D.Phil. / What underlies species distribution and species coexistence has long been of key interest in community ecology. Several methods and theories have been used to address this question. However, it still remains a controversial debate. The recent development of plant DNA barcodes with possibility of merging phylogeny with ecology brings high expectation in uncovering the processes underlying community assemblages. Previous works that used molecular approach in community ecology focused mainly on rainforests. Using a phylogenetic approach, this study brings novel understandings about savanna ecology, especially regarding how megaherbivores impact plant community composition. The Kruger National Park (KNP) is one of the world’s largest reserves, but less studied from a phylogenetic perspective. A DNA database of 445 DNA sequences (plant DNA barcodes, rbcLa + matK) was generated for the woody plants of the KNP. This database proves reliable in reconstructing the phylogeny of Angiosperms of the park. Based on this phylogeny, the present study characterised plant community composition, and investigated how megaherbivores influence this composition. Results indicate that plant communities in the KNP are not neutral, i.e. they are more clustered than expected under various null models. This suggests that ecological forces, most likely habitat filtering may be playing key role in dictating community structure in the KNP. The KNP is well-known for its richness in megaherbivores. The contribution of these animals to the current shape of plant community structures was therefore further investigated. Where megaherbivores have been excluded, plant diversity decreases, but shifts in plant community structure are contingent upon the initial community composition, suggesting that herbivory might be important filter that drives the clustering pattern observed.

A Dissertation submitted to the Faculty of Science in fulfilment of the requirements for the degree: Master of Science In School of Animal, Plants and Environmental Sciences, University of Witwatersrand Johannesburg 2015 / This study aimed to determine the role and relative importance of predator avoidance and nutrition as processes influencing herbivore utilisation of burnt areas, at Satara, Kruger National Park by using variation in fire size and fire season. The driving factors investigated included forage quality, forage quantity, vegetation structure and predation risk. The effect of fire size and season on the landscape was tested by recording grass height, greenness, grass nutrient sampling and visibility. Results showed that fire size and season had a significant impact on the physical environment by decreasing grass/forage quantity post-fire but increasing grass/forage quality. Over time the plots returned to near pre-fire conditions, with forage quantity increasing (grass height and biomass) and forage quality decreasing (nitrogen availability). Visibility increased immediately post-fire, more significantly in regards to herbivores with their heads in a “head-down/grazing” position than in a “head raised/vigilant” position. As with forage quantity and quality, visibility returned to near pre-fire conditions at the end of the study sampling period. The response of animals to these changes in the environment was recorded through dung counts, camera traps and behavioural observations. I found that there was increased use of burnt plots post-fire through comparing herbivore presence on plots burned in different sizes and seasons, day/night utilisation of plots, and using behaviour data I could determine to what extent forage and predation risk were driving the use of burnt areas. I focused on three herbivore species (Aepyceros melampus, Connochaetes taurinus and Equus quagga) which varied in terms of body size digestive systems, forage type, and social behaviour. As expected, I found that predation risk did appear to be more important in driving the behaviour of the smaller-bodied herbivores but both nutrient requirements and susceptibility to predation were shown to play a role in explaining the use of burnt areas for all three herbivore species. Variation between plots in environmental factors such as tree density, forb percentage and distance to water, were found to be not significant and thus did not confound our results

Thesis (M.Sc.)--University of the Witwatersrand, Faculty of Science, 1993.

How so many species coexist while utilizing the same resources is both of ecological interest and important for the management of wildlife communities and parks. This thesis endeavours to understand how grazing herbivores co-exist, with special emphasis on understanding the mechanisms of competition and facilitation over temporal and spatial scales. I compared the dry season use of grasslands, grass species, grass height and grass greenness by white rhino and three other ungulate species, zebra, wildebeest, and impala. I was specifically interested in the extent to which white rhinos, with their capacity to graze both short and tall grass, either competed with or facilitated other grazers. In Chapter Two, I clarify the mechanisms of facilitation and competition in terms of temporal and spatial scales, and discuss why there is little evidence in the literature to support these mechanisms. I conclude that evidence for facilitation through stimulation of grass re-growth during the growing season appears stronger than that for increased resource access through removal of obstructing grass structures during the dormant season. Facilitation may benefit the nutritional gains obtained by certain species in the short term, but these benefits do not appear to be translated into the expected population consequences. In collaboration with co-author Norman Owen-Smith, we suggested this could be due to seasonal tradeoffs between facilitation and competition, as well as to restrictions on the spatial extent of trophic overlap. In Chapter 3 Norman Owen-Smith and I compared the grass height use in relation to body size. We expected that the grass height favoured would increase with the body size of the herbivore species, as suggested from past studies of resource partitioning among large mammalian herbivores. Instead we found that the largest of these species, white rhino, concentrated on the shortest grass, while the smallest species, impala, favoured grass heights intermediate between those grazed by wildebeest and zebra. Results suggest that the scaling of mouth width relative to body size, and hence metabolic demands, is the primary factor governing grass height selection, rather than body size alone. Hence grazing successions governed by body size differences may not be a typical feature of their ecology, contrary to past suggestions. Furthermore, there was considerable overlap in grass height grazed among these four species, indicating that niche separation by grass height is inadequate alone to explain their coexistence. More attention needs to be paid to other aspects such as the grass species selected and habitat structure favoured. Chapter Four compared the overlap in grassland use and grass species use, as well as grass height and grass greenness of swards utilized by the herbivores as the dry season advances. I show that all species prefer grazing lawns during times of abundance, and that zebra leave grazing lawns before other species, and wildebeest leave grazing lawns before white rhino and impala. This suggests zebra and wildebeest may be competitively excluded from grazing lawns through a reduction in grass height, by white rhino and impala, during the dry season. However, white rhino are also potentially the ‘supreme’ facilitator increasing the availability of nutritious grazing lawns, as well as increasing the quality of those lawns through grazing in the wet season. A better understanding of the trade-off between “habitat facilitation” and competitive exclusion by white rhino, allows us to better understand how grazing herbivores co-exist.

No Estado de São Paulo, a Coordenadoria de Defesa Agropecuária CDA, Unidade da Administração Direta, subordinada à Secretaria Estadual da Agricultura, foi criada em 1998. Subseqüentemente a elevada freqüência de surtos de raiva paralítica em herbívoros na parte leste do Estado, incluindo as regiões do Vale do Paraíba e Grande São Paulo, entre 1999 e 2001, obrigou mudanças nas estratégias de controle para a doença, que vinha sendo adotada anteriormente no Estado. Em 2001, a vacinação dos herbívoros domésticos foi declarada obrigatória nas regiões sobre a circunscrição dos Escritórios de Defesa Agropecuária EDAs, localizados nas áreas classificadas como áreas epidêmicas e endêmicas para a raiva e com o reforço das medidas de controle direto contra o Desmodus rotundus, morcegos vampiros, utilizando redes de neblina e vampiricidas (pasta nticoagulante), além do registro e do georreferenciamento para a localização dos abrigos, nas regiões estratégicas em ordem contrária a progressão da doença, causando a redução da população de morcegos vampiros e da circulação viral. Neste estudo, foi considerado o número de herbívoros com raiva com confirmação laboratorial e de acordo com o ano de ocorrência, espécie animal, distribuição por EDAs e por Municípios, providenciados pelo Instituto Pasteur de São Paulo. Com respeito à data de 2001, as porcentagens de vacinação, foram agrupadas sem as divisões por EDAs, devido à ausência de dados. A partir de 2002, os dados foram divididos por EDAs, onde a vacinação foi declarada obrigatória e os dados foram discriminados entre estabelecimentos que vacinaram e herbívoros vacinados contra raiva. O número de morcegos vampiros capturados agrupados por EDAs. Analisando as tendências, indicando os estabelecimentos que vacinaram, herbívoros vacinados contra raiva e a quantidade de morcegos vampiros capturados, foi observado um decréscimo nos casos de raiva em 90% ou mais e com estes resultados, concluiu-se que para o controle da raiva nos herbívoros, as medidas adotadas devem ser focadas, primeiramente na vacinação das espécies alvo e no bom planejamento e organização das medidas de controle diretamente nos principais reservatórios de Desmodus rotundus. / In the State of São Paulo, the Coordenadoria de Defesa Agropecuária-CDA, an administrative unit directly subordinated to the State Secretary of Agriculture was established in 1998. Subsequently, the frequent outbreaks of paralytic rabies in herbivores in Eastern part of the State, including the region of Vale do Paraiba and the Greater São Paulo, between 1999 and 2001, have led to change the strategies for controlling the disease being conducted up to that time in the State. In 2001, the vaccination of domestic herbivores was declared obligatory in regions under the jurisdiction of Escritórios de Defesa Agropecuária - EDAs located in areas classified as rabies epidemic and endemic, and by the reinforcement of the control measures directed against the Desmodus rotundus vampire bats by means of joint effort of EDAs to capture and control the bats using mist nets and the vampiricides (anticoagulant paste), besides the registration, revision and the use of geographical information systems to locate the bats roosts and shelters in strategic regions, in order to interrupt the progression of the disease through the reduction of the bat population density and the viral circulation. In this study, it was considered the number of rabies in herbivores with laboratorial confirmation and according to the year of occurrence, animal species, and distributed by EDAs and by municipalities, provided by the Pasteur Institute-São Paulo. Data regarding 2001, the percentages of vaccination were grouped without the division into EDAs, due to the absence of data. From 2002, the data were divided into EDAs where vaccination has been declared obligatory, and data discriminated into properties that vaccinated and herbivores vaccinated against rabies. The number of captured vampire bats was provided by the respective EDAs. By analyzing the trends indicated by the properties that vaccinated against rabies and the herbivores vaccinated and the amount of vampire bats captured, it was observed a decrease in rabies cases in 90% or more, and with these results we conclude that for the control of rabies in herbivores, the easures adopted must be focused primarily in vaccinating the target animal species and in a well planned and organized measures of control directed to the main reservoirs, i.e., the Desmodus rotundus bats.

This study investigated the effects of a seasonally variable forage resource on herbivore population dynamics. This involved estimating the relative importance of environmental conditions, and the accessible and used forage resources, at different stages of the seasonal cycle to herbivores in different life-stages and at different points in the reproductive cycle. This study was carried out in the Richtersveld region in South Africa, using goats kept by semi-nomadic Nama pastoralists. In the main study site, the Richtersveld National Park (RNP), herd movements follow a general seasonal migratory pattern: herds are based in the riparian zone of the Orange River during the dry season, and on plains away from the river in the wet season. Over 800 uniquely marked female goats in three life-stages (adults, yearlings and kids) were monitored over a three year period (2007 to 2009). These goats were weighed at 2 - 3 month intervals to provide an estimate of body condition. Browse availability in the riparian zone was estimated using measurements at an individual branch-level and a whole tree-level. FPAR satellite imagery was used to estimate forage abundance outside the riparian zone. Goat density was mapped for each week of the study using census data and the herd positions. Goat body condition, survival rates and fecundity rates for each life-stage were modelled as a response to forage availability, density and climatic conditions. The riparian zone in the RNP was found to function as the key resource of the RNP goat population. Forage depletion by goat browsing resulted in a negative feedback on goat body condition. This decline in body condition was directly related to lower adult survival over the dry season. Fecundity was also most influenced by dry season conditions through the negative effect of poor body condition on pregnancy rates and birth rates. Asymmetric competition between life-stages, resulting from the riparian browse profile being depleted from the bottom-up, was predicted to have a strong effect on goat demography by contributing to differences in body condition and survival rates between life-stages. Wet season conditions appeared to have little effect on goat population dynamics, either through increased neonate survival or through a mass carry-over effect influencing dry season survival. Goat body condition and vital rates were compared between the RNP and the neighbouring Kuboes rangeland, which does not have access to the Orange River, to assess the impact of differences in their dry season forage resource. The long-term size and variability of the livestock population in the RNP was also compared with livestock dynamics in Paulshoek, a rangeland 250 km south east of the RNP. The a priori predictions of relative population dynamics in each region, based on perceived differences in the nature of the key resource in each region, were largely supported.

A thesis submitted to the Faculty of Science, University of the Witwatersrand, in fulfilment of the academic requirements for the degree of Doctor of Philosophy. August 2016, Johannesburg / Termites are widely distributed in tropical and subtropical savanna. They are recognised as major ecosystem engineers through their role in nutrient cycling, decomposition, hydrology and alteration of landscape topography with cascading effects manifesting in ecosystem heterogeneity and productivity up the food chains. In this thesis I addressed the effect of geology on termite species diversity, followed by questioning how the different geologies influence the size and spatial distribution of Macrotermes mounds. Furthermore, I explored the effect of termite mounds emanating from different geologies on herbaceous vegetation heterogeneity and finally the effect this heterogeneity has on grazing intensity. Although the diversity of termites has been explored across different environmental gradients such as rainfall, altitude and disturbance, little is known regarding variation in their diversity across landscapes of varying geology. In my quest to understand how varying geology influences the ecology of termites and their functional importance, I sampled granite and basalt for termite diversity using standard transects (100 m x 2 m). I predicted that termite diversity is higher on nutrient-rich geology following the productivity diversity hypothesis. However, both functional and taxonomic diversity were higher on nutrient-poor granite. Twelve species from three subfamilies representing two feeding groups were recorded on granite whereas on basalt only five species from two subfamilies consisting of one feeding group were recorded. Although the influence of Macrotermes mounds on ecosystem heterogeneity has been well studied, little is known on how the environment (geology) and other termite colonies influence size and distribution pattern, despite how these interactions could influence ecosystem functioning. Termite mounds were sampled in 1 km2 plots, four in each geology. Each mound location was recorded using a hand held GPS and structural variables (height and diameter) measured. The data were analysed for spatial distribution of termite mounds using the software Programita. The general distribution pattern of termite mounds (active and inactive mounds combined) was investigated using both the pair correlation function, g(r), and Ripley’s K(r) function. Termite mounds were larger and covered a significant proportion of the landscape on granite compared to basalt. Mounds were generally over-dispersed on granite and randomly distributed on basalt. Mounds covered ~ 6% of the landscape on granite compared with only ~ 0.4% on basalt. These results show that the significance of termites varies across geologies, being more important on nutrient-poor geologies because of their size and a more productive spatial pattern displayed here. The majority of studies testing mound effects on savanna vegetation spatial heterogeneity have been based on single site observations mostly comparing mounds and their paired savanna control plots. Furthermore studies did not consider the spatial effects of mounds with distance into the savanna matrix from mound edge, and this has rarely been tested across landscapes of varying geologies, as well as across mounds of different sizes. Therefore there was a need to explore this in order to broadly understand the functional importance of mounds. I sampled the herbaceous community on and off termite mounds and along distance transects from mounds on nutrient-rich and nutrient-poor geologies. Termite mounds as sources of spatial vegetation heterogeneity was more pronounced on nutrient-poor granite, with larger mounds having greater effect on vegetation composition and diversity than smaller ones. Mounds harboured compositionally different herbaceous plants compared with the savanna matrix on granite whereas there was no difference on basalt. In acknowledging the effect erosion from mounds may have on vegetation heterogeneity, termite mound effect on composition expressed at landscape level based on mound densities recorded in this study was estimated to be 19% of the landscape on granite whereas on basalt, the mounds influenced ~ 0.4% of the landscape. The choice of foraging sites by large herbivores in the landscape is influenced by food quantity, quality, inter and intra-specific competition and predation risk. Termite mounds harbour highly nutritious herbaceous plants compared to the savanna matrix, which makes them preferred foraging sites. Due to very small differences in soil nutrient content between mounds and savanna on basalt, mounds were expected to have little effect on grazing. In line with the set hypothesis termite mounds largely influenced grazing on the nutrient-poor granite and when viewed at landscape scale, based on mound densities and extent of erosion recorded, mounds influenced ~ 28% on granite and only ~ 0.8% on basalt. Overall my study has demonstrated that the significance of termites as ecosystem engineers varies across landscapes of varying geology, being more important on nutrient-poor compared with nutrient-rich geologies. / MT2017

Herbivore abundances are determined by a set of interacting factors that vary among different habitat types. Specifically, herbivore abundances in monocultures and polycultures may be governed by the same set of factors but with varying influences in the different habitats. In addition, monophagous and polyphagous herbivores may respond differently to the same set of influencing factors. I examined several abiotic and biotic factors in manipulated monocultures and polycultures of Borrichia frutescens in a west central Florida salt marsh. The experimental plots differed in both plant diversity and aboveground habitat structure to see how each component of diversity contributed to variability in the abiotic and biotic factors and how those factors were related to differences in herbivore abundances. The monoculture treatment involved clipping all above ground non-host plant material to achieve a host plant monoculture. The polyculture treatments involved pinning all non-host plant material to achieve a polyculture with reduced above ground habitat structure. The second polyculture treatment was a control in which the naturally diverse plots were unmanipulated. Two monophagous and one polyphagous herbivores were chosen for this study because of their abundance and availability in the field. The two monophagous herbivores on the host plant Borrichia frutescens were Pissonotus quadripustulatus (Homoptera:Delphacidae) and Asphondylia borrichiae (Diptera: Cecidomyiidae) both of which have been well studied in the field where the current experiment took place. The polyphagous herbivore was Cyarda acutissima (Homoptera: Flatidae), a poorly known invasive from Cuba. Soil salinity and host plant leaf nitrogen content were the abiotic factors measured. Herbivore abundances, percent egg and gall parasitism by parasitoids, spider abundances on host plant stems and ground spider abundances were the biotic factors measured. Both salinity and host plant leaf nitrogen were significantly different among the different treatments with clipped plots having the highest salinity and leaf nitrogen content. Population densities of both of the monophagous herbivores were not significantly different between treatments. The polyphagous herbivore had significantly higher abundances in the pinned and control plots than in the clipped plots. Stem spider abundances were not significantly different among treatments. Ground spiders, however, were significantly more abundant in control and pinned plots than clipped plots. Parasitism of both monophagous herbivores was not significantly different between treatments but was generally higher in the control plots. The results suggest that for monophagous herbivores bottom-up and top-down factors act antagonistically in monocultures but for the polyphagous herbivore, the presence of multiple host plants is more influential in diverse plots even given the higher abundances of generalist predators.

Interactions between cyanobacteria and herbivorous grazers play an important role in mediating the responses of freshwater phytoplankton assemblages to nutrient enrichment and top-down manipulation. Negative consequences associated with these interactions include dangerous blooms of harmful blue-green algae that have been implicated in the sickness and death of fishes, livestock, and, in extreme cases, humans. Frequently cited mechanisms influencing the interactions between grazers and cyanobacteria include cyanobacterial toxicity and morphology. To tease apart the importance of these mechanisms, I used meta-analysis to quantitatively synthesize the available literature on this topic. In addition, I conducted several experiments using novel techniques to determine the effect that cyanobacterial secondary metabolites from the bloom-forming cyanobacterium,

Thesis (doctoral)--Swedish University of Agricultural Sciences, 2003. / Appendix includes reprints of five manuscripts, four co-authored with others. Includes bibliographical references. Also partially issued electronically via World Wide Web in PDF format; online version lacks appendix.