Effects of Shelterwood Management on Flower-Visiting Insects and Their
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EFFECTS OF SHELTERWOOD MANAGEMENT ON FLOWER- VISITING INSECTS AND THEIR FLORAL RESOURCES by Jessica Erin Fultz A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in Entomology MONTANA STATE UNIVERSITY Bozeman, Montana July 2005 © COPYRIGHT by Jessica Erin Fultz 2005 All Rights Reserved ii APPROVAL of a thesis submitted by Jessica Erin Fultz This thesis has been read by each member of the thesis committee and has been found to be satisfactory regarding content, English usage, format, citations, bibliographic style, and consistency, and is ready for submission to the College of Graduate Studies, Dr. Kevin O’Neill Approved for the Department of Entomology Dr. Gregory D. Johnson Approved for the College of Graduate Studies Dr. Joseph Fedock iii STATEMENT OF PERMISSION TO USE In presenting this thesis in partial fulfillment of the requirements for a master’s degree at Montana State University – Bozeman, I agree that the library shall make it available to borrowers under the rules of the Library. If I have indicated my intention to copyright this thesis by including a copyright notice page, copying is allowable only for scholarly purposes, consistent with “fair use” as prescribed in the U.S. Copyright Law, Requests for permission for extended quotation from or reproduction of the thesis in whole or in parts may be granted only by the copyright holder. Jessica Erin Fultz July 15, 2005 iv ACKNOWLEDGEMENTS Without funding from the United States Forest Service and help from George Markin, Ward McCaughey, and Elaine Sutherland this thesis would not have been possible. Thanks to Dr. Kevin O’Neill for writing the grant proposal, for his guidance, for giving me some freedom, and for taxonomic help with the Sphecidae. Thanks to my graduate committee: Kevin O’Neill, Rich Hurley, Matt Lavin, George Markin, and Bob Peterson for their helping improving my thesis. Thanks to Dr. Richard Miller for the bee and beetle identifications, and for our conversations. Thanks to Ken Aho for his guidance with multivariate statistics, the use of PC-ORD, and his understanding. I also greatly appreciate all the taxonomic help I received from Dr. Matt Lavin (MONT), Dr. Richard Hurley (MTEC), Dr. Mike Ivie (MTEC), and Dr. F. Christian Thompson (SEL). Thanks to Scott Mincemoyer and Jennifer Birdsall for giving me a copy of their soon to be published paper on the vascular plants of the TCEF, Peter Jensen for the first year of data collection, Dr. Robert Peterson for the use of his computer, Dr. Donald Burgess for the use of his phase contrast microscope and digital camera, and to Leon Theroux and Matt Watherspoon for their help around camp. I appreciate the friendly and familial assistance given to me by Art and Sheila Fultz, Alvin and Dena Hoff, the Clark family, the Langel family, Kristy and Judy Mahlum, Amy Myran, Phyllis Roth, Katie Marske, Ken Puliafico, and Taryn Flesjer. Thanks to the cockroaches for being around to inspire and entertain. And finally thanks to the MSU Department of Entomology for being an important part of my undergrad as well as graduate education; I’m sorry you no longer exist. v TABLE OF CONTENTS LIST OF TABLES...........................................................................................................viii LIST OF FIGURES ............................................................................................................ x ABSTRACT...................................................................................................................... xii 1. INTRODUCTION .......................................................................................................... 1 Habitat Disturbance and Pollinator Abundance.............................................................. 3 Silvicultural Practices as Habitat Disturbances .............................................................. 5 Pollination....................................................................................................................... 9 Flower-visiting Insects.................................................................................................. 14 Coleoptera................................................................................................................ 16 Lepidoptera .............................................................................................................. 17 Diptera...................................................................................................................... 18 Hymenoptera............................................................................................................ 19 2. METHODS ................................................................................................................... 23 Study Area .................................................................................................................... 23 Sampling of Floral Resources....................................................................................... 26 Sampling of Flower-visiting Insects ............................................................................. 27 Sampling and Identification of Pollen .......................................................................... 29 Data Analysis................................................................................................................ 30 Abundance, Density, and Diversity ......................................................................... 30 Differences Among Treatments............................................................................... 32 Non-metric Multidimensional Scaling (NMS) ................................................... 32 Multi-Response Permutation Procedure (MRPP)............................................... 35 Association of Floral Resources and Flower-visiting Insects.................................. 37 Mantel Test. ........................................................................................................ 37 3. RESULTS ..................................................................................................................... 38 Floral Resources............................................................................................................ 38 Density and Diversity .............................................................................................. 38 Overall Floral Density......................................................................................... 38 Species Richness (α). .......................................................................................... 39 Shannon-Wiener Index (H’) ............................................................................... 39 Gamma Diversity (γ)........................................................................................... 40 Beta Diversity (γ/α)............................................................................................. 40 Changes in Flower Species ...................................................................................... 42 vi TABLE OF CONTENTS - CONTINUED Unlogged Control................................................................................................ 42 Shelterwood treatments....................................................................................... 42 Meadow Control. ................................................................................................ 43 Differences Among Treatments............................................................................... 47 Flower-visiting Insects.................................................................................................. 52 Abundance and Diversity......................................................................................... 52 Overall Abundance.. ........................................................................................... 52 Species Richness (α) ........................................................................................... 52 Shannon-Wiener Diversity (H’).......................................................................... 53 Gamma Diversity (γ)........................................................................................... 54 Beta Diversity (β)................................................................................................ 54 Changes in Insect Species........................................................................................ 59 Differences Among Treatments............................................................................... 72 Association of Floral Resources and Flower-visiting Insects....................................... 75 Pollen on Insects ........................................................................................................... 76 Quantity and Diversity............................................................................................. 76 Pollen Quantity ................................................................................................... 76 Pollen Type Richness (α).................................................................................... 77 Gamma Diversity (γ)........................................................................................... 78 Beta Diversity (β)................................................................................................ 78 Pollen Types on Families.......................................................................................... 79 Cerambycidae. ...................................................................................................