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Evaluation of the Ecological Impacts of Beaver Reintroduction on Aquatic

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Evaluation of the Ecological Impacts of Beaver Reintroduction on Aquatic Evaluation of the ecological impacts of beaver reintroduction on aquatic systems Alan Law 2014 Submitted to Biological and Environmental Sciences School of Natural Sciences University of Stirling Scotland For the degree of Doctor of Philosophy Supervisor: Dr. Nigel Willby STATEMENT OF ORIGNINALITY I hereby confirm that this PhD thesis is an original piece of work conducted independently by the undersigned and all work contained herein has not been submitted for any other degree. All research material has been duly acknowledged and cited. Signature of Candidate: Alan Law Date: ii GENERAL ABSTRACT The extent and quality of freshwater systems is declining globally. Combined with past drainage, straightening and flow regulation, current systems are often functional but not pristine. Conservation, creation and restoration of freshwater systems is common but requires significant planning, resources and active monitoring and may only be a short-term solution to the long- term problem of destruction and loss of riparian zones. Beavers (Castor spp.) have the ability to create physical and biological habitat heterogeneity through the construction of woody debris dams, thereby restoring lost natural discontinuities in freshwater systems. Beavers may thus offer a natural, more passive solution to the need for wetland restoration or creation and the problem of homogenisation of watercourses. As such, numerous beaver reintroductions and introductions have been undertaken based in part on restoring this lost natural heritage. However, it is crucial to be able to predict the potential effects on existing biota of physical modifications by beavers to ecosystems, especially in the light of further population expansion, whilst also disentangling these effects from other influences, namely herbivory. The impact of beavers on aquatic systems was studied using a combination of field-based surveys and experiments, using aquatic plants and macro-invertebrates as indicators of hydromorphological changes and to quantify the effects of direct foraging. The research presented in this thesis demonstrates beaver adaptive foraging behaviours between terrestrial and aquatic habitats, whilst feeding highly selectively, optimally and opportunistically, using the white water lily (Nymphaea alba) as a model species. The effects of beaver foraging on the aquatic plant resource and diversity was low over short time spans (e.g. 1 year), but when selective foraging was assessed over greater time scales (e.g. 10 years) the effects of foraging were distinct. Significant changes in aquatic plant height, biomass, richness, diversity and composition were observed over this time period due to selective grazing on large rhizomatous species (e.g. Menyanthes trifoliata). These direct effects occurred even though changes in water levels, which are commonly believed to be the main driver of beaver influence on aquatic vegetation, were negligible. In a separate study in Sweden where beavers commonly constructed dams, with ponds then forming upstream, the aquatic plant and coleoptera species iii richness and composition differed in comparison to adjacent non-beaver created wetlands. Therefore, having a range of wetland types in the environment increases physical and biological heterogeneity creating unique niches that are exploited by disparate taxa. The construction of a series of dams within a single reach of stream flowing through a Scottish agricultural landscape also increased physical habitat diversity. Distinctive macroinvertebrate assemblages and modified functional diversity were associated with each dominant habitat type in the stream, resulting in increased landscape scale richness. The findings of this thesis confirm that beaver engineering and foraging has the potential to create unique and highly heterogeneous wetland and stream habitats within landscapes that enhances richness and diversity for multiple species groups. This thesis also supports part of the rationale for the trial reintroduction of beaver to Scotland that beavers can restore degraded habitats. iv ACKNOWLEDGMENTS First and foremost my thanks, respect and appreciation go to my supervisor Nigel Willby whose help was always on hand, continually pushed me in the right direction and made the PhD a thoroughly enjoyable experience. I extend this thanks to Charles Perfect who is incredibly kind, patient and helpful, despite my questions over the years. I also thank David Gilvear who advised me in my first year and helped to obtain funding for this thesis. I thank both Carl Sayer (University College London) and Nils Bunnefeld (University of Stirling) for taking the time to read and examine my thesis. Their constructive comments have significantly improved the quality of work presented herein. I also wish to thank Paul Ramsay, Rosin Campbell-Palmer (Royal Zoological Society of Scotland), all staff at the Scottish Beaver Trial, Simon Jones (Scottish Wildlife Trust), Martin Gaywood (Scottish Natural Heritage), Garth Foster (Aquatic Coleoptera Conservation Trust) and Kevin Jones. The thesis was funded by the University of Stirling’s Horizon studentship which provided structure and services that were essential to the completion of this thesis. Scott Jackson, Ronald Balfour, James Weir and Bill Jamison also provided vital technical support of which I’m extremely grateful. And in no particular order I thank these people who have helped me with my work or kept my brain (semi-)functioning: Colin Bull, Adam Varley, Rab Rouger, Rebecca Barclay, Fiona Thomson, Antoine Keruzoré, Zarah Pattison, Mel Van Niekerk, Anwen Bill, Anna Doeser, Fraser Murdoch, Louis Kitchen, Justyna Olszewska and Willy Yeomans. To Emma, Mum, Graeme, Karen and Daisy who have inspired me to develop a steely work ethic and created a happy, loving home. This thesis is dedicated to James Wiseman (1987- 2013) who left us too soon. v TABLE OF CONTENTS STATEMENT OF ORIGNINALITY .............................................................................................. ii GENERAL ABSTRACT .............................................................................................................. iii ACKNOWLEDGMENTS.............................................................................................................. v TABLE OF CONTENTS ............................................................................................................. vi LIST OF FIGURES .................................................................................................................... ix LIST OF TABLES .................................................................................................................... xiv CHAPTER 1 – General introduction ......................................................................................... 1 1.1 Scientific background............................................................................................... 1 1.2 The return of the beaver .......................................................................................... 3 1.3 Overview of research aims and hypotheses tested ................................................. 12 CHAPTER 2 – Indirect and direct effects of beavers alter richness, composition and diversity of aquatic plants ...................................................................................................... 13 2.1 Introduction ........................................................................................................... 13 2.2 Methods ................................................................................................................ 14 2.3 Results .................................................................................................................. 15 2.5 Discussion ............................................................................................................. 17 2.6 Conclusion ............................................................................................................ 23 CHAPTER 3 – Beavers and lilies: selective herbivory and adaptive foraging behaviour .. 24 3.1 Abstract ................................................................................................................. 24 3.2 Introduction ........................................................................................................... 25 3.3 Methods ................................................................................................................ 27 3.4 Results .................................................................................................................. 31 3.5 Discussion ............................................................................................................. 37 3.6 Conclusion ............................................................................................................ 40 vi CHAPTER 4 – Medium vs. short-term effects of herbivory by Eurasian beaver on aquatic vegetation ................................................................................................................................ 41 4.1 Abstract ................................................................................................................. 41 4.2 Introduction ........................................................................................................... 42 4.3 Methods ................................................................................................................ 45 4.4 Results .................................................................................................................
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