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Drivers of Macrophyte Assemblages in South African Freshwater Systems THESIS Submitted in fulfilment of the requirements for the degree DOCTOR OF PHILOSOPHY at Rhodes University By Grant Douglas Martin January 2013 i Abstract Abstract Potentially damaging submerged invasive freshwater macrophytes have been identified in South African freshwater systems, but have received less attention than their floating counterparts. To ascertain the changes and effects that these species may have on macrophyte ecology, an understanding of the drivers of macrophyte assemblages is essential. The aims of this thesis were to investigate select abiotic and biotic factors driving introduction, establishment and spread of submerged macrophytes in South Africa. Surveys on the status of submerged plant species in South Africa were conducted to find out the distribution and diversity of the species present, imported to, and traded in South Africa. Numerous submerged indigenous and invasive macrophyte locality records were collected during field surveys, of which many were first time records. Pet stores and aquarist trading activities were identified as potential vectors for the spread of submerged macrophytes through online surveys and personal interviews. These results highlighted the potential these species have for continuing to enter, and spread within South African water bodies. Maximum Entropy (MAXENT) is a general-purpose method used to predict or infer distributions from incomplete information, and was used here to predict areas suitable for the establishment of five of these invasive macrophytes. Many systems throughout South Africa, particularly those in the subtropical coastal regions, were found to be climatically suitable for the establishment of Elodea canadensis Michx., Egeria densa Planch., Hydrilla verticillata (L.f.) Royle (all Hydrocharitaceae), Myriophyllum spicatum L. (Haloragaceae), and Cabomba caroliniana Gray (Cabombaceae). i Abstract Despite the high probability of invasion, facilitated by vectors and suitable climate, South Africa’s rich indigenous submerged aquatic flora may be preventing the establishment of these submerged invasive species. Studies on the competitive interactions between a common indigenous submerged macrophytes, Lagarosiphon major (Roxb.) (Hydrocharitaceae) and M. spicatum, an invasive native to Eurasia, were conducted to ascertain which conditions influence competitive superiority. High sediment nutrient conditions significantly increased the growth rate and competitive ability of both species, while clay sediments significantly increased the competitive ability of L. major over M. spicatum, but sandy sediments improved the competitive ability of M. spicatum. These results highlighted the dynamic changes in competition between submerged species driven by abiotic factors, but did not take into consideration the effect that herbivory, a biotic factor, could have on competition between the two species. The effect of herbivory by phytophagous insects of submerged plant species has been regarded as negligible. To find out what this effect is, multiple field surveys were undertaken throughout South Africa to find natural enemies of indigenous Lagarosiphon species with the aim of identifying such species, and quantifying their influence on plant growth dynamics. Several new phytophagous species were recorded for the first time. An ephydrid fly, Hydrellia lagarosiphon Deeming (Diptera: Ephydridae) was ascertained to be the most ubiquitous and abundant species associated with L. major in South Africa. The influence of herbivory by this fly on the competitive ability of L. major in the presence of M. spicatum was investigated using an inverse linear model, which showed that herbivory by H. lagarosiphon reduced the competitive ability of L. major by approximately five times in favour of M. spicatum. This study served to highlight the importance of herbivory as a driver of submerged aquatic plant dynamics. ii Abstract Current ecological theory emphasises the importance of investigating beyond plant-herbivore interactions, by including multitrophic interactions in community dynamics. Therefore, the potential of parasitism by a parasitoid wasp, Chaenusa luteostigma sp. n. Achterberg (Hymenoptera: Braconidae: Alysiinae) on H. lagarosiphon to shift the competitive interactions between the two plant species was also examined. The addition of the parasitoid reduced the effect of herbivory by the fly on L. major by half, thereby shifting the competitive balance in favour of L. major over M. spicatum. This study provides valuable insight into a selection of drivers of submerged macrophyte assemblages of South Africa. It highlights the precarious position of South African freshwater systems with regard to the potential invasion by damaging submerged invasive species. It also provides interesting insights into the effect of competition, herbivory and parasitism on the establishment and spread of species within submerged freshwater systems. Understanding the different influences could assist managers and policy makers to make validated decisions ensuring the integrity of South African freshwater systems. iii Table of Contents Table of Contents Abstract ...................................................................................................................................... i Table of Contents .................................................................................................................... iv List of Tables ............................................................................................................................ x List of Figures ......................................................................................................................... xii Publications arising from this thesis ................................................................................... xix Acknowledgements ................................................................................................................ xx Chapter 1: Introduction .......................................................................................................... 1 1.2 Invasion of freshwater aquatic ecosystems ...................................................................... 1 1.2 Aquatic ecosystems in South Africa ................................................................................ 2 1.3 Introductory paths of aquatic macrophytes ...................................................................... 4 1.4 Drivers of aquatic plant assemblages ............................................................................... 5 1.4.1. Abiotic Drivers ......................................................................................................... 6 1.4.2. Biotic Drivers ........................................................................................................... 7 1.5 Aims ............................................................................................................................... 10 Chapter 2: Description of common invasive and indigenous submerged macrophytes in South Africa ............................................................................................................................ 16 2.1 Introduction .................................................................................................................... 16 2.2 Invasive species .............................................................................................................. 17 2.2.1 Egeria densa Planch (Hydrocharitaceae), dense waterweed ................................... 17 iv Table of Contents 2.2.2 Elodea canadensis Michaux (Hydrocharitaceae), Canadian waterweed ................. 20 2.2.3 Hy drilla verticillata L. (Hydrocharitaceae), hydrilla .............................................. 21 2.2.4 Cabomba caroliniana Grey (Cabombaceae), cabomba, fanwort ............................ 23 2.2.5 Myriophyllum spicatum L. (Haloragaceae) spiked water-milfoil ............................ 24 2.3 Indigenous species.......................................................................................................... 27 2.3.1 Lagarosiphon major (Roxb.) (Ridley) Moss (Hydrocharitaceae), African elodea, oxygen weed and Lagarosiphon muscoides Harv (Hydrocharitaceae). ........................... 27 2.3.2 Stuckenia pectinata (= Potamogeton pectinatus L.) (Potamogetonaceae), sago pondweed .......................................................................................................................... 30 2.4 Discussion ...................................................................................................................... 30 Chapter 3: Pet stores, aquarists and the internet trade as modes of introduction and spread of invasive macrophytes in South Africa ................................................................. 32 3.1 Introduction .................................................................................................................... 32 3.2 Methods & Materials ...................................................................................................... 37 3.2.1 Survey development for pet stores and aquarists .................................................... 37 3.2.2 Selection of test species ........................................................................................... 37 3.3 Results ...........................................................................................................................
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