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Ecological Effects of the Biocontrol Insects, Larinus Planus and Rhinocyllus Conicus, on Native Thistles

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Ecological Effects of the Biocontrol Insects, Larinus Planus and Rhinocyllus Conicus, on Native Thistles ABSTRACT Title: ECOLOGICAL EFFECTS OF THE BIOCONTROL INSECTS, LARINUS PLANUS AND RHINOCYLLUS CONICUS, ON NATIVE THISTLES. Gary Jonathan Dodge Doctor of Philosophy, 2005 Directed By: Professor David W. Inouye Department of Biology Biological control of invasive weeds is, by nature, a delicate balance between introducing effective biological control agents and not introducing another invasive species. A disconcertingly similar suite of traits is used to describe invasive insect species and to identify appropriate biological control agents (or candidates): good control agents and invasive exotic species are good dispersers, they are good colonizers, they have high reproduction rates, and they are suited for broad distribution. It shouldn’t come as a surprise that two previously released weed biological control agents can now be characterized as invasive species. The purpose of this dissertation is twofold: first, it is to explore the ecological relationship between predispersal seed predation and plant population dynamics, and second, it is to elucidate the risks to native plants involved with introduction and redistribution of exotic species. Rhinocyllus conicus and Larinus planus are Eurasian seed-head weevils, introduced and redistributed broadly across the western U.S. to control exotic thistles. Exclusion experiments on native thistles, including one that is rare and imperiled, at sites in Colorado present strong evidence that a decrease in seed production due to herbivory by both R. conicus and L. planus has lead to a reduction in recruitment of the thistles. The density of seedlings in both cases, even in the excluded units, was far below where density dependent effects may play a role in the dynamics of the thistle. Further, a survey of eight western states demonstrates established populations of L. planus and effects on seed production in multiple native species in four states. It also documents the near ubiquity and broad diet breadth of R. conicus. The process for approval of phytophagous biocontrol agents has become more cautious and more efforts are made to prevent nontarget herbivory. Nevertheless, land managers still routinely redistribute previously approved, non-regulated agent insects that appear to pose a higher risk to the native flora. The results of this research will benefit resource managers who wish to consider use of phytophagous insects as biological control agents as well as help ecologists and environmental managers understand the risk probabilities of biological control applications. ECOLOGICAL EFFECTS OF THE BIOCONTROL INSECTS, LARINUS PLANUS AND RHINOCYLLUS CONICUS, ON NATIVE THISTLES By Gary Jonathan Dodge Dissertation submitted to the Faculty of the Graduate School of the University of Maryland, College Park, in partial fulfillment of the requirements for the degree of Doctor of Philosophy 2005 Advisory Committee: Professor David W. Inouye, Chair Professor Robert F. Denno Associate Professor Michele R. Dudash Associate Professor Irwin N. Forseth Professor Douglas E. Gill © Copyright by Gary J. Dodge 2005 Dedication For my wife, Janette. ii Acknowledgements I first and foremost greatly appreciate generous guidance and insight from my advisor and mentor David Inouye and my committee members Doug Gill, Michele Dudash, Irv Forseth, and Bob Denno. Thank you for your support, humor, intellect, and passion for teaching and research. I am also absolutely indebted to Svata Louda whose inspiring research, insight, collaboration, and most generous support were invaluable to this project. I would also like to extend heartfelt thanks to Aaron Inouye, Rebecca Aicher, Cameron Pedersen, Noam Perlmann, Amy Goodstine, Kathy Darrow, Heidi Wenger, Georgia Dendrinos, and Judy Che for their help in the field. David Obenshain, Christopher Cowles, Ben Schapiro, Katryana Hanley-Knutson, Jon Krochmal, Matt Johnson, Emily Hawkins, Kyle Vallecillo, Rubina Rahman, and Silvia Lara were of great assistance in the lab. Eric Coombs from the Oregon Department of Agriculture was of tremendous assistance in conducting and preparing the manuscript for the survey. Site access and permitting was facilitated by Dan Roddy, Ed Delaney, Marie Curtin, Myron Chase, and Ken Stahlnecker of the National Park Service; Gina Ramos and Bob Troiano of the Bureau of Land Management; Liz Hebertson, Sandy Kominsky, and Gay Austin of the Forest Service; Carey Goss of the Fish and Wildlife Service; and Al Cofrancesco from the U.S. Army Corps of Engineers. Finally, I owe multitudes of thanks to my many friends at the Rocky Mountain Biological Laboratory and University of Maryland. iii Funding was provided by the National Science Foundation, the University of Maryland Center for Biodiversity, and the National Park Service. Gracious logistical support was provided by the Rocky Mountain Biological Laboratory and the University of Maryland Department of Biology. iv Table of Contents DEDICATION..............................................................................................................II ACKNOWLEDGEMENTS........................................................................................ III TABLE OF CONTENTS............................................................................................. V CHAPTER 1: COLONIZATION OF AND HERBIVORY ON NATIVE THISTLES BY INTRODUCED BIOCONTROL AGENTS LARINUS PLANUS AND RHINOCYLLUS CONICUS.......................................................................................... 1 Abstract..................................................................................................................... 1 Introduction............................................................................................................... 2 Methods..................................................................................................................... 5 Results..................................................................................................................... 14 Discussion............................................................................................................... 29 Appendices.............................................................................................................. 39 CHAPTER 2: DEMOGRAPHIC EFFECTS OF PRE-DISPERSAL SEED PREDATION BY TWO BIOCONTROL AGENTS ON TRACY’S THISTLE (CIRSIUM UNDULATUM VAR. TRACYI)................................................................. 56 Abstract................................................................................................................... 56 Introduction............................................................................................................. 57 Methods................................................................................................................... 59 Results..................................................................................................................... 69 Discussion............................................................................................................... 83 CHAPTER 3: CONSERVATION IMPLICATIONS OF PREDISPERSAL SEED PREDATION IN ROCKY MOUNTAIN THISTLE (CIRSIUM PERPLEXANS)..... 91 Abstract................................................................................................................... 91 Introduction............................................................................................................. 92 Methods................................................................................................................... 94 Results................................................................................................................... 102 Discussion............................................................................................................. 126 CHAPTER 4: DENSITY EFFECTS ON RECRUITMENT AND SEEDLING ESTABLISHMENT IN TRACY’S THISTLE (CIRSIUM UNDULATUM VAR. TRACYI).................................................................................................................... 135 Abstract................................................................................................................. 135 Introduction........................................................................................................... 136 Methods................................................................................................................. 138 v Results................................................................................................................... 142 Discussion............................................................................................................. 155 CHAPTER 5: PRELIMINARY MATRIX MODEL PROJECTIONS AND ELASTICITY ANALYSIS FOR THREE CIRSIUM SPECIES NATIVE TO COLORADO ............................................................................................................ 157 Abstract................................................................................................................. 157 Introduction........................................................................................................... 158 Methods................................................................................................................. 159 Results................................................................................................................... 169 Discussion............................................................................................................. 174 BIBLIOGRAPHY....................................................................................................
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