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the distribution, abundance and trophic ecology of the fishes of darwin harbour mangrove habitats PDF

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THE DISTRIBUTION, ABUNDANCE AND TROPHIC ECOLOGY OF THE FISHES OF DARWIN HARBOUR MANGROVE HABITATS JULIE M MARTIN B.Sc.(Hons), James Cook University, 1994 A thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy in the Faculty of Education, Health and Science, Charles Darwin University. 2005 I hearby declare that the work herein, now submitted as a thesis for the degree of Doctor of Philosophy, is the result of my own investigations and all references to ideas and work of other researchers have been specifically acknowledged. I hearby certify that the work embodied in this thesis has not already been accepted in substance for any degree and is not being currently submitted in candidature for any other degree. ii ACKNOWLEDGMENTS I would like to thank the many people who gave me inspiration, support, encouragement and assistance during this research. Without them, this project would not have been possible. Most importantly, I thank my husband Andrew for everything, and our children, Manisha and Soni, for their patient, and not so patient, waiting while their mum finished that book. My supervisor, Dr. Keith McGuinness, offered me insight, support and guidance throughout the project. I greatly appreciate his patient reading and constructive comments on all drafts of this thesis. Dr Rik Buckworth, my associate supervisor, was also a source of inspiration, encouragement and friendship. This project was generously supported by two government departments. The Department of Infrastructure, Planning and Environment provided an additional stipend, a boat and project operational costs for three years. The Fisheries Division of the Department of Business Industry and Resource Development gave me in-kind assistance, including (amongst many other things) office space, computing facilities and access to vehicles. I would also like to thank the Centre for Tropical Wetlands Management for their Post Graduate Research Grant of $500. My technical assistant, Paul McHugh, was an inspiration. I know of few other people who would work so tirelessly under such trying conditions (apart from PhD students). His assistance was essential during fieldwork which ranged from wading through knee-deep mangrove mud wielding a chainsaw (and a pistol) in the middle of the wet season to getting up at 3.30 am to go "fishing". Many volunteers were also generous with their assistance in the field work, especially Andrew Buick, Stephanie iii Boubaris, Tony Powell, Patrick Grey, Adrian Donati, Charles Bryce, Richard Sellers, Neil Smit and Roland Griffin. Richard Mounsey, Mac MacCartie, Charles Bryce and Stepanine Boubaris helped design and make trammel nets. Armando Padovan and Rod Metcalfe provided water quality data and assisted with water quality measurements. Thanks also to the many others at the fisheries division for your support and friendship. I learnt a lot and also had fun at the Fisheries Centre, UBC and I much appreciate the help and friendship of Profs. Daniel Pauly, Villy Christensen and Carl Walters, Dr. Reg Watson, Dr. Neil Gribble and Maria Morlin. Thanks also to Drs. Helen Larson, Richard Willan and Chris Glasby at the NT Museum for their help in identifying fish and prey items taken from the stomachs of fish. My thanks should also go to the firearms trainers for trying (in vain) to teach me the art of self-protection. To my colleagues who revel in mud and mangroves - Kristin Metcalfe, Chandra Salgado-Kent, Brad Comley, Grey Coupland, Justin McDonald-it was comforting to know that you were out there too. Kristin also generously provided preliminary data from her PhD project for use in this thesis. There were many others who gave me encouragement and emotional support, thanks especially to my friends Lynne Powell and Gretchen Thombum. Finally I thank my family; Ross and Vic, John and Wendy, my mother Linda Martin, and my parents-in law, Kathleen and Malcolm Buick for their patience and understanding ... and Dad, are you clicking your heels? iv FOR THE FISH Permission to use this image from Patrick Hall and Di Allison, Phish Design Cards, Hobart, Tasmania. v ABSTRACT Darwin Harbour features one of the largest and most diverse mangrove communities in northern Australia. The harbour is also the centre of a thriving recreational fishery. This study examined the role of mangroves in the distribution, abundance and trophic relationships of the fish of Darwin Harbour. Fish were sampled in three different mangrove habitats in three locations in Darwin Harbour for two years, using three methods: trammel nets, light traps and pit traps. Forest structural complexity and aquatic environmental variables were also measured. Diet composition was analysed and trophic groups identified using multivariate techniques. A preliminary trophic model was also developed. At high spring tides, the mangrove forest was used extensively by a wide range of fish. At low tide, only resident species appeared to remain in pools, as pit trap samples comprised the Gobiidae and Pseudomugilidae. Trammel net captures were dominated by the Ariidae, Mugilidae, Clupeidae and Engralulidae families. Fish captured in the light traps were dominated by Engraulidae, Clupeidae, Atherinidae and Gobiidae. Small fish, including juveniles and larvae, were more abundant in the seaward habitats, whereas larger fish, including predatory species, were captured throughout the mangrove forest. This suggests that smaller fish were not seeking refuge from predation in the shallower, more complex habitats. Fish abundance and distribution did not appear to be influenced by small scale variations in structural complexity (stem density and sizes) or mangrove productivity. Instead, large scale differences in forest characteristics, such as mangrove species composition and general topographical features, along with aquatic environmental variables appeared to be more important. vi All habitats appeared to be important feeding areas for fish and seven trophic groups were identified. Three major groups were separated according to feeding mode (nekton, benthic and surface feeders) and within these, seven groups were defined by the main prey items in their diets. The Ecopath with Ecosim trophic model, although preliminary and based on incomplete information, synthesised information about Darwin Harbour from a wide variety of sources, indicated information gaps and identified areas in which future research should be focussed. Some species, such as Toxotes chatareus and two Ariid catfish, were more strongly associated with the higher intertidal habitats. This association may be important ecologically, due to their high levels of predation on the sesarmid crab, Perisesarma darwinensis, which is also found more often in the higher intertidal habitat and is known to consume mangrove leaf litter. Overall, the results show that a diverse array of fish use the mangroves of Darwin Harbour. Some are economically valuable species themselves and many are prey species for fish of recreational importance. There appear to be close trophic links between fish and all habitats and these links must be considered in future management of Darwin Harbour. vii TABLE OF CONTENTS 1. GENERAL INTRODUCTION ...................................................................................... 1 1.1. Background .................................................................................................................... 1 1.2. Mangrove productivity .................................................................................................. 3 1.3. Mangroves as nursery areas ........................................................................................... 5 1.3.1. Mangroves versus other habitats ............................................................................ 5 1.3.2. Habitats within the mangroves .............................................................................. 8 1.3.3. Temporal variation ................................................................................................. 9 1.4. Mangroves of Darwin Harbour .................................................................................... 11 1.5. Project aims and objectives ......................................................................................... 13 1.5.1. Specific objectives ............................................................................................... 14 2. REVIEW OF SAMPLING METHODS AND DESCRIPTION OF STUDY AREA ......................................................................................................................................... 15 2.1. Review of sampling methods ....................................................................................... 15 2.2. Description of study area ............................................................................................. 21 2.2.1. Climate ................................................................................................................. 22 2.2.2. Physical description ............................................................................................. 22 2.3. Study locations ............................................................................................................ 24 2.3.1. Location 1 ............................................................................................................ 25 2.3.2. Location 2 ............................................................................................................ 28 2.3.3. Location 3 ............................................................................................................ 29 2.4. Site Preparation ............................................................................................................ 31 2.4.1. Access .................................................................................................................. 31 2.4.2. Sampling site preparation .................................................................................... 31 2.4.3. Topographic level ................................................................................................ 32 3. HABITAT STRUCTURE AND AQUATIC ENVIRONMENTAL VARIABLES. 33 3.1. Methods ....................................................................................................................... 35 3.1.1. Habitat structure ................................................................................................... 35 3.1.2. Environmental variables ...................................................................................... 36 3 .2. Results ......................................................................................................................... 3 7 3.2.1. Habitat structure ................................................................................................... 37 3.2.2. Environmental variables ...................................................................................... 42 3.3. Discussion .................................................................................................................... 58 3.3.1. Habitat structure ................................................................................................... 58 3.3.2. Environmental variables ...................................................................................... 60 4. A COMPARISON OF FISH CATCHES AT DAWN AND DUSK USING TRAMMEL NETS IN MANGROVE FOREST HABITATS .................................. 65 4.1. Introduction ................................................................................................................. 65 4.2. Methods ....................................................................................................................... 66 4.2.1. Data analysis ........................................................................................................ 67 4.3. Results ......................................................................................................................... 68 4.3.1. Abundance and distribution ................................................................................. 68 4.3.2. Patterns in species composition ........................................................................... 73 4.4. Discussion .................................................................................................................... 73 5. DISTRIBUTION AND ABUNDANCE OF FISH IN THE MANGROVE FORESTS OF DARWIN HARBOUR ........................................................................................... 76 5 .1. Introduction ................................................................................................................. 7 6 5 .2. Methods ....................................................................................................................... 7 8 5.2.1. Sampling design ................................................................................................... 78 5 .2.2. Data analysis ........................................................................................................ 81 5.3. Results ......................................................................................................................... 83 5.3 .1. Abundance and distribution ................................................................................. 8 3 viii 5.3.2. Analysis of abundance ......................................................................................... 89 5.3.3. Diversity .............................................................................................................. 97 5.3.4. Patterns in species composition ........................................................................... 99 5.3.5. Environmental variables .................................................................................... 104 5.4. Discussion .................................................................................................................. 106 5.4.1. Species composition .......................................................................................... 106 5.4.2. Temporal patterns .............................................................................................. 111 5.4.3. Spatial patterns ................................................................................................... 113 5.4.4. Conclusion ......................................................................................................... 118 6. DISTRIBUTION OF LARVAL AND JUVENILE FISH IN MANGROVE HABITATS OF DARWIN HARBOUR ..........................•..•...............•.•...........•....... 120 6.1. Introduction ............................................................................................................... 120 6.2. Methods ..................................................................................................................... 122 6.2.1. Light traps .......................................................................................................... 122 6.2.2. Pit traps .............................................................................................................. 124 6.2.3. Data analysis ...................................................................................................... 125 6.3. Results ....................................................................................................................... 126 6.3.1. Light traps .......................................................................................................... 126 6.3.2. Pit traps .............................................................................................................. 134 6.4. Discussion .................................................................................................................. 142 6.4.1. Lighttraps .......................................................................................................... 142 6.4.2. Pit traps .............................................................................................................. 147 6.4.3. Conclusion ......................................................................................................... 149 7. DIETS AND TROPHIC STRUCTURE OF FISH FROM MANGROVE FORESTS IN DARWIN HARBOUR .......................................................................................... 151 7 .1. Introduction ............................................................................................................... 151 7 .2. Methods ..................................................................................................................... 155 7.2.1. Study sites and sampling methods ..................................................................... 155 7 .2.2. Diet analyses ...................................................................................................... 155 7.2.3. General description of diet.. ............................................................................... 159 7.2.4. Dietary breadth .................................................................................................. 161 7 .2.5. Trophic groups ................................................................................................... 162 7 .2.6. Habitat comparison ............................................................................................ 162 7 .3. Results ....................................................................................................................... 162 7.3.1. Overall diet description ...................................................................................... 162 7.3.2. Sesarrnid predation ............................................................................................ 167 7.3.3. Dietary breadth .................................................................................................. 169 7 .3.4. Ontogenetic changes in diet. .............................................................................. 170 7.3 .5. Trophic groups ................................................................................................... 170 7.3.6. Habitat comparison ............................................................................................ 176 7 .4. Discussion .................................................................................................................. 179 7.4.1. General diet ........................................................................................................ 179 7.4.2. Dietary breadth .................................................................................................. 184 7.4.3. Ontogenetic changes in diet ............................................................................... 185 7 .4.4. Trophic groups ................................................................................................... 186 7 .4.5. Habitat comparison ............................................................................................ 190 7.4.6. Sesarrnid predation ............................................................................................ 191 7.4.7. Conclusion ......................................................................................................... 193 8. TOWARDS A TROPIDC MODEL OF DARWIN HARBOUR .......•..................•. 195 8.1. Introduction ............................................................................................................... 195 8.2. Methods ..................................................................................................................... 197 8.2.1. Parameter estimation .......................................................................................... 198 8.2.2. Missing groups ................................................................................................... 205 8.3. Balancing the model .................................................................................................. 205 8.4. Results and discussion ............................................................................................... 206 ix 9. GENERAL DISCUSSION AND CONCLUSIONS ................................................. 215 9.1. Temporal patterns among fish assemblages .............................................................. 216 9.2. Spatial patterns among fish assemblages ................................................................... 219 9.2.1. Abundance and distribution ............................................................................... 219 9.2.2. Species composition .......................................................................................... 219 9.3. Patterns among fish assemblages in relation to forest structure, aquatic environmental variables and mangrove productivity ................................................................................... 220 9.3.1. Forest area and structural complexity ................................................................ 220 9.3.2. Aquatic environmental variables ....................................................................... 222 9.3.3. Mangrove productivity ...................................................................................... 223 9.4. Diets and trophic ecology of fish assemblages .......................................................... 224 9.5. Habitats within mangroves ........................................................................................ 227 9.6. Conclusion and recommendations ............................................................................. 228 9.6.1. Recommendations .............................................................................................. 229 10. REFERENCES ........................................................................................................... 231 X

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Harbour for two years, using three methods: trammel nets, light traps and pit traps. Forest structural complexity and aquatic environmental variables were also measured. Diet composition was analysed and trophic groups identified using multivariate techniques. A preliminary trophic model was also
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