(Coleoptera: Elateridae) Assemblages in the Acadian
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EFFECTS OF FOREST MANAGEMENT ON CLICK BEETLE (COLEOPTERA: ELATERIDAE) ASSEMBLAGES IN THE ACADIAN FOREST OF MAINE By Shelly L. Thomas B.S. Eastern University, 1995 A THESIS Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy (in Ecology and Environmental Science) The Graduate School The University of Maine May, 2007 Advisory Committee: Robert G. Wagner, Henry W. Saunders Distinguished Professor in Forestry, Advisor William A. Halteman, Associate Professor of Mathematics and Statistics Malcolm L. Hunter, Jr., Libra Professor of Conservation Biology Alan S. White, Professor of Forest Ecology Francis A. Drummond, Professor of Insect Ecology and Insect Pest Management John C. Brissette, Research Forester and Project Leader, USDA Forest Service LIBRARY RIGHTS STATEMENT In presenting this thesis in partial fulfillment of the requirements for an advanced degree at The University of Maine, I agree that the Library shall make it freely available for inspection. I further agree that permission for “fair use” copying of this thesis for scholarly purposes may be granted by the Librarian. It is understood that any copying of publication of this thesis for financial gain shall not be allowed without my written permission. Signature: Date: EFFECTS OF FOREST MANAGEMENT ON CLICK BEETLE (COLEOPTERA: ELATERIDAE) ASSEMBLAGES IN THE ACADIAN FOREST OF MAINE By Shelly L. Thomas Thesis Advisor: Robert G. Wagner An Abstract of the Thesis Presented in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy (In Ecology and Environmental Science) May, 2007 Click beetle (Coleoptera: Elateridae) assemblages were examined in three experiments in the Acadian forest of Maine. First, I used flight intercept traps to compare Elaterid assemblages in stands that developed after clearcut, shelterwood, and selection harvests. Click beetle richness was highest in stands managed under a selection system and lowest in stands regenerated using the clearcut method. The abundance of click beetle species was lower in stands managed by clearcutting than in stands managed using the shelterwood and selection methods. Hardwood basal area was the best environmental predictor for both species richness and species abundance. Second, I examined whether Elaterid assemblages in soil were associated with hardwood (Maple, Birch, Aspen, Blueberry, and Oak) and softwood (Spruce-Fir-Pine, Pine, Hemlock, and Fern-Hemlock) cover types. Elaterid diversity was generally higher in hardwood than softwood stands. Species richness was lowest in the Spruce-Fir-Pine and highest in the Oak cover type. Species abundance was lowest in the Spruce-Fir-Pine and Pine and highest in the Oak, Maple, and Hemlock cover types. Assemblages in hardwood stands were less similar to those in coniferous stands than they were to each other, with assemblages in oak stands being least similar to those in coniferous or other hardwood stands. Four species of click beetle were more abundant in softwood stands, and seven species were associated with increases in specific softwood tree and shrub species. Nine species of click beetle were more abundant in hardwood stands, and fifteen species were associated with increases in specific hardwood tree and shrub species, including six species associated with oak stands. Third, I examined how the species richness, abundance, diversity, and assemblage similarity of click beetles inhabiting coarse woody material (CWM) were affected by gap harvesting and characteristics of the CWM (diameter, degree of decay, and wood type) in Maine’s Acadian forest. Species assemblages varied between harvest treatments, canopy conditions, CWM wood type (hardwood vs. softwood), and especially between CWM decay classes and among diameter classes. Size of harvest gap did not influence the species abundance of click beetles across the small range of gap sizes studied (0.01 to 0.21 ha), and there were few differences between the two harvest treatments. Four of the most common species had higher abundances in closed canopy than harvest gaps. Click beetle species richness and species abundance were higher in CWM that had larger diameters and were more decayed. Click beetle diversity was higher in softwood than hardwood CWM. i ACKNOWLEDGEMENTS I would like to thank my advisor, Bob Wagner; your encouragement, dedication, input, and sense of humor were invaluable and made it possible for me to complete my work. Thank you very, very much! Thank you to my committee members, including Frank Drummond and John Brissette, for your reviews, especially to Mac Hunter for inviting me to your “Mac Meetings,” Al White for your encouragement and time, and most especially to Bill Halteman for the many hours you spent helping me understand my statistical analyses and honing my statistical skills, and going above and beyond the call of duty when it was needed the most. Thank you to Rebecca Holberton and Dave Foster for participating in my comprehensive exams. Thanks to Steve Woods for making it possible for me to study click beetles. Susan Brawley, thank you for your dedication to excellent education, for your hard work at sustaining the GK-12 program, and for helping to make it possible for me to travel back to Kenya. Thank you to Steve Kingsbury for being such a good example of an excellent middle school science teacher. Thanks to the many graduate students who have helped me, including Hanne Jorgensen, Mike Saunders, Tom Woodcock, and the “Mac Meeting” students- Steve Campbell, Fred Beaudry, Dave Patrick, Sean Blomquist, and the late Dave Ververka. Thanks to Dick Dearborne and Charlene Donahue for the use of the MFS insect collection, to Tim Stone for his help with PEF data, and to my many student workers, without whom little could be accomplished. Sincere gratitude to Edward C. Becker and Serge Laplant for correcting my click beetle reference collections. Heartfelt thanks to Joe Sheldon, who guided me into the wonderful world of very small critters and who instigated my thinking about the deep connection between faith ii and ecology. Thanks to the staff and volunteers of Au Sable Environmental Institute, Christians for Biblical Equality, and A Rocha International for your dedication and witness. Thanks to my church families in PA and ME for praying for me through the years; special thanks to Pathway for helping me develop my gifts as the director of the Growth Rings Ministry. My deepest gratitude goes to the many friends who have prayed, laughed, and cried with me during this time, especially Kathy and Chris, Steph O., Traci, Gary, Christi, Jean, Karen, Steph N., Jenny, Megan, Joy, Amy, Fair, and Julie. I would especially like to thank Jack J. Heddon. My favorite memories of these grad school years are inundated with experiences I shared with you. Thank you for your loving friendship, for your enduring patience, for your constant encouragement, and for allowing me to learn from you. I love you, my friend, and I wish you well in all things. Mom and Dad, you have been my inspiration, my shelter, my encouragers, my teachers, and my most rabid fans! Thank you for raising me in a beautiful home, infecting me with your curiosity, passion, and joy, making it possible for me to travel the world, and supporting me through these years. Thank you for introducing me to the Beautiful One and to the Way. I love you both very much! Finally, to the One who was, who is, and who is to come: thank you for your creativity. Please help us to care for your creation and to love as you would have us love- with action, strength, humility, justice, mercy, and hope- and in so doing to create the community you desire. Baruch ata adonai, eloheinu melech ha'olam, borei ha'chipvushit! iii TABLE OF CONTENTS ACKNOWLEDGEMENTS.................................................................................................ii LIST OF TABLES...............................................................................................................x LIST OF FIGURES..........................................................................................................xiv PROLOGUE........................................................................................................................1 Chapter 1. EFFECT OF SYLVICULTURAL METHODS AND ASSOCIATED STAND CHANGES ON CLICK BEETLE (COLEOPTERA: ELATERIDAE) ASSEMBLAGES IN MAINE’S REGION OF THE ACADIAN FOREST.......................6 1.1. Abstract............................................................................................................6 1.2. Introduction......................................................................................................7 1.3. Methods..........................................................................................................10 1.3.1. Study Site........................................................................................10 1.3.2. Measurements.................................................................................13 1.3.2.1. Beetle Sampling...............................................................13 1.3.2.2. Vegetation Sampling.......................................................14 1.3.3. Analytical Approach......................................................................15 1.4. Results...........................................................................................................18 1.4.1. Beetle Taxa.....................................................................................18 iv 1.4.2. Effect of Silvicultural Methods....................................................19