3 



CONTENTS 



Page 



Introduction 5 



Acknowledgments 5 



Methods 6 



Material examined ; . 8 



Historical review of Phoxinus 12 



Phylogenetic relationships of Phoxinus and the related genera 22 



Monophyly of the Hemitremian clade and its position in the family Cyprinidae 22 



Analysis of transformation series in the Hemitremian clade 26 



Phylogenetic relationships of the Hemitremian clade 39 



Discussion . 43 



Non-osteological morphology 44 



External morphology 44 



Intestine and gas bladder , 61 



Osteology 67 



Neurocranium 67 



Viscerocranium 95 



Vertebral column 121 



Pectoral girdle and fin 131 



Phylogenetic relationships of the species of Phoxinus 141 



Phylogenetic relationships 141 



Discussion on the phylogenetic relationships of the species of Phoxinus 144 



Biogeography of Phoxinus 145 



Taxonomy of Phoxinus 150 



Key to the species of Phoxinus 153 



Species accounts 154 



Phoxinus phoxinus (Linnaeus) 154 



Phoxinus brachyurus Berg 161 



Phoxinus issykkulensis Berg 163 



Phoxinus neogaeus Cope 166 



Phoxinus cumberlandensis Starnes & Starnes 171 



Phoxinus tennesseensis Starnes & Starnes 175 



Phoxinus oreas (Cope) 179 



Phoxinus eos (Cope) 183 



Phoxinus erythrogaster (Rafinesque) 188 



Abstract . 1 94 



Literature Cited 196 



Appendices 215 



I. Data matrix I (for Hemitremians) 215 



II. Data matrix II (for Phoxinus) 216 



III. Transformation series used in the analysis of phylogenetic relationships 



within Phoxinus 217 



IV. List of abbreviations 225 



