How Are Plants Responding to a Changing Climate?
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Biodiversity, Conservation and Management 2008-2009 (MSc) How are plants responding to a changing climate? A case study of growth and allocation in Veronica alpina, Viola biflora, Veronica officinalis and Viola palustris in western Norway September 2009 Tessa Bargmann This work is submitted in partial fulfilment of the requirements of the degree of Master of Science University of Oxford, 2009 Candidate number: 264831 Table of Contents List of Figures ......................................................................................................................... iii Abstract ................................................................................................................................. iv Acknowledgements ................................................................................................................. v 1. Introduction ..................................................................................................................... 1 1.1 Climate change and alpine plants ....................................................................................... 1 1.2 The importance of plant traits and allocation for understanding growth and survival ...... 2 1.3 Conservation significance .................................................................................................... 5 1.4 Questions and Aims ............................................................................................................. 6 1.5 Focal species ....................................................................................................................... 7 2. Methods ........................................................................................................................ 11 2.1 Study sites ......................................................................................................................... 11 2.2 Plant collection and Morphometrics ................................................................................. 12 2.3 Data Analysis ..................................................................................................................... 14 3. Results ........................................................................................................................... 15 3.1 Veronica alpina .................................................................................................................. 15 3.2 Viola biflora ....................................................................................................................... 19 3.3 Veronica officinalis ............................................................................................................ 23 3.4 Viola palustris .................................................................................................................... 27 4. Discussion ...................................................................................................................... 31 5. Conclusion ..................................................................................................................... 36 6. References ..................................................................................................................... 37 Appendix ............................................................................................................................... 45 ii Candidate number: 264831 List of Figures Figure 1: Drawing of Viola biflora taken from CLOPLA (Klimešová & Klimeš, 1998) ......................... 8 Figure 2: Drawing of Veronica alpina taken from CLOPLA (Klimešová & Klimeš, 1998) .................... 8 Figure 3: Drawing of Veronica officinalis taken from CLOPLA (Klimešová & Klimeš, 1998) .............. 9 Figure 4: Drawing of Viola palustris taken from CLOPLA (Klimešová & Klimeš, 1998) .................... 10 Figure 5: The SEEDCLIM climate grid in western Norway taken from SEEDCLIM protocol (2009).. 11 Figure 6 a-e: Traits of Veronica alpina plotted against the climate variables precipitation and temperature ..................................................................................................................................... 17 Figure 7 a-c: Allocations of Veronica alpina plotted against the climate variable temperature ..... 19 Figure 8 a-c: Traits of Viola biflora plotted against the climate variables precipitation and temperature ..................................................................................................................................... 20 Figure 9 a-d: Allocations of Viola biflora plotted against the climate variables precipitation and temperature ..................................................................................................................................... 22 Figure 10 a-c: Traits of Veronica officinalis plotted against the climate variables precipitation and temperature ..................................................................................................................................... 24 Figure 11 a-d: Allocation of Veronica officinalis plotted against the climate variables precipitation and temperature .............................................................................................................................. 26 Figure 12 a-b: Traits of Viola palustris plotted against the climate variable precipitation ............. 27 Figure 13 a-e: Allocation of Viola palustris plotted against the climate variables precipitation and temperature ..................................................................................................................................... 29 iii Candidate number: 264831 Abstract Climate change is increasingly affecting organisms around the world. Local extinctions and range changes have been documented over the years, calling for both wide ranging, and detailed studies of the tolerances and adaptations of these species. In this study, plant traits are used to identify these responses in terms of growth and allocation of two alpine plants, Veronica alpina and Viola biflora, as well as two generalists, Veronica officinalis and Viola palustris, which are considered within a climate grid in western Norway. This study shows that these four plant species shift their growth and allocation to different life functions in response to temperature and precipitation increase, and that there are consistencies in responses between species pairs when it comes to allocation. This understanding is then used to discuss the possible implications for the future conservation of alpine plants in the light of climate change. iv Candidate number: 264831 Acknowledgements I owe my deepest thanks to Professor Vigdis Vandvik from the University of Bergen, without whose direction and support I could not have taken part in the SEEDCLIM project. I also want to thank Dr. Olav Skarpaas (University of Oslo), Dr. Kari Klanderud (Norwegian University of Life Sciences), Eric Meineri and Joachim Spindelböck from the University of Bergen for their guidance and discussions. I want to thank Professor Katherine Willis and Professor Robert Whittaker for their early input and suggestions, and Shonil Bhagwat for both his help in the early stages of dissertation planning and for looking over and making useful comments on the first draft. Thank you also to Professor Deborah Goldberg and Emily Farrer from the University of Michigan for their insights. Finally, I want to say thank you to Keno Ferter, Simon Le Mellec and Daja Ferter for their help in the field. v Candidate number: 264831 1. Introduction 1.1 Climate change and alpine plants It is generally accepted by the scientific community that climate is changing at an accelerated rate, and that it is having a profound impact upon plants and animals alike (IPCC, 2007; Parmesan & Yohe, 2003; Root et al., 2003; Walther et al., 2002; IPCC, 2001). The 2001 IPCC report warns that over the last 100 years the global temperature has increased by 0.6oC. However, it is not only the degree of warming during the last century that is alarming, but also the rate of warming, which is currently at its peak (IPCC, 2001). In addition to rising temperatures, changes in precipitation patterns have also been acknowledged. While it is accepted that global climate is changing, it is also important to point out that these changes are not globally uniform, and that some communities are affected more severely, while others remain more resilient during climatic changes (Walther et al., 2002). In addition, it is not only on a coarse scale that climatic impacts differ, but also on a finer scale, where smaller habitat niches exist for a given species. As a result of a shifting climate the geographical ranges of many species have altered, as exemplified by numerous scientific studies (i.e. Moritz et al., 2008; Pauli et al., 2007; Lesica & McCune, 2004; Molau & Alatalo, 1998). Increases in temperature and precipitation have placed additional pressures upon arctic and alpine communities in particular, because unlike their lowland counterparts, there is a limit to how far they can disperse upslope into colder and drier habitats (Roots, 1989). In addition, climate change is predicted to have the largest impact on the biodiversity of extreme environments like alpine and arctic biomes, where small changes in precipitation or temperature will have a proportionally larger effect (Sala et al., 2000). This means that these species face problems such as range contraction, and in the worst case, local extinction. As well as the general upslope