Seasonal changes in the nutrient content of East African grassland vegetation

Significant seasonal changes in the concentrations of N, P, K, and Ca were observed in the herb layer vegetation at Nairobi National Park and Masai Mara Game Reserve in Kenya. These changes were related to the alternation of wet and dry seasons. Live forbs typically had the highest concentrations of all nutrients, especially N and Ca, at both locations. Standing dead and litter compartments showed less seasonal variation and had the lowest concentrations of all nutrients except Ca when compared with those of other compartments. Standing stocks of all nutrients were higher at Nairobi National Park than at Masai Mara due to larger litter and standing dead compartments resulting from lower grazing intensity and a lower frequency of fire. Most of the aboveground nutrients at Nairobi National Park appeared to be recycled through the decomposer pathway, while a large proportion of the standing stock at Masai Mara appeared to be recycled through the excreta of large mammalian herbivores and through fires. The crude protein content of the live grass compartment fell below maintenance levels required for ruminants (5% crude protein) during September 1980 and February 1981 at Nairobi National Park and during June and July 1980 at Masai Mara Game Reserve. Thus, herbivore populations in these preserves may be limited by shortages of nutritionally adequate food during dry seasons, as previously reported for other regions in East Africa.     

Anthropogenic disturbance induces opposing population trends in spotted hyenas and African lions

Large carnivore populations are declining worldwide due to direct and indirect conflicts with humans. Protected areas are critical for conserving large carnivores, but increasing human-wildlife conflict, tourism, and human population growth near these sanctuaries may have negative effects on the carnivores within sanctuary borders. Our goals were to investigate how anthropogenic disturbance along the edge of the Masai Mara National Reserve, Kenya, influences the demography and space-use of two large carnivore species that engage in intense interspecific competition. Here we document, in one disturbed region of the Reserve, a rapid increase in the population size of one large predator, the spotted hyena (Crocuta crocuta), but a striking concurrent decline in numbers of another, the African lion (Panthera leo). Anthropogenic disturbances negatively affected lion populations, and decreasing lion numbers appear to have a positive effect on hyena populations, indicated here by an increase in juvenile survivorship. We also saw an increase in the number of livestock consumed by hyenas. Our results suggest human population growth and indirect effects of human activity along Reserve boundaries may be effecting a trophic cascade inside the Reserve itself. These results indicate both top-down and bottom-up processes are causing a shift in the carnivore community, and a major disruption of guild structure, inside the boundaries of one of the most spectacular protected areas in Africa.     

An assessment of tree availability as a possible cause of population declines in scavenging raptors

Lack of suitable nesting trees is an increasingly common issue for avian conservation given rampant habitat and tree destruction around the world. In the African savannah, habitat loss and particularly tree damage caused by elephants have been suggested as possible factors in the decline of large bird species. Given the recent declines of vultures and other scavenging raptors, it is critical to understand if nest availability is a limiting factor for these threatened populations. Loss of woodland, partially due to elephant populations, has been reported for the Mara‐Serengeti ecosystem. Data on characteristics of trees used for nesting were collected for white‐backed, lappet‐faced, white‐headed vulture, and tawny eagle nests in Masai Mara National Reserve, Kenya. Nest tree characteristics were compared with the distribution of a random subsample of trees to assess nest preferences and determine suitability of available trees. Nearest neighbor distances were estimated as well as availability of preferred nesting trees to determine if tree availability is a limiting factor for tree‐nesting vultures. Tree availability was found to greatly exceed nesting needs for African vultures and tawny eagles. We thus conclude that on a landscape scale, tree availability is not a limiting factor for any of the species considered here (white‐backed, lappet‐faced, white‐headed vultures and tawny eagles).     

Nesting habitat preference of the African White-backed Vulture Gyps africanus and the effects of anthropogenic disturbance

The African White-backed Vulture Gyps africanus is widely distributed across sub-Saharan Africa but populations are in decline. Loss of suitable habitat for foraging and breeding are among the most important causes, and future conservation will require identification of suitable remaining habitat and the threats to it and to the vultures in it. Like many large raptors, African White-backed Vultures have a long breeding cycle and thus spend much of each year near their nest site, but ecological correlates of nest sites have not been quantified for any African vulture species. We use nest-site data for African White-backed Vultures collected during aerial and ground surveys and habitat data derived from a GIS to develop statistical models that estimate the probability of nest presence in relation to habitat characteristics, and test these models against an independent dataset. The models predicted that both direct and indirect disturbance by humans limit the potential distribution. Suitable habitat needs to be identified and receive adequate protection from poaching. Poaching of vultures is thought to be mainly for use in traditional medicine and does not target any particular species, so all vulture species can be considered equally at risk. We predict the likelihood of individuals nesting in currently unprotected areas should they become protected. These predictions show that readily available GIS data combined with relatively simple statistical modelling can provide meaningful large-scale predictions of habitat availability.     

Status of the Mara Woodlands in Kenya

The woodlands of the Masai Mara National Reserve in Kenya have suffered dramatic declines over four decades as a result of elephant and fire pressure. This study examined the current status of woody resources in the Reserve and browse pressure thereon, using both classification (TWINSPAN) and ordination (DCA) techniques. From 333 widespread regular plots used to survey the vegetation, a total of 62 woody species were identified. Thirteen woody habitats were identified on the basis of species composition, varying from species‐rich closed thickets and forest to less diverse open grasslands. A NW/SE stratification of the more open habitats was observed, possibly as a result of differences in soils, rainfall and drainage. Both plant density and diversity were lower than in communal and privately managed areas outside the Reserve. Moreover, browser pressure was substantially higher than that observed previously in the ecosystem, and suggests increased competition for scarcer woody resources within the Reserve. This has implications for the management of the ecosystem as a whole. As woodland and thickets continue to decline, long‐term monitoring should expand to encompass the wider habitat diversity of the open grasslands and unprotected areas where much of the regeneration potential resides.     

Hooded Vulture Necrosyrtes monachus and African White-backed Vulture Gyps africanus nesting at the Olifants River Private Nature Reserve,Limpopo province,South Africa

Basic ecological information is still lacking for many species of African vultures. The Hooded Vulture Necrosyrtes monachus is known as a rare breeding resident in north-eastern South Africa. This study set out to monitor the nests of Hooded Vultures and, secondarily, White-backed Vultures Gyps africanus in the Olifants River Private Nature Reserve over two breeding seasons in 2013 and 2014. A total of 12 Hooded Vulture nests, placed mostly in the tree Diospyros mespiliformis, were found along the Olifants River, with an average inter- nest distance of 0.76 km. Nest success was estimated to be between 0.44–0.89 offspring pair?1 y?1 in 2013 and 0.50–0.67 offspring pair?1 y?1 in 2014, which are the first estimates for Hooded Vultures in South Africa. It is thought that nests of this species have been under-reported due to the fact that they are placed within or below the canopy of densely leafed trees and hence difficult to view from aerial surveys. African White-backed Vultures also bred along the Olifants River, with nests placed in clusters of up to six. Nesting density of this species ranged from about 1.0 to 1.2 nests km?1 and nests were predominantly placed in Ficus sycomorus trees.     

Biomass dynamics of grassland vegetation in Kenya

Seasonal changes in plant biomass in the herb layer were measured at Nairobi National Park and Masai Mara Game Reserve in Kenya from January 1980 to February 1981. Plant biomass fluctuated in response to seasonal rainfall, and live biomass was correlated with rainfall and soil moisture at both locations. Peak values for live biomass at Nairobi National Park occurred at the end of the long rains during the June 1980 sample and ranged from 138 to 197gm^-2. Minimum values for live biomass at Nairobi occurred during a dry season in February 1981 and ranged from 8 to 39 g m^-2. The standing dead and litter compartments were larger than the live compartment during every sample period at Nairobi National Park, and together often comprised more than 80% of the total above-ground biomass. Probably as a result of higher rainfall, peak values for live biomass at Masai Mara Game Reserve were higher than those at Nairobi. Again, peak biomass occurred during June following the long rains, and ranged from 218 to 294 gm^-2. Minimum values for live biomass occurred during February 1981, and ranged from 10 to 48 g m^-2. Standing dead and litter compartments were much smaller than at Nairobi National Park, reflecting more intense herbivore pressure and a greater frequency of fires at Masai Mara compared with that at Nairobi National Park.     

Assessing the accuracy of plotless density estimators using census counts to refine population estimates of the vultures of Kruger National Park

Breeding population estimates for three vulture species in Kruger National Park (KNP), South Africa, were made in 2013 using data from aerial censuses and a plotless density estimator (PDE). PDEs are distance-based methods used to assess sparse populations unsuitable for plot-based methods. A correction factor was applied to the 2013 estimates to reflect the difference between the survey counts and the PDE figures. We flew additional censuses across most of KNP and counted all visible nests to assess the 2013 estimates. Survey counts were within 95% confidence limits of corrected PDE estimates for White-backed Vulture Gyps africanus (count: 892; estimate: 904 [95% CI ±162]), at the limit of confidence for White-headed Vulture Trigonoceps occipitalis (count: 48; estimate: 60 [±13]) and outside confidence limits for Lappet-faced Vulture Torgos tracheliotos (count: 44; estimate: 78 [±18]). Uncorrected PDE estimates accurately reflected White-headed and Lappet-faced Vulture nest counts. The clustered patterns of White-backed Vulture nests and dispersed patterns of White-headed and Lappet-faced Vulture nests offer an explanation for these results and means that corrected PDE densities are inaccurate for estimating dispersed nests but accurate for estimating clustered nests. Using PDE methods, aerial surveys over ～35% of KNP are probably sufficient to assess changes in these vulture populations over time. Our results highlight these globally important breeding populations.     

Nesting distribution of vultures in relation to land use in Swaziland

Three species of vulture (African White-backed, White-headed and Lappet-faced) breed in Swaziland, all of which are threatened within the country. Vulture nests were surveyed using a fix-winged aircraft in low-lying savannas of Swaziland. Nesting was observed in three land use categories: (1) unprotected government cattle ranches, (2) protected cattle ranches, and (3) conservation areas. A total of 248 nests was recorded, of which 240 belonged to the African White-backed Vulture. Nesting densities were highest in conservation areas, an order of magnitude lower on protected cattle ranches and negligible on government ranches. Nests of White-headed Vultures and Lappet-faced Vultures were exclusively located in conservation areas. Nesting densities of African White-backed Vultures in some conservation areas exceeded 260 nests/100 km², which are the highest known densities of this species anywhere in Africa. Nests were almost exclusively located in riparian vegetation, but at Hlane National Park a large proportion of nests were placed in open woodland, possibly as a result of an influx of vultures from adjoining agricultural lands that have only been transformed in recent decades. Where elephants were present in conservation areas, vultures did not nest within their enclosures. The location and density of vulture nests may possibly be used as an indicator of pressure on biological resources in low-lying savannas of Swaziland.     

Comparative changes in density and demography of large herbivores in the Masai Mara Reserve and its surrounding human-dominated pastoral ranches in Kenya

Wildlife habitats in pastoral lands adjoining protected areas in east African savannas are getting progressively degraded, fragmented and compressed by expanding human populations and intensification of land use. To understand the consequences of these influences on wildlife populations, we contrasted the density and demography of 13 wild and three domestic large herbivores between the Masai Mara National Reserve and the adjoining pastoral ranches using aerial surveys conducted in the wet and dry seasons during 1977–2010. Species of different body sizes and feeding styles had different densities between landscapes and seasons. Small-sized herbivores, requiring short, nutritious grasses, and browsers were more abundant in the ranches than the reserve in both seasons. Medium-sized herbivores moved seasonally between landscapes. Larger-bodied herbivores, requiring bulk forage but less susceptible to predation, were more abundant in the reserve than the ranches. The proportions of newborn warthog (Phacochoerus africanus) and juvenile topi (Damaliscus korrigum) were higher in the ranches, with shorter grasses and lower predation risk than in the reserve. These results suggest that pastoral lands adjoining protected areas in African savannas are important as seasonal dispersal and breeding grounds for wild herbivores. However, human population growth and dramatic land use changes are progressively degrading wildlife habitats in pastoral areas, thus restricting the seasonal wildlife dispersal movements between the protected areas and adjoining pastoral lands. Conservation efforts should focus on (1) creating and maintaining functional heterogeneity in protected areas that mimic moderate pastoral grazing conditions to attract small and medium-bodied grazers and (2) securing dispersal areas, including corridors, to ensure continued seasonal large herbivore movements between protected and pastoral systems.     

Hippopotamus and livestock grazing: influences on riparian vegetation and facilitation of other herbivores in the Mara Region of Kenya

Riparian savanna habitats grazed by hippopotamus or livestock experience seasonal ecological stresses through the depletion of herbaceous vegetation, and are often points of contacts and conflicts between herbivores, humans and their livestock. We investigated how hippopotamus and livestock grazing influence vegetation structure and cover and facilitate other wild herbivores in the Mara region of Kenya. We used 5 km-long transects, each with 13 plots measuring 10 × 10 m², and which radiate from rivers in the Masai Mara National Reserve and adjoining community pastoral ranches. For each plot, we measured the height and visually estimated the percent cover of grasses, forbs, shrubs and bare ground, herbivore abundance and species richness. Our results showed that grass height was shortest closest to rivers in both landscapes, increased with increasing distance from rivers in the reserve, but was uniformly short in the pastoral ranches. Shifting mosaics of short grass lawns interspersed with patches of medium to tall grasses occurred within 2.5 km of the rivers in the reserve in areas grazed habitually by hippos. Hence, hippo grazing enhanced the structural heterogeneity of vegetation but livestock grazing had a homogenizing effect in the pastoral ranches. The distribution of biomass and the species richness of other ungulates with distance from rivers followed a quadratic pattern in the reserve, suggesting that hippopotamus grazing attracted more herbivores to the vegetation patches at intermediate distances from rivers in the reserve. However, the distribution of biomass and the species richness of other ungulates followed a linear pattern in the pastoral ranches, implying that herbivores avoided areas grazed heavily by livestock in the pastoral ranches, especially near rivers.     

Coprologic survey of parasites of spotted hyenas (Crocuta crocuta) in the Masai Mara National Reserve,Kenya

Seventy fecal samples from spotted hyenas (Crocuta crocuta) in the Masai Mara National Reserve, Kenya were examined for parasite eggs and oocysts using sugar flotation. A total of nine parasite genera were identified, and all samples were positive for at least one parasite species. Most individuals were infected with Ancylostoma sp. and Spirometra sp., and these species had the highest median intensity of infection. Other parasites identified include Isospora sp., Taeniidae, Spirurida, Toxocara sp., Mesocestoides sp., Dipylidium sp., and Trichuris sp.     

Spatial dynamics of lions and their prey along an environmental gradient

Establishing the ecological determinants of the spatial dynamics of large African savanna mammals is necessary for understanding the cumulative impacts on the suitability and resilience of their natural habitats, of progressive habitat fragmentation and temporal shifts in climate, fire regimes and elephant browsing. Systematic directional gradients and small‐scale spatial dependence were evident in rainfall, herbivore biomass and lion density in the Masai Mara National Reserve during September 1990 to July 1992. Lion density, the biomass of resident and all herbivore species were autocorrelated within 4.3, 4.1 and 3.3 km, respectively, and peaked in areas of relatively low rainfall but higher microtopographic and vegetation heterogeneity and complexity and more diverse drainage systems. Rainfall influenced herbivore biomass and hence lion density non‐linearly and that influence was apparently modified by forage mineral nutrients, structural habitat complexity, degree of drainage and distance to waterpoints. Lion density was related to the biomass of resident herbivores, even after controlling for rainfall and spatial trends. Persistence of the recent decline in Mara woodlands and increasing human population along the reserve–ranch boundary would likely reduce the reserve's lion population because most lions spent the daytime within the vegetation mosaic fringing drainage lines, away from humans and domestic livestock.     

The effects of fire and grazing pressure on vegetation cover and small mammal populations in the Maasai Mara National Reserve

An extensive study of vegetation changes as a consequence of fire and grazing pressure and their effect on small mammal populations inside the Maasai Mara National Reserve, Kenya, was carried out during May–June 1997. Comparison of vegetation maps from 1979 and 1998 suggested that vegetation in 46% of the Reserve area converted from shrubland to grassland, possibly as a result of fire and grazing pressure. We tested the hypothesis that in areas with high fire and grazing impact the population of small mammals was negatively affected. A low density of rodents was recorded in all habitats except in areas of human activity, where artificial resources are constantly present. Capture efforts were unsuccessful in grasslands. Our results confirm those of Norton‐Griffiths (1979) and Dublin (1995) , i.e. that fire and grazing pressure impact the vegetation of the Serengeti–Mara ecosystem and limit the natural regeneration of woodlands. This indirectly affects the small mammal community, which is limited in its long‐term establishment.     

Socio-ecology of Bat-eared foxes (Otocyon megalotis)

Bat-eared foxes were studied for eight months at four sites in Kenya and Tanzania with the main study in the Masai Mara Game Reserve. Eleven foxes were caught and four fitted with radio-collars. Foxes in Masai Mara feed mainly on ants and termites. They showed a marked preference for short grass habitat, and their range expanded when wildebeest lowered grass height by grazing and trampling. Foxes lived in groups (= 2.8) with overlapping home ranges, and relations between groups were usually amicable. Foxes gave birth from September-December and all members of the group provided some care to the cubs.
A review of Bat-eared fox ecology from other studies reveals a consistent picture of this aberrant insectivorous canid. The species foraging strategy involves locating social insects by sound in short grass areas. The species is able to find suitable habitat and multiply rapidly. Although usually living in monogamous pairs, reports of polygyny, and non-reproductive males in groups also exist. Living in groups may increase each fox's chance of feeding on insect swarms, and also provides protection from predators.     

Observations of mixed-species bird flocks at Kichwa Tembo Camp,Kenya

Mixed-species foraging flocks were studied at Kichwa Tembo Camp on the edge of the Masai Mara National Reserve in Kenya between July and September 2004. Observations were made on 29 mixed-species flocks, in which 24 species participated. African Paradise-Flycatcher Terpsiphone viridis, Black-backed Puffback Dryoscopus cubla, Grey-backed Camaroptera Camaroptera brachyura, Collared Sunbird Hedydipna collars and Cabanis's Greenbul Phyllastrephus cabanisi were the most common participants in mixed-species flocks, as well as among the most frequently encountered bird species overall. The Black-backed Puffback was identified as the nuclear species in flocks due to their abundance and frequency with which they were followed by other species. Mixed-species flocks represent another niche dimension in this diverse bird community, but few of these species could be described as flock specialists; most of the birds observed in mixed-species flocks in this study were opportunistic attendant species, including the African Pygmy-Kingfisher Ispidina picta, not previously described as joining mixed-species flocks.     

Dynamics of births and juvenile recruitment in Mara–Serengeti ungulates in relation to climatic and land use changes

Natality and recruitment govern animal population dynamics, but their responses to fluctuating resources, competition, predation, shifting habitat conditions, density feedback and diseases are poorly understood. To understand the influences of climatic and land use changes on population dynamics, we monitored monthly changes in births and juvenile recruitment in seven ungulate species for 15 years (1989–2003) in the Masai Mara Reserve of Kenya. Recruitment rates declined for all species but giraffe, likely due to habitat alteration and increasing vulnerability of animals associated with recurrent severe droughts, rising temperatures, unprecedentedly strong and prolonged El Niño-Southern Oscillation (ENSO) episodes, expansion of settlements, cultivation and human population growth in pastoral ranches adjoining the reserve. Birth rate showed strong and humped relationships with moving averages of monthly rainfall, whereas recruitment responded strongly to cumulative past rainfall. Increasing livestock incursions into the reserve depressed recruitment rate for quarter-grown topi. Expansion of pastoral settlements depressed birth rate in impala, zebra and giraffe. Frequent ENSO-related droughts caused progressive habitat desiccation and hence nutritional shortfalls for ungulates. The responses to climatic, land use and resource influences did not reflect body size, migratory or resident lifestyle, dietary guild, digestive physiology or degree of synchrony of breeding of the ungulate species.     

Cliff characteristics,neighbour requirements and breeding success of the colonial Cape Vulture Gyps coprotheres

The breeding success of endangered colonial nesting species is important for their conservation. Many species of Gyps vultures form large breeding colonies that are the foci of conservation efforts. The Cape Vulture is a globally threatened species that is endemic to southern Africa and has seen a major reduction in its population size (≥ 50% over 48 years). There is evidence that breeding colonies are prone to desertion as a result of human disturbance. Factors that influence the occupancy and breeding success of individual nest‐sites is not fully understood for any African vulture species. We investigated cliff characteristics and neighbour requirements of the Msikaba Cape Vulture colony, a major breeding colony in the southern node of the population in the Eastern Cape, South Africa, together with their nest‐site occupation and breeding success over 13 years. In total, 1767 breeding attempts were recorded. Nest‐sites that had a higher elevation, smaller ledge depth, greater total productivity and were surrounded by conspecifics were more likely to be occupied, although the amount of overhang above the nest was not an important predictor of occupancy. In accordance with occupation, nest‐sites with a smaller ledge depth had higher breeding success; however, nests with a greater overhang were also more successful and height of the nest‐site was not an important predictor of breeding success. The breeding success of a nest‐site in a given year was positively influenced by the number of direct nest neighbours, and nests in the middle of high‐density areas had greater breeding success. This suggests that maintaining a high nest density may be an important consideration if declines of reproducing adults continue. Breeding success declined over the study period, highlighting the effects of a temporal variation or observer bias. Our results identified optimal nest‐site locations (ledge depths of 1 m, at a height of 180 m) and their effects on breeding success. This information can be used for planning reintroduction efforts of the endangered Cape Vulture and for their ongoing conservation.     

Movements of the migratory wildebeest population in the Serengeti area between 1960 and 1973

Data on the monthly distributions of the Serengeti migratory wildebeest population between June 1960 and May 1973 have been analysed. It has been shown that the population performs an annual migration, spending the wet season on the Serengeti plains, then moving west towards Lake Victoria at the beginning of the dry season, and later north to the northern extension of the Serengeti National Park or into the Masai Mara Game Reserve in Kenya. The wildebeest return to the plains as soon as the rain begins, but if this is delayed they may move south-west from the northern areas into the corridor first. Annual differences in the pattern of the migration can be correlated with differences in rainfall. The rainfall affects particularly the timing of the movements of the wildebeest on and off the plains, and the extent to which they utilize the northern areas. A large increase in the size of the population over the years studied has resulted in a greater utilization of the northern areas. It has been shown that about half the total range of the wildebeest population is outside the boundaries of the Serengeti National Park, and it is therefore important that these areas should be protected if such a large population is to survive.     

Challenges and opportunities in population monitoring of cheetahs

Population monitoring is key to wildlife conservation and management but is challenging at the spatial and temporal extents necessary for understanding changes. Noninvasive survey methods and spatial capture–recapture (SCR) models have revolutionized wildlife monitoring by providing the means to acquire data at large scales and the framework to generate spatially explicit predictions, respectively. Despite opportunities for improved monitoring, challenges can remain in the study design and model fitting phases of an SCR approach. Here, we used a search-encounter design with multi-session SCR models to collect spatially indexed photographs and estimate changes in density of cheetahs between 2005 and 2013–2016 in the Masai Mara National Reserve (MMNR) in Kenya. Our SCR models of cheetah encounters suggested little change in cheetah density from 2005 to 2013–2016, with some evidence that density fluctuated annually in the MMNR. The sampling period length (5 vs. 10 months) and timing (early, late, full year) over which spatial encounters were modeled did not alter inferences about density when sample sizes were adequate (>20 spatially distinct encounters). Our average density estimate of ~1.2 cheetahs/100 km² is consistent with the impression that the MMNR provides important cheetah habitat in Africa. During most years, spatial distribution of vegetation greenness (proxy for ungulate habitat quality) accounted for important variation in encounter rates. The search-encounter design here could be applied to other regions for cheetah monitoring. While snapshot estimates of population size across time are useful for wildlife monitoring, open population models may better identify the mechanisms behind temporal changes.