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Comparative Morphology and Evolutionary Relationships of the Sparidae (Teleostei: Percoidei)

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Comparative Morphology and Evolutionary Relationships of the Sparidae (Teleostei: Percoidei) COMPARATIVE MORPHOLOGY AND EVOLUTIONARY RELATIONSHIPS OF THE SPARIDAE (TELEOSTEI: PERCOIDEI) by JULIA JANE DAY A thesis submitted for the degree of Doctor of Philosophy, University of London January 2000 Department of Biology Department of Palaeontology University College London The Natural History Museum Gower Street Cromwell Road London, WC1E6BT London, SW7 5BD ProQuest Number: 10611135 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a com plete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest ProQuest 10611135 Published by ProQuest LLC(2017). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States C ode Microform Edition © ProQuest LLC. ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106- 1346 A b s t r a c t The comparative morphology of the family Sparidae is described comprehensively for the first time and is used to formulate character data for phylogenetic analysis. The data is found to be particularly character rich in areas such as the braincase, jaws and gill arches. Phylogenetic analysis, using PAUP* was performed in order to resolve the evolutionary relationships for 29 sparid genera and representatives sparoid families: Centracanthidae; Lethrinidae; Nemipteridae. Parsimony analysis of this data yielded three equally parsimonious trees. The Sparidae constitute a monophyletic group, with the inclusion of Centracanthidae, which is embedded within cladistically derived sparids. A grouping of derived sparids are found to be reasonably well supported when judged by Bremer support and bootstrapping, while relationships amongest those taxa more basal are found to be weakly supported. Further analysis of the data is assessed from 1) character quality through application of Le Quesne probabilities; 2) data partitioning; 3) influence of outgroups; 4) effects of ordering and 5) recoding, using non-additive binary coding. These analyses also support a hypothesis of relationships amongest derived sparids that is both well supported and resolved. However, the relationships of basal sparids are sensitive to these analyses, suggesting that not much confidence may be placed in the revealed theories of their interrelationships. The conflict between alternative trees reflects the high levels of homoplasy, which is not uncommon for percoid data sets. The geographic distribution is explained using three methods of cladistic biogeography, based on irreversible and reversible methods of ancestral area analysis and dispersal-vicariance analysis. The Indo-Pacific is identified as the most likely ancestral area for the Sparidae. Reconstruction of the evolution of feeding strategies among sparids suggests that there is a progressive transition from generalist to specialized feeders with four assemblages recognized. The diversification of feeding strategies within the Sparidae may have had important consequences for the evolution of the group which is discussed. Fossil sparid material from the Early to Middle Eocene is redescribed and included in the Recent matrix for further phylogenetic analyses. Comparison of the fossil material warrants the erection of a new genus and species, Ellaserrata monksi and a new genus Abromasta microdon is erected for Pagellus microdon. A minimum age of origin for the group can be postulated at 55Ma. PREFACE I declare that this dissertation is the result of my own work and includes nothing that is the outcome of work done in collaboration. No part of this work has been submitted for a degree, diploma or any other qualification at any other University. Julia J. Day iii ACKNOWLEDGEMENTS I am indebted to my supervisors, Michael Coates and Peter Forey for providing me with the opportunity to work within the field of systematic biology and for their continued support and encouragement. I would particularly like to express my thanks to Peter, for devoting time and energy into critically reviewing my work. I gratefully acknowledge the receipt of a BBSRC grant 96700156 which was the primary source of funding for this project, in addition to the Margaret- Brown Scholarship from University College London. I also thank the Systematics Association and the American Museum of Natural History for providing additional support for travel. This project as been carried out in the Department of Palaeontology, at The Natural History Museum, London, and for this I thank Robin Cocks and his successor Stephen Donovan for making me welcome here. I extend my sincere gratitude to all those at the NHM and UCL who provided help and encouragement over the past years, in particular; Claudia Amphlett, Helen Chatterjee, Sian Evans, Charlotte Jeffrey, Paul Jeffrey, David Lees, Claire Mellish, Neale Monks, Brian Rosen, Phil Rye, Marcello Ruta, Steve Salisbury, Jackie Skipper, Andrew Smith, John Stuart, Jon Todd, Paul Taylor, Mike Weedon, Sally Young. I would particularly like to thank Mark Wilkinson, Marcello Ruta and David Gower for discussion and their invaluable help into the theory and practice of cladistics and Neale Monks for Mac support. I would also like to thank the members of the fish research group of the Department of Zoology, NHM, in particular, Tony Gill, Ollie Crimmen and Sean Davidson for help with finding and preparing specimens. Thanks also to the NHM’s library services and photographic units, especially Phil Crabb and Nick Hayes. I am grateful to David Bellwood who brought the subject matter of this thesis to my attention and to David Johnson for his encouragement with pursuing this project. I also benefited from David’s capacious knowledge in interpreting the anatomy of gill arches, not to mention the provision of some indispensable dissection equipment. My work on the collections at the following institutions was made possible by Barbara Brown at The American Museum of Natural History; David Catania at the Californian Academy of Sciences; George Burgess at the Florida Museum of Natural History, Gainesville; Roberto Zorzin and Chiara Sorbini at the Museo Civico di Storia Naturale, Verona, Italy and Daniel Goujet at the Museum National d’histoire Naturelle, Paris, whom I thank for their help. I am also deeply grateful to the JLB Smith Institute of Ichthyology, South Africa, for the donation of specimens to the Natural History Museum, and in particular to Andrew Bentley who organised this for me. Thank you to all my friends, in particular Charlo, Claudia, Helen and Sarah for their much needed support. My special thanks to Mark for things more than I can say, but above all for being a wonderful friend. And to my parents, for their unending support and encouragement. I owe you all, thank you so much. iv Contents CONTENTS Title page i Abstract ii Preface iii Acknowledgements iv Contents v List of figures ix List of plates xii List of tables xiv CHAPTERS: 1. INTRODUCTION General introduction 1 Overview of current problems on percoid systematics 2 Sparid systematics 3 Aims of study 5 Methods 7 Materials 8 Anatomical nomenclature 9 Abbreviations of anatomical terminology 11 2. COMPARATIVE MORPHOLOGY OF THE SPARIDAE Introduction 19 Part 1 Osteology of the basal sparid Dentex dentex 20 Systematic zoology 20 Description 21 Cranium 21 v Contents Post-cranium 3 3 Part 2 Comparative morphology 3 7 Cranium 3 7 Post-cranium 5 1 Discussion 5 4 Character anaylsis 5 6 Data matrix 61 3. PHYLOGENETIC RELATIONSHIPS WITHIN THE SPARIDAE Introduction 98 Cladistic theory 9 9 Character partioning and coding in cladistic analysis 1 01 Character coding 103 Selection of taxa and outgroups 106 Part 1 Phylogenetic analysis 107 Character distribution 108 Missing values 108 Safe taxonomic reduction 110 Tree statistics 110 Tree topology 112 Character optimization 112 Monophyly of Sparoidea 112 Monophyly of Sparidae 113 Tree support 121 Inclusion of Centracanthidae within the Sparidae 123 Support for the phylogenetic hypothesis 123 Part 2 Further analyses of the data 124 Character quality 124 Data partitions 126 Partition homogeneity test 128 Deletion of outgroup taxa 129 Phylogenetic analysis using ordered characters 131 Effects of coding in phylogenetic inference 132 Discussion 135 4. PHYLOGENETIC BIOGEOGRAPHY AND ECOLOGY OF THE SPARIDAE Introduction 157 Part 1 Phylogenetic perspectives on biogeography 158 Contents Background to cladistic biogeography 158 Delimitation of geographic regions 159 Geographic distributions 159 Ancestral area 161 Ancestral area analysis using irreversible parsimony 161 Ancestral area analysis using reversible parsimony 162 Dispersal-vicariance analysis (DIVA) 163 Discussion 165 Part 2 Phylogenetic perspectives on feeding strategies 166 General 166 Taxon-feeding cladogram 167 Morphology and diet 168 Habitat associations 169 Discussion 170 5. FOSSIL RECORD AND EVOLUTIONARY HISTORY OF THE SPARIDAE Introduction 178 Part 1 Comparative morphology of fossil sparids 179 Previous work 179 Criteria used to identify fossil sparids 179 Overview of the fossil record 179 Systematic Palaeontology 180 The Bolca sparids 180 Sparnodus 181 Sparnodus vulgaris 182 ?Sparnodus micracanthus 188 Validity of Sparnodus vulgaris 189 Ellaserrata monksi (gen. et sp. nov.) 189 Abromasta microdon (gen. nov.) 192 Sciaenurus bowerbanki 195 General remarks 202 Part 2 Influence of fossils on Recent classification 204
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