream, ceased in 1981 and the densities of all species on the alluvium declined in 1982. Rainfall was low in 1982 and very low in 1983 and 1984. The density of buffalo in the study area declined, owing to high mortality and a change in habitat utilization (buffalo herds spent more time in the south of the park). Densities of other species were high during the drought, but the densities of waterbuck, kudu, eland, bushbuck and warthog declined in 1985. Zebra density declined on the ecotone, but not on the alluvium. The densities of eland and kudu increased in 1986 and 1988, respectively, but the densities of the other species were still low in 1989. Impala density on the alluvium in October was related to rainfall during the preceding wet season and was high in drought years. Elephant density declined after a management cull. Rhinoceros density decreased by > 90% during the 1985/6 wet season, as a result of poaching and captures. The long-term decline in the densities of grazers was probably due primarily to the cessation of flooding by the Zambezi River, rather than to low local rainfall in some years. During the drought, the large grazers (e.g. buffalo) died before the smaller grazers (e.g. warthog). Waterbuck density declined less than the density of other grazers, because waterbuck could cross to vegetated sandbanks in the Zambezi River.     

Buffalo, bush meat, and the zoonotic threat of brucellosis in Botswana

Background

Brucellosis is a zoonotic disease of global importance infecting humans, domestic animals, and wildlife. Little is known about the epidemiology and persistence of brucellosis in wildlife in Southern Africa, particularly in Botswana.

Methods

Archived wildlife samples from Botswana (1995–2000) were screened with the Rose Bengal Test (RBT) and fluorescence polarization assay (FPA) and included the African buffalo (247), bushbuck (1), eland (5), elephant (25), gemsbok (1), giraffe (9), hartebeest (12), impala (171), kudu (27), red lechwe (10), reedbuck (1), rhino (2), springbok (5), steenbok (2), warthog (24), waterbuck (1), wildebeest (33), honey badger (1), lion (43), and zebra (21). Human case data were extracted from government annual health reports (1974–2006).

Findings

Only buffalo (6%, 95% CI 3.04%–8.96%) and giraffe (11%, 95% CI 0–38.43%) were confirmed seropositive on both tests. Seropositive buffalo were widely distributed across the buffalo range where cattle density was low. Human infections were reported in low numbers with most infections (46%) occurring in children (<14 years old) and no cases were reported among people working in the agricultural sector.

Conclusions

Low seroprevalence of brucellosis in Botswana buffalo in a previous study in 1974 and again in this survey suggests an endemic status of the disease in this species. Buffalo, a preferred source of bush meat, is utilized both legally and illegally in Botswana. Household meat processing practices can provide widespread pathogen exposure risk to family members and the community, identifying an important source of zoonotic pathogen transmission potential. Although brucellosis may be controlled in livestock populations, public health officials need to be alert to the possibility of human infections arising from the use of bush meat. This study illustrates the need for a unified approach in infectious disease research that includes consideration of both domestic and wildlife sources of infection in determining public health risks from zoonotic disease invasions.     

Abundance and distribution of mammals in Katavi National Park, Tanzania

Ground transects were used to determine densities of 24 larger mammals in Katavi National Park. The Park consists of miombo woodland habitat and two seasonal lakes. Mammalian biomass was extremely high due primarily to large numbers of buffalo. The highest mammal densities were found around Lake Chada to the south-east of the Park. Contrary to earlier reports, species' densities did not differ significantly between dry, wet and intermediate seasons, suggesting that, aside from elephant and warthog, mammals did not enter or leave the Park in large numbers. Similarly, aside from zebra and waterbuck, sightings of species in different vegetation types did not change markedly between seasons. Thus, seasonal lakes in Katavi National Park support very high local concentrations rather than acting as dry season refuges for mammals outside the Park.     

Does competition regulate ungulate populations? Further evidence from Serengeti,Tanzania

Summary Changes in populations of several ungulate species in the Serengeti-Mara region of East Africa over the past 30 years suggest several hypotheses for their regulation and coexistence. Recent censuses in the 1980s have allowed us to test the hypotheses that: (1) there was competition between wildebeest (Connochaetes taurinus) and Thomson's gazelle (Gazella thomsoni). This predicted that gazelle numbers should have declined in the 1980s when wildebeest were food limited. Census figures show no change in gazelle numbers between 1978 and 1986, a result contrary to the interspecific competition hypothesis; (2) wildebeest and African buffalo (Syncerus caffer) populations were regulated by intraspecific competition for food. Since both populations reached food limitation in the 1970s, the hypothesis predicted that the populations should have been stable in the 1980s. The results confirm these predictions for wildebeest and the buffalo population in the Mara reserve. In the Serengeti the buffalo population declined 41% over the period 1976–1984. The decline was not evenly distributed over the park, some areas showing an 80–90% decline, others no change or an increase in numbers. The decline was associated with proximity to human habitation; (3) an outbreak of the viral disease, rinderpest, in 1982 may have been the cause of the drop in buffalo population. Blood serum samples to measure the prevalence of antibodies were collected from areas of decreasing, stable and increasing populations. If rinderpest was the cause of decrease there should be a negative relationship between the prevalence of rinderpest and the instantaneous rate of increase (r). The results showed no relationship. We conclude that rinderpest was not the major cause of the drop in buffalo numbers. Elephant (Loxodonta africana) numbers dropped 81% in Serengeti in the period 1977–1986. In the Mara there was little change. The evidence suggests that extensive poaching in northern and western Serengeti during 1979–1984 accounted for the drop in both elephant and buffalo numbers.     

Changes in large herbivore populations across large areas of Tanzania

We collated aerial census data collected during the late 1980s to early 2000s for large herbivore populations in eight large census zones in Tanzania. Of the ungulate populations that showed significant changes in densities at the start versus end of this decade, most declined; very few populations increased significantly. Thomson's gazelle, Grant's gazelle, hartebeest, reedbuck, roan antelope, sable antelope, warthog and zebra, for example, declined in over 50% of the zones where they were surveyed. Interestingly, small‐bodied species fared particularly poorly in many census zones, whereas elephant and giraffe generally fared well across the country. Most populations of all herbivores declined in some portions of the country (e.g. Burigi‐Biharamulo, Katavi, Greater Ruaha and Tarangire census zones). These surveys suggest that, even in a country renowned for its protected areas and conservation commitment, some large herbivore populations need more conservation attention in order to remain stable.     

Large herbivores in Upper Lupande Game Management Area, Luangwa Valley, Zambia

Upper Lupande Game Management area in Luangwa Valley, Zambia currently supports good populations of sixteen large and medium-sized herbivores. Impala was the most abundant constituting 34% of the total population estimates, followed by buffalo (19%), zebra (12%), wildebeest (10%), warthog (8%), elephant (6%) and hippopotamus (3%). Other species made up the remaining 8%. The survey zone contained a herbivore biomass of 10,796.18 times 10³ kg. Elephants contributed 50.2% and buffalo 25.1% of the total herbivore biomass of the area. The estimated maximum sustainable off-take potential in Upper Lupande was 1110.14 times 10³kg per annum. The distribution of most of the species studied suggested favourable recruitment status. There were species-specific distribution patterns and densities in the survey zone.     

Anti-Trypanosoma brucei activity of nonprimate zoo sera

Constitutive anti-Trypanosoma brucei subsp. brucei S 427 clone 1 and 22 activities were evaluated in sera from 22 species of nonprimate mammals. The sera fell into 5 categories. Sera from Cape buffalo, giraffe, and greater kudu showed a concentration-dependent inhibition of replication of the 2 clones of organisms, which was dependent on the presence of xanthine oxidase. Sera from warthog and springbok also severely limited trypanosome replication but lacked xanthine oxidase. Their antitrypanosome activity was inactivated by heating at 56 C for 30 min but not affected by absorbing with trypanosomes at 4 C. Sera from lion and leopard showed a concentration-dependent inhibition of the growth of T. brucei S427 clone 1 organisms, but not clone 22 organisms. These sera lacked xanthine oxidase. Their anti-T. brucei S 427 clone 1 activity was inactivated by heating at 56 C for 30 min but not removed by absorbing with trypanosomes. Serum from Grant's gazelle prevented replication of both T. brucei clones, lacked xanthine oxidase, and was not affected by heating at 56 C. Sera from waterbuck, Thompson's gazelle, sitatunga, Cape hartebeeste, gerenuk, Grant's zebra, cow, several cat, cougar, bobcat, and domestic cat were fully supportive of trypanosome replication irrespective of concentration tested up to a maximum of 48% v/v in culture medium. Sera from different individuals of the same mammal species had similar effects on trypanosomes, and samples collected from the same individual at different times also had similar activities indicating species-specific stable expression, or lack thereof, of constitutive serum antitrypanosome components.     

Relationships among physical traits,age and social status in Thomson's and Grant's gazelles

Territorial male Thomson's and Grant's gazelles on a commercial game ranch near Nairobi, Kenya, were older, heavier and possessed longer horns than bachelor conspecifics. Among territorial Thomson's gazelles, males accompanied by females were older than solitary individuals but differed little in frame and horn dimensions. Small sample sizes precluded similar analyses for Grant's gazelles. These results suggest the importance of size, age and perhaps experience in determining social status.     

Interactions among host diet, nutritional status and gastrointestinal parasite infection in wild bovids

In this study, I explored the interactions among host diet, nutritional status and gastrointestinal parasitism in wild bovids by examining temporal patterns of nematode faecal egg shedding in species with different diet types during a drought and non-drought year. Study species included three grass and roughage feeders (buffalo, hartebeest, waterbuck), four mixed or intermediate feeders (eland, Grant's gazelle, impala, Thomson's gazelle) and two concentrate selectors (dik-dik, klipspringer). Six out of the nine focal species had higher mean faecal egg counts in the drought year compared to the normal year, and over the course of the dry year, monthly faecal egg counts were correlated with drought intensity in four species with low-quality diets, but no such relationship was found for species with high-quality diets. Comparisons of dietary crude protein and faecal egg count in impala showed that during the dry season, individuals with high faecal egg counts (> or =1550 eggs/g of faeces) had significantly lower crude protein levels than individuals with low (0-500 eggs/g) or moderate (550-1500 eggs/g) egg counts. These results suggest that under drought conditions, species unable to maintain adequate nutrition, mainly low-quality feeders, are less able to cope with gastrointestinal parasite infections. In particular, during dry periods, reduced protein intake seems to be associated with declining resilience and resistance to infection.     

Foraging behaviour of wild impala (Aepyceros melampus) and Burchell's zebra (Equus burchelli) in relation to sward height

Foraging behaviour plays a key role in the interaction between herbivores and vegetation, their predominant food source. Understanding this interaction is crucial to providing information that is useful for conservation of herbivores. The objective of this study was to determine how sward height influences functional response and movement patterns of free ranging wild impala and zebra at the Kenya Wildlife Service Training Institute. The study was conducted for 3 months. Sward height is an important parameter that indicates how intensive a sward has been grazed and it influences intake rate through its effect on bite size. Bite size, instantaneous intake rate, specific mass intake rate and feeding station interval for impala and zebra increased with sward height. Sward height in combination with an animal's sex was found to have a profound effect on specific mass intake rate in impala. Zebra had a longer feeding station interval and lower stepping rate in tall swards compared to impala. Despite differences in their specific body mass and digestive strategies, impala and zebra maximized their intake rates in tall swards as a trade‐off among the swards. Tall swards are therefore critical in the study area and should be protected from bush encroachment which is a persistent problem.     

Changing vulnerability to predation related to season and sex in an African ungulate assemblage

The consequences of predation for prey population dynamics depend on the extent to which this mortality is predisposed by malnutrition or senescence, or additive in the sense that animals that would otherwise not have died at that time were killed. In places lacking effective predators, few adult ungulates die during the summer or wet season months when food is plentifully available. Hence the seasonal distribution of predator kills as well as the age and sex classes of the prey indicates the extent to which malnutrition contributes to mortality as well as other influences on vulnerability. Using records of animal deaths assembled over 35 years in South Africa's Kruger National Park, these patterns were investigated for 12 ungulate species forming the prey of lions, and for three other large predators with respect to one prey species. Buffalo, kudu and giraffe were more strongly represented in kills made during the late dry season, while wildebeest and zebra made relatively greater contributions during the wet season. Impala, waterbuck, warthog and rarer antelope species became more prominent in kills during transitional periods between seasons. Five prey species showed an elevation in representation of males in lion kills during the mating season, as well as impala for all predator species. Females were more prominently represented in kills during the time of late gestation and parturition for three prey species. Hence reproductive activities as well as changing vegetation cover and food resources affected vulnerability to predation. Shifts in susceptibility to predation over the seasonal cycle corresponded with rainfall‐related variation in the annual representation of these ungulate species in lion kills. The availability of vulnerable prey species, age and sex classes at different stages of the seasonal cycle helps maintain a high abundance of lions. These factors contribute to the strong additive impact that predation has had on the abundance of some of these ungulate populations.