Master Thesis the Influence of Moose (Alces Alces) On
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MASTER THESIS THE INFLUENCE OF MOOSE (ALCES ALCES) ON NUTRIENT DYNAMICS IN A FEN ECOSYSTEM KASSIOPEIA DEVRIENDT SUPERVISOR: RUDY VAN DIGGELEN MASTER THESIS THE INFLUENCE OF MOOSE (ALCES ALCES) ON NUTRIENT DYNAMICS IN A FEN ECOSYSTEM Author: Kassiopeia Devriendt - University of Antwerp (UA) Supervisor: Prof. Dr. Rudy Van Diggelen - University of Antwerp (UA) Co-supervisor: Dr. Dries Kuijper – Mammal Research Institute (MRI), Polish Academy of Sciences Co-supervisor: Prof. Dr. hab. Bogumiła Jędrzejewska - Mammal Research Institute (MRI), Polish Academy of Sciences Left: Catchment area of Biebrza National Park (source: see references). Right: Moose in the brushwood (photo: Sanne Dirickx). Faculty of Biology Master thesis handed in to achieve the Masters degree of Biology with graduating option ‘Ecology and Environment’. University of Antwerp Mammal Research Institute (Polish Academy of Sciences) 2011 - 2012 “First, therefore I will speak of the Elke, which the Savages call a Mose: it is a very large Deare, with a very faire head, and a broad palme, like the palme of a fallow Deares horne, but much bigger, and is 6 footewide between the tipps, which grow curbing downwards: he is of the bignesse of a great horse”. Samuel Champlain, 1603 (in: Merril, 1920) “By coincidence she saw water in a lake, in the deepest of the valley, where farmers were collecting rushes, invading the marsh Latona went down, kneeled and leaned on the soil to take and drink from the water” (r5, Metamorphoses VI, 339-38) TABLE OF CONTENTS ABSTRACT…………………………………………………………………………….…………………..….p.1 GENERAL INTRODUCTION.....…………………………………………………….……………………...p.1 AIM OF THE STUDY………………………………………………………………….……………………..p.3 GENERAL HYPOTHESES ……..…………………………………………………….…………………..…p.3 STRUCTURE OF THE STUDY………………………...…………………………….…………………..….p.3 NUMERICAL ANALYSES STUDY SITE STUDY SPECIES PART 1: REMOVAL OF NUTRIENTS BY MOOSE Introduction Method…………………………………………………………………………….…………………..….p.6 1.1 Diet of moose in summer 1.1.1A Transect data 1.1.1B Microscopic diet analysis 1.1.2 Diet selectivity by moose 1.1.3 Comparison of the moose diet with other ungulate species 1.1.4 Comparison of the methods 1.1.5 Is there a link of browsing frequency with plant height and water level? 1.2 Place of nutrient removal 1.3 Amount of nutrients removed by moose 1.3.1 Moose density 1.3.2 Biomass removal by moose 1.3.3 Amount of nutrients removed from the browse species A. Salix cinerea B. Other deciduous trees C. Sedges D. Herbs Results………………………………………………………………………………………………….….p.14 1.1 Diet of moose in summer 1.1.1 Diet composition 1.1.2 Diet selectivity of moose 1.1.3 Comparison of the moose diet with other ungulate species 1.1.4 Comparison of the methods 1.1.5 Is there a link of browsing frequency with plant height and water level? 1.2 Place of nutrient removal 1.3 Amount of nutrients removed by moose Discussion……………………………………………………………………………..………………..….p.25 1.1 Diet of moose in summer 1.1.1A Plant category proportions 1.1.1B Species contributing to the moose diet and selectivity 1.1.2 Comparison of the moose diet with other ungulate species 1.1.3 Comparison of the methods 1.1.4 Is there a link of browsing frequency with plant height and water level? 1.2 Place of nutrient removal 1.3 Amount of nutrients removed by moose Conclusion……………………………………………..………………………………………………......p.29 PART 2: NUTRIENT INPUT IN THE ECOSYSTEM BY MOOSE Introduction Method……………………………………………..……………………………………………………....p.30 2.1 Place of nutrient release by moose 2.1.1 Pellet density per structural vegetation type 2.1.1A Transect data 2.1.1B Plot data 2.1.2 Link of pellet deposition with moose paths and rest places 2.2 Amount of nutrient input 2.2.1 Nutrient concentrations in moose and ungulate pellets 2.2.2 Amount of pellets 2.3 Amount of nutrient input 2.3.1 Pellet deposition 2.3.2 Urine deposition Results……………………………………………..………………………………………………..............p.33 2.1 Place of nutrient input 2.1.1 Pellet densities 2.1.2 Link of pellet deposition with moose paths and rest places 2.2 Amount of nutrient input 2.2.1 Nutrient concentrations in moose and ungulate pellets 2.2.2 Amount of pellets 2.3 Amount of nutrient input 2.3.1 Pellet deposition 2.3.2 Urine deposition Discussion……………………………………………..………………………………………………........p.36 2.1 Amount of nutrient input Conclusion……………………………………………..…………………………………………………...p.37 PART 3: NUTRIENT UPTAKE BY PLANTS Introduction Method……………………………………………..………………………………………………….…....p.38 3.1 Nutrient content of Salix cinerea 3.1.1 Nutrient (N and P) content of leaves and twigs per age category 3.1.2 Nutrient content in relation to twig diameter 3.2 Estimation of yearly nutrient uptake of Salix cinerea Results……………………………………………..…………..…………………………………………....p.41 3.1 Nutrient content of Salix cinerea 3.1.1 Nutrient (N and P) content of leaves and of twigs per age category 3.1.2 Prediction of nutrient content by twig diameter 3.2 Estimation of yearly nutrient uptake of Salix cinerea Discussion…………………………………………..……………………………………………………....p.42 3.1 N and P content in willows 3.2 Estimation of yearly nutrient uptake of Salix cinerea Conclusion……………………………………………..……………………………………………….......p.45 GENERAL DISCUSSION ON NUTRIENT DYNAMICS: SYNTHESIS OF PART 1, 2 AND 3 1. Do moose cause a redistribution or a recycling of nutrients?……………………………………….........p.46 2. Nutrient process rates by moose in comparison to willow requirements………………………...….........p.46 3. Do moose suppress or stimulate shrub growth? ……….............................................................................p.49 GENERAL CONCLUSION……………………………………………………………..………………....p.51 ACKNOWLEDGEMENTS REFERENCES……………………………………………………………………………………………...p.53 APPENDIX I: Maps of the study site APPENDIX II: Illustrations on the method of part 1 and 2 APPENDIX III: Tables on the calculation of nutrient removal by moose APPENDIX IV: Diet preferences linked to plant height and water level APPENDIX V: Statistical analyses of forage differences in structural vegetation types APPENDIX VI: Table on the diet of moose APPENDIX VII: Winter diet of Moose APPENDIX VII: Calculation of the yearly nutrient uptake by existing vegetation APPENDIX IX: Shrub encroachment in BNP APPENDIX X: Reference collection microscopic diet analysis ABSTRACT This study investigates the influence of moose (Alces alces), a large herbivore, on nutrient dynamics in an undisturbed fen ecosystem. This is of particular interest as mineralisation rates in fen systems are low, offering the potential to moose to enhance the nutrient turnover by short-cutting the decomposition. A nutrient balance of nitrogen (N), the limiting nutrient in Biebrza National Park (BNP), and phosphorus (P) is developed, in order to examine the relevance of moose activities concerning nutrients in the ecosystem. The comparison of the amount of moose’s nutrient input with mineralisation rates and atmospheric nutrient deposition is in this respect of major importance. Furthermore, it is investigated whether or not moose re-allocate nutrients by foraging and depositing excretion in different habitats, as often done by herbivores. In addition, the estimated yearly nutrient uptake of the vegetation – affected by an enhanced nutrient turnover – is compared with the total nutrient input. Finally, I discuss whether moose, by suppression or stimulation, may cause a shift in specific plant communities or species. This study indicates that moose have a substantial impact on N dynamics, specifically in undrained sites. Moose cause no nutrient redistribution but generate a local recycling of nutrients in the brushwood, dominated by willows and birches, instead. As a consequence, moose may stimulate brushwood expansion, in spite of the mechanical damage caused. This stimulatory effect contrasts with the often supposed role of moose in suppressing forestation. As this study demonstrates, moose may not be considered as an appropriate restoration tool to suppress ongoing shrub encroachment in BNP. GENERAL INTRODUCTION Nutrient dynamics imply fluxes of nutrients moving from one compartment to another (Fig.1), the rate of which is described by nutrient turnover rate (Begon et al., 2009). At low turnover rates, nutrients are essentially stored for a long time, at high rates they quickly become available. At low turnover rates, herbivores have potentially the largest effect on nutrient dynamics (Sjögersten et al., 2010). Examples could include nutrient cycling in undisturbed fen ecosystems, because these are characterised by a very low mineralisation rate (Van der Linden et al., 1996; Riutta, 2008; Olde Venterink et al., 2009). Consequently, external inputs may change the cycling rate but not the cycle itself (Bokdam, 2001). A fen is partly fed by groundwater and partly by surface water (Vandenbussche et al., 2002). The water level is at or very close to the surface level throughout the year (Hermy et al., 2004), leading to anoxic conditions (Van der Linden et al., 1996) and a very low decomposition rate (Chapin et al., 2003). As a consequence, in the long term, nutrients are sequestrated in persistent organic material, called peat (Chapin et al., 2003; Riutta, 2008). Herbivores can accelerate nutrient cycling directly by short-cutting the decomposition process (Van der Wal & Brooker, 2004). They transfer nutrients from their forage into urinary and faecal excretion. Nutrient recycling by this process is normally much faster than the turnover rate of litter (Van der Wal & Brooker, 2004). Moreover, selective foraging by large herbivores may result in a redistribution of nutrients