The Biodiversity and Metabolism of Peatland Pools
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THE BIODIVERSITY AND METABOLISM OF PEATLAND POOLS JEAN MARIA BEADLE Submitted in accordance with the requirements for the degree of Doctor of Philosophy The University of Leeds School of Geography September 2015 The candidate confirms that the work submitted is her own, except where work which has formed part of jointly authored publications has been included. The contribution of the candidate and the other authors to this work has been explicitly indicated below. The candidate confirms that appropriate credit has been given within the thesis where reference has been made to the work of others. Chapter 2 of this thesis is based on: Beadle Jeannie M, Brown Lee E, Holden Joseph (2015). Biodiversity and ecosystem functioning in natural bog pools and those created by rewetting schemes. WIREs Water, 2: 65-84. doi: 10.1002/wat2.1063. The paper was written by the candidate but review comments from both co-authors were included in the submitted version. This copy has been supplied on the understanding that it is copyright material and that no quotation from the thesis may be published without proper acknowledgement. © 2015 The University of Leeds and Jean Maria Beadle Acknowledgements I am especially thankful to my supervisors, Dr. Lee Brown and Professor Joseph Holden, for their invaluable guidance, encouragement and support with all aspects of my research. I am also extremely grateful to Simon Wightman and Gareth Edwards, my supervisors from the Royal Society for the Protection of Birds (RSPB), and to Steve Garnett from the RSPB for all his help with site selection and access at Geltsdale RSPB Reserve. I am indebted to Rachel Gasior, David Ashley, Martin Gilpin and the rest of the laboratory staff in the School of Geography, University of Leeds, for their patience and help with all aspects of laboratory work. I am also especially thankful to Ed Turner for his kind permission to use the Forsinard depth profile data analysed in Chapter 6. Special mention must also be made of Chris Carter, for identifying and producing remarkable images of algae found in the pools. Thanks must also go to the following people for verifying my invertebrate identifications; Garth Foster (Coleoptera), Steve Brooks from the Natural History Museum (Chironomidae) and Lynne Speight from APEM Ltd. (all other taxa). Also, for their help in organising site access, I am grateful to Andy Lloyd and Alistair Lockett (Peatscapes), Peter Christopherson and Astrid Hanlon (Yorkshire Peat Partnership), Chris McCarty (Natural England), Jeremy Greensides and Lindsay Waddell (Raby Estates), Peter Welsh (National Trust), Jonny Wildman (Dinsdales) and Duncan Hutt (Northumberland Wildlife Trust). Thanks must also go to Lorna Sherrin, Rob Rose and Beverley Dodd (ECN) for their help with Moor House site access and weather data, and to Annika Kristofferson for providing the Abisko weather data. This research was funded by a Natural Environment Research Council (NERC) studentship (NE/J50001X/1) with CASE support from the RSPB for which I am grateful. The fieldwork in Abisko was funded by an INTERACT Transnational Access grant to Dr. Lee Brown and Dr. Graeme Swindles. Finally, I would like to dedicate this thesis to Rob, whose unwavering love and warped sense of humour have kept me sane, and to my family, especially my wonderful Mum and Dad, Jean and George Beadle, whose love and support is beyond measure and means more than I can ever hope to express. Abstract Knowledge of the ecology and ecosystem functioning of pools created by the peatland restoration method of drain-blocking is limited. This thesis provides the most comprehensive study of these peatland pool ecosystems to date. Data were collected to investigate the spatial and temporal distribution of macroinvertebrate communities in both drain-blocked and naturally-occurring pools on areas of blanket bog in northern England. Corresponding environmental data were analysed to identify any mechanisms underpinning biodiversity metrics. The metabolism (i.e. rates of photosynthesis and respiration) of drain-blocked pools was estimated using the diel dissolved oxygen (DO) method. Drain-blocked pools were found to house macroinvertebrate communities similar to those in natural pools, including taxa characteristic of acid-mire pool habitats, thus providing suitable habitat for peatland aquatic taxa. There was some evidence that pools in reprofiled drains were more biodiverse than L-shaped pools. New pools were colonised quickly with taxon richness peaking in pools aged five to eight years, and pools aged 15 years housing some fauna not found in younger pools. Results suggested that gamma diversity would be best encouraged by creating a range of pool sizes with diverse vegetation composition. Drain-blocked pools were found to be strongly heterotrophic, with high rates of respiration driven by elevated levels of dissolved organic carbon and the relatively large sediment-water interface. Water temperature and DO profiles obtained from pools in England, Scotland and Sweden revealed severe diurnal stratification regimes, with implications for sampling methodologies, gas fluxes and carbon cycling. This research will help to inform future drain-blocking schemes and also has relevance for studies analysing the biogeochemistry and biota of other shallow waterbodies. List of Abbreviations DO dissolved oxygen DOC dissolved organic carbon EC electrical conductivity GPP gross primary production MOB methane oxidising bacteria NEP net ecosystem production POC particulate organic carbon R respiration TDS total dissolved solids Glossary Alpha diversity The diversity of a single site (local species pool). Autotrophic A system which fixes more carbon via photosynthesis than it respires. An organism capable of photosynthesis. Beta diversity The difference in species composition between sites. Dystrophic Having brown acidic water that is low in oxygen and supports little life, owing to high levels of dissolved humus. Eutrophic Rich in nutrients and so supporting a dense plant population, the decomposition of which may be harmful to biota by depriving it of oxygen. Gamma biodiversity Diversity measured at a landscape scale (the regional species pool). Heterotrophic A system which releases more carbon via respiration that it fixes via photosynthesis. An organism which cannot fix carbon for itself and relies on external sources. Humic substances A major component of the DOC in peatland aquatic systems; an organic compound of decaying organic matter made up of humic and fulvic acids. Minerotrophic Fed by water which has flowed over or through rocks or other minerals, often acquiring dissolved chemicals which raise the nutrient levels and reduce the acidity. Oligotrophic Relatively poor in plant nutrients but often containing abundant oxygen in the deeper parts. Ombrotrophic Fed exclusively by precipitation TABLE OF CONTENTS Chapter 1: Introduction ................................................................................................................ 1 1.1 Research context ........................................................................................................... 1 1.2 Research aims ............................................................................................................... 4 1.3 Thesis structure ............................................................................................................. 5 Chapter 2: Biodiversity and ecosystem functioning in natural bog pools and those created by rewetting schemes ........................................................................................................................ 9 2.1 Introduction .................................................................................................................. 9 2.2 Natural pools ............................................................................................................... 14 2.2.1 Formation, distribution and morphology ........................................................... 14 2.2.2 Primary producers ............................................................................................... 15 2.2.3 Macroinvertebrates ............................................................................................ 17 2.2.4 Amphibians ......................................................................................................... 20 2.3 Pools created by restoration measures ...................................................................... 21 2.3.1 Background ......................................................................................................... 21 2.3.2 Primary producers ............................................................................................... 24 2.3.3 Macroinvertebrates ............................................................................................ 25 2.3.4 Amphibians ......................................................................................................... 27 2.4 Environmental influences on macroinvertebrate community composition .............. 27 2.4.1 Pool size / permanence ....................................................................................... 28 2.4.2 Habitat heterogeneity ......................................................................................... 32 2.4.3 Pool age ............................................................................................................... 34 2.4.4 Dispersal of aquatic invertebrates .....................................................................