Distributional niche of relatively rare sable antelope in a South African savanna: habitat versus biotic relationships

The geographic distribution of a species is governed by climatic conditions, topography, resources and habitat structure determining the fundamental niche, while the local distribution expressed via home range occupation may be compressed by biotic interactions with competitors and predators, restricting the realised niche. Biotic influences could be especially important for relatively rare species. We investigated how rainfall, geology, land type and abundance of other ungulate species serving as competitors or prey for predators contributed to the patchy distribution of sable antelope herds within Kruger National Park. Data were provided by annual aerial surveys of ungulate populations conducted between 1978 and 1988. Sable herds were more commonly present on granitic and sandstone substrates than on more fertile basalt. They occurred both in the moist south‐west and dry north of the park. They were most abundant in sour bushveld and mopane savanna woodland, and mostly absent from knob thorn‐marula parkland. The presence of sable was negatively associated with high concentrations of impala and wildebeest, less consistently related to the abundance of zebra, and positively associated with the occurrence of buffalo herds. Best supported models included the separate effects of the most abundant grazers along with land type. Interspecific relationships seemed more consistent with vulnerability to predation as the underlying mechanism restricting the distribution of sable herds than with competitive displacement. Sable favoured land types distinct from those where wildebeest, the most preferred prey of lions, and impala, numerically the most important resident prey species, were most abundant. Hence the risk of predation, associated with habitat conditions where abundant prey species are most concentrated, can exert an overriding influence on the distribution of rarer species in terms of their home range occupation.     

Spatial partitioning of the soil water resource between grass and shrub components in a West African humid savanna

Most savanna water balance models assume water partitioning between grasses and shrubs in a two-layer hypothesis, but this hypothesis has not been tested for humid savanna environments. Spatial partitioning of soil water between grasses and shrubs was investigated in a West African humid savanna by comparing the isotopic composition (oxygen-18 and deuterium) of soil water and plant stem water during rainy and dry conditions. Both grass and shrub species acquire most of their water from the top soil layer during both rainy and dry periods. A shift of water uptake pattern towards deeper horizons was observed only at the end of the dry season after shrub defoliation. The mean depth of water uptake, as determined by the isotopic signature of stem water, was consistent with grass and shrub root profiles and with changes in soil water content profiles as surveyed by a neutron probe. This provides evidence for potentially strong competition between shrubs and grasses for soil water in these humid savannas. Limited nutrient availability may explain these competitive interactions. These results enhance our understanding of shrub-grass interactions, and will contribute to models of ecosystem functioning in humid savannas.     

Belowground competitive suppression of seedling growth by grass in an African savanna

Coexistence of N₂-fixing legumes and non-legume trees with grasses in African savannas results in intense competition between these life-forms. We hypothesised that belowground competition might induce different nutritional constraints in N₂- versus non-N₂-fixing species. A field (Hluhluwe-imFolozi nature reserve, South Africa) competition experiment with two N₂-fixing legume species (Acacia burkei and Acacia karroo) and two non-N₂-fixing species (Schotia brachypetala and Spirostachys africana) both with clipped grass and without grass was established. Plants were supplied with no fertilizer, or generous amounts of fertilizer (200?kg?N?ha^?1, 100?kg P₂O₅ ha^?1, 7.1?kg K₂O ha^?1) supplied as either 28?C10 (N?CK), P or a combination of these fertilizers (NPK). Regularly clipped grass suppressed growth (by more than 90?%) of both N₂- and non-N₂-fixing seedlings equally. Biomass accumulation of seedlings grown with grass and the grasses themselves responded positively to NK and/or NPK, but not P, although P-fertilization did have effects on foliar [N] and ??¹⁵N values of trees and grasses showing that plants accessed the fertilizer. Tree ??¹⁵N values and foliar [N] were also modified by NPK, demonstrating access to fertilizer. However, the ameliorative effects of NPK on grass competition-induced biomass suppression were only partial. This may be due to ??non-resource competition?? (i.e. root gaps) imposed by dense grass roots. The fact that nutrients were able to partially ameliorate the effects of grass competition, however, indicates that such ??non-resource competition?? may be partially overcome by even more generous supply of fertilizers.     

Seasonal distribution of topsoil ammonium and nitrate under legume-grass and grass swards

Losses of soil N through leaching and N₂ fixation by legumes often are related to soil nitrate concentration. The seasonal distribution of soil ammonium and nitrate concentrations under ungrazed legume-grass and grass swards were evaluated on two experiments that were established in 1983 (Exp. 1) and in 1984 (Exp. 2). Treatments were white clover (Trifolium repens L.) (WC), red clover (Trifolium pratense L.) (RC), and birdsfoot trefoil (Lotus corniculatus L.) (BT), each grown with tall fescue (Festuca arundicacea Schreb.) (TF) at two legume proportions, and a pure stand of TF. The concentrations of both forms of N were measured in the top 20-cm layer during 2 years in Exp. 1 and for 1 year in Exp. 2. The concentrations of nitrate and ammonium were least in winter and spring, and greatest in summer. The concentration of nitrate for the mixtures decreased in the order WC-TF, RC-TF, and BT-TF in both summers of Exp. 1 but there were no mixture differences in Exp. 2. The concentration of soil ammonium was not affected by the treatments applied. We conclude that the concentration of soil nitrate usually was small for these swards but became greater and often dependent on species and legume proportion during summer. The concentration of soil ammonium also was greater in summer but was not affected by species or legume proportion. Journal of Paper no. J.-13359 of the Iowa Agric. and Home Econ. Exp Stn., Ames. Project 2281. Supported in part by the Facultad de Agronomía, Montevideo, Uruguay. Journal of Paper no. J.-13359 of the Iowa Agric. and Home Econ. Exp Stn., Ames. Project 2281. Supported in part by the Facultad de Agronomía, Montevideo, Uruguay.     

Condensed tannins deter feeding by browsing ruminants in a South African savanna

Summary The palatability of 14 species of woody plant was assessed for three species of browsing ruminant, namely kudus, impalas and goats. Results show that palatability was most clearly related to leaf contents of condensed tannins. The effect was a threshold one, with all plants containing more than 5% condensed tannins being rejected as food during the wet season period. In contrast palatability was not influenced by concentrations of protein-precipitating polyphenols, and only weakly related to contents of nitrogen, phosphorus, cations, fibre components and other secondary metabolites. Insect herbivory shows a different pattern. These findings support the hypotheses that (i) condensed tannins function to protect plant cell walls against microbial attack; (ii) hydrolyzable tannins function to inactivate the digestive enzymes of insect herbivores. Large mammalian herbivores are influenced by condensed tannins due to their dependance upon microbial fermentation of plant cell walls for part of their energy needs.     

Herbivore and nutrient control of lawn and bunch grass distributions in a southern African savanna

In African savannas, bottom-up soil nutritional factors and top-down herbivory have both been suggested to control the distribution patterns of bunch and lawn grasses. We tested the separate and combined roles of these factors on grassland distribution in a South African savanna by focusing on three main objectives, namely (1) are grazing patches and lawns restricted to specific soils or sites, (2) does herbivory alter rates of nutrient cycling which facilitates lawns and (3) are there any differences in foliage quality between lawn and bunch grass-dominated sites that might influence animals in choosing to feed on lawns? We set up ten sites along a grazing gradient in the Hluhluwe-iMfolozi Park in KwaZulu-Natal, South Africa. At these sites, we measured total soil nutrients, soil nutrient turnover rates and grass foliage nutrient concentrations. We found that in the Hluhluwe-iMfolozi Park the spatial distribution of lawn and bunch grassland types appears to be an animal-driven phenomenon and not dependent on specific soil properties. The short-structured and distinct species found in lawns do not appear to be restricted to nutrient-enriched patches. However, the grasses of these lawns had significantly higher nutrient concentrations, in their foliage, which might explain the high attraction to these patches by herbivores. We also did not find any animal-induced stimulation of nutrient cycling rates that are often associated with lawn grass species.     

Bush encroachment under three contrasting land-use practices in a mesic South African savanna

This study determined the effects of land-use practice had on the rate and extent of bush encroachment in a mesic savanna in KwaZulu-Natal, South Africa. Changes in woody cover were measured for 1 km² sites in areas under communal, commercial and conservation land-use systems for the period between 1937 and 2000. Land users from each area were interviewed to gain the histories of each area and to determine how the changes in woody cover had impacted them and whether anything was being done to counteract the spread of trees and shrubs on their land. Bush encroachment occurred across all three of the land-use types in the 67-year period between 1937 and 2000. The results showed that land-use practice had enormous impacts on the process of bush encroachment. The communal site showed a decrease in grass (21%) and tree (5%) cover and an increase in shrub cover (13%). At the commercial site, there was a considerable decrease in grass cover (46%) and moderate increase in shrub cover (10%) and a massive increase in tree cover (36%). The area under conservation showed a substantial decrease in grass cover (47%), a slight decrease in shrub cover (19%) and a massive increase in tree cover (66%). The perceived causes of these changes were fairly similar amongst the different land users. The changes were mostly not perceived to be a problem for the communal land users. The main advantages mentioned were increased woody resources for building and firewood and increased browse availability. The commercial and conservation land users perceived the changes to have significant negative connotations including the loss of grazing land and biodiversity and secondary invasion of encroached areas by alien plant species. Despite these perceptions, very little has been done to combat bush encroachment in the commercial and conservation land use systems.     

Maintenance of ladybeetle diversity across a heterogeneous African agricultural/savanna land mosaic

The importance of agricultural land for maintaining indigenous biodiversity is an important contemporary issue. A large agricultural estate in Swaziland, which is a mosaic of citrus orchards, exotic tree windbreaks, indigenous riparian vegetation and savanna was investigated. The distribution of larval and adult coccinellids within the habitat types, and the relative influences of temperature, relative humidity, ground insolation and prey presence was assessed over a 2-year period. The highest coccinellid densities and species richness occurred within the orchards. Riparian habitats on the boundaries of the estate were closer to the orchard habitat in species composition than was the natural savanna. Not surprisingly, the presence/absence of prey was the most important variable which determined coccinellid distribution and density in the agricultural landscape. Relative humidity, temperature and ground insolation had no significant effect on coccinellid densities. Some species, such as the exotic Cheilomenes lunata, and the indigenous Scynmus sp. and Nephus sp. were widely dispersed and occurred in all habitats, while the economically important but exotic species, Rodolia cardinalis was only within the citrus orchards. Overall, the coccinellid assemblage was not a good indicator of habitat type. Nevertheless, a variety of habitat types (i.e. a heterogenous landscape) appear to be essential for the survival of most coccinellids. Indeed, the heterogeneous agricultural and natural land mosaic provides maximum plant diversity and hence coccinellid diversity, thus increasing the potential for natural control of pest prey species in the orchards, while at the same time maximizing survival of indigenous coccinellids.     

Transformation of thicket to savanna reduces soil quality in the Eastern Cape, South Africa

Xeric succulent thicket in the Eastern Cape, South Africa has been used for farming goats since the early 1900s. This habitat is characterised by a dense cover of the succulent bush Portulacaria afra and by a warm, semi-arid climate with evenly distributed annual rainfall of 250–400 mm. Heavy browsing by goats results in the loss of P. afra and transforms the thicket to an open savanna dominated by annual grasses. Eight fence-line comparisons between thicket and savanna were used to investigate differences in soil quality associated with the vegetation change. Composite soil samples were taken to a depth of 10 cm from 1 ha plots on either side of the fence-line. Associated with the change from thicket to savanna, a significant decrease (paired t-test, P < 0.05) was found in total C (respective means of 5.6 vs. 3.0%), total N (0.33 vs. 0.24%), labile C (2.8 vs. 1.5%), CO₂ flux (1.9 vs. 0.5 µmol m^–2 s^–1), soil respiration in the laboratory (144 vs. 79 ng C kg^–1 s^–1), (NH₄)OAc-extractable Mg (55 vs. 28 mmol_c kg^–1), and laboratory infiltration rate (51 vs. 19 mm h^–1). In the same direction there was a similarly significant increase in modulus of rupture (16 vs. 34 kPa), water-soluble Ca (2.3 vs. 3.4 mmol_c kg^–1) and pH (6.7 vs. 7.7). The soil C content of 5.6% in thicket is surprisingly high in this warm, semi-arid climate and suggests that the dense P. afra bush strongly regulates soil organic matter through microclimate, erosion control, litter quantity and perhaps chemistry. Savanna soils had a greater tendency to crust (as evident in a lower rate of laboratory infiltration and greater modulus of rupture) than thicket soils. This was attributed to their lower organic matter content, which probably reduced aggregate stability. Savannas are likely to be more prone to runoff and erosion not only because of sparser vegetation but also because of a decline in soil quality.     

Ecology of spiders inhabiting abandoned mammal burrows in South African savanna

Summary We marked the sites chosen by 338 foundress queens of two desert ant species (Veromessor pergandei and Myrmecocystus flaviceps) and monitored changes in the spacing of both species and the foraging activity at V. pergandei young nests. Although the long established colonies of both species tend intraspecifically toward regular dispersion, queens of both species were intraspecifically clumped. After 3 months, when the first workers emerged, the young colonies (reduced to a total of 42 colonies) were randomly spaced intraspecifically. We also followed the spatial patter of queens with respect to established colonies of both species. Queens founded nests away from the nests of all established colonies on the site. After three months, the young colonies were dispersed away from conspecifics only. During June through August 1986, we censused the number of foragers at the surviving V. pergandei nests. Young colonies that were more active also tended to be far from established conspecifics in July and August. There was no correlation of forating activity with distance to heterospecific established colonies in any month. These results indicate that established conspecifics may reduce the survivorship of young ant colonies.     

Frequent fire affects soil nitrogen and carbon in an African savanna by changing woody cover

When tropical and sub-tropical ecosystems burn, considerable amounts of N present in the biomass fuel may be released. This ultimately results in a loss of fixed N to the atmosphere. It is often assumed that this volatilization loss of N with frequent fire will result in a reduction of plant-available N and total system N. By changing the amount of woody biomass fire may, however, also have indirect effects on N and C dynamics. Here we consider the effects of 50 years of frequent fire on total soil N and soil organic C (SOC) and total soil N in a mesic savanna in the Kruger National Park, South Africa. We also determine how changes in woody biomass may affect total soil N and SOC. We measured soil and fine root N and C concentrations as well as total soil N and SOC pools in four burning treatments, including fire exclusion, of a long-term fire experiment. Our results show that regardless of soil depth, fire treatment had no significant effect on total soil N and SOC. Our results also show that under trees total soil N and SOC concentrations of the surface soil increase, and pools of N and SOC increase to a depth of 7 cm. However, the extent to which soil N and C dynamics differed under canopies and away from canopies was dependent on fire treatment. Our results show that the effect of fire on soil N and C is mediated both through the indirect effect of changes in woody cover and the direct effects of fire (volatilization losses of nutrients). We suggest that woody thickening in this mesic savanna will have pronounced effects on long-term N and C dynamics.     

Community structure and species composition along a disturbance gradient in a communally managed South African savanna

The woody and herbaceous vegetation was investigated around four rural settlements in the savanna area of the eastern Transvaal Lowveld, South Africa. At each settlement three transects radiating out from the settlement, representing a gradient from high to low disturbance, were sampled for community structure and species composition. In general, attributes of woody community structure (density, height, biomass, basal area and diversity) were negatively related to increasing disturbance. Herbaceous cover responded positively. Although individual woody species exhibited a range of responses to disturbance, overall species compositional changes were not related directly to the intensity of disturbance. Individual woody species were classified into behavioural species response groups according to their response along the disturbance gradient. Local wood harvesters demonstrated marked selection for particular species and size classes, which should have disproportional impacts on community structure. However, proportional size class distributions were little altered along the gradient.     

Biomass and production estimates of a fish community in a small South African estuary

The fish community of the small (17·5 ha) intermittently open East Kleinemonde estuary was sampled between 1994 and 1997 to estimate population size, standing stock, growth and productivity. The estuarine-spawning species were numerically more abundant ( n c . 750 000) but due to their small size contributed only 11·7% to the total biomass. The total annual productivity of all fishes in the estuary ( n c . 890 000), with a standing stock of 28·44 g m ⁻², was calculated at 55·89 g m⁻² year⁻¹. The small sparid Rhabdosargus holubi with a production estimate of 41·35 g m⁻² year⁻¹ accounted for <74% of the total fish production in this estuary.     

Seasonal variation in the diet of South African Galagos

The feeding behavior and diet of two species of bushbaby, Galago senegalensisand Galago crassicaudatus,in South Africa were examined in relation to seasonal changes in climate. The populations studied were allopatric, but both fed predominantly on Acaciagum and invertebrates. Data on their diet were collected by direct observation of a radiocollared female of each species and by analysis of fecal samples. Differences in diet were found between the species within seasons and within species between the seasons. Both spent more time gum-feeding in winter than in summer. Prey size and type differed between the species in summer but converged in winter. The quantity of insects taken was fairly similar between seasons for G. senegalensisbut dropped considerably for G. crassicaudatusin winter. These observations are interpreted in terms of interspecific differences in feeding strategies, which are considered to be a function of the difference in body size of the two galago species.     

Leaf gas exchange characteristics and water- and nitrogen-use efficiencies of dominant grass and tree species in a West African savanna

Whereas leaf gas exchange properties are important to assess carbon and water fluxes in ecosystems worldwide, information of this type is scarce for savanna species. In this study, gas exchange characteristics of 2 C₄ grass species (Andropogon canaliculatus and Hyparrhenia diplandra) and 2 C₃ tree species (Crossopteryx febrifuga and Cussonia arborea) from the West-African savanna of Lamto (Ivory Coast) were investigated in the field. Measurements were done in order to provide data to allow the parameterization of biochemically-based models of photosynthesis (for C₄ and C₃ plant metabolic types) and stomatal conductance ; and to compare gas exchange characteristics of coexisting species. No systematic difference was found between grass and tree species for reference stomatal conductance, under standard environmental conditions, or stomatal response to incident light or vapour pressure deficit at leaf surface. Conversely, grass species displayed higher water (1.5-2 fold) and nitrogen (2-5 fold) photosynthetic use efficiencies (WUE and NUE, ratio of net photosynthesis to transpiration and leaf nitrogen, respectively). These contrasts were attributed to the CO₂ concentrating mechanism of C₄ plants. When looking within plant life forms, no important difference was found between grass species. However, significant contrasts were found between tree species, Cussonia showing higher NUE and reference stomatal conductance than Crossopteryx. These results stress the need to account for functional diversity when estimating ecosystem carbon and water fluxes. In particular, our results suggest that the tree/grass ratio, and also the composition of the tree layer, could strongly affect WUE and NUE at the ecosystem scale in West African savannas.