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Phylogeography of Smallmouth Bass (Micropterus Dolomieu)

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Phylogeography of Smallmouth Bass (Micropterus Dolomieu) PHYLOGEOGRAPHY OF SMALLMOUTH BASS (MICROPTERUS DOLOMIEU) AND COMPARATIVE MYOLOGY OF THE BLACK BASS (MICROPTERUS, CENTRARCHIDAE) WILLIAM CALVIN BORDEN Bachelor of Science in Zoology University of Toronto Toronto, Ontario 1991 Master of Science in Zoology University of Toronto Toronto, Ontario 1996 Submitted in partial fulfillment of requirements for the degree DOCTOR OF PHILOSOPHY IN REGULATORY BIOLOGY at the CLEVELAND STATE UNIVERSITY May, 2008 This dissertation has been approved for the Department of Biological, Geological, and Environmental Sciences and for the College of Graduate Studies by Date: Dr. Miles M. Coburn, Department of Biology, John Carroll University Co-advisor Date: Dr. Robert A. Krebs, BGES/CSU Co-advisor Date: Dr. Clemencia Colmenares, The Cleveland Clinic Advisory Committee Member Date: Dr. F. Paul Doerder, BGES/CSU Advisory Committee Member Date: Dr. Harry van Keulen, BGES/CSU Internal Examiner Date: Dr. Mason Posner, Department of Biology, Ashland University External Examiner One fish two fish red fish blue fish. Black fish blue fish old fish new fish. This one has a little star. This one has a little car. Say! what a lot of fish there are. — Dr. Seuss, One fish two fish red fish blue fish ACKNOWLEDGMENTS Many people were involved in the completion of this work. Firstly, numerous groups and individuals provided tissue samples of smallmouth bass and other fishes. They include J. McClain, A. Bowen, and A. Kowalski (USFWS – Alpena, MI); L. Witzel, A. Cook, E. Arnold, and E. Wright (Ontario Ministry of Natural Resources – Lake Erie Management Unit); J. Hoyle (OMNR – Lake Ontario Management Unit); M. Nadeau and D. Gonder (OMNR – Upper Great Lakes Management Unit); K. Schmidt, B. Stefanoky, N. Proulx, and J. Lehner (MnDNR); T. Burzynski (WDNR); J. Johnson, S. DeWitt, J. Diemond, M. Werda, K. Glomski, and D. Fielder (MiDNR – Alpena Fishery Research Station); E. Braun, A. Grier, M. Burlingame, and S. Sheldon (InDNR); K. Kayle, J. Deller, C. Knight, T. Bader, A. Ford (Ohio Division of Wildlife – Fairport Harbor Unit); R. Knight, J. Tyson, and G. Emond (ODW – Sandusky Unit); M. Greenlee (ODW); D. Einhouse, W. Pearsall, J. Evans, and S. Cornett (NYDSEC); B. Crump and R. Standage (Ouachita NF); B Zawiski and S. Taylor (Ohio EPA); M. Austin (Ohio Department of Transportation); R. Stein (Ohio State U); R. Butryn and R. Gonser (Indiana State U); G. Steinhart (Lake Superior State U); A. Echelle (Oklahoma State U); M. Coburn and L. Kousa (John Carroll U); R. Draper (The Ohio Smallmouth Alliance); B. Whiting (Trust Lands Office); A. Thomas, M. Lyons, J. Holt, A. Ford, B. Dunagan, T. Bowens, M. Jedlicka, O. Lockhart, J. Giboney, T. Marth, and D. Kiss (Cleveland State U); C. Landreville; J. Graham; M. Smith; P. and C. Hughes. In particular, the Fairport Harbor staff (ODW) provided samples that formed the bulk of chapters 2 and 3 and offered assistance at every step of the project. I sincerely appreciate their contributions and wish them to understand that this project was doomed without their support. I hope that this work in some way benefited their needs. Several additional groups invited me to join them in the field (MiDNR, USFWS, InDNR, NYSDEC). I enjoyed the opportunity and challenge of integrating academic and fisheries interests and needs and look forward to maintaining this connection in the future. Financial support to conduct field and lab work was provided by the Department of Biological, Geological, and Environmental Sciences and the College of Science (CSU), an Ohio Sea Grant (R/RL-5 to C. Stepien, U of Toledo), a Doctoral Research Expense Award (CSU), research funds of Miles Coburn and Bob Krebs, and personal funds. In addition, BGES provided a salary through a teaching assistantship for four years, although at times I wish they had not!! The morphological work was largely self-funded. Chris Sheil (John Carroll U) provided space, equipment, and critiques of my illustration skills, or lack thereof. His Scientific Illustration course provided much of the incentive to finish this part of the dissertation. Specimens for dissection and clearing and staining were graciously provided by R. Robins, T. Vigliotti, L. Page, and J. Albert (FLMNH), R. Winterbottom (ROM), B. Kuhajda (UAIC), J. Williams, S. Smith, and S. Jewett (USNM), N. Rios and H. Bart, Jr. (TU), D. Catania (CAS), T. Cavender and M. Kibbey (OSUM), D. Nelson (UMMZ), G. Steinhart and R. Stein (OSU), B. Shaner, P. Landford, and J. Biagi (GADNR), G. Garrett (TXDP&W), and J. Williams (USGS-FL). As much as I evolved throughout this process, so did my committee and advisors. I used an assortment of advisors at various times; each voice contributed something to the final product. They include John Holcomb (Mathematics Department at CSU), Ralph Gibson (BGES), Joe Keiper (CMNH), Kevin Kayle and John Deller (ODW – Fairport Harbor), and Michael Tevesz (Center for Sacred Landmarks at CSU). Clemencia Colmenares (Cleveland Clinic Foundation) and Harry van Keulen (CSU) were always enthusiastic and provided sound advice from science to politics to straw-house building. Mason Posner (Ashland U) was kind enough to join the committee as an external reviewer. However, several people carried the bulk of the responsibility with respect to my intellectual and academic development. Rick Winterbottom (ROM) understood my need to return to school and provided encouragement during the early years in particular when I was in doubt. Michael Gates (CSU) also provided an ear and sage advice when I contemplated leaving the program. During this time, Miles Coburn stepped forward to be an advisor. He was soon joined by Bob Krebs, and collectively they were a voice of logical reason and continual support while giving me plenty of freedom to make my own trail, and perhaps more importantly, make my own mistakes. Through it all, they encouraged my scholarship, protected my opportunity to carve out a niche in icththyology, and defended me when necessary while often taking the brunt of sharp criticism for their efforts. Finally, Paul Doerder was first a fellow hiker and later a committee member, but always an ardent supporter and defender. In many ways, the fact that he was not a committee member allowed us to discuss topics that exclude traditional advisors and students. As a consequence, I received an education in the politics of academia. One trait they share is a quiet and reserved demeanor that comes in part with a self-confidence of person and an expertise of field. This quality amongst others, made it enjoyable to work with each of them. Suffice it to say, I am indebted to them. A number of graduate students provided plenty of distractions from the work that should have been done. Alex, Toriano, Jenny, and Melissa were former labmates and all have moved on to bigger and better things. Congratulations. Finally, my parents financially supported me during semesters when I chose not to TA and were always confident that I would make the right decisions for me. Of course, they shared much of my anxiety and stress. Former graduate students from U of Toronto reminded me of the light on the other side. The Toronto gang kept me grounded and offered assistance in disposing of bodies. Amongst my biggest Canadian supporters is Cathy who saw me through the rough times and just wanted me to finish. In Cleveland, Annmarie and Francis (the legacy of CESTP is alive and well) did their best to break the last straw while simultaneously providing a welcome relief from the grind of school. To all I say “Thank you” and hope that I can repay your efforts and time. PHYLOGEOGRAPHY OF SMALLMOUTH BASS (MICROPTERUS DOLOMIEU) AND COMPARATIVE MYOLOGY OF THE BLACK BASS (MICROPTERUS, CENTRARCHIDAE) WILLIAM CALVIN BORDEN ABSTRACT Biogeographic patterns were investigated within a single species and among its congenerics. Smallmouth bass (Micropterus dolomieu) were characterized by portions of the cytochrome b gene and the control region although bass in Lake Erie were also evaluated using eight nDNA microsatellites. Overall, little population structure was observed among sites in Lake Erie which was surprising given that most male smallmouth bass exhibit nest site fidelity. But, these lake bass were divergent from bass residing in nearby tributaries indicating that lacustrine bass are reproductively isolated from riverine bass even though both groups may spawn in the same rivers. Adjacent riverine populations have also diverged from each other. Smallmouth bass from 71 sampling sites spanning their native distribution were analyzed to examine post-glaciation colonizing routes from glacial refugia into the Great Lakes. Phylogenetic analyses indicated that ancestral haplotypes exist in today’s Interior and Eastern Highlands, and the tree structure supported sequential colonization events northward with a loss of diversity in more distant populations. Western rivers and the upper Great Lakes were characterized by one phylogroup that viii was distinct from bass occupying the Ouachita Highlands. Bass from Lake Ontario, Georgian Bay, and the St. Lawrence River were characterized largely by just a single, derived haplotype of unknown geographical origin. The two central lakes, Huron and Erie, were composed of multiple phylogroups indicating numerous source populations and access routes. The unique biogeography of smallmouth bass and its congenerics (Micropterus, Centrarchidae) was addressed using comparative morphology to complement the molecularly based phylogeographic component. The genus’s greatest diversity is centered in the southeastern USA although several outliers are restricted to watersheds in Texas or the Interior Highlands. Myological variation was quantified for all black bass, but characters suitable for phylogenetic analysis were minimal. Observed variation was characterized primarily as unique abnormalities, often within a single specimen, or mimicking variants that were shared irregularly among species. The lack of significant myological variation may be correlated with the low degree of ecological, anatomical, and life history diversity among black bass.
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