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The Pennsylvania State University The Graduate School College of Agricultural Sciences EPIGEAL INSECT DIVERSITY AND DYNAMICS IN AGROECOSYSTEMS ADOPTING TRANSGENIC CROPS A Thesis in Entomology by Timothy W. Leslie © 2007 Timothy W. Leslie Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy December 2007 The thesis of Timothy W. Leslie was reviewed and approved* by the following: Shelby J. Fleischer Professor of Entomology Thesis Adviser Chair of Committee David J. Biddinger Research Associate in Entomology Mary E. Barbercheck Professor of Entomology David A. Mortensen Professor of Crop and Soil Sciences Gary W. Felton Department Head, Professor of Entomology * Signatures are on file in the Graduate School ABSTRACT Using data from five years of field experiments, we assessed the influence of insecticidal transgenic crop technologies on non-target epigeal insect diversity in both maize and diversified vegetable agroecosystems. Concurrently, we considered the relative influence of crop species, crop rotations, and proximity to forested field margins on these insect communities. Our approach differed from other transgenic non-target studies by emphasizing species-level identification of a functionally diverse subset of the epigeal insect community, notably Coleoptera, with extensive analysis of Carabidae, to maximize the biological and functional resolution of the data. We examined population dynamics as well as community response using current statistical approaches in the fields of biodiversity sciences and community ecology. Insecticide inputs were lower in transgenic crops due to the efficient control of important pest taxa, which served to sustain epigeal biodiversity. In maize agroecosystems, no differences were found between the coleopteran communities in conventional and transgenic insect control practices where transgenic field corn contained a neonicotinoid seed treatment. Both management tactics negatively influenced coleopeteran communities compared to a control. Dominant members of the carabid community exhibited temporal partitioning of resources. One species, Harpalus pensylvanicus, consistently exhibited higher activity-densities in the control fields, and was identified as a possible bio-indicator. Despite management influences, differences between crop species and the effect of crop rotations was often sufficient to mask any treatment effects. Carabid communities were significantly different between maize and a forested field margin, and shifts in community structure along a transect were most rapid at the grassy margin between the two habitats. Carabidae communities were more diverse in the forest edge, where plant diversity was also highest. Despite strong habitat associations, most abundant carabids were found in both forest and maize. Differences between carabid communities in maize fields at varying distances from a forested margin suggested landscape level influences. In conclusion, the spatial and temporal heterogeneity of agroecosystems in the northeastern U.S. seemed to promote on-farm iii insect diversity, although non-target effects from transgenics coupled with neonicotinoid seed treatments and conventional insect control methods were discernible from a control. iv TABLE OF CONTENTS LIST OF FIGURES ......................................................................................................... vii LIST OF TABLES ........................................................................................................... xii ACKNOWLEDGEMENTS ............................................................................................ xiv CHAPTER 1: INTRODUCTION .......................................................................................1 Literature Cited .......................................................................................................7 CHAPTER 2: TRANSGENES SUSTAIN EPIGEAL INSECT BIODIVERSITY IN DIVERSIFIED VEGETABLE FARM SYSTEMS .....................................................10 Abstract .................................................................................................................10 Introduction ............................................................................................................11 Methods .................................................................................................................14 Results ...................................................................................................................17 Discussion .............................................................................................................21 Literature Cited .....................................................................................................27 CHAPTER 3: CARABIDAE POPULATION DYNAMICS AND TEMPORAL PARTITIONING: RESPONSE TO COUPLED NEONICOTINOID- TRANSGENIC TECHNOLOGIES ..................................................................................45 Abstract .................................................................................................................45 Introduction ...........................................................................................................46 Methods .................................................................................................................50 Results ...................................................................................................................53 Discussion .............................................................................................................55 Literature Cited .....................................................................................................62 v CHAPTER 4: COLEOPTERAN COMMUNITY DYNAMICS IN MOSAICS OF TRANSGENIC MAIZE ....................................................................................................75 Abstract .................................................................................................................75 Introduction ...........................................................................................................76 Methods .................................................................................................................78 Results ...................................................................................................................85 Discussion .............................................................................................................89 Literature Cited .....................................................................................................98 CHAPTER 5: CARABIDAE COMMUNITY DYNAMICS ACROSS AN AGRICULTURE-FOREST ECOTONE ........................................................................110 Abstract ...............................................................................................................110 Introduction .........................................................................................................111 Methods ...............................................................................................................114 Results .................................................................................................................120 Discussion ...........................................................................................................126 Literature Cited ...................................................................................................133 CHAPTER 6: CONCLUSIONS .....................................................................................149 Literature Cited ...................................................................................................159 APPENDIX A: Coleopteran species collected in maize at Penn State Research Farms in Rock Springs, PA during 2003 and 2004 ........................................................162 APPENDIX B: Carabidae species (including county records) collected from field experiments at Penn State Research Farms in Rock Spring, PA from 2001–2005 ........163 vi LIST OF FIGURES Figure 2.1: Species rarefaction curves for epigeal beetles and ants found in transgenic and isoline diversified vegetable farm systems consisting of sweet corn, potatoes, and acorn squash in 2001, and sweet corn and potatoes in 2002 (based on 50 permutations of data) .............................................................38 Figure 2.2: Species rarefaction curves for epigeal beetles and ants found in transgenic and isoline sweet corn, potato and acorn squash in 2001, and sweet corn and potatoes in 2002 (based on 50 permutations of data) .....................39 Figure 2.3: Species rarefaction curves for Carabidae found in transgenic and isoline diversified vegetable farm systems consisting of sweet corn, potatoes, and acorn squash in 2001, and sweet corn and potatoes in 2002 (based on 50 permutations of data) ..........................................................................40 Figure 2.4: Species rarefaction curves for Carabidae found in transgenic and isoline sweet corn, potato and acorn squash in 2001, and sweet corn and potato in 2002 (based on 50 permutations of data) ..................................................41 Figure 2.5: Rank abundance (on a log10 scale) of Carabidae in transgenic and isoline diversified farm systems in 2001 and 2002 ..................................................42 Figure 2.6: Rank abundance of Staphylinidae in transgenic and isoline diversified farm systems in 2001