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The Pennsylvania State University The Graduate School Department of Entomology LIFE HISTORY OF PTEROSTICHUS MELANARIUS (COLEOPTERA: CARABIDAE) AND ITS IMPORTANCE FOR BIOLOGICAL CONTROL IN FIELD CROPS A Thesis in Entomology by Anna Katherine Busch © 2016 Anna Katherine Busch Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science August 2016 ii The thesis of Anna K. Busch was reviewed and approved* by the following: John F. Tooker Associate Professor of Entomology and Extension Specialist Thesis Advisor Mary E. Barbercheck Professor of Entomology Heather D. Karsten Associate Professor of Crop Production/Ecology Gary W. Felton Professor of Entomology Head of the Department of Entomology *Signatures are on file in the Graduate School iii ABSTRACT Pterostichus melanarius (Coleoptera: Carabidae) is a Palearctic generalist predator native to Europe that was introduced to North America in the 1920’s and has since become widespread in the United States. This beetle is non-native to the U.S., yet appears to have little to no negative effect on native carabids and is a critical natural enemy in agricultural systems, especially because it feeds on the invasive gray garden slug Deroceras reticulatum. My thesis begins with a review that compiles in one publication the details of P. melanarius distribution, life history, diet, and factors that influence its populations. I used ecological niche modeling to determine the current potential range of P. melanarius in the United States and found that annual mean temperature and precipitation of the driest month were the most important factors to determine probable habitat suitability. My second chapter investigates the effects of crop rotation and integrated pest management on pests and natural enemies in no-till cropping systems in central Pennsylvania. Overall, I found that diversified rotations that implemented continuous crop cover with perennial forages and cover crops and incorporated IPM enhanced the abundance of natural enemies early in the growing season and can suppress pests to similar levels attained by simple rotations managed with Bt hybrids, pesticidal seed treatments, and prophylactic pesticide application. I also found that protecting and increasing carabid abundance, especially P. melanarius, is important for slug control and can be achieved by reducing tillage, increasing crop species diversity, and using integrated pest management. Lastly, I examined the predator-prey relationship between ground beetles and slugs by investigating non-consumptive effects of P. melanarius on D. reticulatum behavior and physiology. I found that combined physical and chemical cues of P. melanarius induced fear behavior in D. reticulatum that resulted in increased hiding and decreased foraging by D. iv reticulatum. Due to the ineffectiveness of radioimmunoassays, I could not determine a physiological fear response. Nevertheless, there appear to be non-consumptive effects, which suggest that the pest management value of ground beetle predators such as Pterostichus melanarius may be underestimated in agricultural systems. v TABLE OF CONTENTS LIST OF FIGURES ................................................................................................................. vii LIST OF TABLES ................................................................................................................... ix ACKNOWLEDGEMENTS ..................................................................................................... x Chapter 1 Life History, Biology, and Ecological Niche Modeling of Pterostichus melanarius in the United States ....................................................................................... 1 Introduction ...................................................................................................................... 1 Taxonomy ........................................................................................................................ 2 Current distribution .......................................................................................................... 3 Predicted range of P. melanarius in the U.S. ........................................................... 4 Ecological niche modeling results ............................................................................ 5 Habitat and range expansion .................................................................................... 8 Biology and lifecycle ....................................................................................................... 9 Adult diet .................................................................................................................. 10 Predation and parasitism of P. melanarius ............................................................... 11 Agricultural practices ....................................................................................................... 12 Tillage ....................................................................................................................... 12 Cover crops and intercropping ................................................................................. 13 Insecticides ............................................................................................................... 14 Conclusion ........................................................................................................................ 16 References ........................................................................................................................ 17 Tables ............................................................................................................................... 25 Figures .............................................................................................................................. 26 Chapter 2 Effects of rotation diversity and integrated pest management on pests and natural enemies in no-till maize ....................................................................................... 29 Introduction ...................................................................................................................... 29 Materials and methods ..................................................................................................... 33 Study site .................................................................................................................. 33 Maize establishment and damage assessments ........................................................ 34 Slug activity-density ................................................................................................. 35 Crimson clover versus red clover cover crop ........................................................... 35 European corn borer damage assessments ............................................................... 36 Seasonal potato leafhopper abundance .................................................................... 36 Activity-density of arthropod predators ................................................................... 37 Seasonal predation .................................................................................................... 37 Statistical methods .................................................................................................... 38 Results .............................................................................................................................. 39 Maize establishment and damage assessments ........................................................ 39 Slug activity-density ................................................................................................. 40 Slug damage to maize seedlings grown for grain .................................................... 41 vi Crimson clover versus red clover cover crop ........................................................... 41 European corn borer damage assessments ............................................................... 41 Seasonal potato leafhopper abundance .................................................................... 42 Activity-density of arthropod predators ................................................................... 43 Seasonal predation .................................................................................................... 45 Discussion ........................................................................................................................ 45 Conclusion ........................................................................................................................ 51 References ........................................................................................................................ 52 Tables ............................................................................................................................... 58 Figures .............................................................................................................................. 63 Chapter 3 Non-consumptive effects of a ground beetle predator Pterostichus melanarius on the gray garden slug Deroceras reticulatum ............................................................... 71 Introduction ...................................................................................................................... 71 Materials and methods ..................................................................................................... 74 Collecting and maintaining slugs