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Arthropod and Plant Diversity in Maize Agro-Ecosystems of South Africa

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Arthropod and Plant Diversity in Maize Agro-Ecosystems of South Africa Arthropod and plant diversity in maize agro-ecosystems of South Africa M Botha 21044082 Thesis submitted for the degree Philosophiae Doctor in Environmental Sciences at the Potchefstroom Campus of the North- West University Promoter: Prof SJ Siebert Co-promoter: Prof J van den Berg May 2017 PREFACE Studying the diversity of living organisms is truly a humbling experience. Nowhere is this complexity more apparent than in two of the largest and most ecologically important groups of organisms in terrestrial ecosystems, vascular plants and their associated invertebrates. It is estimated that at least half of all recorded macroscopic species worldwide are plants and the arthropods that feed on them. All we can hope to accomplish with this work is to present a mere snapshot in time and space of the dynamic plant and invertebrate communities living at the crop-rangeland interface, an immensely complex system governed by both anthropogenic and natural factors. Nevertheless, with this and on-going future research we may systematically build a more complete picture of anthropogenic impacts on the natural habitat and in doing so, ensure a more sustainable and diverse agricultural landscape. Naturally, this study would not have been possible without several individuals and organisations which I would like to thank for their exceptional contributions towards the completion of this thesis. Firstly, I want to thank my supervisors, Prof. Stefan Siebert and Prof. Johnnie van den Berg for their inspiration, guidance and tireless dedication to this work. I also thank our statistician, Dr. Suria Ellis for conducting the majority of our statistical analyses as well as her valuable advice and assistance throughout this project. Also, a special thanks to Dr. Niels Dreber (University of Göttingen, Germany), Dr. Frances Siebert (North-West University) and Prof. Braam van Wyk (University of Pretoria) for valuable discussions on the topic of plant functional traits and assistance with statistical analyses. Of course, a fundamental aspect of this study was the identification of an overwhelming number of collected specimens. As such, the Pretoria National Herbarium (PRE) is acknowledged for assistance in the identification of unknown plant specimens as well as the Biosystematic division of the Agricultural Research Council (ARC) and the Ditsong Museum in Pretoria for assistance in the identification of arthropod predators. A special thanks to Prof. Ansie Dippenaar-Schoeman and her team for the tedious task of identifying the myriad of spider specimens to species level. Also, I would like to thank all my friends, family and colleagues for their moral support, encouragement and keeping me (reasonably) sane throughout the ups and downs of this journey. A special thanks to Marié du Toit for designing our study area maps and to Bheki Maliba, Dennis Komape and Bianca Greyvenstein for their significant contributions towards data collection in the field. Finally, the financial assistance of the National Research Foundation (NRF) towards this research is hereby acknowledged. Opinions expressed and conclusions arrived at, are those of the author and are not necessarily to be attributed to the NRF. Additional financial support was provided by GenØk-Centre of Biosafety, Norway, Norad project GLO-3450. i Declaration I, Monique Botha, declare that the work presented in this PhD thesis is my own work, that it has not been submitted for any degree or examination at any other university, and that all the sources I have used or quoted have been acknowledged by complete reference. ______________________________________________ Me. M. Botha (Student) ______________________________________________ Prof. S.J. Siebert (Supervisor) ______________________________________________ Prof. J. van den Berg (Co-supervisor) ii ABSTRACT Agricultural intensification in the twentieth century has led to rapid biodiversity decreases on farmland. In sub-Saharan Africa, where rapid population increases and high direct dependence on natural resources coincide, biodiversity loss due to land-use change is of particular concern. Stock-grazing and dryland crop agriculture are two prominent and growing land-uses in the Grassland and Savanna Biomes of South Africa. Maize (Zea mays L.) represents the most important grain crop, with an approximate annual production of 128 million tons of maize grain on approximately 31 million hectares of land. Understanding what effect farmland management regimes have on the complexity and interactions of biota in remnant semi-natural ecosystems is a necessary step towards a sustainable future for biodiversity in agro-ecosystems. However, there has not been a considerable effort to understand the effects of these agricultural disturbances on species, structural, or functional diversity in South Africa’s grassy biomes. The research project described in this thesis aimed to address the knowledge gap regarding biodiversity of maize agro-ecosystems in the Grassland and Savanna Biomes of South Africa by providing insight into the observational patterns of taxonomic and functional diversity, compositional structure and diversity relationships of two major groups of biota (vascular plants and plant-associated arthropods) in relation to an agricultural disturbance gradient at regional and local scales. Surveys were conducted in six provinces of South Africa, namely North-West, Mpumalanga, KwaZulu-Natal, Limpopo, Free State and the Eastern Cape. The transformation of semi- natural grassland and savanna into maize fields resulted in severely decreased species diversity, functional diversity and abundance as well as marked changes in species composition of plants and arthropods. However, there was no evidence for reduced levels of species diversity, functional diversity or trait abundance of plants and arthropods at medium disturbance intensity marginal vegetation (30-100 m from the maize field edges) compared to low-disturbance intensity rangelands. The pattern was consistent across the Grassland and Savanna Biomes. This suggests that the possible disturbance effects of maize fields do not have considerable negative effects on either the diversity or species assemblages of plant and arthropod communities at ≥30 m from the area of active cultivation. Uncultivated semi-natural vegetation of the Grassland and Savanna Biomes had distinct arthropod assemblages although these distinctions were better explained by geographical position than by plant features such as tree and grass cover. There was also evidence for positive relationships between low-growing (>2m) plant species and arthropod richness, diversity and abundance in maize fields and in uncultivated vegetation. The patterns recorded in this study suggest that crop field margins ≥ 30 m from the site of active cultivation are valuable conservation sites for the continued persistence of beneficial species and functional diversity of non-crop plants and arthropods within the agricultural environment. Key terms: agricultural disturbance gradient; alpha-diversity; beta-diversity; corn (Zea mays); functional diversity; Grassland; insects; plant-arthropod diversity relationships; Savanna iii OPSOMMING ʼn Toename in intensiewe kommersiële landbou-bestuurspraktyke in die twintigste eeu het gelei tot 'n verlies aan biodiversiteit in landbou-ekostelsels. Hierdie grootskaalse verlies aan biodiversiteit is veral prominent in sub-Sahara-Afrika as gevolg van die gekombineerde effekte van eksponensiële bevolkingsgroei en direkte afhanklikheid van natuurlike hulpbronne. Die Grasveld en Savanna Biome van Suid-Afrika word merendeels benut vir veeproduksie en gewasverbouing. Mielies (Zea mays L.) word beskou as die belangrikste graangewas, met ʼn jaarlikse opbrengs van ongeveer 12 miljoen ton wat op ongeveer 2,5 miljoen hektaar landbougrond geproduseer word. Om ʼn volhoubare toekoms vir biodiversiteit in landbou- ekosisteme te verseker, is dit noodsaaklik om kennis te dra van die effek van landbou-bestuurspraktyke op die kompleksiteit en interaksies van biota in oorblywende natuurlike habitatte. Tot dusver is daar is egter nog geen betekenisvolle poging aangewend om die gevolge van landbou-versteurings vir spesie-, en funksionele diversiteit in Suid-Afrika te bestudeer nie. Die navorsingsprojek wat uiteengesit is in hierdie proefskrif was gemik daarop om hierdie gaping in die literatuur aan te spreek deur inligting te verskaf aangaande die diversiteitspatrone (spesie asook funksioneel) en spesiesamestellings van plante en insekte langs ʼn mielieland-bufferstrookgradiënt, eerstens tussen ses verskillende provinsies in en tweedens tussen twee verskillende biome (grasveld en savanna). Opnames is gedoen in die Noordwes-, Mpumalanga-, KwaZulu-Natal-, Limpopo-, Vrystaat- en Oos-Kaap provinsies van Suid-Afrika. Resultate het daarop gedui dat die getal individue, spesiediversiteit en funksionele diversiteit van plante en insekte beduidend laer was in mielielande as in die aangrensende onbewerkte natuurlike veld, en dat die spesiesamestelling ook aansienlik verskil het tussen hierdie twee habitatte. Daar was egter geen aanduiding dat die teenwoordigheid van mielielande ʼn beduidende negatiewe effek gehad het op die getal individue, spesiediversiteit of funksionele diversiteit van plante en insekte in die bufferstrook (30-100 m vanaf die mielielande) nie. Beide die Grasveld en Savanna Biome het dieselfde patrone getoon. Hierdie resultate dui daarop dat die effek van versteurings geassosieer met die landbouaktiwiteite waarskynlik nader as 30 m van die
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