The study was initially a long term study of the ecology of small mammals at Mlawula Nature Reserve in the eastern part of Swaziland. Due to the drought and dwindling numbers of rodents in the reserve the study was redirected to determining the factors resulting in the low numbers. The impacts of food and cover and grazing by the larger ungulates were studied. In addition, the age class distribution and gastro-intestinal parasites of small mammals were studied. A study of the small mammals in the Siphiso Valley of Mlawula Nature Reserve was conducted over four years from August 2000. The population density, biomass and composition of the small mammal community in the area were studied. The community comprised of Mus minutoides A. Smith, 1834, Mastomys natalensis (A. Smith, 1834), Lemniscomys rosalia (Thomas, 1904), Crocidura hirta Peters, 1952, Steatomys pratensis Peters, 1846 and Graphiurus murinus (Desmarest, 1822). Mus minutoides was the dominant species with pregnant females caught from November to May. Species richness varied significantly with the time of the year. The biomass, density and numbers of small mammals were low and by the end of the second year of the study, small mammal density was close to zero. Mastomys natalensis from a Middleveld study site, Luyengo, Swaziland were used to study the age structure of the population by means of eye lenses. The eye lens to age (in days) curve determined by Leirs (1994) was applied. A large percentage of M. natalensis in winter (June) were 2 months old while in spring/summer (October to March) the population consisted mainly of 3 month old specimens. A very low number of specimens were older than 4 months. This suggests a high mortality/removal rate of the young especially in the winter months. The impact of grazing pressure and rainfall on small mammal densities were investigated. High grazing pressure by ungulates rendered the habitat unsuitable for small mammals as it removed cover and encouraged colonization by alien invasive plant species. This effect was exacerbated by diminishing and unpredictable rains, such that mild grazing pressure negatively impacted on small mammal communities and on individual species. When the small mammals disappeared from the study site, M. natalensis was reintroduced to determine the factors that led to the disappearance. Supplementary food resulted in the longest persistence of the reintroduced mice while the impact of additional cover was small. Predation was likely responsible for the rapid decline of the reintroduced mice. Small mammals were examined for ectoparasites and gut parasites as these were thought to negatively impact on their physiology and reproduction. Ectoparasites collected included the ticks Ixodes sp. and Boophilus sp., the mite Allodermanyssus sp. and another species of mite. The gastrointestinal tracts contained the helminths Syphacia sp., Heligmonina sp., Trichuris sp., Protospirura sp., two unidentified nematode species and different cestode species. A new species of heligmosomoid nematode is described and named. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2007.

The changes in plant quality and availability in space and time present a substantial problem to mammalian herbivores. As a result, these herbivores need to alter their foraging behaviour to maximize their energy gain at both small (plant level) and large (landscape level) scales. A megaherbivore, the African elephant (Loxodonta africana), has been shown to be selective in its foraging choices at both of these scales. Furthermore, the ratio of palatability:defences (e.g. fibre and total polyphenols) has been highlighted as an important determinant of habitat selection in elephants. The elephants in Ithala Game Reserve (IGR) frequently leave IGR during the wet season and forage outside the reserve. However, they predominantly feed on the low-nutrient granite soils of the reserve and return to a high-nutrient area with dolerite soils during the dry season. In an attempt to understand these seasonal movements, I focused on how the small-scale foraging decisions of the elephants lead to large-scale seasonal movements in IGR, KwaZulu-Natal. Plant availability was determined seasonally for seven target species across four areas in the reserve. Crude protein, fibre, energy and total polyphenols as well as the ratios of palatability:digestion-reducing substances were analysed in the wet and dry seasons. All factors and their interactions were significant in a MANOVA. Consequently, I employed a dimension-reducing Principal Components Analysis (PCA) to better understand the factors of greatest importance. The PCA highlighted four of the six most important factors to be the ratios of palatability:digestion-reducing substances. The other two important variables were total polyphenols (negative effect) and crude protein (positive effect). At small spatial scales, I found that the elephants were selective in their decisions, especially during the dry season. For example, the increased inclusion of the principal tree species Acacia nilotica from 2.9% in the wet season to 39.3% during the dry season appears to be a result of a decline in total polyphenols and fibre during the dry season. At large spatial scales, the elephants moved back into IGR from the low-nutrient granite soils in the east in response to an increase in forage quality in the west as the quality declined in the east at the same time. However, it is unclear as to why the elephants are leaving the reserve during the wet season. Some possible explanations for this are discussed.Key-words: acid detergent fibre, crude protein, Loxodonta africana, neutral detergent fibre, total polyphenols, plant part quality. / Thesis (M.Sc.)-University of KwaZulu-Natal, Pietermaritzburg, 2013.

As the dry season progresses across southern Africa, the availability and quality of food declines for large herbivores. Female white rhinos compensate for these declines by expanding and/or shifting their home ranges. These changes may be to incorporate habitat types that contain high quality food or quite simply more food. To determine the factors that drive these seasonal changes in home ranges, I focused on dry season changes in the availability and quality of grass in habitats utilised by white rhinos in the Ithala Game Reserve, South Africa. I expected that if food quality was the main driver, white rhinos would follow optimal foraging principles and incorporate habitat types with the highest nutritional quality into their dry season home ranges. Alternatively, due to their large body size (>1000 kg) and thus ability to survive on low quality food, they may rather incorporate habitat types with high food availability. In contrast to previous studies, I found that during the dry season female white rhinos did not increase the size of their home ranges, but rather shifted their home range boundaries. This resulted in individuals increasing the amount of Bushveld and decreasing the amount of Wooded Grasslands within their dry season home ranges. When I explored the different factors that could explain these patterns, I found that changes in the crude protein content of grass was the key factor driving the incorporation and exclusion of habitat types in the home ranges. During the dry season, white rhinos incorporated the habitat that had the smallest seasonal reduction in crude protein content, while excluding the one with the largest decrease in crude protein. As a result, my results suggest that the search for high quality best explains the seasonal home range shifts of female white rhinos in the Ithala Game Reserve. / Thesis (M.Sc.Ecology)-University of KwaZulu-Natal, Pietermaritzburg, 2013.

Pollinators are believed to have played a key role in the radiation of flowering plants. The Grant-Stebbins model of pollinator-mediated speciation, in which evolutionary shifts between pollinators result in phenotypic diversification and enforce reproductive isolation, is one of the most compelling hypotheses for the rapid diversification of angiosperms. A key principle in this model is that plant pollination systems tend towards specialization, resulting in convergent suites of floral traits (syndromes) associated with particular types of pollinators. However, the expectation of pollination system specialization is not always supported by ecological data and has also been questioned on theoretical grounds. In this thesis, I examine pollination by Hemipepsis spider-hunting wasps (Hymenoptera, Pompilidae, Pepsinae) and use this system to address questions about levels and proximal mechanisms of floral specialization, floral shifts and convergent evolution of floral traits. Specialized pollination by Hemipepsis wasps is a newly described pollination system within the angiosperms. I document pollination by these wasps for the first time in 15 South African grassland plant species, including two species of Eucomis (Hyacinthaceae) and 13 asclepiads (Apocynaceae: Asclepiadoideae). In one of the asclepiads, Xysmalobium undulatum, I describe a bimodal pollination system involving both Hemipepsis wasps and a cetoniine beetle. I also describe an unusual and potentially antagonistic pollination mechanism whereby wasps are systematically dismembered during the insertion of pollinia in the two asclepiads Pachycarpus asperifolius and P. appendiculatus. I have used these and previous case studies to establish the existence of a new pollination guild, consisting of at least 21 plant species (across 10 genera and three families), that are reliant on four functionally similar species of Hemipepsis wasp for pollination. Plants in the guild are distributed throughout the moist grasslands of eastern South Africa and flower from September through until early May, peaking in December/January. The Hemipepsis-wasp pollination guild is characterized by high levels of functional specialization (17 of the 21 known guild members are pollinated exclusively by Hemipepsis wasps), despite the absence of morphological adaptations to prevent non-pollinating insects from accessing nectar. I used field and laboratory based experiments to explore the function of floral traits in enforcing specialization. These showed that Hemipepsis wasps primarily use scent, rather than visual cues, to locate flowers, but I was unable to firmly identify specific compounds responsible for the attraction of these wasps (compounds that elicited antennal responses in preliminary GC-EAD experiments did not attract wasps in bioassays). The chemical composition of the floral scents of guild members was examined for 71 individuals representing 14 species in addition to previous studies, and found to comprise complex blends of volatiles (usually containing between 30 and 50 compounds), typically dominated by aliphatics and monoterpenes with small amounts of aromatics. I also showed that the floral colours of guild members are similar to background vegetation, suggesting that floral colours are adapted for crypsis to avoid detection by non-pollinating insects. Palatability choice experiments with honeybees showed that non-pollinating insects find the nectars of at least three of the asclepiad guild members distasteful. Plants in this guild thus appear to achieve specialization through biochemical filters (scent as an attractant and differentially palatable nectar) and cryptic coloration. Pollinator-mediated convergence in floral traits is the fundamental basis for pollination syndromes, but has seldom been rigorously analyzed. Flowers in the Hemipepsis-wasp pollination guild share several qualitative traits, including dull greenish- or brownish-white colour, often with purple blotches, exposed sucrose dominant nectar with a relatively high sugar concentration (typically over 50% sugar by weight) and a sweet/spicy fragrance to the human nose. To test for convergent evolution in guild members, I compared scent, nectar and colour traits of guild members to those of congeners with different pollinators. Although traits often differed between guild members and their congeners, I found little evidence for overall convergence in floral scent profiles and nectar properties, but floral colours in the guild were significantly closer to the colour of background vegetation than those of congeners. At this stage, the lack of knowledge about specific floral volatiles that influence Hemipepsis-wasp behaviour and secondary nectar constituents that limit non-pollinator visits makes it difficult to identify the extent of biochemical convergent evolution within the guild. The directions and functional traits involved in evolutionary transitions between pollination by Hemipepsis wasps and other vectors are currently difficult to ascertain as there is limited phylogenetic data for the plant families concerned. In the genus Eucomis, fly and Hemipepsis-wasp pollinated species are very similar in floral morphology and colour, but differ strongly in floral scent. Using manipulative field experiments in conjunction with detailed analyses of colour, scent and morphology, I was able to show that a shift between wasp and fly pollination could be induced simply by manipulating oligosulphides in the scent emission from inflorescences. When considered in combination with other experiments highlighting the importance of scent as a pollinator attractant for all guild members, this suggests that scent properties may have played a key role in the evolutionary transitions between pollination by Hemipepsis wasps and other vectors. This research has established that pollination by Hemipepsis spider-hunting wasps is more geographically and phylogenetically widespread than was previously known, and has confirmed that these wasps are important and consistent pollinators in southern African grassland ecosystems. I have shown that a distinct guild of plants is specialized for pollination by these wasps. The high levels of specialization within this guild highlight the effectiveness of biochemical filters and cryptic coloration in limiting the spectrum of flower visitors. The major challenge ahead will be to identify the floral volatiles that attract Hemipepsis wasps and the non-sugar constituents that make the nectars of some guild members differentially palatable. These would both contribute greatly to our understanding of floral specialization and the mechanisms involved in the radiation of the angiosperms. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2011.