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The Ecology of Freshwater Zooplankton in the Hawkesbury-Nepean River, with Special Reference to Community Structure and Grazing

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The Ecology of Freshwater Zooplankton in the Hawkesbury-Nepean River, with Special Reference to Community Structure and Grazing THE ECOLOGY OF FRESHWATER ZOOPLANKTON IN THE HAWKESBURY-NEPEAN RIVER, WITH SPECIAL REFERENCE TO COMMUNITY STRUCTURE AND GRAZING Tsuyoshi Kobayashi This thesis is submitted for the degree of Doctor of Philosophy at the University of New South Wales June 1998 "I hereby declare that this submission is my own work and that, to the best of my knowledge and belief, it contains no material previously published or written by another person nor material which to a substantial extent has been accepted for the award of any degree or diploma of a university or other institute of higher learning, except where due acknowledgment is made in the text." 7~· T. Kobayashi CERTIFICATE OF ORIGINALITY I hereby declare that this submission is my own work and to the best of my knowledge it contains no materials previously published or written by another person, nor material which to a substantial extent bas been accepted for the award of any other degree or diploma at UNSW or any other educational inslitution, except where due acknowledgementis made in the thesis. Anycontnbution made to the research by others, with whom I have worked at UNSW or elsewhere, is explicitly acknowledged in the thesis. I also declare that the intellectual content of this thesis is the product of my own work, except to the extent that assistance from others in the project's design and conception or in style, presentation and linguistic expn:ssion is acknowledged. (Signed) ······························�········-····· .. ··················•·········· Contents CONTENTS i List of Figures V List of Tables vii Acknowledgments viii Abstract ix Page Chapter 1 General Introduction 1 1.1. Ecological studies of freshwater river zooplankton 2 1.2. Ecological studies of freshwater river zooplankton in Australia 3 1.3. Present study 4 1.4. Hawkesbury-Nepean River 5 Chapter 2 Taxonomic Composition, Richness and Density: Comparison with Other Rivers 7 2.1. Summary 8 2.2. Introduction 9 2.3. Materials and methods 10 2.3.1. Study sites 10 2.3.2. Zooplankton sampling 11 2.3.3. Subsampling and counting 11 2.3.4. Species identification 12 2.3.5. Environmental variables 12 2.3.6. Statistical Analyses 13 2.4. Results 13 2.4.1. Environmental variables 13 2.4.2. Zooplankton 14 i) Taxonomic composition 14 ii) Density and relative abundance 15 iii) Correlation between river environmental variables and community densities 15 2.5. Discussion 16 2.5.1. Taxonomic richness and composition 16 2.5.2. Density and relative abundance 18 2.5.3. Correlation between river environmental variables and community densities 19 2.5.4. Community structure ofriver zooplankton 20 Chapter 3 Patterns in the Daytime Vertical Distribution of Dominant River Microzooplankton 22 3. I. Summary 23 Contents 3.2. Introduction 24 3.3. Materials and methods 25 3.3.1. Study site 25 3.3.2. Zooplankton collection, subsampling and counting 25 3.3.3. Statistical analyses 26 i) Overall difference in vertical distribution 26 ii) Correlation of relative densities at 1 m depth (RD,m, %) with river flow rate 27 3.4. Results 27 3.4.1. Overall difference in density between the two depths 27 3.4.2. Spearman's rank correlation of RD,m with river flow rate 28 3.5. Discussion 28 Chapter4 Associations Between River Environmental Variables and Zooplankton Body Masses 31 4.1. Summary 32 4.2. Introduction 33 4.3. Materials and methods 34 4.3.1. Study site 34 4.3.2. Zooplankton sampling, subsampling, identification and counting 35 4.3.3. Biomass and mean body mass 35 4.3.4. Environmental variables 36 4.3.5. Statistical analyses 36 4.4. Results 36 4.4.1. Flow, turbidity, temperature and chlorophyll a 36 4.4.2. Mean body mass and total biomass 3 7 4.4.3. Simple correlation 38 4.4.4. Partial correlation 38 4.5. Discussion 39 Chapter 5 Community Grazing in situ: Importance of Microzooplankton 42 5 .1. Summary 43 5.2. Introduction 44 5.3. Materials and methods 46 5 .3. l. Study site 46 5.3.2. Grazing experiments 47 5.3.3. Determination of zooplankton density 49 5.3.4. Determination of zooplankton biomass 49 5.3.5. Temperature and chlorophyll a 50 5.3.6. Search of predictive models 50 11 Contents 5.4. Results 52 5.4.1. Temperature and chlorophyll a 52 5.4.2. Zooplankton biomass 52 5.4.3. Community grazing rates 53 5.4.4. Predictive models based on total biomass 53 5.4.5. Predictive models based on taxonomic composition 54 5.5. Discussion 55 5.5.1. Grazing impact 55 5.5.2. Grazing models 58 Chapter 6 Community Structure: Seasonal Variation, Overall Pattern and Functional Aspects 62 6.1. Summary 63 6.2. Introduction 64 6.3. Materials and methods 65 6.3.1. Study site 65 6.3.2. Zooplankton sampling, subsampling, identification and counting 65 6.3.3. Estimate of density and body mass 65 6.3.4. Density-size distribution 65 6.3.5. Weight-specific grazing rate and mean body mass 66 6.4. Results 67 6.4.1. Density-size distribution 67 6.4.2. Mean body mass and grazing rate 68 6.5. Discussion 69 6.5.1. Seasonal variation 69 6.5.2. Overall pattern 70 6.5.3. Conversion to biomass-size distribution 71 6.5.4. Mean body mass and grazing rate 72 Chapter 7 Conclusions 75 Chapter 8 References 81 Appendix I Ecological Studies of Freshwater Zooplankton in Rivers, Published After 1970 99 Appendix II A Review of Radiotracer Methods for Zooplankton Grazing Measurements 103 Appendix III Data 110 1. Physico-chemical variables 111 2. Zooplankton community densities 115 111 Contents 3. Vertical density of dominant zooplankton taxa at North Richmond 116 4. Biomass and mean body size of zooplankton at North Richmond 126 5. Community grazing rate at North Richmond 127 6. Density-size distributions of zooplankton at North Richmond 128 7. Parameters oflog-linear model for density-size distributions of zooplankton at North Richmond 142 lV Figures and Tables List of Figures Facing Page I.I. Relative trend of ecological studies of freshwater zooplankton in rivers, published after 1970 2 1.2. Schematic relationship between chapters in the present thesis 4 1.3 Hawkesbury-Nepean River 5 2.1. Temporal variation in flow rate over Penrith weir and temperature in the Hawkesbury-Nepean River 13 2.2. Density and percentage composition of the zooplankton community in the Hawkesbury-Nepean River 15 2.3. Box plots of taxonomic richness of zooplankton in rivers 17 2.4. Relationship between maximum densities of planktonic rotifers and crustaceans in rivers 18 3.1. Examples of seasonal pattern in the vertical distribution ofzooplankton at North Richmond 27 3.2. Examples of overall pattern in the vertical distribution of zooplankton at North Richmond 27 3.3. Significant correlation ofrelative densities at a depth of 1 m with river flow rate at North Richmond 28 3.4. Overall relationship between river flow rate and relative densities at a depth of 1 m for rnicrozooplankton at North Richmond 28 4.1. River environmental conditions at North Richmond 36 4.2. Zooplankton populations at North Richmond 37 4.3. Zooplankton community at North Richmond 37 4.4. Simple correlations of mean zooplankton body masses with river environmental variables and with zooplankton biomasses at North Richmond 38 4.5. Relationship between mean body mass of copepods and river flow rate at North Richmond 39 5.1. Community biomass of zooplankton at North Richmond 52 5.2. Percentage composition of zooplankton biomass at North Richmond 53 V Figures and Tables 5.3. Zooplankton community grazing rates at North Richmond 53 5.4. Correlation of community grazing rate with total biomass of zooplankton at North Richmond 53 6.1. Examples of pattern in the density-size distribution of the zooplankton community at North Richmond 68 6.2. Parameters of log2-log2 regression of density against mean body mass at North Richmond 68 6.3. Overall pattern in density-size distribution of the zooplankton community between March 1992 and April 1993 68 6.4. Relationship between mass-specific grazing rate and mean body mass of the zooplankton community at North Richmond between July 1992 and April 1993 68 6.5. Relationship between community grazing rate and mean body mass of the zooplankton at North Richmond between July 1992 and April 1993 69 6.6. Overall pattern in biomass-size distribution of the zooplankton at North Richmond between March 1992 and April 1993 71 Vl Figures and Tables List of Tables Facing Page 2.1. Summary statistics for environmental variables in the Hawkesbury-Nepean River 13 2.2. Zooplankton taxa in the Hawkesbury-Nepean River 14 2.3. Densities of common zooplankton taxa in the Hawkesbury-Nepean River 14 2.4. Simple correlation coefficients between environmental variables and community density of zooplankton in the Hawkesbury-Nepean River 16 3.1. Comparison of overall mean density of dominant zooplankton between 1 m and 4 m depths at North Richmond 27 4.1. Dry weights of zooplankton taxa at North Richmond 37 4.2. Partial correlations between river environmental variables and mean body masses of populations and of the community of zooplankton and their respective biomasses at North Richmond 38 5.1. Phytoplankton taxa at North Richmond between September 1992 and April 1993 46 5.2. Predictive models of community grazing rate at 1 m depth, based on total zooplankton biomass, temperature and chlorophyll a 53 5.3. Predictive models of community grazing rate at 4 m depth, based on total zooplankton biomass, temperature and chlorophyll a 54 5.4.
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