Biodiversity of Minnesota Caddisflies (Insecta: Trichoptera)
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Conservation Biology Research Grants Program Division of Ecological Services Minnesota Department of Natural Resources BIODIVERSITY OF MINNESOTA CADDISFLIES (INSECTA: TRICHOPTERA) A THESIS SUBMITTED TO THE FACULTY OF THE GRADUATE SCHOOL OF THE UNIVERSITY OF MINNESOTA BY DAVID CHARLES HOUGHTON IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY Ralph W. Holzenthal, Advisor August 2002 1 © David Charles Houghton 2002 2 ACKNOWLEDGEMENTS As is often the case, the research that appears here under my name only could not have possibly been accomplished without the assistance of numerous individuals. First and foremost, I sincerely appreciate the assistance of my graduate advisor, Dr. Ralph. W. Holzenthal. His enthusiasm, guidance, and support of this project made it a reality. I also extend my gratitude to my graduate committee, Drs. Leonard C. Ferrington, Jr., Roger D. Moon, and Bruce Vondracek, for their helpful ideas and advice. I appreciate the efforts of all who have collected Minnesota caddisflies and accessioned them into the University of Minnesota Insect Museum, particularly Roger J. Blahnik, Donald G. Denning, David A. Etnier, Ralph W. Holzenthal, Jolanda Huisman, David B. MacLean, Margot P. Monson, and Phil A. Nasby. I also thank David A. Etnier (University of Tennessee), Colin Favret (Illinois Natural History Survey), and Oliver S. Flint, Jr. (National Museum of Natural History) for making caddisfly collections available for my examination. The laboratory assistance of the following individuals-my undergraduate "army"-was critical to the processing of the approximately one half million caddisfly specimens examined during this study and I extend my thanks: Geoffery D. Archibald, Anne M. Christensen, Mark L. Galatowitsch, Karen A. Egerman, Christina. C. Fenendael, Peter A. Gillis, Kim Ha, Arrianne S. Haughland, Travis J. Ling, Emily A. Malcolm, Nathan J. O'Neil, Ryan C. Stephen, Jennifer M. Zaspel and Jessica L. Zeglin. Primary funding for this research came from a U.S. Environmental Protection Agency Science to Achieve Results Fellowship, and substantial support from the Minnesota Nongame Wildlife Tax Checkoff and Minnesota State Park Nature Store Sales i 3 through the Minnesota Department of Natural Resources' (MNDNR) Natural Heritage and Nongame Research Program. Special thanks are due to Richard J. Baker, MNDNR, for assistance with the latter funding source. Further support came from a Doctoral Dissertation Special Grant from the Graduate School, University of Minnesota (UM); several grants from the Dayton and Wilkie Fund, Bell Museum of Natural History, UM; grants from the Chiang Travel Fund, Department of Entomology, UM; and from the University of Minnesota Insect Museum. I appreciate the time and technical expertise of Joel Chirhart and Bob E. Murzyn, Minnesota Pollution Control Agency, for assisting with the analysis of landuse data. I, likewise, thank Jay C. Bell and Dan R. Wheeler, Department of Soil, Water, and Climate, UM, for doing the same with soil data. Thanks to Ed Quinn, MNDNR, Division of Parks and Recreation, for providing a permit to collect in state park habitats. Thanks to Matt Carroll, David A. Etnier, Elizabeth Etnier, Khamal Ghandi, and Robert Suranyi and for providing me lodging and hospitality while in the field. Additional thanks are due to David A. Etnier for guiding me to remote collecting sites in the Boundary Waters Canoe Area Wilderness, and for taking it upon himself to identify the specimens that we collected. Finally, I thank my colleagues at Insect Museum for support and sanity maintenance, especially Roger J. Blahnik, M. Lourdes Chamorro, Michelle A. DaCosta, Fernando Munoz-Quesada, Henrique Paprocki, Michael Saboren, Rebecca B. Simmons, and Jennifer M. Zaspel. I further thank my friends and family from outside the world of entomology for continually reminding me that the world does not revolve around caddisflies. Someday I may believe them. ii 4 ABSTRACT Over 300,000 caddisfly specimens were examined based on 317 light trap samples collected during 1999-2001, and from museum records. Two hundred-eighty four caddisfly species were determined to occur in Minnesota, representing 20 families and 74 genera. The relative occurrence in different regions, habitat types, and levels of upstream disturbance is documented for each species. Detrended Correspondence Analysis and a UPGMA dendogram of caddisfly relative abundance data from 58 Minnesota watersheds delineated five regions of caddisfly biodiversity. Caddisfly species richness and diversity were significantly highest in the Lake Superior and Northern regions, lowest in. the Northwestern and Southern regions, and intermediate in the Southeastern region. These determined regions had nearly double the classification strength of a priori ecological regions or watersheds in partitioning the natural variation of the caddisfly fauna. Classification strength also decreased with increasing spatial scale and decreasing taxonomic resolution. Canonical Correspondence Analysis found that temperature, percentage of disturbed habitat, and stream gradient were related to caddisfly species composition, although the correlation between temperature and disturbed habitat made determination of the relative importance of those variables difficult. Caddisfly species richness correlated negatively with percentage of disturbed upstream habitat for small and medium-sized streams in the Northern, Northwestern, and Southern regions. Change in the composition of trophic feeding groups based on habitat type generally followed a pattern predicted by the river continuum concept in the Lake Superior, Northern, and Southeastern regions, whereas fine particle filtering collectors iii 5 dominated all sizes of lakes and streams in the Northwestern and Southern regions. Binomial regression analysis determined three fine particle filtering collectors as indicators of habitat disturbance-particularly that of lakes and small streams- independent of other environmental variables. Although the absence of historical data makes it difficult to separate the relative importance of natural and anthropogenic factors, loss of caddisfly biodiversity and homogenization of feeding ecology has probably occurred in at least the Northwestern and Southern regions due to human disturbance. With baseline data now in place, potential future changes can be evaluated with greater confidence. iv 6 TABLE OF CONTENTS Acknowledgments ...........................................................................................…........i Abstract ................................................................................................................….iii Table of Contents………………………………………………………………..…..v List of Tables……………………………………………………………………….vii List of Figures....................................................................................………………ix CHAPTER 1. Introduction Importance of Biodiversity Research ............................................................ ……..1 Overview of the Trichoptera . .........................................................................…….2 Past Caddisfly Taxonomic Research ..............................................................…….3 Study Objectives ............................................................................................…….5 Literature Cited ..............................................................................................…….7 Figures............................................................................................................ ……15 CHAPTER 2. Delineation and Characterization of Minnesota Caddisfly Regions Abstract ............................................................................................................. 21 Introduction ....................................................................................................... 22 Materials and Methods ...................................................................................... 24 Results................................................................................................................ 33 Discussion ...................................................................................................... …39 Literature Cited ................................................................................... ...........….53 Tables .................................................................................................. ...........….71 Figures.................................................................................................. ...........….76 v 7 CHAPTER 3. Evaluation of Minnesota Geographic Classifications Based on Caddisfly Data Abstract .................................................................................................................... 100 Introduction ............................................................................................................. 100 Materials and Methods ............................................................................................ 104 Results and Discussion ............................................................................................ 106 Literature Cited ....................................................................................................... 114 Table ........................................................................................................................ 121 Figures .....................................................................................................................