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The Influence of Climate and Atmospheric C02 On THE INFLUENCE OF CLIMATE AND ATMOSPHERIC C02 ON RADIAL GROWTH IN PSEUDOTSUGA MENZIESII AND PINUS PONDEROSA OVER THE 20™ CENTURY A thesis Presented to The Faculty of Graduate Studies of The University of Guelph by VANESSA STRETCH In partial fulfillment of the requirements for the degree of Master of Science July, 2011 © Vanessa Stretch, 2011 Library and Archives Bibliotheque et 1*1 Canada Archives Canada Published Heritage Direction du Branch Patrimoine de Pedition 395 Wellington Street 395, rue Wellington OttawaONK1A0N4 OttawaONK1A0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-82786-4 Our file Notre reference ISBN: 978-0-494-82786-4 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par I'lnternet, preter, telecommunication or on the Internet, distribuer et vendre des theses partout dans le loan, distribute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non­ support microforme, papier, electronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extraits substantiels de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission. In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondaires ont ete enleves de thesis. cette these. While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. 1+1 Canada ABSTRACT THE INFLUENCE OF CLIMATE AND ATMOSPHERIC C02 ON RADIAL GROWTH IN PSEUDOTSUGA MENZIESII AND PINUS PONDEROSA OVER THE 20Tff CENTURY Vanessa Stretch Advisor: University of Guelph, 2011 Dr. Ze'ev Gedalof This study illustrates changes in residual radial growth of Pseudotsuga menziesii (Douglas-fir) and Pmus ponderosa (ponderosa pine) in western North America over the past century. Using 382 tree-ring site chronologies, residual growth was statistically examined for evidence of direct CO2 fertilization and indirect changes in climatic sensitivities. Results demonstrate that a direct CO2 fertilization effect is unlikely in these species in western North America. Instead, a significant restriction in radial growth over the last century has been noted and associated with changing relationships to seasonal temper­ ature and precipitation parameters. Specifically, precipitation is more often becoming less limiting and temperature more limiting. These results may sug­ gest a change in the water-use efficiency of these species, a potential indirect implication of increased atmospheric CO2. These findings suggest that with a continually changing climate, the radial growth of P. menziesii and P. pon­ derosa in western North America will not respond uniformily and future forest ecosystem dynamics may be difficult to predict. ACKNOWLEDGMENTS First, I have to thank my advisor, Ze'ev Gedalof. For all of the times I felt uneducated and completely overwhelmed, your belief in me and constant assurance that I could do this pushed me through. Thank you for making me use MATLAB and making me think in ways I never thought I could. Oh, and thanks for sending me to so many conferences! I look forward to working with you again next year. Also, I have to thank my advisory committee member, Aaron Berg, for all of his time and revisions. I must thank the Natural Sciences and Engineering Research Council, Le Fonds Quebecois de la Recherche sur la Nature et les Technologies, the University of Guelph, and Ze'ev Gedalof for the funding to support this research. I'd also like to thank any and all dendrochronologists who have contributed data to the ITRDB, and particularly those who provided me with metadata. This thesis would not be possible without the sharing of data and knowledge within this community. It is one of which I am proud to now be a part. A special thanks my CEDaR labmates: Graham, Vesta, Kerry, and Rob. An extra special shoutout goes to Kerry for dragging me out of the lab and up to the 'Gonq and for being the best Habs fan in Ontario! I want to thank all of my fellow Hutt Dogs. You guys are the best! Thanks specifically for T&T, all the beer and gin, the Hutt Dog sports and outings, and for keeping me sane throughout this process. A further thanks to the entire Geography department at the University of Guelph. You have all made Guelph feel like i home. Also, I need to thank my family for supporting me throughout my lengthy university career. I am convinced you have no idea what I am studying, but your unconditional support is still very much appreciated. Erinn and Lisa, thanks for being the bestest friends I could have ever asked for. I could have never made the move and done this without your support! And lastly, Rashaad. Thank you for listening to me complain about my thesis (mostly MATLAB), for forcing me to work on days when I didn't feel like working, for teaching me LaTeX, for making all of my beautiful maps, and for being the most supportive person I know! I look up to you. n CONTENTS Table of Contents iii List of Figures vi List of Tables vii 1 Introduction 1 1.1 Background 1 1.2 Problem Statement 2 1.3 Goals &; Objectives 2 1.4 Thesis Outline 3 2 Literature Review 4 2.1 Introduction 4 2.2 Climate Change & CO2: A Historical Evaluation 5 2.2.1 Climatic Shifts over Time 5 2.2.2 Greenhouse Gases & Carbon Dynamics 6 2.3 Plant Physiology & C02 9 2.3.1 Photosynthesis & Carbon Sequestration 10 2.3.2 Acclimation & Downregulation 14 2.3.3 Water-Use Efficiency 16 2.3.4 Between-Species Differences 17 2.4 Possible Research Methodologies 18 2.4.1 Growth Chambers & Greenhouses 18 2.4.2 Open-Top Chambers 19 2.4.3 Free-Air Carbon Enrichment (FACE) Sites 21 2.4.4 Dendrochronology 23 2.5 Conclusion: Uncertainties & Knowledge Gaps 25 111 3 Research Design & Methods 28 3.1 Introduction: Research Design 28 3.2 Study Region 28 3.2.1 Pseudotsuga menziesii (P. menziesii) 28 3.2.2 Pinus ponderosa (P. ponderosa) 31 3.3 Data Compilation & Formatting 33 3.3.1 Raw-ring Width Data 33 3.3.2 Metadata (Site Factor Data) 34 3.3.3 Climate Data 34 3.4 Removal of Known Growth Factors 35 3.4.1 Tree-age 35 3.4.2 Climate 36 3.5 Analysis of Residual Chronologies 38 3.6 Climate Re-analysis 38 3.7 Sensitivity Analysis 39 3.8 Conclusion 40 4 Results: Chronology Analyses 41 4.1 Introduction 41 4.2 Results 41 4.2.1 Study Sites 41 4.2.2 Climatically Sensitive Chronologies 42 4.2.3 Residuals 44 4.2.4 Spatial Patterns in Residuals 47 4.2.5 Trends in Climate-related Growth 47 4.2.6 Sensitivity Analysis 50 4.3 Conclusion 56 5 Analysis & Discussion 58 5.1 Scholarly Discussion of Results 58 5.1.1 C02 Fertilization? 58 5.1.2 Implications of a Changing Climate 60 5.1.3 Other Impacts on Growth 64 5.2 Conclusion 66 iv 6 Summary &z Conclusions 67 6.1 Overview of Findings 67 6.2 Future Forest Dynamics 69 6.3 Potential Future Research 70 6.4 Conclusions 71 Works Cited 72 A Site-specific Climate &z Residual Analysis Results 84 B Site-specific Sensitivity Results 104 v LIST OF FIGURES 2.1 The role of CO2 in plant productivity 10 2.2 The specificity of RuBisCO 13 3.1 Natural distribution of P. menziesii 29 3.2 Natural distribution of P. ponderosa 32 4.1 Map of study sites in western North America 42 4.2 Frequency of climatic variables included in AIC best-fit models .... 44 4.3 % Explained variance from multiple linear regression 45 4.4 Trends in residuals of P. menziesii 48 4.5 Trends in residuals of P. ponderosa 49 4.6 Differences in climate parameter correlations 51 4.7 Frequency of significant decreases in radial sensitivity to climate pa­ rameters 53 4.8 Frequency of significant increases in radial sensitivity to climate pa­ rameters 54 4.9 Frequency of significant changes in direction of radial sensitivity to climate parameters 55 4.10 The spatial distribution of sites with changing climatic sensitivities . 56 VI LIST OF TABLES 2.1 C02 budgets from 1750 to 1990 9 2.2 Internal & external biological factors influencing photosynthesis ... 11 2.3 Advantages of free-air carbon enrichment (FACE) studies 22 3.1 Climatic data for regional subdivisions of the range of P.menziesii . 30 4.1 Proportion of climatically sensitive sites 43 4.2 Trends in residual chronologies 46 4.3 Species differences in residual trends 46 4.4 Significant trends in climate-related growth 50 A.l Site-specific climate & residual analysis results 86 B.l Site-specific sensitivity results 106 Vll CHAPTER 1 INTRODUCTION 1.1 Background The responses of forests to post-industrial increases in atmospheric CO2 remain under considerable debate.
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