Comparative Impact of Invasive Alien Trees and Vineyards on Arthropod Diversity in the Cape Floristic Region, Western Cape by Rembuluwani Norman Nicholas Magoba Dissertation presented in fulfilment of the requirements for the degree PhD in Entomology at the University of Stellenbosch Promoter: Prof. Michael John Samways Faculty of AgriSciences Department of Conservation Ecology and Entomology & Centre for Invasion Biology December 2010 Comparative Impact of Invasive Alien Trees and Vineyards on Arthropod Diversity in the Cape Floristic Region, Western Cape Declaration By submitting this thesis/dissertation electronically, I declare that the entirety of the work contained therein is my own, original work, and that I have not previously in its entirety or in part submitted it for obtaining any qualification. Rembuluwani Norman Nicholas Magoba (Preferred name: Rembu) Copyright © 2010 Stellenbosch University All rights reserved ii Comparative Impact of Invasive Alien Trees and Vineyards on Arthropod Diversity in the Cape Floristic Region, Western Cape GENERAL ABSTRACT Invasive alien trees (IATs) and agriculture are rapidly altering the capacity of ecosystems to provide a variety of essential services, with decreasing habitat quality having an adverse effect on arthropod biodiversity. Although both agricultural intensification and IATs have an impact on ecosystems, it is unclear which of these two is currently the most severe. Here I compare the influences of vineyards and IATs on arthropod diversity in the Cape Floristic Region (CFR) from two perspectives. Firstly, insect diversity from four different vegetation types (native fynbos, IATs, cleared invasive alien trees (CIATs) and vineyards), and their ecotones were assessed. Secondly, soil compaction in the different vegetation types was assessed and compared to determine how soil compaction relates to arthropod species richness and abundance. Surface-active arthropods were used for these evaluations, as they were considered to be the most abundant, yet sensitive groups for these comparative assessments. Pitfall trapping, which has been widely used for such comparative studies, was the method of choice. Additionally, as a result of incidental wildfires at some of the study sites (IATs and fynbos vegetation), it was decided opportunistically to assess recovery of arthropod diversity following the impact of fire. The results categorically indicate that IATs had by far the most significant impact, in comparison with vineyards, in reducing arthropod diversity and changing assemblage composition. Both IATs and vineyards, not surprisingly, supported lower arthropod diversity compared to fynbos, while CIATs supported arthropod species richness comparable to that of fynbos. However, different invertebrate species responded differently to IAT invasion, with some species even adapted to conditions in IAT patches. Environmental factors such as leaf litter and soil compaction were strongly correlated with arthropod species richness. However, other factors, such as soil moisture content, were not correlated with species richness, despite varying significantly between different vegetation types. IATs were found to reduce soil compaction, while vineyards increased soil compaction. The ecotone between adjacent land-use types was important for sustaining high arthropod species richness, particularly for the CIATs/fynbos ecotone, followed by iii Comparative Impact of Invasive Alien Trees and Vineyards on Arthropod Diversity in the Cape Floristic Region, Western Cape the interface between CIATs/vineyard and vineyard/fynbos. Some species even preferred ecotones. Surface-active invertebrate species richness declined substantially immediately after fire in both IATs and fynbos. Nevertheless, recovery was rapid, with some species recolonizing or re-establishing within three months. Indeed, both species richness and abundance were high following fire. Fire had no significant impact on arthropod assemblage within IATs, despite having a serious impact in the fynbos where a different arthropod assemblage was observed after fire. The recovery of the historic assemblages may be dependent on both the pre-fire composition of the habitat and its adjacent vegetation. Effective co-operation among conservation biologists, farmers, and landowners to actively plan the future of CFR landscape biodiversity is required. iv Comparative Impact of Invasive Alien Trees and Vineyards on Arthropod Diversity in the Cape Floristic Region, Western Cape ALGEMENE OPSOMMING Indringer uitheemse bome (IUB) en landbou bedrywighede is vinnig besig om die kapasiteit van ekosisteme vir die voorsiening van „n verskeidenheid van noodsaaklike dienste, met „n afname in habitatkwaliteit en „n nadelige uitwerking op artropood biodiversiteit, te verander. Alhoewel beide, landbou intensifikasie en IUB‟s, ‟n impak op ekosisteme uitoefen, is dit onduidelik watter van hierdie twee huidig die mees ernstige een is. In hierdie studie vergelyk ek die invloed van wingerde en IUB‟s op artropood diversiteit in die Kaapse Floristiese Streek (KFS) vanuit twee perspektiewe. Eerstens, insekdiversiteit van vier verskillende vegetasietipes (natuurlike fynbos, IUB‟s, verwyderde indringer uitheemse bome (VIUB‟s) en wingerde) en hulle ekotone is bepaal. Tweedens, is grondverdigting in en van die verskillende vegetasietipes ondersoek en vergelyk om te bepaal hoe grondverdigting verband hou met artropood spesies rykheid en volopheid. Oppervlak-aktiewe artropode is vir hierdie bepalings gebruik aangesien hulle beskou word as die mees volopste, maar ook mees sensitiewe groepe vir hierdie vergelykende bepalings. Pitvalvangste, wat alom gebruik word vir sulke vergelykende studies, was die verkose werkswyse. Bykomend, as gevolg van onbeplande veldbrande in sommige studie gebiede (IUB‟s en fynbos plantegroei), is oppertunisties van die geleentheid gebruik gemaak om die herstel van artropood diversiteit na die impak van brand te bepaal. Die resultate toon duidelik aan dat IUB‟s, in vergelyking met wingerde, by verre die mees beduidende impak gehad het deur ‟n verlaging van artropood diversiteit en veranderende groeperingssamestellings. Beide, IUB‟s en wingerde, het, nie onverwags nie, in vergelyking met fynbos, laer artropood diversiteit ondersteun, terwyl VIUB‟s artropood spesies verskeidenheid, vergelykbaar met dié van fynbos, ondersteun het. Verskillende invertebrate spesies het egter verskillend gereageer op IUB indringing, en sommige spesies het selfs by toestande in IUB kolle aangepas. Omgewingsfaktore soos blaarafval en grondverdigting was sterk gekorrelleer met artropood spesies rykheid. Ander faktore, soos grondvoginhoud, was egter nie in ooreenstemming met spesies rykheid nie, ten spyte van ‟n beduidende wisseling tussen verskillende vegetasietipes. IUB‟s het grondverdigting verminder, terwyl by wingerde grondverdigting toegeneem het. v Comparative Impact of Invasive Alien Trees and Vineyards on Arthropod Diversity in the Cape Floristic Region, Western Cape Die ekotone tussen aanliggende grondgebruike was belangrik om hoë artropood spesies rykheid, veral by die VIUB‟s/fynbos ekotoon, gevolg deur die kontaksones tussen VIUB‟s/wingerd en wingerd/fynbos, te onderhou. Sommige spesies het selfs ekotone verkies. Oppervlak-aktiewe invertebraat spesies rykheid het onmiddelik na brand in beide, IUB‟s en fynbos, wesenlik afgeneem. Tog was herstel vinnig, met sekere spesies wat binne drie maande teruggekeer of hulle hervestig het. Inderdaad was beide, spesies rykheid en volopheid, hoog na ‟n brand. Brand het geen noemenswaardige invloed op artropood groeperings binne IUB‟s gehad nie, ten spyte van ‟n ernstige impak in fynbos waar ‟n verskillende artropood groepering na ‟n brand waargeneem is. Die herstel van die historiese groeperings van beide, die voor-brand samestelling van die habitat en sy aangrensende plantegroei, mag afhanklik wees. Effektiewe en aktiewe samewerking tussen bewaringsbioloë, boere, en grondeienaars om die toekoms van KFS landskap biodiversiteit te beplan, word benodig. The above translation of abstract in Afrikaans was done by Prof. Henk Geertsema, a retired entomologist at Stellenbosch University. vi Comparative Impact of Invasive Alien Trees and Vineyards on Arthropod Diversity in the Cape Floristic Region, Western Cape Table of Contents Declaration ....................................................................................... ii GENERAL ABSTRACT ......................................................................................... iii ALGEMENE OPSOMMING .................................................................................... v Table of Contents .......................................................................... vii List of Figures ................................................................................. xi List of Tables ............................................................................... xvii List of Appendices ......................................................................... xx Acknowledgements ..................................................................... xxii Chapter 1 – General Introduction .................................................... 1 Biodiversity crisis ...................................................................................................... 1 Impact of alien organisms ........................................................................................ 1 Impact of agriculture ................................................................................................2