The Phylogeography and Conservation of the Brassy Minnow, Hybognathus Hankinsoni
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THE PHYLOGEOGRAPHY AND CONSERVATION OF THE BRASSY MINNOW, HYBOGNATHUS HANKINSONI by DAMON M. NOWOSAD B.Sc., University of Victoria, 1998 B.Ed., University of British Columbia, 2002 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES (Zoology) THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) March 2011 ©Damon M. Nowosad, 2011 Abstract Brassy minnow, Hybognathus hankinsoni, is a little-studied cyprinid fish with highly disjunct distributions in western Canada. Phylogeographic scenarios on the origins of brassy minnow in British Columbia (BC) were explored using two mitochondrial loci (cyt b and ND4) that were sequenced for up to 32 localities. This revealed an approximate ‘east-west’ geographic split, suggesting that BC populations are likely post-glacial colonists from the Mississippi-Missouri refugium. However, certain ‘eastern’ populations exhibited incongruences between gene and species trees, suggesting complex evolutionary histories in brassy minnow. Sampling brassy minnow within BC revealed highest catch rates in the Interior of the province, and a year-long survey conducted in the Lower Mainland (n = 60 sites), revealed that brassy minnow abundances were highest at Westham Island. A logistic-regression habitat model was performed incorporating seven physical habitat parameters over 37 sites, identifying conductivity as a near significant parameter for predicting brassy minnow presence. Additionally, in the Lower Mainland, eight invasive species constituted 28 % of the relative abundance of species sampled. Compared to historical records from the University of BC Fish Museum, sites exhibited a significant decline in the number of cypriniform species, including brassy minnow, but showed no significant change in the number of invasive species present. A detrended correspondence analysis (DCA) of species abundance across sites suggested brassy minnow overlapped most closely with two invasive species: bullfrogs, Lithobates catesbieanus, and brown bullhead, Ameiurus nebulosus. To explore the impact of brown bullhead on brassy minnow, pools with and without adult brown bullhead were compared at Tamboline Slough, Westham Island, and showed significant differences in the native fish species abundances across pools, including brassy minnow. Additionally, growth experiments were conducted over 90 days with brassy minnow, young-of-year brown bullhead, and redside shiner, Richardsonius balteatus, kept in all possible combinations for a total of four treatments per species. Treatments showed that brassy minnow were the only species to exhibit weight loss and mortality when with other species. In addition to providing insights into the biogeography, ecology, and conservation implications of brassy minnow, my thesis provides a quantitative baseline for invasive species found within the Lower Mainland. ii TABLE OF CONTENTS ABSTRACT...............................................................................................................................ii TABLE OF CONTENTS..............................................................................................................iii LIST OF TABLES......................................................................................................................vii LIST OF FIGURES...................................................................................................................viii ACKNOWLEDGEMENTS...........................................................................................................xi CHAPTER 1: GENERAL INTRODUCTION AND BACKGROUND .................................................... 1 1.1 Postglacial dispersal and (re)colonization of Canadian freshwater fishes .......................... 1 1.2 Conservation of freshwater fishes in Canada .................................................................... 2 1.3 Brassy minnow (Hybognathus hankinsoni) ....................................................................... 4 1.4 General thesis objectives and overview ............................................................................ 6 CHAPTER 2: THE PHYLOGEOGRAPHY AND ORIGINS OF H. HANKINSONI IN BC ......................... 7 2.1 Introduction .................................................................................................................... 7 2.1.1 The unusual and disjunct distribution of brassy minnow in BC ......................................... 7 2.1.2 Possible origins of brassy minnow in BC, and using molecular markers to test hypotheses on population movements ...................................................................................... 9 2.1.3 Mitochondrial (mtDNA) loci as molecular markers ......................................................... 11 2.1.4 General phylogeny of Hybognathus ................................................................................. 12 2.1.5 Objectives ......................................................................................................................... 13 2.2 Materials and methods .................................................................................................. 15 2.2.1 The Lower Mainland and lower Fraser River ................................................................... 15 2.2.2 Obtaining and collecting samples outside the Lower Mainland ...................................... 15 2.2.3 Genetic analyses - sequencing mitochondrial regions..................................................... 16 2.2.4 Phylogenetic analyses - gene tree re-constructions ........................................................ 17 2.3 Results ........................................................................................................................... 18 2.3.1 Cyt b and ND4 mitochondrial gene trees ......................................................................... 18 2.4 Discussion ...................................................................................................................... 21 iii 2.4.1 An approximate ‘east-west’ split in brassy minnow mtDNA gene trees ......................... 21 2.4.2 The origins of brassy minnow in BC and the Lower Mainland ......................................... 23 2.4.3 Confounded cyprinids - genetic polymorphisms and gene tree issues ........................... 24 2.5 Conclusions ................................................................................................................... 26 CHAPTER 3: TOWARDS A PREDICTIVE HABITAT MODEL FOR H. HANKINSONI ........................ 34 3.1 Introduction .................................................................................................................. 34 3.1.1 General Hybognathus conservation - vulnerable life histories? ...................................... 34 3.1.2 The overall distribution of brassy minnow in Canada, and the highly disjunct BC populations ................................................................................................................................ 35 3.1.3 Patchy local abundances of brassy minnow, and the value of a predictive habitat model ................................................................................................................................................... 36 3.1.4 Objectives ......................................................................................................................... 37 3.2 Materials and methods .................................................................................................. 38 3.2.1 Sampling in the Lower Mainland ..................................................................................... 38 3.2.2 One year minnow trap survey .......................................................................................... 38 3.2.3 Catch-per-unit effort ........................................................................................................ 39 3.2.4 Relative abundance and sampling autocorrelations........................................................ 39 3.2.5 Measuring site habitat attributes .................................................................................... 40 3.2.6 Use of logistic regression for predictive habitat models ................................................. 41 3.3 Results ........................................................................................................................... 42 3.3.1 Catch-per-unit-effort in BC for brassy minnow ................................................................ 42 3.3.2 Seasonal and regional brassy minnow relative abundances in the Lower Mainland ...... 42 3.3.3 Sites with positive temporal autocorrelations for brassy minnow sampling .................. 43 3.3.4 A predictive habitat model for brassy minnow ................................................................ 44 3.4 Discussion ...................................................................................................................... 44 3.4.1 Distribution and catch-per-unit-effort for brassy minnow in BC ..................................... 44 iv 3.4.2 Relative abundance, persistence, and regional habitat use of brassy minnow in the Lower Mainland......................................................................................................................... 48 3.4.3 Predicting brassy minnow presence ................................................................................ 52 3.5 Conclusions ..................................................................................................................