Soil Ecology of the Exotic Dune Grass Leymus Arenarius
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University of Louisville ThinkIR: The University of Louisville's Institutional Repository Electronic Theses and Dissertations 5-2018 Soil ecology of the exotic dune grass Leymus arenarius. Matthew L. Reid University of Louisville Follow this and additional works at: https://ir.library.louisville.edu/etd Part of the Botany Commons, and the Terrestrial and Aquatic Ecology Commons Recommended Citation Reid, Matthew L., "Soil ecology of the exotic dune grass Leymus arenarius." (2018). Electronic Theses and Dissertations. Paper 2995. https://doi.org/10.18297/etd/2995 This Doctoral Dissertation is brought to you for free and open access by ThinkIR: The University of Louisville's Institutional Repository. It has been accepted for inclusion in Electronic Theses and Dissertations by an authorized administrator of ThinkIR: The University of Louisville's Institutional Repository. This title appears here courtesy of the author, who has retained all other copyrights. For more information, please contact [email protected]. SOIL ECOLOGY OF THE EXOTIC DUNE GRASS LEYMUS ARENARIUS By Matthew L. Reid B.A. Hendrix College, 2009 M.S. University of Louisiana at Monroe, 2013 A Dissertation Submitted to the Faculty of the College of Arts and Sciences of the University of Louisville in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in Biology Department of Biology University of Louisville Louisville, Kentucky May 2018 SOIL ECOLOGY OF THE EXOTIC DUNE GRASS LEYMUS ARENARIUS By Matthew L. Reid B.A. Hendrix College, 2009 M.S. University of Louisiana at Monroe, 2013 A Dissertation Approved on April 9th, 2018 By the Following Dissertation Committee ________________________________ Dissertation Director Sarah Emery ________________________________ Margaret Carreiro ________________________________ Susanna Remold ________________________________ Stephen Yanoviak ________________________________ Phoebe Zarnetske ii ACKNOWLEDGMENTS I would like to begin by thanking my fellow lab members, Jeff Masters, Brad Gottshall, Andrea Howes, Kyle Anderson, Erin Kinnetz, Binod Baysal, Aaron Sexton, Emily Pennington, and Sarah Benton for assistance with lab work and reviewing grant proposals and manuscripts. I also thank Andrea Howes for assistance with field work. Thanks also to the Michigan Department of Natural Resources, Grand Traverse Regional Land Conservancy, and Ottowa County Parks for allowing me to conduct research on their properties. Funding for the projects described here were provided by the Garden Club of America, Kentucky Academy of Sciences, Sigma Xi, and the University of Louisville. My wife Sarah Reid provided constant support over the duration of this process. Finally, I thank my mentor, Sarah Emery, and all of my committee members for their input and guidance throughout this process. iii ABSTRACT SOIL ECOLOGY OF THE EXOTIC DUNE GRASS LEYMUS ARENARIUS Matthew L. Reid April 9, 2018 Exotic plants can alter the structure and function of native communities both above- and belowground, which is particularly important in primary successional sand dune ecosystems where plant-soil interactions are critical drivers of successional dynamics. In Great Lakes sand dunes, the native foundation grass Ammophila breviligulata dominates early in succession until an accumulation of detrimental soil organisms causes its die-back of the grasses. Die-back of this native foundation species can be slowed by mutualistic interactions between plants and mycorrhizal fungi. Changes to the plant community resulting from invasion, and subsequent effects on the soil community have the potential to disrupt the successional dynamics of dune systems. Leymus arenarius is a dune-building grass native to northern European sand dunes, and is currently invading Great Lakes dunes. This dissertation examines the effects of L. arenarius invasion on soil community structure, multi-trophic interactions within the soil, and plant community development. In a greenhouse study manipulating the presence/absence of soil organisms, I found evidence of altered multi-trophic interactions. Specifically, arbuscular mycorrhizal fungi provided greater protection from plant-parasitic nematodes to L. arenarius, resulting in weaker biomass reductions relative to A. breviligulata. In a field survey, I found that L. arenarius had little effect on the iv structure of soil nematode communities and soil bacterial functional diversity relative to uninvaded areas. In a growth chamber experiment, I found that direct competitive effects from L. arenarius were generally similar to effects from A. breviligulata, for later successional native species. However, L. arenarius more strongly suppressed growth of secondary invaders, relative to suppression by A. breviligulata. Soil conditioning effects from L. arenarius were present, but were generally weak suggesting that indirect effects of invasion on soil community composition are less important than direct competitive effects. Experimentally-derived differences in multi-trophic interactions between L. arenarius and A. breviligulata suggested that the invasion of L. arenarius into communities dominated by A. breviligulata would result in altered plant-soil interactions. However, the invasion of L. arenarius did not result in large impacts on plant and soil communities. Overall, these results suggest that L. arenarius has relatively high functional similarity to its native counterpart, A. breviligulata. v TABLE OF CONTENTS PAGE ACKNOWLEDGMENTS ................................................................................................. iii ABSTRACT ....................................................................................................................... iv LIST OF TABLES ........................................................................................................... viii LIST OF FIGURES ........................................................................................................... ix CHAPTER I: INTRODUCTION .........................................................................................1 STUDY SYSTEM ..........................................................................................................3 ORGANIZATION OF DISSERTATION ......................................................................4 CHAPTER II: NATIVE AND EXOTIC FOUNDATION GRASSES DIFFER IN TRAITS AND RESPONSES TO BELOWGROUND TRI-TROPHIC INTERACTIONS ................................................................................................................7 SUMMARY ....................................................................................................................7 INTRODUCTION ..........................................................................................................8 MATERIALS AND EMTHODS ..................................................................................12 RESULTS .....................................................................................................................17 DISCUSSION ...............................................................................................................19 CHAPTER III: DUNE SOIL COMMUNITIES ASSOCIATED WITH CLIMATIC AND EDAPHIC FACTORS, NOT EXOTIC PLANT PRESENCE ..........................................31 SUMMARY ..................................................................................................................31 INTRODUCTION ........................................................................................................32 MATERIALS AND METHODS ..................................................................................35 RESULTS .....................................................................................................................40 DISCUSSION ...............................................................................................................42 CHAPTER IV: NATIVE AND EXOTIC PIONEER GRASSES HAVE SIMILAR EFFECTS ON SECONDARY SUCCESSION ON MICHIGAN DUNES .......................54 SUMMARY ..................................................................................................................54 INTRODUCTION ........................................................................................................55 MATERIALS AND METHODS ..................................................................................58 RESULTS .....................................................................................................................62 DISCUSSION ...............................................................................................................65 vi CHAPTER V: SUMMARY AND FUTURE DIRECTIONS ............................................75 SUMMARY ..................................................................................................................75 FUTURE DIRECTIONS ..............................................................................................76 REFERENCES ..................................................................................................................79 APPENDIX I .....................................................................................................................94 APPENDIX II ....................................................................................................................98 APPENDIX III .................................................................................................................100 CURRICULUM VITA ....................................................................................................104