Influence of Switchgrass Ecotype, Cultivar and Planted Stand Diversity on Herbivores, Natural Enemies, and Biological Control in Bioenergy Cropping Systems
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INFLUENCE OF SWITCHGRASS ECOTYPE, CULTIVAR AND PLANTED STAND DIVERSITY ON HERBIVORES, NATURAL ENEMIES, AND BIOLOGICAL CONTROL IN BIOENERGY CROPPING SYSTEMS By Marissa K. Schuh A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of Entomology – Master of Science 2016 ABSTRACT INFLUENCE OF SWITCHGRASS ECOTYPE, CULTIVAR AND PLANTED STAND DIVERSITY ON HERBIVORES, NATURAL ENEMIES, AND BIOLOGICAL CONTROL IN BIOENERGY CROPPING SYSTEMS By Marissa K. Schuh Switchgrass is a perennial, C4 grass that has emerged as a model bioenergy crop for a large portion of the US. Because of the potential for switchgrass to occupy many acres, understanding how cultivar and planted stand diversity choices could impact insects and ecosystem services is key. To investigate how cultivars differed the preference of herbivores, fall armyworm was chosen to represent how a generalist, chewing herbivore. Two life stages were used to measure establishment, consumption levels, and life history traits on different cultivars of switchgrass. These experiments revealed that the lowland ecotype supported lower levels of feeding and tended to slow development. The second experiment investigated how herbivores, natural enemies, and biological control were impacted by different switchgrass cultivars and planted stand diversities. Switchgrass was established both as different cultivar monocultures and in mixtures with grasses and forbs. Sweep samples were used to collect arthropods and egg cards were used as sentinel prey to measure predation levels. While there were differences between sampling years, generally upland ecotypes supported a greater abundance of herbivores, natural enemies, and ultimately higher levels of biological control. The effect of planted stand diversity was more mixed. Diverse plots hosted more herbivores in both years, but more variable natural enemy numbers and no differences in biological control. Clearly, choices about which switchgrass cultivar and seeding diversity to establish can impact herbivores, arthropod communities, and potentially ecosystem services. ACKNOWLEDGEMENTS I will be forever grateful to Dr. Doug Landis for funding this project and for offering his guidance and wisdom in regards to both science and life. I would also like to thank my committee members Carolyn Malmstrom and Rufus Isaacs for their input and assistance. This work was funded by USDA NIFA grant “Control and Mitigation of Generalist Pests in Perennial Grass-Dominated Bioenergy Landscapes” (USDA NIFA Sustainable Biofuels Program Award #2011-67009-30137; Malmstrom, Isaacs, Landis). I would also like to thank the members of the Landis lab. Julia Perrone laid the groundwork for so many elements of this project, provided assistance in the field, and prevented things from getting “out of control” (in a bad way). Dr. Christie Bahlai served as an ad hoc advisor and statistical wizard. Dr. Aaron Fox and Dr. Bill Wills also gave me much appreciated advice and encouragement in multiple stages of this process. Dan Gibson, Sara Hermann, and Andrew Myers provided camaraderie throughout the writing of this thesis. Finally, thank you to Lindsey Hawkins, Jessica Kalin, Erin Forster, and the other undergraduate students who tromped through wet switchgrass, wrangled worms, and counted insects during the duration of this project. To my fellow graduate students in the Department of Entomology at Michigan State, thank you for the good company. To Aubrey McElmeel, thank you for always being available to help me unpack graduate school life and listen to my mystified ramblings on switchgrass genetics. To my family, who has always supported my endeavors, thank you for continuing to do so from 600 miles away. To Aubrey Drake Graham, RuPaul Charles, and many others, thank you for bringing frivolity and joy to this process. iii Finally, I would like to thank Stan Karas for his constant love and companionship during this process. Thank you for helping me both buckle down and escape, for coming in on weekends to weed plots or scan leaves, and for being the silver lining of this whole process. iv TABLE OF CONTENTS LIST OF TABLES ........................................................................................................................ vii LIST OF FIGURES ....................................................................................................................... ix KEY TO ABBREVIATIONS ....................................................................................................... xii CHAPTER 1: LITERATURE REVIEW .........................................................................................1 Abstract ................................................................................................................................2 Biofuels in the United States ................................................................................................3 Government Policies ................................................................................................3 Current Sources: First-Generation Biofuels.............................................................3 Second-Generation Biofuels ................................................................................................6 Advantages ...............................................................................................................6 Potential Biomass Sources in the Upper Midwest ...................................................7 Perennial Crops as Dedicated Biomass Crops .........................................................8 Switchgrass ........................................................................................................................10 Basic Information...................................................................................................10 Breeding Efforts, Cultivar Varieties, and Production ............................................12 Herbivores and Natural Enemies in Switchgrass ...............................................................16 Switchgrass-Specific Pests.....................................................................................16 Generalist Herbivores ............................................................................................17 Natural Enemies .....................................................................................................18 Factors Impacting Pest and Natural Enemy Communities ....................................19 Thesis Objectives ...............................................................................................................21 CHAPTER 2: EFFECT OF SWITCHGRASS ECOTYPE AND CULTIVAR ON ESTABLISHMENT, CONSUMPTION LEVEL, AND DEVELOPMENT OF A GENERALIST HERBIVORE ................................................................................................................................22 Abstract ..............................................................................................................................23 Introduction ........................................................................................................................24 Materials and Methods .......................................................................................................28 Plants ......................................................................................................................28 Insects ....................................................................................................................29 Neonate Establishment and Feeding Trial .............................................................29 Late Instar Feeding Trial ........................................................................................30 Image Processing ...................................................................................................31 Statistical Analysis .................................................................................................31 Results ................................................................................................................................33 Neonates .................................................................................................................33 Late Instar Larvae ..................................................................................................34 Discussion ..........................................................................................................................37 Neonates .................................................................................................................37 Late Instar Larvae ..................................................................................................39 v Plant Factors Relevant to Both Larval Ages ..........................................................40 Conclusions ........................................................................................................................45 Acknowledgements ............................................................................................................46 CHAPTER 3: INFLUENCE OF SWITCHGRASS CULTIVAR, ECOTYPE, AND PLANTED STAND DIVERSITY ON INSECT COMMUNITIES AND BIOLOGICAL CONTROL SERVICES .................................................................................................................47 Abstract ..............................................................................................................................48 Introduction