Ecological Aspects of the Macroinvertebrates Associated with Two Submersed Macrophytes in Lake Kariba
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ECOLOGICAL ASPECTS OF THE MACROINVERTEBRATES ASSOCIATED WITH TWO SUBMERSED MACROPHYTES IN LAKE KARIBA By Crispen Phiri Thesis Presented for the Degree of DOCTOR OF PHILOSOPHY in the Department of Zoology UNIVERSITY OF CAPE TOWN FEBRUARY 2010 DECLARATION ECOLOGICAL ASPECTS OF THE MACROINVERTEBRATES ASSOCIATED WITH TWO SUBMERSED MACROPHYTES IN LAKE KARIBA I CRISPEN PHIRI hereby a) grant the University of Cape Town free licence to reproduce the above thesis, in whole or in part, for the purpose of research; b) declare that: i. the work contained in this thesis is my own unaided work, both in concept and execution, and that apart from the normal guidance from my supervisor, I have received no assistance ii. neither the substance nor any part of the thesis has been submitted in the past, or is being, or is to be submitted for a degree in the University or any other University iii. data presented here are original, and any other sources of data are fully acknowledged. SIGNED: ________________________________ DATE: _________________________________ TABLE OF CONTENTS ABSTRACT ………………………………………………………………………………… ix ACKNOWLEDGEMENTS ………………………………………………………………… xi CHAPTER 1 GENERAL INTRODUCTION 1.1 Introduction ………………………………………………………………….... 1 1.2 Aquatic Macrophytes…………………………………………………………. 2 1.3 Effect of Macrophytes on Water Physicochemical Conditions…………... 3 1.4 Organisms Associated with Aquatic Macrophytes………………………… 5 1.4.1 Periphyton…………………………………………………………………....... 5 1.4.2 Vertebrates…………………………………………………………………….. 6 1.4.3 Epiphytic invertebrates …………………………………………………….... 6 1.5 Factors Structuring Invertebrate Assemblages in Aquatic Ecosystems… 7 1.5.1 Effect of habitat structure on aquatic invertebrates……………………….. 8 1.5.1.1 The habitat heterogeneity hypothesis…………………………………….... 8 1.5.1.2 Aquatic macrophytes and habitat heterogeneity………………………….. 8 1.5.1.3 Habitat heterogeneity and aquatic macrophytes in Lake Kariba………... 10 1.5.2 Biotic interactions and invertebrate assemblages in aquatic ecosystems............................................................................................... 10 1.5.2.1 Predatoy-prey interactions…………………………………………………... 10 1.5.2.1(i) Fish predation……………………………………………………………….... 10 1.5.2.1(ii) Invertebrate predation……………………………………………………….. 12 1.5.2.2 Other biotic factors…………………………………………………………… 13 1.5.3 Habitat complexity, predator-prey relations and macroinvertebrate assemblages associated with Lagarosiphon and Vallisneria……………. 15 1.6 Effect of Climatic Conditions on Aquatic Invertebrates…………………… 15 1.6.1 Temperature: effect on aquatic invertebrates……………………………... 15 1.6.1.1 Climate variability hypothesis and aquatic invertebrates……………….... 16 1.6.2 Other physicochemical characteristics and macroinvertebrate assemblages…………………………………………………………….......... 17 1.6.3 Water physicochemical condition and epiphytic macroinvertebrate assemblages in Lake Kariba……………………………………………….... 18 1.7 Summary………………………………………………………………………. 19 1.8 Justification of the Study ……………………………………………………. 19 i 1.9 Objectives of the Study ……………………………………………………… 22 1.10 Organization of the Thesis ………………………………………………….. 22 CHAPTER 2 STUDY AREA, MATERIALS AND METHODS 2.1 Introduction …………………………………………………………………… 24 2.2 Study Area ……………………………………………………………………. 24 2.2.1 Lake Kariba …………………………………………………………………… 24 2.2.2 General climatic conditions in the vicinity of Lake Kariba………………… 26 2.2.3 Hydrological and limnological characteristics of Lake Kariba……………. 26 2.2.4 Aquatic macrophytes of Lake Kariba: a historical account………………. 27 2.2.5 Description of Lagarosiphon and Vallisneria………………………………. 30 2.2.5.1 Lagarosiphon………………………………………………………………….. 30 2.2.5.2 Vallisneria…………………………………………………………………….... 31 2.2.6 Shallow inshore fish community of Lake Kariba…………………………... 31 2.3 Materials and Methods……………………………………………………….. 32 2.3.1 Invertebrates associated with Lagarosiphon and Vallisneria: effect of plant morphological complexity and water physicochemical properties… 32 2.3.1.1 Site characterisation………………………………………………………….. 32 2.3.1.2 Water physical and chemical characteristics………………………………. 33 2.3.1.3 Epiphytic invertebrates……………………………………………………….. 34 2.3.1.4 Data analysis………………………………………………………………….. 35 2.3.2 Temporal variation and the effect of selected water physical and chemical parameters on macroinvertebrates associated with Lagarosiphon………………………………………………………………….. 36 2.3.2.1 Water physical and chemical characteristics………………………………. 36 2.3.2.2 Macroinvertebrate community structure……………………………………. 36 2.3.2.3 Data analysis………………………………………………………………….. 37 2.3.3 Body size distribution, biomass estimates and life histories of common insect taxa associated with Lagarosiphon................................................ 38 2.3.3.1 Insect body-size measurement and estimation of biomass…………….... 38 2.3.3.2 Data analysis………………………………………………………………….. 39 2.3.4 Effect of plant density on macroinvertebrates associated with Lagarosiphon………………………………………………………………..... 39 2.3.4.1 Measurement and characterisation of vegetation density………………... 39 ii 2.3.4.2 Water physicochemical assessment……………………………………….. 40 2.3.4.3 Macroinvertebrate sampling…………………………………………………. 40 2.3.4.4 Data analysis………………………………………………………………….. 40 2.3.5 Aquatic insects associated with Lagarosiphon and Vallisneria in fishless ponds……………………………………………………………..................... 41 2.3.5.1 Description of ponds………………………………………………………….. 41 2.3.5.2 Culture and maintenance of plants in ponds………………………………. 42 2.3.5.3 Macroinvertebrate sampling and processing…………………………........ 42 2.3.5.4 Water physicochemical assessment……………………………………….. 42 2.3.5.5 Data analysis………………………………………………………………….. 43 2.3.6 The effect of fish predation on macroinvertebrates associated with Lagarosiphon and Vallisneria: a pilot study in small ponds…………….... 43 2.3.7 An assessment of short-term survivorship of Ischnura (Odonata: Coenagrionidae) naiads at different densities of juvenile Oreochromis niloticus……………………………………………………………………….... 44 2.3.7.1 Vegetation density……………………………………………………………. 45 2.3.7.2 Ischnura naiad size classes and density…………………………………… 45 2.3.7.3 Fish (Oreochromis niloticus) densities……………………………………… 46 2.3.7.4 Recording naiad survival…………………………………………………….. 46 2.3.7.5 Data analysis………………………………………………………………….. 47 CHAPTER 3 INVERTEBRATES ASSOCIATED WITH Lagarosiphon AND Vallisneria: EFFECT OF PLANT MORPHOLOGICAL COMPLEXITY AND WATER PHYSICOCHEMICAL PROPERTIES 3.1 Introduction ………………………………………………………………….... 48 3.2 Materials and Methods……………………………………………………….. 49 3.3 Results ……………………………………………………………………….... 50 3.3.1 Site characterisation and water physicochemical variation among sites.. 50 3.3.2 Spatial distribution of Lagarosiphon and Vallisneria…………………….... 50 3.3.3 Spatial aspects of invertebrate assemblages associated with Lagarosiphon and Vallisneria……………………………………………...... 51 3.3.4 Effect of water physicochemical variables on macroinvertebrate assemblage structure……………………………………………………….... 51 3.3.5 Comparative assessment of epiphytic invertebrates associated with Lagarosiphon and Vallisneria………………………………………………. 52 iii 3.4 Discussion …………………………………………………………………….. 68 3.4.1 Physicochemical environment and its effect on epiphytic invertebrates... 68 3.4.2 Comparative assessment of invertebrate assemblages on Lagarosiphon and Vallisneria……………………………………………….. 70 3.4.3 Size class distribution of chironomids and two mayfly genus associated with Lagarosiphon and Vallisneria………………………………………….. 76 3.4.4 Conclusions……………………………………………………………………. 77 CHAPTER 4 TEMPORAL VARIATION AND THE EFFECT OF WATER PHYSICOCHEMICAL CHARACTERISTICS ON MACROINVERTEBRATES ASSOCIATED WITH Lagarosiphon 4.1 Introduction ………………………………………………………………….... 79 4.2 Materials and Methods……………………………………………………….. 80 4.3 Results ……………………………………………………………………….... 81 4.3.1 Water chemistry …………………………………………………………….... 81 4.3.1.1 Relationships among water variables………………………………………. 85 4.3.2 Macroinvertebrates associated with Lagarosiphon ………………………. 87 4.3.3 Temporal variation of common macroinvertebrate taxa …………………. 92 4.3.3.1 Turbellaria……………………………………………………………………… 92 4.3.3.2 Naididae……………………………………………………………………….. 92 4.3.3.3 Gastropoda……………………………………………………………………. 92 4.3.3.4 Hydracarina and Conchostraca……………………………………………... 93 4.3.3.5 Ostracoda……………………………………………………………………… 93 4.3.3.6 Ephemeroptera……………………………………………………………….. 94 4.3.3.7 Odonata……………………………………………………………………...... 98 4.3.3.8 Hemiptera & Trichoptera…………………………………………………….. 98 4.3.3.9 Diptera…………………………………………………………………………. 99 4.3.4 Temporal variation in total macroinvertebrate abundance……………….. 100 4.3.5 Relationships among invertebrate taxa…………………………………….. 101 4.3.6 Relationships between macroinvertebrate taxa and water physicochemical variables …………………………………………............. 104 4.3.7 Univariate analysis of macroinvertebrate community structure ……….... 106 4.3.8 Non-parametric multivariate analysis of macroinvertebrate community structure ……………………………………………………………………….. 112 iv 4.4 Discussion …………………………………………………………………….. 118 4.4.1 Invertebrates associated with Lagarosiphon in shallow waters of Lake Kariba…………………………………………………………………………... 119 4.4.2 Temporal aspects in physicochemical conditions…………………………. 121 4.4.3 Effect of water physiochemical characteristics on invertebrates associated with Lagarosiphon………........................................................ 121 4.4.3.1 Effects of water temperature on invertebrates ……………………………. 122 4.4.3.2 Effects water level on invertebrates………………………………………… 126 4.4.3.3 Effects of other water physicochemical variables………………………….