The Roles of Forest Fragments and an Invasive Shrub in Structuring Native Bee Communities and Pollination Services in Intensive Agricultural Landscapes
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MIAMI UNIVERSITY The Graduate School Certificate for Approving the Dissertation We hereby approve the Dissertation of Michael John Minnick Candidate for the Degree DOCTOR OF PHILOSOPHY ______________________________________ Thomas O. Crist, Director ______________________________________ David J. Berg, Reader ______________________________________ Amélie Y. Davis, Reader ______________________________________ David L. Gorchov, Reader ______________________________________ Jing Zhang, Graduate School Representative ABSTRACT THE ROLES OF FOREST FRAGMENTS AND AN INVASIVE SHRUB IN STRUCTURING NATIVE BEE COMMUNITIES AND POLLINATION SERVICES IN INTENSIVE AGRICULTURAL LANDSCAPES by Michael J. Minnick This dissertation examines how an invasive woody plant, Lonicera maackii, temporally and spatially structures native bee communities of forest-edge habitat in agricultural landscapes. In Chapter 1, I measured bee species composition and pollination services ≤200 m from isolated forest patches in response to L. maackii flower removals. Removing flowers released a subset of small-bodied bees and increased pollination services after two years. Pollination services provisioned by large-bodied and generalist bee species (e.g. Bombus spp) increased when nearby plants were adjacent to intact L. maackii flowers. Findings suggest that L. maackii flowers suppress one component of the bee community and attract another to the forest patch that increases usage of the adjacent crop fields. In Chapter 2, I compared two components of the bee community and their responses to L. maackii density, floral resources of the forest patch, and the surrounding landscape. Bees sampled in pan traps were typically small, specialized, and responded to local patch features. Bees sampled in vane traps were larger in body size, social, and responded to landscape composition 3 km from the forest patch. These findings suggest that L. maackii floral resources support weaker foragers within the forest patch as well as larger bees that forage throughout the landscape. Both components of the bee community responded to tree community composition and were vertically stratified in the tree canopy. In Chapter 3, I measured bee diversity and community composition at different vertical strata in response to L. maackii density and flowering period as well as floral resource availability of woody plants. I found that L. maackii supports a component of the vertically stratified bee community which changes interactions with floral resources of the native woody vegetation at different vertical strata. Collectively, my studies demonstrate that L. maackii structures forest-edge bee communities through mechanisms involving functional and life history traits of individual bee species. Therefore, in my Conclusion Chapter, I developed a synthetic model that assigned an Agricultural Landscape Response Index for Bees (ALRIB) value between 0 and 1 to bees of each species that corresponds with their likelihood of responding to the forest patch as an island or as one land cover type within a broader mosaic of different resources. I conclude that L. maackii invasion into forest fragments within intensively managed agricultural landscapes filters the bee community in favor of species that use its floral resources and exhibits an overall homogenizing effect on species diversity. THE ROLES OF FOREST FRAGMENTS AND AN INVASIVE SHRUB IN STRUCTURING NATIVE BEE COMMUNITIES AND POLLINATION SERVICES IN INTENSIVE AGRICULTURAL LANDSCAPES A DISSERTATION Presented to the Faculty of Miami University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Department of Biology by Michael J. Minnick The Graduate School Miami University Oxford, Ohio 2020 Dissertation Director: Thomas O. Crist © Michael John Minnick 2020 TABLE OF CONTENTS General Introduction .................................................................................................................................. 1 1 Introduction .................................................................................................................................... 1 2 Bee Pollination and Invasive Plants.................................................................................................. 3 3 Bee Diversity and Modern Agricultural Landscapes ......................................................................... 6 4 References .................................................................................................................................... 13 Chapter 1: Bee communities and pollination services in adjacent crop fields following flower removal in an invasive forest shrub ............................................................................................................................ 26 1 Abstract ......................................................................................................................................... 26 2 Introduction .................................................................................................................................. 28 3 Methods ........................................................................................................................................ 30 3.1 Study Design ......................................................................................................................... 30 3.2 Bee Community .................................................................................................................... 32 3.3 Sentinel Flower Visitation ..................................................................................................... 33 3.4 Sentinel Fruit Production ...................................................................................................... 33 3.5 Analyses ................................................................................................................................ 33 4 Results ........................................................................................................................................... 37 4.1 Bee community & functional traits ....................................................................................... 37 4.2 Bee visits to sentinel plants................................................................................................... 38 4.3 Sentinel plant production ..................................................................................................... 38 5 Discussion...................................................................................................................................... 39 5.1 Bee community & functional traits ....................................................................................... 39 5.2 Bee visits to sentinel plants................................................................................................... 41 5.3 Sentinel plant production ..................................................................................................... 42 6 References .................................................................................................................................... 46 Chapter 2: Community responses of native bees to landscape composition depend on bee functional traits and seasonal floral resource availability .......................................................................................... 74 1 Abstract ......................................................................................................................................... 74 2 Introduction .................................................................................................................................. 76 3 Materials and Methods ................................................................................................................. 78 3.1 Site Selection ........................................................................................................................ 78 3.2 Bee Community .................................................................................................................... 79 3.3 Local Floral Resource Availability .......................................................................................... 79 iii 3.4 Landscape Features .............................................................................................................. 80 3.5 Analyses ................................................................................................................................ 81 4 Results ........................................................................................................................................... 83 4.1 Communities ........................................................................................................................ 83 4.2 Bee Community Composition ............................................................................................... 83 4.3 Habitat Patch and Landscape Composition Effects ............................................................... 84 4.4 Seasonal Effects and L. maackii Density ................................................................................ 87 5 Discussion........................................................................................................................................... 88 5.1 Bee Responses to Resolution and Scale ................................................................................. 88 5.2 Seasonal Resource Availability: Flowers, Lonicera