Causes and Consequences of Prolonged Dormancy: Why Stay Belowground?
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University of Montana ScholarWorks at University of Montana Graduate Student Theses, Dissertations, & Professional Papers Graduate School 2010 Causes and consequences of prolonged dormancy: Why stay belowground? Jennifer R. Gremer The University of Montana Follow this and additional works at: https://scholarworks.umt.edu/etd Let us know how access to this document benefits ou.y Recommended Citation Gremer, Jennifer R., "Causes and consequences of prolonged dormancy: Why stay belowground?" (2010). Graduate Student Theses, Dissertations, & Professional Papers. 175. https://scholarworks.umt.edu/etd/175 This Dissertation is brought to you for free and open access by the Graduate School at ScholarWorks at University of Montana. It has been accepted for inclusion in Graduate Student Theses, Dissertations, & Professional Papers by an authorized administrator of ScholarWorks at University of Montana. For more information, please contact [email protected]. CAUSES AND CONSEQUENCES OF PROLONGED DORMANCY: WHY STAY BELOWGROUND? By JENNIFER ROBIN GREMER B.S. in Environmental Studies, University of California, Santa Barbara, 2002 Dissertation presented in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Organismal Biology and Ecology The University of Montana Missoula, MT Fall 2010 Approved by: Perry Brown, Associate Provost for Graduate Education Graduate School Anna Sala, Co-Chair Organismal Biology and Ecology Elizabeth Crone, Co-Chair Wildlife Biology Ragan Callaway Organismal Biology and Ecology Lila Fishman Organismal Biology and Ecology Thomas H. DeLuca School of Environment, Natural Resources and Geography Bangor University, United Kingdom Gremer, Jennifer, Ph.D., Fall 2010 Organismal Biology and Ecology Causes and consequences of prolonged dormancy: Why stay belowground? Chairpersons: Dr. Anna Sala and Dr. Elizabeth E. Crone Prolonged dormancy is a stage in which mature plants fail to resprout during the growing season and instead remain alive belowground. Though it is relatively common, the causes and consequences of this intriguing stage have remained elusive. In this dissertation, I investigate the causes and consequences of prolonged dormancy in a long lived perennial herb, Astragalus scaphoides . First, I use a combination of demography and ecophysiology to study the proximate mechanisms associated with prolonged dormancy. Analysis of a long-term demographic dataset indicates that both endogenous factors (e.g. age, condition, and history) and exogenous factors (e.g. climate and spatial variation) are associated with dormancy. I then investigate the association between stored resources and dormancy. My results indicate that individual plants with low levels of stored available carbon are more likely to enter prolonged dormancy. Surprisingly, individuals increased their mobile carbon concentrations while dormant, presumably by remobilizing structural carbon into mobile forms. Since stored resources integrate past conditions and performance with current state, these results can explain why some individuals remain belowground while others emerge to grow and reproduce. I used matrix models to examine the ultimate causes and consequences of prolonged dormancy. I found evidence that prolonged dormancy acts as a conservative strategy that allows plants to avoid the risk of a variable environment. Further, my results demonstrate that intermediate levels of dormancy result in the highest fitness advantage. Finally, I measured the trade-offs associated with emerging during times of environmental stress. Although plants showed remarkable physiological tolerance to stress, stress led to demographic costs. Therefore, prolonged dormancy is shown to be a beneficial strategy in a variable environment. Together, my research identifies both the proximate causes of prolonged dormancy, as well as the ultimate consequences of remaining belowground during the growing season. Therefore, my research not only identifies why some plants go dormant while others emerge, but also explains the prevalence of this intriguing life stage in the life histories of so many perennial plants. ii ACKNOWLEDGEMENTS As I began graduate school, I was asked why I needed two advisors. My answer was simple; I was attempting to bridge the fields of demography and ecophysiology, and so I had advisors with strengths in each discipline. Since then, I have realized that, as mentors, Elizabeth Crone and Anna Sala complement each other in many more ways than just their fields of study. I am greatly indebted to both of them for their contagious enthusiasm, companionship, and guidance through the last several years. I am a better ecologist because of them. I am also extremely grateful to Peter Lesica, who started collecting data on Astragalus scaphoides many years ago. Because of his efforts, I was able to address questions that would not have been possible within the scope of a dissertation. I thank my committee, Ray Callaway, Lila Fishman, and Tom DeLuca, for feedback and guidance. I never encountered a closed door when I needed their help. I would also like to thank Doug Emlen for hours of discussion about my project, and his incredible enthusiasm about disappearing plants. I have many people to thank for assistance in the field and in the lab. Kelly Hopping, Beth Miller, Andrew Gundlach, Jennifer Williams, Martha Ellis, Justin Keller, and Beth Roskilly helped me find, count, and, sometimes, dig up plants in the field. I would especially like to thank Nora Lahr, Beth Miller, and Kelly Hopping for teaching me how to analyze stored resources in the lab and, at times, for running analyses for me. I would like to thank both of my lab groups, the Crone lab and the Sala lab, for their feedback and comments on endless drafts of presentations, proposals, and manuscripts. I would also like to thank them for their support and friendship over the years. Funding was provided by a National Science Foundation EPSCoR fellowship, P.E.O. Scholar Award, the Nicholas Memorial Scholarship, and the Jack E. Schmautz Scholarship, as iii well as support from the University of Montana. Parts of this research were supported by an NSF LTREB grant to Elizabeth Crone, and an NSF Grant to Elizabeth Crone and Anna Sala (NSF DEB 02-40963). Finally, I am deeply grateful for the support of my family and friends. My sister, Nicole Martin and my parents, Mike and Kathy Gremer, have encouraged me, held me up, and kept me going whenever I needed them. I cannot express my gratitude for their boundless love and support. I am also indebted to my friends both in Missoula and farther away, with whom I have shared countless hours of companionship, laughter, and adventure. I dedicate this dissertation to my Grandma Kay and Grandpa John, who always had confidence in me and told me “to find what I love and make a living doing it.” iv TABLE OF CONTENTS ABSTRACT………………………………………………………………………………………ii ACKNOWLEDGEMENTS………………………………………………………………………iii LIST OF TABLES……………………………………………………………………………….vii LIST OF FIGURES…………………………………………………………………………..…viii GENERAL INTRODUCTION AND OVERVIEW…………………………………………….1 CHAPTER 1 Patterns of prolonged dormancy in Astragalus scaphoides : Inference from long term monitoring ............................................................................................. 9 Introduction ................................................................................................................................. 9 Field Methods ............................................................................................................................ 11 Analyses and results .................................................................................................................. 13 Discussion ................................................................................................................................. 20 Literature Cited ......................................................................................................................... 24 Figure Legends .......................................................................................................................... 31 CHAPTER 2 Disappearing plants: Why they hide and how they return. .......................................................... 40 Abstract ..................................................................................................................................... 40 Introduction ............................................................................................................................... 41 Methods ..................................................................................................................................... 43 Results ....................................................................................................................................... 47 Discussion ................................................................................................................................. 49 Literature Cited ......................................................................................................................... 53 Figure Legends .......................................................................................................................... 58 CHAPTER 3 Are dormant plants hedging their bets? Demographic consequences of prolonged dormancy in variable environments ................................................................................................................... 60 Abstract