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The Influence of Exotic Plant Invasion on Native Bee Abundance and Demography, and the Pollination and Abundance of Native Plants

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The Influence of Exotic Plant Invasion on Native Bee Abundance and Demography, and the Pollination and Abundance of Native Plants University of Montana ScholarWorks at University of Montana Graduate Student Theses, Dissertations, & Professional Papers Graduate School 2013 The influence of exotic plant invasion on native bee abundance and demography, and the pollination and abundance of native plants Jennifer Palladini The University of Montana Follow this and additional works at: https://scholarworks.umt.edu/etd Let us know how access to this document benefits ou.y Recommended Citation Palladini, Jennifer, "The influence of exotic plant invasion on native bee abundance and demography, and the pollination and abundance of native plants" (2013). Graduate Student Theses, Dissertations, & Professional Papers. 10621. https://scholarworks.umt.edu/etd/10621 This Dissertation is brought to you for free and open access by the Graduate School at ScholarWorks at University of Montana. It has been accepted for inclusion in Graduate Student Theses, Dissertations, & Professional Papers by an authorized administrator of ScholarWorks at University of Montana. For more information, please contact [email protected]. THE INFLUENCE OF EXOTIC PLANT INVASION ON NATIVE BEE ABUNDANCE AND DEMOGRAPHY, AND THE POLLINATION AND ABUNDANCE OF NATIVE PLANTS By Jennifer Dawn Palladini B.S. College of Charleston, South Carolina, 1999 M.A. Humboldt State University, California, 2002 Dissertation Presented in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Biology and Ecology The University of Montana Missoula, Montana Spring 2013 Approved by: Dr. John Maron, Chair, Division of Biological Sciences Dr. Ragan M. Callaway Division of Biological Sciences Dr. Winsor Lowe Division of Biological Sciences Dr. Elizabeth Crone Harvard Forest Dr. Monica Geber Department of Ecology and Evolutionary Biology UMI Number: 3568125 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. UMI 3568125 Published by ProQuest LLC (2013). Copyright in the Dissertation held by the Author. Microform Edition © ProQuest LLC. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, MI 48106 - 1346 ABSTRACT Palladini, Jenniter, PhD 2013 Chairperson: Dr. John L. Maron Flowering plants and bees are fundamentally linked. Bees rely wholly on floral resources for food and many native plants rely on bees to assist in reproduction. Despite this fundamental connection, how plants and their bee pollinators influence one another’s abundance remains unclear. Also unclear is how invasion by exotic flowering plants disrupts bee- native plant relationships. I examined how the invasive plant, Euphorbia esula (Euphorbiaceae) influenced the native annual forb, Clarkia pulchella (Onagraceae) by altering pollinator visitation (i.e., indirect effects) and through direct resource competition. I observed pollinator visitation and estimated the extent to which plant reproduction was limited by pollen receipt (i.e., pollinator visits) in invaded and native-dominated sites. Though pollinator visitation was strongly reduced in invaded sites, plants were only weakly pollen limited. In contrast, E. esula removal experiments indicated that the effect of resource competition on C. pulchella fitness was very strong. Seed addition experiments indicated that changes in fecundity resulting from ameliorating competition for pollinators could influence future plant abundance, however these effects were dwarfed by the effects of resource competition. To determine if strong reductions in pollinator visitation in invaded sites was mirrored by changes in entire bee communities, I quantified bee communities in sites dominated by E. esula and native-dominated sites. Bee abundance, richness and diversity were reduced in invaded sites. Thus, though native bees foraged on E. esula flowers, the net effect of invasion on bee communities was negative. However, because bees are highly mobile, patterns of forager abundance in relation to floral abundance may be misleading. Such patterns may be the result of bees moving to high resource patches rather than any fundamental change in bee abundance based on altered demographic processes. To examine this issue, I quantified the influence of floral resources and plant invasion on the demography of the native solitary bee, Osmia lignaria (Megachilidae). I placed nest blocks and preemergent cocoons in 27 sites that varied along a floral resource gradient and in extent of invasion, and found that nesting and fecundity were positively correlated with the abundance and richness of the native forb community and that nesting decreased with increasing exotic forb species richness. Despite increased parasitism in high-resource native-dominated sites, a marginally positive relationship between native forb species richness and bee population growth rates remained, suggesting that floral resources can positively influence bee demography. Finally, via a literature review, I examined whether excessive harvesting of pollen by bees has led to the evolution of chemical defense of pollen. While bees are generally perceived as mutualists, they also act as herbivores, collecting substantial quantities of pollen to rear larvae, and the loss of pollen to consumption by bees has the potential to decrease plant fitness. Many morphological features of flowers are interpreted to serve in defense of pollen from excessive harvesting, and there is growing evidence that pollen can contain secondary compounds that may be an additional means of pollen defense. ii TABLE OF CONTENTS Chapter 1. Indirect competition for pollinators is weak compared to direct resource competition: pollination and performance in the face of an invader 1 Abstract 2 Introduction 3 Methods 6 Results 11 Discussion 13 Acknowledgements 17 Literature Cited 18 Figures 27 Appendix 30 Chapter 2. Exotic plant invasion reduces abundance, richness and diversity of native bees in Montana grasslands 31 Abstract 32 Introduction 33 Methods 35 Results 40 Discussion 41 Acknowledgements 45 Literature Cited 46 Figures 56 Appendix 60 Chapter 3. Demographic responses of a solitary bee to floral resource gradients created by native and invasive plants 62 Abstract 63 Introduction 64 Methods 68 Results 75 Discussion 77 Literature Cited 84 Figures 92 Appendices 95 Chapter 4. Pollen defense: Do plants chemically protect pollen from consumption by bees? 100 Abstract 101 Introduction 102 Bees sequester substantial amounts of pollen 103 iii Pollen consumption and plant reproduction 105 Plant defense of pollen 106 Toxic pollen 108 Toxic pollen and specialization 111 Benefits to plants 114 Future research priorities 117 Literature Cited 122 Tables 133 iv CHAPTER ONE INDIRECT COMPETITION FOR POLLINATORS IS WEAK COMPARED TO DIRECT RESOURCE COMPETITION: POLLINATION AND PERFORMANCE IN THE FACE OF AN INVADER 1 Abstract Invasive plants have the potential to reduce native plant abundance through both direct and indirect interactions. Direct interactions, such as competition for soil resources, and indirect interactions, such as competition for shared pollinators, have been shown to influence native plant performance, however, we know much less about how these interactions influence native plant abundance in the field. While direct competitive interactions are often assumed to drive declines in native abundance, an evaluation of their influence relative to indirect mechanisms is needed to more fully understand invasive plant impacts. We quantified the direct effects of resource competition by the invasive perennial forb, Euphorbia esula (Euphorbiaceae), on the recruitment, subsequent performance, and ultimate adult abundance of the native annual, Clarkia pulchella (Onagraceae). We contrast these direct effects with those that indirectly resulted from competition for shared pollinators. Although E. esula dramatically reduced pollinator visitation to C. pulchella , plants were only weakly pollen-limited. Pollen supplementation increased the number of seeds per fruit from 41.28 to 46.38. Seed addition experiments revealed that the impacts of ameliorating pollen limitation only increased potential recruitment by 12.3%. In contrast, seed addition experiments that ameliorated direct competition with E. esula resulted in an increase in potential future recruitment of 574%. Our results show that while the indirect effects of competition for pollinators can influence plant abundance, its effects are dwarfed by the magnitude of direct effects of competition for resources. 2 Introduction Invasive plants often have well documented negative impacts on native plant abundance (reviews by Levine et al. 2003; Vilá et al. 2011). These impacts are most commonly thought to derive from invasives outcompeting natives for resources such as nutrients or light (Melgoza et al. 1990; D'Antonio and Mahall 1991; Gorchov and Trisel 2003; Vilá and Weiner 2004; Combs et al. 2011), however, indirect interactions between invasives and natives, which can be mediated by soil microbes (Klironomos 2002; Stinson et al. 2006), herbivores (White et al. 2006; Meiners 2007; Orrock et al. 2008), or pollinators (Bjerknes et al. 2007; Morales and Traveset 2009) may also play a role. A key question concerns what the relative strength of
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