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Duckweed Relationships The Ecology and Palaeoecology of Diatom – Duckweed Relationships Thesis submitted for the degree of Doctor of Philosophy University College London by David Emson Department of Geography University College London and Department of Botany Natural History Museum February, 2015 1 I, David Emson 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. David Emson 2 Abstract This thesis focuses on the ecology and palaeoecology of diatom-duckweed relationships and utilises a combined experimental, ecological and palaeoecological approach. In particular, the study sought to determine the potential of the epiphytic diatom Lemnicola hungarica to be utilised as a proxy indicator of past dominance of duckweed (Lemna) in small ponds. To this end, contemporary sampling of epiphytic diatom assemblages from a variety of macrophytes (including multiple samples of free-floating plants) were collected from around the world and analysed for diatom epiphytes. In this study, even despite significant environmental gradients, L. hungarica showed a significant association with free-floating plants (including Lemna spp.) as did Sellaphora seminulum. To determine whether this relationship might be used to infer Lemna- dominance in sediment cores, diatom assemblages were analysed in surface sediments from English Lemna and non-Lemna covered ponds and in a core from a pond (Bodham Rail Pit, eastern England) known to have exhibited periods of Lemna-dominance in the past. In both cases, the data suggested that both L. hungarica and S. seminulum were excellent predictors of past Lemna-dominance. Finally, to infer the consequences of Lemna-dominance for the long-term biological structure and ecosystem function of the Bodham Rail Pit, the sedimentary remains of diatoms, plant pigments, and plant and animal macrofossils were enumerated from two sediment cores. These stratigraphic data were compared with the diatom Lemna-indicator metric which indicated three distinct Lemna cycles. Sediment core analyses suggested major compositional, structural and ecological changes brought about by the Lemna cycles, especially in the submerged macrophyte community and in fish-invertebrate relationships. These data reveal that duckweed proliferation, often brought about by eutrophication and terrestrialisation in ponds, can result in dramatic ecological changes due to a strong physical ecosystem engineering effect. 3 Acknowledgements I would like to start by sincerely thanking my patient and supportive supervisors, Carl Sayer and Helen Bennion for their unstinting belief, guidance and unswerving direction both from the head and just as importantly from the heart. I will never be able to thank them enough, not just for their academic prowess but also because of their extraordinary and uplifting human qualities particularly in my hour of need during those dark and troubled times. This research project was, and forever will be, my personal therapy and I’ve had two of the best ‘therapists’ a person could ever wish for. Thank you for being there and thank you for acting as a bridge between me and my mental health media work! My late wife Daksha showed me that what you do for a living is important, but how you do it is even more important and Carl and Helen are the epitome of this sentiment. I wish to express my gratitude to all those good folk in the department of Geography, ECRC and ENSIS for their unconditional support, advice and inspiration particularly those humble legends Rick Battarbee, and Hilary and John Birks. For me, an honour and a humbling privilege and I hope that I managed to glean some of their words of wisdom. Admiral Nelson acknowledged that the real strength of a ship was the people that manned the ship and for me, the people of the good ship ‘HMS Geography’ were a towering strength as they taught me how to sail and to navigate palaeoecological waters properly. Frankly, no laboratory then no research, and I would like to say a really big and heartfelt thank you to the often unsung heroes of the laboratory Janet Hope, Tula Maxted, ‘the Colonel’ Ian Patmore, and Kevin ‘Brummie’ Roe for teaching me the ropes and smoothing out those experimental wrinkles upon my brow with suggestions and advice that money cannot buy. Thank you for that very personal touch. Also, a big thank you to Miles Irving and Cath D’Alton for providing maps so quickly and so professionally from their drawing office cabin. Another big thank you to Lady P (aka Helen Greaves) for making available the water chemistry plot covering those magical years of monitoring the old Rail Pit. A huge debt of gratitude to Pat Cubitt the erstwhile landowner and farmer for making the Bodham Rail Pit feel like it was my own garden pond. My sincere thanks to Jorge ‘Bear’ Salgado and Gizem Bezirci for expertise in coring and air-guitars! 4 I am indebted to Tom Davidson for all times sake and for the opportunity to visit NERI, Silkeborg, Denmark, and grateful thanks to Hugh Clout and the Mead Scholarship for kindly providing the funds to visit NERI; Suzanne McGowan (University of Nottingham) for so much more than her renowned expertise in pigment analyses; Eileen Cox and Elliot Shubert (NHM) for letting me loose in the Botany lab and SEM suite; ENSIS for financial support; Ben ‘Guru’ Goldsmith, Ewan ‘Newt’ Shilland, Anson Mackay, Simon Turner, Peter Langdon, Roger Flower and Patrick Rioual for providing data and samples with a nod and a wink. My grateful thanks to Handong Yang and Neil Rose for the radiometric analysis and to Martin ‘Time-Team’ Bridge for dendrochronological and brachiation expertise in oak trees! Also, a very warm appreciation to Paul Wood and Lynda Howard (Loughborough University), Hilary Birks (University of Bergen) and Pauline Lang (SEPA) for their expertise in the identification of various biological bits and pieces. A special many ta’s to Kevin Ruddy and Maria Asunción Ramos Cruz for providing (intact) macrophytes from Thailand and Peru. A special thanks to Gordon Copp and Serhan Tarkan (CEFAS), and ‘Lord’ Keith Wesley for teaching me all about our crucial crucians. A very heartfelt and sincere thank you to June and Derek Sayer for making their home feel like a home from home to me…and for the other many waifs and strays from UCL. I owe a huge debt of gratitude to the late and great Frank Round for his letters of support, encouragement and inspiration. Last but certainly not least, I would like to thank everyone past and present at the ECRC for simply being whom and what you are; your wonderful, warm, considerate and respectful presence shown towards the old hippie will be forever cherished; and I will forever carry those magical times that we all shared in the research microscope room, the laboratories and on fieldwork with me and beyond. A special ‘nod and a wink’ to Camel and Andrew Latimer as your friendship and musical genius kept my body and soul together as you helped me traverse those harsh and unforgiving deserts of my life, and in particular for ‘Spirit of the Water’ and ‘For Today’. 5 I dedicate this research to the loving memories of my father Raymond, my mother Margaret, my wife Daksha and my little bundle of joy, my baby daughter Freya. Their souls in heaven are stars at night; they will guide me on my way before we meet again…another day. 6 CONTENTS TITLE 1 ABSTRACT 3 ACKNOWLEDGEMENTS 4 DEDICATION 6 TABLE OF CONTENTS 7 LIST OF TABLES 14 LIST OF FIGURES 16 Chapter 1 – Introduction. .. 22 1.1 Research background. 22 1.2 Small lowland ponds. 23 1.2.1 Definition. 23 1.3 Pond or lake: does size make a difference? . 23 1.4 Pond size and conservation value. 25 1.5 Pond and shallow lake ecology, eutrophication and alternative stable states. 27 1.6 Floating plant dominance as an ecological stable state. 29 1.7 Ecosystem engineers. 30 1.7.1 Definition. 30 1.7.2 Effects of ecosystem engineers. 30 1.8 Invasive aquatic plants. 31 1.8.1 Biological invasions and ecosystem engineering. 31 1.8.2 Invasive free-floating plants. 32 1.8.3 Lemna minuta Kunth. 33 1.9 Tracking ecological change in shallow lakes and ponds. 34 1.10 Lemna-epiphytic diatom history. 37 1.10.1 Diatoms and their importance in the aquatic environment . 37 7 1.11 Epiphytic diatom community structure. 38 1.11.1 Relationship between epiphytic diatoms and aquatic macrophytes. 39 1.11.2 Physical and chemical hypotheses of periphyton and substrate. 43 1.11.3 Artificial substrata and epiphyton. 44 1.12 Overall aims and specific research questions to be addressed. 45 1.13 Structure and outline of thesis. 47 Chapter 2 – Pilot study sites and diatom analysis methods. 52 2.1 Introduction. 52 2.1.1 Pilot study sites. 52 2.2 Methods. .. 56 2.2.1 Macrophyte sampling. 56 2.2.2 Epiphytic diatom slide preparation. 57 2.2.3 Diatom slide preparation from surface sediments. 59 2.2.4 Diatom counts. 59 2.2.5 Water chemistry. 60 2.2.6 Numerical methods. 60 2.3 Availability of supporting data. 62 Chapter 3 – Is there a reliable host macrophyte-diatom association between the Lemnaceae and Lemnicola hungarica: developing a novel approach for inferring past pond ecology?. 68 3.1 Introduction. 68 3.1.2 Host macrophyte and epiphytic diatom specificity. 68 3.2 Lemnicola hungarica and duckweed. 69 3.3 Aims and methods. 73 3.3.1 Aims. 73 3.3.2 Methods. 73 3.4 Results. 75 3.4.1 Freshwater macrophytes and associated epiphytic diatom floras. 75 3.4.2 Relationships between contemporary epiphytic diatoms and macrophyte habitat.
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