Plant Finding Behavior by Insects and Biological
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PLANT FINDING BEHAVIOR OF PHYTOPHAGOUS INSECTS AND BIOLOGICAL CONTROL OF AQUATIC PLANTS A dissertation submitted to Kent State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy by Justin L. Reeves December 2010 Dissertation written by Justin L. Reeves B.A., Western State College of Colorado, 2006 Ph.D., Kent State University, 2010 Approved by Patrick D. Lorch , Chair, Doctoral Dissertation Committee Mark W. Kershner , Members, Doctoral Dissertation Committee Ferenc A. de Szalay _ Marilyn A. Norconk _ Accepted by James L. Blank , Chair, Department of Biological Sciences John R. D. Stalvey , Dean, College of Arts and Sciences ii TABLE OF CONTENTS LIST OF FIGURES………………………………………………….…………………...vi LIST OF TABLES……………………………………………………………………...viii ACKNOWLEDGMENTS………………………………………………………….……..x CHAPTER I. Introduction………………………………………………………….….…1 II. Biological control of invasive aquatic and wetland plants by arthropods: a meta-analysis of data from the last three decades…………...………..…9 Abstract…………………………………………………………....9 Introduction .………………………………………………….….10 Methods…………………………………………………………..13 Literature Search…………………………………………13 Data Extraction…………………………………………..14 Analyses………………………………………………….18 Results…………………………………………………………....22 Discussion ...………...…………………………………………..31 Acknowledgments ……………………………………………….37 References………………………………………………………..38 III. Biological control of Eurasian watermilfoil by Euhrychiopsis lecontei: assessing efficacy and timing of sampling……………………………….47 Abstract…………………………………………………………..47 Introduction……………………………………………………....48 Methods…………………………………………………………..50 Results and Discussion ………………………………………….57 Acknowledgments………………………………………………..65 References…………………………..……….……….…………..65 iii IV. Vision is important for plant location by the phytophagous aquatic specialist, Euhrychiopsis lecontei Dietz (Coleoptera: Curculionidae)…..69 Abstract…………………………………………………………..69 Introduction………………………………………………….…...70 Methods…………………………………………………….…….73 Importance of light in plant location……………………..77 Weevil attraction to plants in vials……………………….78 Visual plant differentiation………………………………79 Effect of water turbidity on plant location……………….80 Results……………………………………………………………81 Importance of light in plant location……………………..81 Weevil attraction to plants in vials……………………….83 Visual plant differentiation………………………………83 Effect of water turbidity on plant location……………….85 Discussion………………………………………………………..85 Acknowledgments…………………………………………….….89 References………………………………………………………..89 V. Visual plant differentiation by the milfoil weevil, Euhrychiopsis lecontei Dietz (Coleoptera: Curcuionidae)………………………………………..93 Introduction………………………………………………….…...93 Methods……………………………………………………….….94 Results and Discussion…………………………………………..96 Acknowledgments…………………………………………….….99 References………………………………………………………100 VI. Visual active space of the milfoil weevil, Euhrychiopsis lecontei Dietz (Coleoptera: Curcuionidae)……………………………………………..101 Abstract…………………………………………………………101 Introduction……………………………………………………..102 Methods…………………………………………………………104 Visual active space…………………………………..….104 Effect of turbidity on visual active space……………….108 Results…………………………………………………………..110 Visual active space……………………………………...110 Effect of turbidity on visual active space……………….113 Discussion………………………………………………………113 Acknowledgments…………………………………………....…120 References………………………………………………………120 iv VII. Vision should not be overlooked as an important host-plant detection and selection mechanism for phytophagous insects………………………...124 Abstract………………………………………………………....124 Introduction……………………………………………………..125 Assumption 1: Vision, in general, is not an important host location mechanism because insect visual acuity is poor……………………………………………..127 Assumption 2: Vision is only used when appropriate chemical cues are detected……………………………….132 Assumption 3: Insects cannot visually differentiate plant species………………………………………….134 Assumption 4: Color is the only important visual stimulus for phytophagous insects…………………………..135 Underexplored areas of study for phytophagous insect visual ecology…………………………………………………........139 Conclusions……………………………………………………..144 Acknowledgments...………………………………………….…145 References………………………………………………………145 VIII. Conclusions……………………………………………………………….163 APPENDIX I. A method for growing legumes with and without root nodules for studying nodule-attacking Rivellia (Diptera: Platystomatidae)……….171 Introduction……………………………………………………..171 Methods ….……………………………………………………..173 Results and Discussion………………………………………....174 Acknowledgments………………………………………………177 References………………………………………………………177 APPENDIX II. Reprint permissions for published chapters………………………......178 v LIST OF FIGURES CHAPTER II. Biological control of invasive aquatic and wetland plants by arthropods: a meta-analysis of data from the last three decades. Fig. 2.1. Categories used to organize papers from literature search and their overall proportion of the entire set of papers……………………………...15 Fig. 2.2. Log response ratio effect size estimates for biological control agent analysis………………………………………………………………….…24 Fig. 2.3. Log response ratio effect size estimates for one vs. two agent use analysis……………………………………………………………….........26 Fig. 2.4. Log response ratio effect size estimates for observational (no control group; initial value used as control) vs. experimental (true control group) study analysis…………………………………………….………………....27 Fig. 2.5. Log response ratio effect size estimates for lab vs. field study analysis…………………………………………………………………..…28 Fig. 2.6. Log response ratio effect size estimates for plant variable measured analysis…………………………………………………………………..…29 Fig. 2.7. Log response ratio effect size estimates for replicate type analysis……..................................................................................................30 CHAPTER III. Biological control of Eurasian watermilfoil by Euhrychiopsis lecontei: assessing efficacy and timing of sampling. Fig. 3.1. Geographic distribution of the Michigan and Wisconsin lakes included in this study……………………………………………………………...…52 Fig. 3.2. Changes in proportional plant density [(final plant density – initial plant density) / initial plant density] at lake-specific control sites vs. treatment sites……………………………………………………….............……..…58 Fig. 3.3. The relationship between final EWM densities (stems/m2) and the number of days passed between initial and final surveys for control and treatment sites………………………………………………………....61 vi Fig. 3.4. Average proportional EWM density change by lake and year at control vs. treatment sites………………………..……………………63 CHAPTER IV. Vision is important for plant location by the phytophagous aquatic specialist, Euhrychiopsis lecontei Dietz (Coleoptera: Curculionidae). Fig. 4.1. Photograph of an adult Euhrychiopsis lecontei head. …………….…....74 Fig. 4.2. Grid used to track weevil movement in experimental arenas...…….…..76 Fig. 4.3. Results of light importance experiment………………………...………82 Fig. 4.4. Results of weevil attraction to plants in vials experiment………...……84 CHAPTER VI. Visual active space of the milfoil weevil, Euhrychiopsis lecontei Dietz (Coleoptera: Curculoionidae). Fig. 6.1. Cross section and dimensions of trough used for these experiments....105 Fig. 6.2. Results of pooled active space replicates………………………...…...112 Fig. 6.3. Results of turbidity experiment………………………...……………..115 Fig. 6.4. Conceptual model of plant finding by E. lecontei………………….…118 vii LIST OF TABLES CHAPTER II. Biological control of invasive aquatic and wetland plants by arthropods: a meta-analysis of data from the last three decades. Table 2.1. Results of meta-analyses performed in this study…………………....23 Appendix 2.A. Biological control agents and their target plants used in the literature search for this study…………………………………...41 Appendix 2.B. Papers used in the above analyses……….………………………44 CHAPTER III. Biological control of Eurasian watermilfoil by Euhrychiopsis lecontei: assessing efficacy and timing of sampling. Table 3.1. Lakes used in this analysis, their locations (county, state), surface area, average and maximum depths, and mean/range of final weevil densities (# / stem) at treatment sites..……………………………………...…..53 CHAPTER VI. Visual active space of the milfoil weevil, Euhrychiopsis lecontei Dietz (Coleoptera: Curculionidae). Table 6.1. Pooled visual active space replicates showing how many weevils swam toward the vial with a plant stem (out of 40), how many of these contacted the vial first, and whether they contacted the vial or trough wall first significantly more often…………………………………..111 Table 6.2. Turbidity results showing how many weevils swam toward the vial with a plant stem (out of 25), how many of these contacted the vial first, and whether they contacted the vial or trough wall first significantly more often……………………………………….…....114 viii CHAPTER VII. Vision should not be overlooked as an important host-plant detection and selection mechanism for phytophagous insects. Appendix 7.A. Bibliographic information for references on phytophagous insect visual ecology…………………………………………..158 ix ACKNOWLEDGMENTS This dissertation is dedicated to my family: Pat, Andrea, and Candice Reeves. Their love and support have allowed me to successfully pursue my interest in biology since I was a child. I would never have gotten this far if I did not have such great parents. Special thanks are extended to my advisor, Pat Lorch, whose advice, guidance, and willingness to let me pursue my own interests allowed me to keep motivated until the end. I also thank