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Population Ecology of Eriophorum Latifolium, a Clonal Species in Rich Fen Vegetation

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Population Ecology of Eriophorum Latifolium, a Clonal Species in Rich Fen Vegetation Anders Lyngstad Population Ecology of Eriophorum latifolium, a Clonal Species in Rich Fen Vegetation Thesis for the degree of Philosophiae Doctor Trondheim, October 2010 Norwegian University of Science and Technology Faculty of Natural Sciences and Technology Department of Biology NTNU Norwegian University of Science and Technology Thesis for the degree of Philosophiae Doctor Faculty of Natural Sciences and Technology Department of Biology © Anders Lyngstad ISBN 978-82-471-2332-4 (printed ver.) ISBN 978-82-471-2333-1 (electronic ver.) ISSN 1503-8181 Doctoral theses at NTNU, 2010:179 Printed by NTNU-trykk Synopsis PREFACE It was the spring of 2005, and it was the right time to move onwards. A position as a research fellow working with long-term time series at the Museum of Natural History and Archaeology at the Norwegian University of Science and Technology (NTNU) was announced, and a PhD-project was developed with the studies of former haymaking lands at Sølendet and Tågdalen nature reserves as a starting point. Work began in earnest in January 2006, and continued at an ever increasing pace until July 2010, when all the parts of the thesis were finally completed and assembled. The study was financed by NTNU, and was carried out at the Museum of Natural History and Archaeology and the Institute of Biology, both NTNU. I am deeply grateful to my main supervisor Professor Asbjørn Moen at the Museum of Natural History and Archaeology, and my co-supervisor Associate Professor Bård Pedersen at the Institute of Biology. This project rests on the long-term studies of rich hayfens that were initiated by Asbjørn 40 years ago, and that are still ongoing, much due to his continuous effort. Without Asbjørn, there simply would be no thesis. Bård has been an equally important factor (to put it statistically) to the end result, especially in the long process of turning vague ideas into scientific questions. The quality of this thesis has been hugely improved by his lucid, analytical and clear thinking. I thank you both, for the opportunity, the guidance and the support. The collaboration with Radboud University Nijmegen, Institute for Water and Wetland Research, Department of Experimental Plant Ecology in The Netherlands has been particularly fruitful. I’m grateful for the kind reception during my visits there, and I want to thank Professor Hans de Kroon and, of course, Post. Doc. Eelke Jongejans. Eelke introduced me to, and guided me patiently through, the bewildering world of matrix models. Many others have been involved at different stages or in different parts of the project. Lester Rocha at the Institute of Biology has been invaluable in teaching me statistical methods, and also in how to successfully fight R. My long-term colleague Dag-Inge Øien has been, and still is, vital to the long-term studies, and his contribution is manifest throughout the thesis. I also want to thank Nina Sletvold, Line Johansen, Sverre Lundemo and Sølvi Wehn for help and good advice. The field work could not have been completed without the conscientious and patient help of Eva Ulvan, Ane Fremstedal, Sigrid Hansen and Christine Halseth. You managed to make measuring ”grass” quite fun! Credit is also due to James Speed and Richard Binns for proofreading manuscripts and correcting my English. I want to thank my family for helping out, Jan Eirik and Beate for their friendship, and Tonje for taking care of me (and feeding me), especially during the last months of writing. Finally, thank you Arvid and Sondre for reminding me there are things in life more important than a thesis. i Synopsis Synopsis CONTENTS Preface…………………………………………………………………………………………….………….……….…….. i Contents…………………………………………………………………………………………..……….……..………… iii List of papers……………………………………………………………………………….……………………………. 1 Declaration of contributions…………………………………………………………………………. 1 General introduction…………………………………………………………………………………………………. 2 Aims…………………………………………………………………………………………………………….……………. 6 Study species……………………………………………………………………………………………………….……. 7 Study areas………………………………………………………………………………………………….…..……........ 10 Sølendet Nature Reserve………………………………………………………….…..……….…….… 10 Tågdalen Nature Reserve……………………………………………………………...………………. 11 Vegetation regions and climate of Sølendet and Tågdalen nature reserves..… 14 Materials and methods……………………………………………..…….…………….……………….….….……. 23 Paper I. Rich fen vegetation and hay crop……………………………………….…………… 23 Paper II. Size of E. latifolium ramets…………………………………………..…….…..……… 25 Paper III. Population dynamics of E. latifolium………………………………………...…. 25 Paper IV. Flowering time series in E. latifolium………………………………………...… 28 Results and discussion………………………………………………………………………...………….…...……. 31 The vegetation and hay crop in boreal rich fen………………………………….………..... 31 The impact of long-term mowing on ramets of E. latifolium…………………..…… 32 Population dynamics of E. latifolium….………………………………………………..….…… 33 The impact of climate, mowing and cost of reproduction in E. latifolium….... 35 Conclusions and further studies……………………………………………………………………………...… 38 References…………………..……………………………………………….………………………….……………….... 40 Papers I-IV iii Synopsis Synopsis LIST OF PAPERS This thesis is based on four papers. I) Lyngstad, A., Moen, A. & Øien, D.-I. 2010. Rich fen vegetation and hay crop on traditionally used outlying land in central Norway. (manuscript) II) Lyngstad, A. 2010. Opposite effects of mowing on size of Eriophorum latifolium ramets in rich fen carpet and lawn communities. (manuscript) III) Lyngstad, A., Rocha, L. & Jongejans, E. 2010. Population dynamics of Eriophorum latifolium in boreal rich fens vary with mowing regime and plant community. (manuscript) IV) Lyngstad, A., Moen, A. & Pedersen, B. 2010. Effects of mowing and climate on flowering in Eriophorum latifolium in boreal rich fens, results from long- term monitoring. (manuscript) Declaration of contributions The ideas for the papers were conceived and developed jointly by Lyngstad and the supervisors Moen and Pedersen. Papers I and IV are based on long-term data (1967- 2008) collected, assembled and maintained by Moen with the help of numerous co- workers. Øien and Lyngstad has participated in this task since 1992 and 1999 respectively. Papers II and III are based on a field experiment planned by Lyngstad, Moen and Pedersen, and Lyngstad carried out the field work. The analyses of classification and ordination in paper I were chiefly done by Moen and Øien, whereas the analyses of hay crop were carried out by Lyngstad. Moen and Lyngstad have written the major parts of paper I. The modelling approach in Paper III was envisioned and worked out by Jongejans and Lyngstad, and data analyses was done by Lyngstad in collaboration with Rocha. Model development in Paper IV was done by Pedersen, and statistical analyses performed by Lyngstad. Lyngstad wrote papers III and IV with contributions from the co-authors. 1 Synopsis GENERAL INTRODUCTION Boreal rich fens Fens are mires that are influenced by both precipitation and water from the mineral soil. There are three local vegetation gradients used to differentiate fen vegetation in the Fennoscandian tradition of mire studies. The poor – rich gradient reflecting the pH and mineral concentration, the mire expanse – mire margin gradient (often) reflecting the peat depth, and the hummock – mud bottom gradient reflecting the groundwater level (e.g. Tuomikoski 1942, Sjörs 1948, Ruuhijärvi 1960, Persson 1961, Eurola 1962, Malmer 1962a, b). The poor – rich gradient in fens separates poor fen, intermediate fen, moderately rich fen and extremely rich fen (Du Rietz 1949, Sjörs 1983), poor fen has pH between 4 and 5.5, intermediate and moderately rich fen between 5 and 7, while extremely rich fen has pH from 6.8 to 8 (Rydin & Jeglum 2006). Moderately and extremely rich fen lawns in boreal areas in Norway often have Campylium stellatum as the dominant species in the bottom layer and Trichophorum cespitosum ssp. cespitosum as the dominant species in the field layer. Other abundant species are e.g. Thalictrum alpinum, Eriophorum latifolium, Molinia caerulea and Scorpidium cossonii (Moen 1990). Some exclusive extremely rich fen species in boreal Norway are Dactylorhiza lapponica, Listera ovata, Carex atrofusca, C. capillaris, C. capitata, C. hostiana, Kobresia simpliciuscula, Schoenus ferrugineus, Barbilophozia rutheana, Fissidens adianthoides and Palustriella commutata (Moen 1990: 197, Fremstad 1997). Mowing in outlying lands Traditional mowing of outlying lands to make hay for winter fodder has ceased in Scandinavia (Emanuelsson 2009), leading to successional changes in the plant cover. The upper boreal rich fens were traditionally mowed every second year using a scythe, and the typical hay crop (see Material and methods, Paper I) was 100-150 g/m2. Studies in boreal hay fens in central Norway (Moen 1990, Aune et al. 1994, 1996, Moen et al. 1999) have shown a reduction in the hay crop as well as the above- and below-ground biomass of the field layer with increasing mowing frequency. An increase in the ratio of the above-ground/below-ground biomass shows that mowing forces the plants to mobilise resources from their below-ground organs (Fitter 1986, Aune et al. 1996). The 2 Synopsis succession in formerly mown boreal grasslands and fens is slow, and recommenced mowing has largely resulted in changes in species’ abundance, whereas turnover of species has been minor. However, the changes have been more pronounced in the relatively more productive grassland and fen margin vegetation than in open fen areas
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