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Groundwater Ecology: Invertebrate Community Distribution Across the Benthic, Hyporheic and Phreatic Habitats of a Chalk Aquifer in Southeast England

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Groundwater Ecology: Invertebrate Community Distribution Across the Benthic, Hyporheic and Phreatic Habitats of a Chalk Aquifer in Southeast England Groundwater Ecology: Invertebrate Community Distribution across the Benthic, Hyporheic and Phreatic Habitats of a Chalk Aquifer in Southeast England Jessica M. Durkota Thesis submitted in partial fulfilment of the requirements for the degree of Doctor of Philosophy University College London Declaration of the Author I, Jessica M. Durkota, confirm that the work presented in this thesis is my own. Where information has been derived from other sources, I confirm that this has been indicated in the thesis. This work was undertaken with the partial support of the Environment Agency. The views expressed in this publication are mine and mine alone and not necessarily those of the Environment Agency or University College London (UCL). 2 Abstract Groundwater is an important resource for drinking water, agriculture, and industry, but it also plays an essential role in supporting the functioning of freshwater ecosystems and providing habitat for a number of rare species. However, despite its importance, groundwater ecology often receives little attention in environmental legislation or research. This study aims to improve our understanding of the organisms living in groundwater-dependent habitats and the influence of environmental conditions on their distribution. Invertebrate communities occurring in the benthic, hyporheic and phreatic habitats were surveyed at twelve sites over four years across the Stour Chalk Block, a lowland catchment in southern England. A diverse range of stygoxenes, stygophiles and stygobionts, including the first record of Gammarus fossarum in the British Isles, were identified using morphological and molecular techniques. The results indicate that under normal conditions, each habitat provided differing environmental conditions which supported a distinctive invertebrate community. While the community recorded in the benthic habitat was characterised by a diverse assemblage of surface water species typical of Chalk streams, the phreatic community comprised a small number of exclusively crustacean stygofauna (such as Niphargus kochianus and Crangonyx subterraneus) and the hyporheic habitat supported a mixture of surface and groundwater species. Surprisingly, the results indicate that some species, such as Agapetus fuscipes (normally considered a surface water taxon), move into the hyporheic habitat in a predictable, seasonal pattern, potentially in response to grazing opportunities. However, the results collected during the high and low flow events which occurred during this study also show the widespread movement of multiple species (such as Gammarus pulex and Niphargus fontanus) between habitats in response to environmental disturbance. Collectively, these results reflect the movement of fauna longitudinally, laterally and vertically over time throughout the catchment, as though along a continuum rather than between three separate habitats. This suggests that our conceptualisation of lotic functioning should be expanded to better integrate the contribution from groundwater. The approach taken by this study provides a greater understanding of the full diversity of aquatic invertebrates within this catchment and the way in which their distribution fluctuates across habitats. This study is one of the first to concurrently assess invertebrate distribution across the benthic, hyporheic and phreatic habitats; in addition, the relatively frequent and long-term sampling approach also facilitated a more detailed temporal assessment of these communities. A greater understanding of the distribution and requirements of the fauna inhabiting groundwater-dependent habitats, and their response to environmental change is essential for the conservation of these species and management of lotic ecosystems. 3 Acknowledgements Over the course of my study I have received support and encouragement from a number of individuals, only some of whom it is possible to give particular mention to here: Foremost I would like to thank the UCL community for their advice and support throughout this study: Julian Thompson and Roger Flower for supervising this work (and assistance with freeze coring); Jan Axmacher, Helen Bennion, Anson Mackay and Carl Sayer for their encouragement and advice; Julia Day for the use of her lab and her support in undertaking the molecular analyses; Janet Hope and Tula Maxted for their support in the geography lab and especially to Ian Patmore for his help in fabricating a Bou-Rouch pump from a poor sketch on the back of a napkin; Miles Irving in the Drawing Office for his unfailing assistance in producing diagrams and posters; and to my graduate cohort, James Cheshire, Miriam Fernandez, Tori Gay, William Gray, Caitlin Gutierrez, Charlotte Hall, Regan Koch, Dan Lewis, Luca Marazzi, Luke Mitchell, Matt Owen and Emma Wiik for their support, kindness and good humour throughout this process. I would also like to thank the wider research community, specifically Anne Robertson, Rachel Stubbington, Tim Johns and Paul Wood for their advice (and enthusiasm) on sampling hyporheic environments; Michael Dobson, Terry Gledhill, Lee Knight, Peter Shaw and colleagues at the Environment Agency for their assistance in the verification of specimen; Jane Newcombe for her help in analysing the microbial samples; Bill Beaumont for his help in facilitating the freeze-core sampling; and Rosie Blackman for her collaboration in drafting the Gammarus fossarum manuscript. I would further like to acknowledge my funders, the Environment Agency, UCL Department of Geography and UCL Graduate School for the opportunities they provided. Finally, I would like to thank my friends and family for their support (and patience) over the past eight years. 4 Table of Contents 1 Review of Groundwater Ecology............................................................................. 10 1.1 Introduction ........................................................................................................... 11 1.2 Groundwater Dependent Habitats........................................................................ 11 1.3 Groundwater Ecology ........................................................................................... 13 1.4 Environmental Conditions in the Groundwater Environment............................... 20 1.5 Aims and Objectives ............................................................................................. 27 1.6 Thesis Structure.................................................................................................... 29 2 Experimental Design and Methods ......................................................................... 30 2.1 Introduction to Experimental Design and Methodology ....................................... 31 2.2 Study Area ............................................................................................................ 32 2.3 Methodological Approach ..................................................................................... 42 3 Environmental Conditions Across the Three Habitats......................................... 54 3.1 Introduction ........................................................................................................... 55 3.2 Physiochemical Environmental Conditions .......................................................... 55 3.3 Chemical Environmental Conditions .................................................................... 69 3.4 Physical Environmental Conditions ...................................................................... 84 3.5 Discussion and Summary of Environmental Conditions ...................................... 92 4 Distribution of Biological Communities Across the Three Habitats .................. 99 4.1 Introduction ......................................................................................................... 100 4.2 Communities Recorded in Benthic Habitats ...................................................... 100 4.3 Communities Recorded in Hyporheic Habitats .................................................. 117 4.4 Communities Recorded in Phreatic Habitats ..................................................... 133 4.5 Invertebrate Community Distribution Across Three Habitats ............................ 148 5 Community Response to High and Low Flow Disturbances ............................. 154 5.1 Introduction ......................................................................................................... 155 5.2 Invertebrate Community Response to High Flows ............................................ 155 5.3 Invertebrate Community Response to Low Flows ............................................. 163 5.4 Summary............................................................................................................. 175 6 Morphological and Molecular Taxonomy............................................................. 177 6.1 Introduction ......................................................................................................... 178 6.2 The Gammaride .................................................................................................. 178 6.3 Identification Methods and Results .................................................................... 180 5 6.4 Discussion .......................................................................................................... 185 6.5 Summary ............................................................................................................ 187 7 Summary, Conclusions and Implications for Future Management
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