RESPONSES OF OAKS TO MAMMAL AND INSECT HERBIVORY A dissertation submitted to Kent State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy by Cynthia Lynn Perkovich May 2021 © Copyright All rights reserved Except for previously published materials Dissertation written by Cynthia Lynn Perkovich B.S., Kent State University, 2013 M.S., University of Nebraska-Lincoln, 2016 Ph.D., Kent State University, 2021 Approved by David Ward, Ph.D. , Chair, Doctoral Dissertation Committee Oscar Rocha, Ph.D., Members, Doctoral Dissertation Committee Matthew Lehnert, Ph.D. Don Cipollini, Ph.D. Edgar Kooijman, Ph.D. Accepted by Laura G. Leff, Ph.D. , Chair, Department of Biological Sciences Mandy Munro-Stasiuk, Ph.D. , Interim Dean, College of Arts and Sciences TABLE OF CONTENTS TABLE OF CONTENTS ............................................................................................................. iii LIST OF FIGURES .......................................................................................................................vi LIST OF TABLES .........................................................................................................................ix ACKNOWLEDGEMENTS .......................................................................................................... xi I. INTRODUCTION................................................................................................. 1 REFERENCES..................................................................................................... 14 II. HERBIVORE-INDUCED DEFENSES ARE NOT UNDER PHYLOGENETIC CONSTRAINTS IN THE GENUS QUERCUS (OAK): PHYLOGENETIC PATTERNS OF GROWTH, DEFENSE, AND STORAGE ................................ 25 ABSTRACT .......................................................................................................... 25 INTRODUCTION ................................................................................................ 27 METHODS ........................................................................................................... 31 RESULTS ............................................................................................................. 37 DISCUSSION ....................................................................................................... 51 CONCLUSIONS................................................................................................... 58 REFERENCES ..................................................................................................... 60 III. ABOVEGROUND HERBIVORY CAUSES BELOWGROUND CHANGES IN TWELVE OAK (QUERCUS) SPECIES: A PHYLOGENETIC ANALYSIS OF ROOT BIOMASS AND NUTRIENT RE-ALLOCATION ................................. 80 ABSTRACT .......................................................................................................... 80 INTRODUCTION ................................................................................................ 82 iii METHODS ........................................................................................................... 85 RESULTS ............................................................................................................. 92 DISCUSSION ..................................................................................................... 105 CONCLUSIONS................................................................................................. 112 REFERENCES ................................................................................................... 114 IV. DIFFERENTIATED DEFENSE SYNDROMES IN RESPONSE TO VARYING HERBIVORE PRESSURES: OAK TREES INCREASE DEFENSES IN RESPONSE TO INSECTS AND DECREASE NUTRITIVE QUALITY IN RESPONSE TO DEER ....................................................................................... 130 ABSTRACT ........................................................................................................ 130 INTRODUCTION .............................................................................................. 132 METHODS ......................................................................................................... 135 RESULTS ........................................................................................................... 141 DISCUSSION ..................................................................................................... 151 CONCLUSIONS................................................................................................. 155 REFERENCES ................................................................................................... 157 V. PERIODICAL CICADAS INCREASE DEFENSES IN NORTH AMERICAN FOREST TREES: BEFORE, DURING, AND AFTER A MASS OUTBREAK ............................................................................................................................. 168 ABSTRACT ........................................................................................................ 168 INTRODUCTION .............................................................................................. 170 METHODS ......................................................................................................... 172 iv RESULTS ........................................................................................................... 175 DISCUSSION ..................................................................................................... 186 CONCLUSIONS................................................................................................. 190 REFERENCES ................................................................................................... 192 VI. PROTEIN:CARBOHYDRATE RATIOS IN THE DIET OF GYPSY MOTH LYMANTRIA DISPAR AFFECT ITS ABILITY TO TOLERATE TANNINS ............................................................................................................................ .201 ABSTRACT ........................................................................................................ 201 INTRODUCTION .............................................................................................. 203 METHODS ......................................................................................................... 205 RESULTS ........................................................................................................... 208 DISCUSSION ..................................................................................................... 214 CONCLUSIONS................................................................................................. 219 REFERENCES ................................................................................................... 221 VII. CONCLUSIONS AND RECOMMENDATIONS FOR FUTURE RESEARCH ............................................................................................................................. 229 OVERVIEW ....................................................................................................... 229 SUMMARY AND FUTURE DIRECTIONS ..................................................... 230 REFERENCES ................................................................................................... 239 v LIST OF FIGURES Figure 1. The phylogenetic relationships of the 12 Quercus species used for analysis of growth, defense, and nutrient reallocation in response to simulated herbivory ......................................... 32 Figure 2. Diagram of the treatments applied to the Quercus saplings with varying location and intensity of simulated herbivory ................................................................................................... 33 Figure 3. Trade-offs between Quercus constitutive traits and phylomorphospace projections of the Quercus phylogeny ................................................................................................................. 42 Figure 4. Individual Quercus species apical shoot relative growth rates plotted by location and intensity of simulated herbivory. .................................................................................................. 45 Figure 5. The mean condensed tannin concentrations of individual Quercus species in response to location of simulated herbivory (apical vs auxiliary) in each treatment................................... 47 Figure 6. The average leaf shape factor for all simulated herbivory treatments, plotted against the Quercus phylogenetic tree to show phylogenetic relationships. ................................................... 49 Figure 7. The phylogenetic relationships of the 12 Quercus species used for analysis of belowground responses to simulated herbivory. ........................................................................... 86 Figure 8. Diagram of the treatments applied to the Quercus saplings with varying location and intensity of simulated herbivory. .................................................................................................. 88 Figure 9. Pattern of increased investment by Quercus species in belowground biomass relative to aboveground regrowth in response to varying location and intensity of simulated herbivory. .... 95 vi Figure 10. Patterns of investment by Quercus species in belowground biomass in response to differing locations and intensities of simulated herbivory. ........................................................... 96 Figure 11. Investment of oak saplings in coarse and fine root biomass in response to differing