Effects of the Spatial Heterogeneity Formed by Ambrosia Dumosa on Individual and Population Growth of the Invasive Annual Grass Schismus Barbatus
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EFFECTS OF THE SPATIAL HETEROGENEITY FORMED BY AMBROSIA DUMOSA ON INDIVIDUAL AND POPULATION GROWTH OF THE INVASIVE ANNUAL GRASS SCHISMUS BARBATUS. DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of the Ohio State University By Susana Rodríguez-Buriticá ***** The Ohio State University 2009 Dissertation Committee: Approved by: Professor Maria M. Miriti, Adviser Professor Ralph Boerner Adviser Professor Elizabeth Marschall Evolution, Ecology and Organismal Biology Graduate Program Professor Karen Goodell ABSTRACT Natural ecosystems are exposed to numerous factors that determine their structure and dynamics. Although change is typically inherent in to biological systems, very few factors change a natural ecosystem as drastically as the arrival and successful establishment of non-native species. The impact of the Mediterranean grass (Schismus barbatus) in the southwestern U.S. exemplifies such dynamics. In my dissertation, I empirically and theoretically explore two components of Schismus invasion. First, native shrubs may promote Schismus invasion if Schismus benefits from the favorable growth conditions that shrubs provide. Second, Schismus may threaten ecosystem level properties if Schismus precludes establishement of shrubs. Thus Schismus may compromise the maintenance of Islands of fertility that favor growth of native plant community. To meet these objectives, I explore Schismus performance with respect to the location and size of one of the most abundant native shrubs in the Sonoran and Mojave Deserts, the Bur-sage shrub, Ambrosia dumosa. As a first approximation to the study system, I describe the natural distribution of Schismus with respect to the location of native shrubs (Chapter 1). Then, I examine factors that control Schismus-Ambrosia interaction using a three-year field experiment (Chapter 2), and subsequent ii characterization of soil nutrients changes around Ambrosia shrubs in the context of Schismus invasion (Chapter 3). I found that Schismus barbatus responds to the spatial heterogeneity formed by Ambrosia. This response is strongly controlled by water availability (Chapters 1 and 2). Ambrosia creates a dichotomous landscape formed by nutrient rich areas below their canopies and nutrient poor areas at inter-shrub spaces (Chapter 2). In this landscape, performance of Schismus is favored around Ambrosia during years with good precipitation. Nevertheless, at any given growing season, shading and water interception by shrub canopies are the most influential factors on Schismus performance. Given that germination was consistently low below Ambrosia canopies, the consequences of Schismus-Ambrosia interactions might not be as straightforward as predicted by the individual effect of Ambrosia on Schismus. In fact, Ambrosia might actually reduce realized Schismus densities. Conversely, inter-specific competition with Schismus regulates Ambrosia seedling survival. Thus, high-density stands of Schismus might generate a negative feedback on the dynamics of favorable areas associated with shrubs; ultimately limiting Schismus abundance. I design a modeling framework to test this hypothesis and present several predictions about the long-term demographic consequences of reciprocal interactions between Schismus and Ambrosia (Chapter 4). iii TO MY FAMILY iv ACKNOWLEDGMENTS I want to thank my adviser, Maria Miriti for her personal and academic support during all these years. Maria has taught me the value of integrating simple ecological questions into a wider theoretical framework. Her sense of perspective has enormously improved my ability to interpret ecological phenomena. I thank my committee members, R. Boerner, E. Marschall, and K. Goodell for their support at all times during the development of my dissertation. Their critical comments on my research proposal greatly improved the coherence and feasibility of this project. In addition, their advice during specific stages of the dissertation helped me fulfilled all the objectives I initially defined. I especially want to thank H. Howe for his comments on several aspects of this dissertation. His experience helped me to identify key issues in my study system. I am thankful with my lab mates who during these years have helped at different stages of my research. In particular, I thank E. Quintero, K. Mulac, H. Sweigard, and N. Woods, for their willingness to help me with intense field work, and their useful comments on several chapters presented here. In addition, I want to thank A. Campbell, S. Harms, and M. Clark for their contribution during the analysis of field samples. v I thank all the people that in one way or another were involved during the field work of this project. I want to thank P. Sethi, and E. Hooper and all the personnel of the Joshua Tree National Park who helped me to solve incredible logistic challenges; especially T. Ledoux and G. Graham. I had nice academic stimulation during my PhD. in the EEOB Department. I particularly want to thank, J. Huang, J. Miesel, and A. McKinney, for their comradeship. Funding for my dissertation was provided by The Ohio State University, The Joshua Tree National Park, and The Ecological Society of America. I am extremely grateful for all my friends in Columbus from whom I received invaluable support during all these years. Thanks to Maria Teresa Beltran, Rosario Cortez, Lina Juswara, and Christine Johnson, who patiently listened and comforted me when I was discouraged. Thanks to Carla Giai, Jonathan Hall, Hala Zahreddine, and Ranjan Shrestha for their support and friendship. I thank Agusti Muñoz for his encouragement and companionship, especially during my first three years in Columbus. I especially want to express my gratitude to Rosario Barbieri, who showed me the greatness of sincere friendship. I would not have pursued a Ph.D. in ecology without the constant and unconditional encouragement of my friends from Colombia, and my family from almost everywhere. I am thankful to Yolima, Talia, Julia, and Alejandro for their friendship. From my family back in Colombia, I received permanent inspiration and support; in particular from my cousin Marta Isabel. Here in the U.S., my family has always supported my endeavors; I am especially thankful to my grandparents vi Carlos y Leonor, my aunt Ines, and my cousin Anita. My parents, my sister, and my brother are always with me despite the distance. vii VITA October 11, 1976………………………..............Born – Bogotá, Colombia 2001.…………………………………….............. B.Sc. Biology, Universidad Nacional de Colombia, Bogotá 2002-Present……………………………………..Graduate Teaching and Research Associate, Department of EEOB, Ohio State University. PUBLICATIONS 1. Rodriguez-Buritica, S. and M. Miriti. “Biting the hand that feeds: The invasive grass Schismus barbatus (Poaceae) is facilitated by, but reduces establishment of the native shrub, Ambrosia dumosa (Asteraceae)”. Journal of Vegetation Science, in press, (2009). 2. Miriti, M., S. Rodriguez-Buritica, J.S. Wright, and H.F. Howe. “Episodic death across species of desert shrubs”. Ecology, 88: 32-36, (2007). 3. Rodriguez-Buritica, S., M.A. Orjuela R., and G. Galeano G. “Demography and life history of Geonoma orbignyana: an understory palm used as foliage in Colombia”. Forest Ecology and Management, 211: 329-340, (2005). viii FIELDS OF STUDY Major field: Evolution, Ecology, and Organismal Biology ix TABLE OF CONTENTS Abstract ............................................................................................................................................ ii Acknowledgments ............................................................................................................................ v Vita ................................................................................................................................................. viii List of Tables .................................................................................................................................. xiii List of Figures ................................................................................................................................. xvi Chapters: Introduction ....................................................................................................................................... 1 1. Bitting the hand that feeds: the invasive grass schismus barbatus (poaceae) is facilitated by, but reduces establishment of the native shrub, ambrosia dumosa (asteraceae) ........... 6 1.1. Introduction ........................................................................................................................ 6 1.2. Methods ............................................................................................................................. 9 1.3. Results ............................................................................................................................. 14 1.4. Discussion ........................................................................................................................ 22 2. Role of facilitation and temporal environmental fluctuations on the persistence of the exotic grass Schismus barbatus (Poaceae) in a heterogeneous environment. ............................ 27 2.1. Introduction ...................................................................................................................... 27 x 2.2. Methods ..........................................................................................................................