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Evaluation of a Plant-Herbivore System In EVALUATION OF A PLANT-HERBIVORE SYSTEM IN DETERMINING POTENTIAL EFFICACY OF A CANDIDATE BIOLOGICAL CONTROL AGENT, CORNOPS AQUATICUM FOR WATER HYACINTH, EICHHORNIA CRASSIPES A thesis submitted in fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY of RHODES UNIVERSITY by ANGELA BOWNES December 2008 Abstract Water hyacinth, Eichhornia crassipes Mart. Solms-Laubach (Pontederiaceae), a free- floating aquatic macrophyte of Neotropical origin, was introduced into South Africa as an ornamental aquarium plant in the early 1900’s. By the 1970’s it had reached pest proportions in dams and rivers around the country. Due to the sustainability, cost efficiency and low environmental risk associated with biological control, this has been a widely used method in an attempt to reduce infestations to below the threshold where they cause economic and ecological damage. To date, five arthropod and one pathogen biocontrol agents have been introduced for the control of water hyacinth but their impact has been variable. It is believed that their efficacy is hampered by the presence of highly eutrophic systems in South Africa in which plant growth is prolific and the negative effects of herbivory are therefore mitigated. It is for these reasons that new, potentially more damaging biocontrol agents are being considered for release. The water hyacinth grasshopper, Cornops aquaticum Brüner (Orthoptera: Acrididae), which is native to South America and Mexico, was brought into quarantine in Pretoria, South Africa in 1995. Although the grasshopper was identified as one of the most damaging insects associated with water hyacinth in its native range, it has not been considered as a biocontrol agent for water hyacinth anywhere else in the world. After extensive host-range testing which revealed it to be safe for release, a release permit for this candidate agent was issued in 2007. However, host specificity testing is no longer considered to be the only important component of pre-release screening of candidate biocontrol agents. Investigating biological and ecological aspects of the plant-herbivore system that will assist in determination of potential establishment, efficacy and the ability to build up good populations in the recipient environment are some of the important factors. This thesis is a pre-release evaluation of C. aquaticum to determine whether it is sufficiently damaging to water hyacinth to warrant its release. It investigated interactions between the grasshopper and water hyacinth under a range of nutrient conditions found in South African water bodies as well as the impact of the grasshopper on the competitive performance of water hyacinth. ii Both plant growth rates and the response of water hyacinth to herbivory by the grasshopper were influenced by nutrient availability to the plants. The ability of water hyacinth to compensate for loss of tissue through herbivory was greater under eutrophic nutrient conditions. However, a negative linear relationship was found between grasshopper biomass and water hyacinth performance parameters such as biomass accumulation and leaf production, even under eutrophic conditions. Water hyacinth’s compensatory ability in terms of its potential to mitigate to detrimental effects of insect feeding was dependent on the amount of damage caused by herbivory by the grasshopper. Plant biomass and the competitive ability of water hyacinth in relation to another free- floating aquatic weed species were reduced by C. aquaticum under eutrophic nutrient conditions, in a short space of time. It was also found that grasshopper feeding and characteristics related to their population dynamics such as fecundity and survival were significantly influenced by water nutrient availability and that environmental nutrient availability will influence the control potential of this species should it be released in South Africa. Cornops aquaticum shows promise as a biocontrol agent for water hyacinth but additional factors that were not investigated in this study such as compatibility with the South African climate and the current water hyacinth biocontrol agents need to be combined with these data to make a decision on its release. Possible management options for this species if it is to be introduced into South Africa are discussed. iii Table of contents Title page………………………………………………………………………….……….i Abstract……………………………………………………………………………..……..ii Table of Contents………………………………………………………………………....iv List of Figures……………………………………………………………………... …...vii List of Tables……………………………………………………………………….…...xiv Acknowledgements………………………………………………………………….......xv Chapter 1: Introduction 1.1 General Introduction…………………………………………………………..2 1.2 The plant-herbivore system……………………...……………………….........7 1.2.1 Water hyacinth, Eichhornia crassipes ………………………………….7 1.2.2 The water hyacinth grasshopper, Cornops aquaticum …………………8 1.3 The South African water hyacinth biological control programme………..…10 1.4 Recent developments in biological control…………………………………..13 1.4.1 Pre-release efficacy testing……………………………..……………...13 1.4.2 Ecology of the plant-herbivore system………………………………...15 1.5 Thesis outline………………………………………………………………...16 Chapter 2: Influence of nitrate and phosphate levels occurring in South African water bodies on the potential of Cornops aquaticum to growth and productivity of water hyacinth 2.1 Introduction………………………………………………………………….19 2.2 Materials and Methods………………………………………………………24 2.2.1 Nutrients …………………………………………………………...24 2.2.2 Experimental design ……………………………………………….26 2.2.3 Statistical analysis …………………………………………………27 2.3 Results………………………………………………………………………..28 2.3.1 Nitrogen and phosphorus levels of water hyacinth leaves 3, 4 and 5 from chemical analysis and evaluation of the relationship between leaf nitrogen content (%) and water nitrate levels (mgL -1)……………………………………………………..28 2.3.2 Effect of water nutrient treatment on the wet weight of water hyacinth leaves of plants grown at high, medium and low nutrient levels ………………………………………………………..29 2.3.3 Effect of nutrients and herbivory on plant biomass ……………......30 2.3.4 Effect of nutrients and herbivory on plant growth parameters ……………………………………………………………….32 iv 2.3.5 Effect of nutrients and herbivory on leaf, ramet and flower production …………………………………..……………….36 2.3.6 Effect of nutrient treatment on feeding damage and leaf preference ………………...…………………………………………40 2.4. Discussion…………………………………………………………………...41 Chapter 3: Performance of Cornops aquaticum in response to variation in water hyacinth plant quality as influenced by environmental nitrogen (N) availability 3.1 Introduction…………………………………………………………………..52 3.2 Materials and Methods……………………………………………………….57 3.2.1 Effect of plant nutrient levels on Cornops aquaticum survival, fecundity and body size ………………………………….……..57 3.2.1.1 Trial 1 – Experimental design………………….………...………57 3.2.1.2 Trial 2 – Experimental design……………………………………58 3.2.1.3 Statistical analysis………………………………………………..58 3.2.2 Effect of plant nutrient levels on survival and feeding and development rates of Cornops aquaticum nymphs …………………...….59 3.2.2.1 Experimental design………………………………...…………....59 3.2.2.2 Statistical analysis………………………………………………..60 3.3 Results………………………………………………………………………..61 3.3.1 Effect of nutrient levels on survival, fecundity and body size ………………………………………………………………….61 3.3.2 Effect of nutrient treatment on survival and development rates of C. aquaticum ………………………...…………………………..66 3.3.3 Effect of nutrient treatment on feeding rates of Cornops aquaticum nymphs …………………………………………..……..….... 69 3.4 Discussion……………………………………………………………………72 Chapter 4 Investigating the response of water hyacinth to herbivory by different densities of male and female Cornops aquaticum under eutrophic water nutrient conditions 4.1 Introduction………………………………………………………………….85 4.2 Materials and Methods………………………………………………………91 4.2.1 Experimental design ……………………………………………….91 4.2.2 Statistical analysis ……………………………………………...….92 4.3 Results………………………………………………………………………..93 4.3.1 Effect of insect treatment on plant biomass ………………………..93 4.3.2 Effect of insect treatment on leaf parameters… …………………...96 4.3.3 Effect of insect treatment on plant growth and productivity parameters ………………………………………………….99 4.3.4 Effect of insect treatment on defoliation ………………….............105 4.3.5 Effect of insect treatment on leaf area damage …………………106 4.4 Discussion…………………………………………………………………..107 v Chapter 5: Inter- and intraspecific competitive interactions between water hyacinth and water lettuce as influenced by Cornops aquaticum herbivory 5.1 Introduction………………………………………………………………....121 5.2 Materials and Methods……………………………………………………...126 5.2.1 Experimental design ………………………………………………………126 5.2.2 . Statistical analysis ………………………………………………........….127 5.3 Results………………………………………………………………………128 5.3.1 Impact of herbivory on competition ……………………………... 128 Water hyacinth………………………………………………….128 Water lettuce……………………………………………………132 5.3.2 Impact of herbivory on plant biomass (mean end-weights) ……....134 5.3.3 Impact of herbivory and competition on water hyacinth biomass (total end-weights) …………………………………………….135 5.4 Discussion…………………………………………………………………………..137 Chapter 6: General discussion ………………………………………………………..144 References ……………………………………………………………………………...157 vi List of figures Figure 2.1 Relationship between water nitrate levels (mgL -1) and mean nitrogen content of leaves 3, 4 and 5 of water hyacinth plants grown at high, medium and low nutrient levels (n = 15/leaf position/nutrient treatment)…......……………………………………………………………...29 Figure
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