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Abstract Alliaria Petiolata (Garlic ABSTRACT ALLIARIA PETIOLATA (GARLIC MUSTARD) RESPONSE TO HERBICIDE AND JUNE PRECIPITATION, AND SUBSEQUENT EFFECTS ON THE FOREST FLOOR COMMUNITY by Wendy Wenger Hochstedler The impact of invasive plant species on native plants is largely assumed to be negative, but supporting evidence is sparse. We examined the long-term effects of herbicide on Alliaria petiolata and the subsequent effects on the plant community in southwestern Ohio. November herbicide application effectively killed A. petiolata, but did not reduce recruitment; spring densities of A. petiolata rosettes were not lower in sprayed plots. Only modest differences were noted in forest floor vegetation, suggesting A. petiolata rosettes competed with other plant species. We tested the hypothesis that higher June precipitation promotes rosette growth and survival with a rain shelter experiment. The three different water treatments affected soil moisture, but not A. petiolata growth or survival. Dry treatments may not have replicated drought years based on water availability measurements. June precipitation is probably not a reliable predictor of A. petiolata rosette survival in years with above average precipitation. ALLIARIA PETIOLATA (GARLIC MUSTARD) RESPONSE TO HERBICIDE AND JUNE PRECIPITATION, AND SUBSEQUENT EFFECTS ON THE FOREST FLOOR COMMUNITY A Thesis Submitted to the Faculty of Miami University in partial fulfillment of the requirements for the degree of Master of Science Department of Botany by Wendy Wenger Hochstedler Miami University Oxford, Ohio 2006 Advisor ____________________________________ David L. Gorchov Reader _____________________________________ Martin Henry H. Stevens Reader _____________________________________ Michael A. Vincent Table of Contents Introduction 1 Literature Cited 4 Chapter 1: Alliaria petiolata response to herbicide and subsequent effects 7 on the forest floor community in a deciduous forest in southwest Ohio Abstract 7 Introduction 8 Methods 10 Results 14 Discussion 16 Literature Cited 22 Chapter 2: The effects of June precipitation on Alliaria petiolata growth, 47 density and survival Abstract 47 Introduction 47 Methods 50 Results 53 Discussion 54 Literature Cited 56 Conclusion 68 Literature Cited 73 Appendices: Appendix A. Map of Hueston Woods State Park 75 Appendix B. Map of Hueston Woods Nature Preserve, study sites, and plots 76 Appendix C. Sample size for analyses 77 Appendix D. NMDS stress as a function of dimensionality 78 Appendix E. Growth forms of all taxa 79 Appendix F. Mean 2004 peak percent cover 80 Appendix G. Mean 2005 peak percent cover 83 Appendix H. Multiple response permutation procedure results 86 Appendix I. Mean percent cover of each growth form 89 Appendix J. Soil properties 92 Appendix K. Common species observed within precipitation study plots 93 Appendix L. Response variables of A. petiolata and soil water contents 94 Appendix M. Nutrient concentration of water added to plots 96 Appendix N. Pairwise comparisons of soil water content 97 Appendix O. Shelter microclimate measurements 98 Appendix P. Comments on rain shelter design and use 102 ii List of Tables Chapter 1 Table 1. Survival of A. petiolata from Oct. to May each year 27 Table 2. Effect of treatment on adult A. petiolata cover 28 Table 2. Univariate repeated measures ANOVA of May A. petiolata rosette cover 29 Table 4. Mean plant species richness, 2000–2005 30 Table 5. Shannon-Wiener plant diversity indices, 2000–2005 31 Table 6. Effect of treatment on peak cover 32 Table 7. Contingency table of Podophyllum peltatum presence, old-growth stand 33 Table 8. Contingency table of Podophyllum peltatum presence, second-growth stand 33 Table 9. Contingency table of Stellaria media presence, old-growth stand 34 Chapter 2 Table 1. A. petiolata response variables in dry, average and wet treatments 61 List of Figures Chapter 1 Figure 1. Mean May A. petiolata adult and rosette cover 35 Figure 2. Mean May A. petiolata adult and rosette density 36 Figure 3. NMDS ordination , 2001–2005 37 Figure 4. Percent cover of spring perennials, old-growth stand 43 Figure 5. Percent cover of spring perennials, second-growth stand 44 Figure 6. Percent cover Podophyllum peltatum 45 Figure 7. Percent cover Stellaria media 46 Chapter 2 Figure 1. Fixed-location rain shelter with sample plot and moisture barrier 62 Figure 2. Calibration of percent and gravimetric soil water content 63 Figure 3. Ambient precipitation, throughfall, and water applied to treatments 64 Figure 4. Weekly soil water content 65 Figure 5. Soil moisture release curve for Russell-Miamian silt loam soils 66 Figure 6. Effect of June water treatments on mean rosette survival 67 iii Acknowledgments To the following people I am greatly indebted for assistance with the work presented here and for my sanity over the past two years. My advisor Dr. David Gorchov, Associate Professor of Botany, Miami University, has provided invaluable guidance at all stages of this project, including his superlative editing skills which have raised the standard of my writing and made completion of the thesis possible. Brad Slaughter, Adriane Carlson, and Lauren Saunders provided data in addition to what I collected for this research, and Brad helped with plant identification and data collection. Hank Stevens, Assistant Professor of Botany, MU, guided my persistent questions about R programming, provided code for problematic analyses in Chapter 1, and loaned field equipment. Michael Vincent, Curator, Willard Sherman Turrell Herbarium, MU, provided help with plant identification, and Drs. Stevens and Vincent served on my Thesis Committee, providing suggestions for improving this body of work. The Gorchov lab members provided computer advice, valuable comments on manuscripts, and consistent sources of laughter. Field assistants Mark Hochstedler, Elizabeth Valentine, Katy Levings, Jessica Hoisington, Jeremy Ash, Susan Sprunt, Melanie Link-Perez, Xanic Rondon, Greg Osborn, and Erica Cunningham put their life on the line so I would not face the feral hogs of Hueston Woods alone. The Ecology Program at Miami University provided a Research Assistantship, and the Ecology Research Center and staff allowed this research and provided logistical assistance. I thank other labs for their involvement in Chapter 2: Mike Vanni’s lab for water analyses, Jon Costanzo and Scott Johnston for weather data, and Alfred Conklin at Wilmington College, Wilmington, OH for soils advice and use of pressure plate extractors. Botany office staff Barb Wilson and Vickie Sandlin are simply amazing individuals without whom we grad students might wander aimlessly. I am grateful to Wendy Cass, Botanist at Shenandoah National Park, for her enthusiasm and professional mentorship, as well as to plant ecologists Lorna Harder, Clair Mellinger, and Kenton Brubaker for formative educational experiences. Botany comrades, my volleyball team, June Thompsen and Nancy Mumaw supported me with friendship, made me exercise, and provided tasty food. I thank my family and most iv importantly, Mark Hochstedler, for being impromptu field assistant, personal chef and chauffeur, and for believing in me through it all. Thank you. Funding for this research was provided by MU Department of Botany Academic Challenge, Hudson Garden Club, Ohio Biological Survey, and Garden Club of Ohio. I thank the staff at Hueston Woods State Park and the Ohio Department of Natural Resources for allowing me to conduct this research, and Shenandoah National Park for granting me days of leave to begin data collection. v INTRODUCTION Invasions by non-native species are of great concern to biologists, land managers and conservationists. While invasive species are considered to be the second leading cause of biodiversity loss in the United States (Wilcove 1998), claims that invasive plant species cause declines and extinctions of native species is often speculative and evidence is sparse (Davis 2003, Gurevitch and Padilla 2004). For example, most studies documenting compositional and diversity impacts by introduced plant species compare invaded and uninvaded areas and are correlational in nature (Levine et al. 2003). Regardless, an estimated $120 billion in environmental damages and losses are caused by invasive species each year in the United States (Pimentel et al. 2005) and additional knowledge of invasive ecology and impacts on native species is needed to make effective management decisions. To quantify the impacts of an introduced plant species of the eastern deciduous forest floor community, we report here on a long-term field assessment of the competitive ability of Alliaria petiolata (M. Bieb.) Cavara and Grande (Brassicaceae, garlic mustard). Alliaria petiolata is an introduced biennial plant of Eurasian origin which is pollinated by generalist species (Anderson et al. 1996, Cruden and McClain 1996), produces as many as 15,000 seeds per m2 (Anderson et al. 1996), and has a viable seedbank for up to five years (Baskin and Baskin 1992). Because individuals are capable of self-pollination (Anderson et al. 1996, Cruden and McClain 1996), one individual can found a population (Cavers et al. 1979, Baskin and Baskin 1992, Nuzzo 1993, Byers and Quinn 1998) and efforts to control A. petiolata populations require extensive and long- term removal within a target area to be effective (Baskin and Baskin 1992, Nuzzo 1993). Alliaria petiolata populations extend throughout the northern midwest and eastern half of North America and southern Canada (Nuzzo 1991, Welk et al. 2002). Due to its establishment over a wide area
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