(Cyprinidae: Cypriniformes) with a Description of a New Species from the Tennessee River Drainage, Tennessee

University of Tennessee, Knoxville TRACE: Tennessee Research and Creative Exchange Doctoral Dissertations Graduate School 5-2001 Phylogenetic systematics and zoogeography of the nearctic members of the minnow genus Phoxinus (Cyprinidae: Cypriniformes) with a description of a new species from the Tennessee River drainage, Tennessee Christopher Eugene Skelton University of Tennessee Follow this and additional works at: https://trace.tennessee.edu/utk_graddiss Recommended Citation Skelton, Christopher Eugene, "Phylogenetic systematics and zoogeography of the nearctic members of the minnow genus Phoxinus (Cyprinidae: Cypriniformes) with a description of a new species from the Tennessee River drainage, Tennessee. " PhD diss., University of Tennessee, 2001. https://trace.tennessee.edu/utk_graddiss/6383 This Dissertation is brought to you for free and open access by the Graduate School at TRACE: Tennessee Research and Creative Exchange. It has been accepted for inclusion in Doctoral Dissertations by an authorized administrator of TRACE: Tennessee Research and Creative Exchange. For more information, please contact [email protected]. To the Graduate Council: I am submitting herewith a dissertation written by Christopher Eugene Skelton entitled "Phylogenetic systematics and zoogeography of the nearctic members of the minnow genus Phoxinus (Cyprinidae: Cypriniformes) with a description of a new species from the Tennessee River drainage, Tennessee." I have examined the final electronic copy of this dissertation for form and content and recommend that it be accepted in partial fulfillment of the equirr ements for the degree of Doctor of Philosophy, with a major in Ecology and Evolutionary Biology. David A. Etnier, Thomas G. Hallam, Major Professor We have read this dissertation and recommend its acceptance: Arthur C. Echternacht, Gary F. McCracken, Paul W. Parmalee Accepted for the Council: Carolyn R. Hodges Vice Provost and Dean of the Graduate School (Original signatures are on file with official studentecor r ds.) To the Graduate Council: We are submitting herewith a dissertation written by Christopher Eugene Skelton entitled "Phylogenetic Systematics and Zoogeography of the Nearctic Members of the Minnow Genus Phoxinus (Cyprinidae: Cypriniformes) with a Description of a New Species from the Tennessee River Drainage, Tennessee". We have examined the final copy of this dissertation for form and content and recommend that it be accepted in partial fulfillment of the requirements for the degree of Doctor of Philosophy, with a major in Ecology and Evolutionary Biology. ~avidA./~· Etnfer,1lz/4' Major Prof:tf/7 Thomas G. Hallam, Department Head We have read this dissertation ommend its acceptance: C. C. Echtemacht Gary F. cCraci&n / Paul W. Parmalee Accepted for the Council: PHYLOGENETIC SYSTEMATICS AND ZOOGEOGRAPHY OF THE NEARCTIC MEMBERS OF THE MINNOW GENUS PHOXINUS (CYPRINIDAE: CYPRINIFORMES) WITH A DESCRIPTION OF A NEW SPECIES FROM THE TENNESSEE RIVER DRAINAGE, TENNESSEE A Dissertation Submitted for the Doctor of Philosophy Degree The University of Tennessee, Knoxville Christopher Eugene Skelton May 2001 11 ACKNOWLEDGMENTS I would like to thank my major professor, Dr. David Etnier for suggesting this study and teaching me untold numbers of things about many subjects. I thank Arthur C. Echtemacht, Gary F. McCracken, and Paul W. Parmalee for participating on my committee and providing helpful comments on my dissertation. For field assistance I thank Dick Bryant, Pat Ceas, Deanna Daniels, Roland Kays, J.D. Freye, Maureen Hays, Steve Holdeman, Clinton Jones, Charlie Saylor, Ed Scott, Chris Paxton, and Wayne Starnes. For genetics help I am grateful to Stan Guffey and John Young. For help obtaining financial support during various phases of the study I thank Richard Biggins, Charles Saylor, and Rick Bivens. I am especially grateful to Rick Bivens, Bart Carter, and Carl Williams of the Tennessee Wildlife Resources Agency (TWRA) for providing field and photography support. Bob Muenchen and Barry Hart assisted with statistical work. These past years would have been much less fun without the razor sharp wit of Bo Baxter, Brian Evans, Steve Fraley, Barry Hart, Steve Holdeman, Mark Hughes, Barron Moody, and Chris Paxton. Collections were made under permits issued by the TWRA to D. Etnier and C. Skelton. Laboratory protocols were reviewed and passed by the Office of Laboratory Animal Care (Permit #723). 111 ABSTRACT Questions regarding the taxonomy and systematics of the Nearctic Phoxinus (Cyprinidae: Cypriniformes) were investigated in this study. First, a new member of the genus, Laurel Dace, was described from the Tennessee River drainage, Tennessee. This distinctive minnow is one of seven described North American Phoxinus species and one of four that occurs in Tennessee. It differs from all congeners by having the combination of two uninterrupted black lateral stripes, and development of black pigment on the underside of the head of nuptial males. Based on coloration and tuberculation patterns, the Laurel Dace is hypothesized to be part of a monophyletic group comprised of P. areas, P. tennesseensis, and P. cumber/andensis. The Laurel Dace appears to be very rare; it is currently known from only six localities on the Walden Ridge portion of the Cumberland Plateau. Human activities such as coal mining, agriculture, and timbering threaten its future existence. A second objective was to assess the phylogenetic relationships of the Nearctic Phoxinus. Five data sets were employed to this end. The first consisted of allele frequencies calculated from gene products resolved with starch gel electrophoresis, and the second of 30 morphological characters. The third was a subset of the morphological data set exclusive of all characters construed as being related to feeding. The fourth and fifth data sets employed a combination of the allele frequency data and the two morphological data sets, respectively. There was little congruence between results generated with morphological data sets and the discretely coded allozyme data set. The lV latter data set produced poorly resolved results and was not useful for phylogenetic analysis. However, a UPGMA phenogram based on Rogers genetic similarity was highly congruent with a tree produced from the morphological data set exclusive of feeding characters. The relationships indicated by the UPGMA phenogram and the morphological data set were both considered viable hypotheses. Strong evidence supported a clade consisting of P. areas, P. tennesseensis, P. cumber/andensis, and Laurel Dace. Of the remaining Nearctic Phoxinus, P. erythrogaster is sister to this clade or sister to P. eos. Phoxinus neogaeus is the basal member of the genus. Distributions of Phoxinus species are hypothesized to be influenced in large part by Pleistocene glaciation and stream capture events. V TABLE OF CONTENTS PART 1: INTRODUCTION GENERAL BACKGROUND .............................................. 2 OBJECTIVES .......................................................... 4 LITERATURE CITED ................................................... 6 PART 2: A NEW DACE OF THE GENUS PHOXINUS (CYPRINIDAE: CYPRINIFORMES) FROM THE TENNESSEE RIVER DRAINAGE, TENNESSEE INTRODUCTION ...................................................... 10 MATERIALS AND METHODS ........................................... 11 RESULTS ............................................................ 13 Holotype ........................................................ 14 Allotype ........................................................ 14 Paratopotypes .................................................... 14 Paratypes . 14 Non-type Material ................................................ 15 Diagnosis . 16 Description ...................................................... 16 Coloration ...................................................... 17 Tuberculation .................................................... 19 Variation ....................................................... 20 Comparisons .................................................... 22 DISCUSSION ......................................................... 23 Relationships and Biogeography ..................................... 23 Distribution ..................................................... 24 Behavior, Ecology, and Life History .................................. 25 Conservation Status ............................................... 27 MATERIAL EXAMINED ................................................ 30 LITERATURE CITED .................................................. 33 APPENDICES ......................................................... 36 VI PART 3: PHYLOGENETIC SYSTEMATICS AND ZOOGEOGRAPHY OF THE NEARCTIC MEMBERS OF THE MINNOW GENUS PHOXINUS (CYPRINIDAE: CYPRINIFORMES) INTRODUCTION ...................................................... 63 MATERIALS AND METHODS ........................................... 64 Outgroup Choice ................................................. 65 Allozyme Methods ................................................ 66 Phylogenetic Methods ............................................. 68 RESULTS ............................................................ 69 Allozyme Analyses ............................................... 69 Phylogenetic Analyses ............................................. 71 DISCUSSION ......................................................... 73 Allozyme Analyses ............................................... 73 Phylogenetic Analyses ............................................

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