The study was conducted in the Eastern Kalahari sandveld of Botswana on a shrub savanna vegetation type dominated by Terminalia sericea / Boscia albitrunca / Grewia flava and Dichrostachys cinerea woodland. It was initiated in an attempt to determine the impact of livestock grazing within piospheres on soil nutrients, range condition and the influence of season on forage quality and diet composition of livestock. The study was centered around the water points and conducted on both free range grazing and controlled conditions. Measurements on soil and vegetation attributes were recorded at particular points along the transects from the water point. Vegetation and livestock diets were measured seasonally over a period of two years. Chemical analyses of soil revealed the low background fertility of the Kalahari sandveld. The impact of dung and urine on soil chemistry was localized in the area immediate to the vicinity of the water point. Phosphorus, pH and cation exchange capacity were the most responsive attributes to variation along the transect from the water point. High livestock units carried at any particular borehole had an influence on the level of soil nutrient status. Management plans should aim at a more even spread of nutrients by improving the distribution of water points. The zonation of vegetation along the transect from water point reflected the type of management, indicating different class of range condition which can be used in range evaluation and planning. Heavy grazing pressure and trampling in the vicinity of the water point kills sensitive perennial grasses resulting in a zone dominated by annual plants. High amount of available biomass were recorded during summer and autumn and low biomass occurred in spring. The 3 - paddock system produced less biomass compared to other systems, while biomass of palatable species was favored by the 9 - paddock system. Forage utilization was higher following drought years, when grazing pressure was concentrated on reduced forage availability. Utilization of forage was greatest in spring and lowest in summer. Utilization along the transect from water did not taper off until after 4000m from the water point in the free range grazing situation suggesting that forage availability was limiting factor, while in the controlled conditions the influence of grazing tapered off at 1200m. Piosphere size as determined by the distance livestock can travel was greater in the free range grazing management area than in the controlled management conditions. Canopy volumes and leaf dry mass values reflect quantitative variations in the contribution of relatively small number of woody species. In general, both leaf volume and leaf mass decrease with the increase in distance from water due to the reduced plant density. Grewia flava and D. cinerea contributed substantially to the total leaf dry mass within the height below 2m. Leaf dry mass above 2m was largely contributed by Acacia gerrardii, T. sericea and B. albitrunca. Woody species diversity increased with the increase in distance from the water point. High density of G. flava was concentrated to the immediate vicinity of the water points, while species such as Bauhinia petersiana and Croton gratissimus occurred only at further distances from water. There was no clear pattern in the density distribution of D. cinerea along the transect from the water point. Plants exhibit variations in the concentration of nutrients between species and season. High levels of crude protein, phosphorus and low crude fibre content occurred in summer for most species and the opposite was observed during winter or spring. Crude protein and phosphorus during dry periods were believed to be the limiting nutrients in maintaining nutritional quality in grazing animals. Mature forage generally is deficient, and may require supplementation of crude protein or phosphorus. Crude protein, phosphorus and crude fibre were not influenced by the grazing systems. The nutrient enrichment through cattle dung and urine in the vicinity of the water point was reflected in the forage nutrient content. The micro - histological technique proved to be a useful tool for estimating the botanical composition of livestock diets. The technique, however, under-estimates the forbs in the diet of livestock. Diets of cattle were dominated by grasses all year round with a high proportion of woody plants occurred during the spring when available herbaceous biomass was low. Diet of goats was 72% and 82% browse in summer and spring, respectively. Competition for herbaceous plants was high between cattle and sheep. Seasonal species diversity was high in summer and lowest in spring. Goats are more diverse in their diets compared to cattle or sheep which have a strong similarity in their diets. / Dissertation (PhD(Plant Production and Soil Science: Pasture Science))--University of Pretoria, 2002. / Plant Production and Soil Science / unrestricted

This thesis explains two aspects of animal spatial foraging behaviour arising as a direct consequence of animals' need to drink water: the concentration of animal impacts, and the response of animals to those impacts. In semi-arid rangelands, the foraging range of free-ranging large mammalian herbivores is constrained by the distribution of drinking water during the dry season. Animal impacts become concentrated around these watering sites according to the geometrical relationship between the available foraging area and the distance from water, and the spatial distribution of animal impacts becomes organised along a utilisation gradient termed a "piosphere". During the dry season the temporal distribution of the impacts is determined by the day-to-day foraging behaviour of the animals. The specific conditions under which these spatial foraging processes determine the piosphere pattern have been identified in this thesis. At the core of this investigation are questions about the response of animals to the heterogeneity of their resources. Aspects of spatial foraging are widely commented on whilst explaining the consequences of piosphere phenomena for individual animal intake, population dynamics, feeding strategies and management. Implicated are our notions of optimal foraging, scale in animal response, and resource matching. This thesis addressed each of these. In the specific context of piospheres, the role of energy balance in optimal foraging was also tested. Field experiments for this thesis showed a relationship between goat browsing activity and measures of spatial impact. As a preliminary step to investigating animal response to resource heterogeneity, the spatial pattern of foraging behaviour/impacts was described using spatial statistics. Browsing activity varied daily revealing animal assessment of the spatial heterogeneity of their resources and an energetic basis for foraging decisions. This foraging behaviour was shown to be determined by individual plants rather than at larger scales of plant aggregation. A further experiment investigated the claim that defoliation has limited impact on browser intake rate, suggesting that piospheres may have few consequences for browser intake. This experiment identified a constraining influence of browse characteristics at the small scale on goat foraging by relating animal intake rate to plant bite size and distribution. Computer simulation experiments for this thesis supported these empirical findings by showing that the distribution of spatial impacts was sensitive to the marginal value of forage resources, and identified plant bite size and distribution as the causal factors in limiting animal intake rate in the presence of a piosphere. As a further description of spatial pattern, piospheres were characterised by applying a contemporary ecological theory that ranks resource patches into a spatial hierarchy. Ecosystem dynamics emerge from the interactions between these patches, with piospheres being an emergent property of a natural plant-herbivore system under specific conditions of constrained foraging. The generation of a piosphere was shown to be a function of intake constraints and available foraging area, whilst piosphere extent was shown to be independent of daily energy balance including expenditure on travel costs. A threshold distance for animal foraging range arising from a hypothesised conflict between daily energy intake and expenditure was shown not to exist, whereas evidence for an intermediate distance from water as a focus for accumulated foraging activity was identified. Individual animal foraging efficiency in the computer model was shown to be sensitive to the piosphere, while animal population dynamics were found to be determined in the longer term by dry season key resources near watering points. Time lags were found to operate in the maintenance of the gradient, and the density dependent moderation of the animal population. The latter was a direct result of the inability of animal populations to match the distribution of their resources with the distribution of their foraging behaviour, because of their daily drinking requirements. The result is that animal forage intake was compromised by the low density of dry season forage in the vicinity of a water point. This thesis also proposes that piospheres exert selection pressures on traits to maximise energy gain from the spatial heterogeneity of dry season resources, and that these have played a role in the evolution of large mammalian herbivores.