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δ¹³C and stomatal density variability in modern and fossil leaves of key plants in the western United States Item Type text; Dissertation-Reproduction (electronic) Authors Van de Water, Peter Kent Publisher The University of Arizona. Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction or presentation (such as public display or performance) of protected items is prohibited except with permission of the author. Download date 03/10/2021 21:13:32 Link to Item http://hdl.handle.net/10150/289058 INFORMATION TO USERS This manuscript has t>een repfoduced fRxn the microfilm master. UMI fUms the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may t>e any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. 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Ml 48106-1346 USA 800-521-0600 NOTE TO USERS Page(s) not included in the original manuscript are unavailable from the author or university. The manuscript was microfilmed as received. 138 This reproduction is the best copy available. UMI 6"C AND STOMATAL DENSITY VARL\B1LITY IN MODERN AND FOSSIL LEAVES OF KEY PLANTS IN THE WESTERN UNITED STATES by Peter Kent Van de Water Copyright O Peter Kent Van de Water 1999 A Dissertation Submitted to the Faculty of the DEPARTMENT OF GEOSCIENCES In Partial Fulfillment of the Requirements For the Degree of DOCTOR OF PHILOSOPHY In the Graduate College THE UNIVERSITY OF ARIZONA 1999 UMI Number 9957980 Copyright 1999 by Van de Water, Peter Kent All rights reserved. UMI^ UMI Microform9957980 Copyright 2000 by Bell & Howell Information and Learning Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. Bell & Howell Information and Leaming Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, Ml 48106-1346 2 THE UNIVERSITY OF ARIZONA « GRADUATE COLLEGE As members of the Final Examination Committee, we certify that we have read the dissertation prepared by PETER KENT VAN DE WATER entitled 6 13c AND STOKATAI- DFWSTTY vartartt ttv tw MnnFWW Awn Fn<;<;TT LEAVES OF KEY PLANTS IN THE WESTERN UNITED STATES and recommend that it be accepted as fulfilling the dissertation requirement; for the Degree of Doctor of Philosophy Date Date t<iAA^/99 Date Date llll'r/'i'f • Date Final approval and acceptance of this dissertation is contingent upon the candidate's submission of the final copy of the dissertation to the Graduate College. I hereby certify that I have read this dissertation prepared under my direction and recommend that it be accepted as fulfilling the dissertation requirement.// ^ Dissertation Director Steven W. Leavitt Date 3 STATEMENT BY AUTHOR This dissertation has been submitted in partial fulfillment of requirements for an advanced degree at the University of Arizona and is deposited in the University Library to be made available to borrowers under rules of the Library. Brief quotations fi'om this dissertation are allowable without special permission, provided that accurate acknowledgment of source is made. Requests for permission for extended quotation from or reproduction of this m^u^ppt in yl^le or in part may be granted by the copyright holder. ^J^ SIGNED: 4 ACKNOWLEDGMENTS I am deeply indebted to the many individuals who have brought me to the point of writing this document. My interests in paleoenvironments and the earth sciences in general were inspired by associates of the Desert Laboratory at Tumamoc Hill, the Laboratory of Tree-ring Research, and the department of Grosdences, at the University of Arizona. These include my doctoral committee consisting of Steven W. Leavitt, Julio L. Betancourt, Jay Quade, Austin Long and David G. Williams. I have had the good fortune to be associated with committee members that have accommodated my every need. Additional thanks are given to those who got me through graduate school. These include the Tumamoc Hill Gang; Jan Bowers, Ana MacKay, Paul Martin, Kik Moore, Mary Ellen Morbeck, Donna Opocensky, Betsy Pierson, Jan Price, and Jack Wolf. I am especially indebted to Peter J. Mehringer, Jr. who opened the door to secrets of the past and became a lifelong friend. Similariy, Bob Miller, showed me that I could comprehend the deep history of geology and have fun doing it. I was similarly influenced by Peter E. Wigand and Saxon Sharpe at the Desert Research Laboratory in Reno Nevada. I am especially grateful to Mary Kay O'Rourke who believed in my ability to analyze pollen grains and provided me with an income throughout my graduate student career. Numerous colleagues helped my completion of tasks and ultimately facilitated my doctoral work. These include Luis Fernandez, Kira Flanagan, Anne Hyvarinnenin, Nicos Melitos, and Seumas Rogan. Funding was provided through NSF grant (EAR-9418268) to Steven W. Leavitt and Julio L. Betancourt. Field and laboratory assistance was provided by a number of people and is greatly appreciated. They include; Larissa Aumand, Angela Barclay, Mark Betancourt, Gabriel Cisneros, Tom Harian, Camille Holmgren, Mika Jensen, Tom Jones, Danielle Kane, Kerrie Kuehn, Claudio Latorre, Melanie Lenart, Vada Maryol, Leslie Parkinson, Jon and Lisa Pedicino, Nicole Ricketts, Kate Rylander, Staffan Schorr, Mike Wall and Edward Wright. Access to the mass spectrometer was provided by Austin Long and Dave Dettman. Midden material was graciously provided by Tom Van Devender, GeofiTSpaulding, and Peter Wigand. Discussions concerning the use of carbon isotopes has been greatly facilitated by James Ehleringer, Hyrum Johnson, and Larry Toolin. Finally, I thank my family. Without the love and guidance of my parents, and the inspiration that they provided throughout my life, this dissertation would not have been possible. I also owe special thanks to my sister who has always had the ability to remind me of what's important and how foolish we all look at times. DEDICATION This dissertation is dedicated to Lisa who has instructed me on the fine art of life. 6 TABLE OF CONTENTS LIST OF ILLUSTRATIONS 9 LIST OF TABLES 10 ABSTRACT 11 INTRODUCTION 13 Atmospheric Carbon Dioxide 16 Plant Physiology 27 Stable Carbon Isotopes 33 Stomatal Density 41 Stomatal variability 44 Intra-plant stomatal density 44 Inter-plant stomatal density 46 Dissertation Format 53 PRESENT STUDY 56 Introduction 57 Sampling Strategy 57 Modem transects 57 Modem transect sample collection 65 Herbarium sample collection 67 Packrat middens 67 Packrat midden collection 73 Methods 75 Sample analysis 75 Isotopic analysis 75 Stomatal measurements 79 Results 82 Future Research Directions 89 REFERENCES CITED 91 APPENDIX A 114 Abstract 115 Introduction 117 Methods 120 Results 124 Discussion 129 Conclusions 135 References Cited 137 7 TABLE OF CONTENTS - Continued APPENDIX B 143 Abstract 144 Introduction 145 Methods 151 Results and Discussion 160 Modem transects 160 Herbarium material 167 Packrat middens 170 Conclusions 176 References Cited 179 APPENDIX C 185 Abstract 186 Introduction 188 Geographic Setting 193 Methods 195 Results 199 Discussion 214 Cj Gymnosperms 222 Cj Angiosperms 224 C4 Angiosperms 226 CAM Angiosperms 228 Conclusions 231 References Cited 233 APPENDIX D 241 Abstract 242 Introduction 244 Stable Carbon Isotopes and Plant Physiology; Theory 247 Eco-physiology parameters 250 Site Description and Methods 252 Laboratory Methods 254 Modem Midden Comparisons with the Surrounding Plant Communities . 257 Results and Discussion 258 C3 Gymnosperms 258 C4 Angiosperms 267 CAM Angiosperms 269 Conclusions 270 Appendix 273 References Cited 281 8 TABLE OF CONTENTS - Continued APPENDIX E 287 Abstract 288 Introduction 290 Background 292 Modem Samples 298 Paclcrat Nfidden Samples 299 Methods 302 Results and Discussion 304 Conclusions 322 References Cited 323 9 LIST OF ILLUSTRATIONS Figure l.I, Historic Atmospheric CO, 17 Figure 1.2, Reconstructed Atmospheric COj 18 Figure 1.3, Reconstructed Atmospheric 6"C 23 Figure 1.4, Modeled Carbon Assimilation 28 Figure 2.1, Monthly mean climatic &ctors 62 Figure 2.2, Utah climate gradients 64 Figure 2.3, New Mexico climate gradients 68 Figure 2.4, Isotope Standards 77 10 LIST OF TABLES Table 2.1, Plant species analyzed 58 Table 2.2, Plants collected along each elevational gradient 59 Table 2.3, Utah climate stations used to calculate climatic gradients 63 Table 2.4, New Mexico climate stations used to c&!culate climatic gradiems .. 66 Table 2.5, Collection data for herbarium samples 70 Table 2.6, Collection data for midden nuterial 76 11 ABSTRACT During the last deglaciation, 15,000 to 12,000 calendar years ago, global warming and wholesale shifts in regional precipitation patterns produced dramatic changes in vegetation worldwide. Paleobotanical records, namely pollen and macrofossils, have been used not only to reconstruct shifts in plant distributions and abundances, but also to quantify changes in temperature and precipitation amounts or seasonality. In addition to climatic change, during the fiill glacial period atmospheric COj values had dropped 30% to 200 ppmv compared to the Holocene, preindustrial value of280 ppmv.