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The Impact of Land Use on Invertebrate Assemblages in the Succulent Karoo, South Africa

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The impact of land use on invertebrate assemblages in the Succulent Karoo, South Africa. By ‘Makebitsamang Constance Nchai Thesis presented in partial fulfilment of the requirements for the degree of Master of Science (Conservation Ecology) at the University of Stellenbosch. Supervisor: Dr. Cornelia B. Krug Co-supervisor: Prof. M. J. Samways December 2008 Abstract The Succulent Karoo biodiversity hotspot is threatened by pressure caused by increasing human populations and its associated land use types. Land use is primarily focussed on agriculture, with livestock grazing as a dominant land use in the region. Cultivation is also practiced along the major perennial rivers, and in drier areas, where this largely depends on rainfall. Only about seven percent of the biome is formally protected, and this area substantially under-represents the biodiversity of the Succulent Karoo and does not incorporate key ecological processes and biodiversity drivers. Therefore, there is urgent need for outside reserve conservation initiatives, whose success depend on understanding the ecosystem function of the Succulent Karoo. This study aimed to determine the impacts of heavy grazing, light grazing and cultivation (in a 30-year old fallow field) on assemblages of ground-dwelling and flying invertebrates. Seasonal assemblage changes were also determined. Vegetation structure and composition were determined using the line-intercept method to determine if vegetation patterns explain patterns in invertebrate assemblages. Abandoned fields harbour the lowest number of plant species, and these together with the heavily grazed sizes are dominated by a high cover of Galenia africana (Aizoaceae). Lightly grazed sites have the highest structural complexity, with a high cover of succulents and non-succulent perennials. After the winter rains, annual plants occupy most of the bare ground in heavily grazed and previously cultivated sites. Seasonal changes in assemblages of ground-dwelling and flying invertebrates were determined by sampling during the four seasons at the same localities. Results of pitfall traps sampling for ground-dwelling invertebrates and coloured pan traps for flying invertebrates showed that overall species richness and diversity peaked in spring for flying invertebrates while peaks in richness for ground-dwelling invertebrates were in summer, with no difference in overall diversity. Overall abundance for ground-dwelling invertebrates was highest in summer and lowest in winter. Ground-dwelling invertebrate fauna was dominated by Formicidae and Araneae. Grazing and cultivation lead to skewed community composition of ground-dwelling invertebrates which favours disturbance tolerant and generalist species such as Anoplolepis steingroeveri (Forel). The lightly grazed areas favoured the specialised ant taxa and predators. Also, a trend for flying ii invertebrates was that a high number of specialists and a high functional diversity occurred on the lightly grazed sites. Pollinator assemblages differed between seasons. In autumn and spring, the highest richness and abundances of pollinators were found in cultivated and heavily grazed sites. In summer, abundances of pollinators were highest in grazed sites and lowest on the cultivated fields. Flying invertebrate indicator species differ between seasons for each land use types, and these seasonal indicator assemblages are largely dependent on climatic factors while land use indicator assemblages are determined by vegetation structure and composition. Formicidae and Araneae are the only indicators for ground-dwelling invertebrate species. For conservation purposes, the results show that a large number of invertebrate species can be obtained by having a mosaic of different land use types, as these tend to have different seasonal assemblages for different taxa. iii Opsomming Die Sukkulente Karoo bioom se biodiversiteit word al hoe meer bedreig deur die ewig stygende druk vanaf menslike populasies en hul gepaardgaande grondgebruik tipes. Grondgebruik in dié area fokus hoofsaaklik op landbou, waar veebeweiding die dominante grondgebruik is. Gewasproduksie kom wel voor langs al die hoof aanhoudend vloeiende riviere, asook in sommige droër areas, maar laasgenoemde is grootliks afhanklik van reënval. Slegs sowat sewe persent van die bioom word amptelik beskerm, en selfs dié area ervaar wesenlike onderverteenwoordiging van Sukkulente Karoo biodiversiteit en neem ook nie sleutel ekologiese prosesse en biodiversiteit aandrywers in ag nie. Buite-reservaat bewarings inisiatiewe word dus dringend benodig, en die sukses van hierdie inisiatiewe sal afhang van ‘n werklike begrip rondom ekosisteem funksie in die Sukkulente Karoo. Hierdie studie mik om die impakte van swaar beweiding, ligte beweiding en gewasproduksie (in ‘n 30-jaar oue braakland) op versamelings van grondlewende en vlieënde ongewerweldes te bepaal. Seisoenale versameling veranderinge was ook bepaal. Plantegroei struktuur en samestelling was bepaal d.m.v. die lyn-onderskep metode. Hiermee kon bepaal word of patrone in plantegroei die patrone waargeneem in invertebraat versamelings verduidelik. Verlate velde huisves die laagste getal plant spesies, en word, soos die swaar beweide velde, gedomineer deur ‘n hoë bedekking Galenia africana (Aizoaceae). Lig beweide liggings het die hoogste strukturele kompleksiteit gehad, met ‘n hoë bedekking sukkulente asook nie-sukkulente meerjariges. Na winter reëns het eenjarige plante meeste van die kaal grond in swaar beweide velde en voorheen bewerkte velde beset. Seisoenale veranderinge in versamelings van grondlewende en vlieënde ongewerweldes was bepaal deur proefnemings oor al vier seisoene by dieselfde liggings. Resultate verkry van pitval strikke vir grondlewende ongewerweldes asook gekleurde pan strikke vir vlieënde ongewerweldes, wys dat algehele spesies rykheid en diversiteit vir vlieënde ongewerweldes ‘n piek bereik in die lente, terwyl grondlewende ongewerwelde spesies rykheid eers piek in somer, met geen verskil in algehele diversiteit nie. Algehele volopheid van grondlewende ongewerweldes was die hoogste in somer en laagste in iv winter. Grondlewende ongewerwelde fauna was gedomineer deur Formicidae en Araneae. Beweiding en gewasverbouing het gelei tot ‘n skewe gemeenskapsamestelling van grondlewende ongewerweldes – wat steuring tolerante en algemene spesies soos Anoplolepis steingroeveri (Forel) bevoordeel. Die lig beweide areas bevoordeel gespesialiseerde mier taksons en predators. Ook vir vlieënde ongewerweldes was daar ‘n hoë aantal spesialis spesies en hoë funksionele diversiteit op lig beweide velde. Bestuiwer versamelings het verskil tussen seisoene. In herfs en lente was die hoogste spesies rykheid en volopheid van bestuiwers gevind in bewerkte en swaar beweide liggings. In somer was volopheid van bestuiwers die hoogste in beweide liggings, en die laagste in bewerkte lande. Vlieënde ongewerwelde indikator spesies verskil tussen seisoene vir elke grondgebruik tipe, en hierdie seisoenale indikator spesies versamelings is grootliks afhanklik van klimaats faktore, terwyl grondgebruik indikator versamelings bepaal word deur plantegroei struktuur en samestelling. Formicidae en Araneae is die enigste indikators vir grondlewende ongewerwelde spesies. Vir bewarings doeleindes: resultate wys dat ‘n groot aantal ongewerwelde spesies kan verkry word deur ‘n mosaïek van verskillende grondgebruik tipes te hê. Hierdie mosaïek grondgebruik benadering neig om verskillende seisoenale versamelings vir verskillende taksons te huisves. v Acknowledgements Firstly, I wish to thank my supervisor, Dr. Cornelia B. Krug for her unending support, dedication, patience and willingness to help at all times. I am deeply indebted to Prof. Timm Hoffman for identification of plants, Prof. Henk Geertsema for identification of flying insects, Carmen Boonzaaier for ant identification, Patrick Raevell for identification of bugs, Dr. Pia Addison for her comments on my draft version and my co-supervisor Prof. Michael Samways for comments on the draft. It would be unrealistic of me to fail to acknowledge the assistance of Dr. Alicia Timm, Corey Bazelet and Emile Bredenhand for their additional help in identification of insects. The same gratitude also goes to Sara Haveron, for her unending help during field work. I am also grateful to my colleagues; James, Hérve, Elvis, Paul, Pierre and Emelie for continued friendship, help and continued times of laughter. Thanks to Lebohang Sekhokoane and Donald Modiko-Iponga for their logistic support. I thank the Lesotho Government, as well as the South African Department of Science and Technology BIOTA programme, GUN 2070538 to Dr. C. B Krug, for financial assistance. A word of thanks goes also to all Paulshoek community members, for their friendly and warm welcome in Paulshoek. Field assistants and Mariana Lot; thank you very much for many hours of help in the field. Thanks to my parents and my sisters for their support and taking care of the two little ones while I was away. To my husband Tlali, you deserve the biggest word of thanks for your support, patience, understanding and unending words of encouragement and for believing in me leaving the family at the most difficult time of our life. Thanks to Kebitsamang and ‘Matlhaku, who have always wondered where their mum was when they most needed her. Above all, I thank God almighty for helping me to be where I am today. Without him, I could not have made it. vi Table of contents DECLARATION...............................................................................................................
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    UNIVERSITAT DE BARCELONA FACULTAT DE FARMÀCIA DEPARTAMENT DE PRODUCTES NATURALS, BIOLOGIA VEGETAL I EDAFOLOGIA LABORATORI DE BOTÀNICA Sistemàtica i filogènia d’Artemisia i gèneres relacionats: una aproximació citogenètica i molecular amb especial èmfasi en el subgènere Dracunculus Jaume Pellicer Moscardó Barcelona, 2009 UNIVERSITAT DE BARCELONA FACULTAT DE FARMÀCIA DEPARTAMENT DE PRODUCTES NATURALS, BIOLOGIA VEGETAL I EDAFOLOGIA PROGRAMA DE DOCTORAT: MEDICAMENTS, ALIMENTACIÓ I SALUT BIENNI 2004-2006 Sistemàtica i filogènia d’Artemisia i gèneres relacionats: una aproximació citogenètica i molecular amb especial èmfasi en el subgènere Dracunculus Memòria presentada per Jaume Pellicer Moscardó per a optar al títol de Doctor per la Universitat de Barcelona Dr. Joan Vallès Xirau Dra. Teresa Garnatje Roca Jaume Pellicer Moscardó Barcelona, 2009 Aquest treball ha estat desenvolupat gràcies a una beca predoctoral FPI i tres borses de viatge per a estades breus concedides pel Ministeri de Ciència i Innovació. El finançament de les recerques ha anat a càrrec dels projectes dels Ministeris de Ciència i Tecnologia i Ciència i Innovació CGL2004-04563-C02- 02/BOS, CGL2007-64839-C02-01-01/BOS i CGL2007-64839-C02-01- 02/BOS, així com dels suports de la Generalitat de Catalunya a Grups de Recerca de Catalunya 2005/SGR/00344 i 2009/SGR/00439. Als meus pares i a la meva germana, naturalment Bé haguera estat trobar-nos a l’alçada impossible de la poesia de les nostres fonts, rierols i senderes, de la prosa plena i saborosa dels nostres vells màrgens, obagues i marjals, del ritme suau i primitiu dels nostres platjars i dunars, de la rudesa i cruor dels nostres cingles i runars, dels clímax dels nostres cims i abims.