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Taxonomic Review, Phylogeny, and Geographical Population Structure of the Sawfishes (Chondrichthyes, Pristiformes) Vicente Vieira Faria Iowa State University

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Taxonomic Review, Phylogeny, and Geographical Population Structure of the Sawfishes (Chondrichthyes, Pristiformes) Vicente Vieira Faria Iowa State University Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations 2007 Taxonomic review, phylogeny, and geographical population structure of the sawfishes (Chondrichthyes, Pristiformes) Vicente Vieira Faria Iowa State University Follow this and additional works at: https://lib.dr.iastate.edu/rtd Part of the Molecular Biology Commons, and the Zoology Commons Recommended Citation Faria, Vicente Vieira, "Taxonomic review, phylogeny, and geographical population structure of the sawfishes (Chondrichthyes, Pristiformes)" (2007). Retrospective Theses and Dissertations. 15506. https://lib.dr.iastate.edu/rtd/15506 This Dissertation is brought to you for free and open access by the Iowa State University Capstones, Theses and Dissertations at Iowa State University Digital Repository. It has been accepted for inclusion in Retrospective Theses and Dissertations by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. Taxonomic review, phylogeny, and geographical population structure of the sawfishes (Chondrichthyes, Pristiformes) by Vicente Vieira Faria A dissertation submitted to the graduate faculty in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Major: Ecology and Evolutionary Biology Program of Study Committee: Gavin Naylor, Co-major Professor John Nason, Co-major Professor Dean Adams Bonnie Bowen Gregory Courtney Colin Simpfendorfer Iowa State University Ames, Iowa 2007 Copyright © Vicente Vieira Faria, 2007. All rights reserved UMI Number: 3259448 UMI Microform 3259448 Copyright 2007 by ProQuest Information and Learning Company. All rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. ProQuest Information and Learning Company 300 North Zeeb Road P.O. Box 1346 Ann Arbor, MI 48106-1346 ii In memory of Fernando Faria Missing you dad. iii I dedicate this work to Patricia and Raul Eliete, Ana, and Guilherme iv TABLE OF CONTENTS ABSTRACT v CHAPTER 1. GENERAL INTRODUCTION 1 Thesis organization 2 CHAPTER 2. TAXONOMIC REVIEW OF THE SAWFISHES (CHONDRICHTHYES, PRISTIDAE) 3 Abstract 3 Introduction 4 Material & Methods 7 Historical account 17 Species account 41 Key to the species of Pristidae 77 References 79 Figures 91 Tables 106 CHAPTER 3. SAWFISH MOLECULAR PHYLOGENY 131 Abstract 131 Introduction 132 Material & Methods 135 Results 141 Discussion 146 References 150 Figures 158 Tables 165 CHAPTER 4. INFERENCES ON THE GEOGRAPHICAL POPULATION STRUCTURE OF ENDANGERED SAWFISHES 170 Abstract 170 Introduction 171 Material & Methods 174 Results 178 Discussion 185 References 193 Figures 201 Tables 205 Appendix 210 CHAPTER 5. GENERAL CONCLUSIONS 215 Future Research 217 ACKNOWLEDGMENTS 218 v ABSTRACT Sawfishes (Pristidae) are considered endangered species worldwide, but their conservation and management have been undermined due to poor understanding of the group’s natural taxonomy. The main purpose of this work was to review the taxonomy of sawfishes based on historical taxonomic literature supplemented with empirical observations on morphology, genetics, and geographic distribution. A molecular phylogenetics study based on both mitochondrial and nuclear genes was carried out to estimate the evolutionary interrelationships among the different species of sawfishes. Seven distinct living species are recognized based on external morphology and DNA sequence comparisons. Four are distributed in the Indo-West Pacific, two in the Atlantic, and one in the East Pacific. The phylogeny obtained suggests Anoxypristis cuspidata (Latham 1794) is sister to monophyletic Pristis. The later is further divided into two clades: (1) P. clavata Garman 1906 basal to sister species P. zijsron Bleeker 1851 and P. pectinata Latham 1794, and (2) P. zephyreus Jordan & Starks 1895 basal to sister species P. microdon Latham 1794 and P. perotteti Müller & Henle 1841. Population structure within species was investigated based on mitochondrial DNA sequence data. Results indicate that Western and Eastern Atlantic populations of P. pectinata and P. perotteti should be viewed as separate units for conservation purposes. The pattern for the Indo-West Pacific was less clear. It is not possible to invoke a mechanism of isolation that applies to all species. Nevertheless, a sub-division of sawfish populations into Indian and West Pacific conservation units is consistent with both the patterns of nucleotide variation and variation in rostral tooth count. vi Pristis pristis (Linneaus 1758) cannot be assigned to a single species. The nominal species P. pristis has historically been associated with features from several different species and as such is a chimaeric taxon that does not exist in nature. This has caused confusion in the past and in the present time. Herein, I propose the name P. pristis be suppressed. 1 CHAPTER 1. GENERAL INTRODUCTION The modern sawfishes first appear as fossils during the Eocene, 56 million years ago. They are characterized by a distinctive long snout with rostral teeth on the border. All sawfishes are classified in the family Pristidae of the order Pristiformes. Sawfishes can attain gigantic sizes: up to at least 6 m in total length. They are distributed in tropical and sub- tropical seas. They frequent bays and estuaries, and can penetrate great distances into rivers, far from tidal influences. Sawfish populations have declined globally due to overfishing and habitat destruction. Currently, all sawfish species are listed on the World Conservation Union’s (International Union for the Conservation of Nature) Red List of Threatened Species as Critically Endangered. Sawfish taxonomy is among the most confused of all of the chondrichthyans. Currently seven nominal species are recognized: the knifetooth sawfish A. cuspidata, the smalltooth group, including: P. pectinata, P. zijsron, P. clavata, and the largetooth group: P. perotteti, P. microdon and P. pristis. However, this number may vary between four and ten. Sawfishes are morphologically conserved, which makes identification of species within each group very difficult. Most species lack type material and most of material available for comparison in collections is composed of isolated rostra. In terms of nomenclature, the more stable taxa are P. clavata and P. zijsron. As for P. pectinata, this has been considered a circumglobal species, but some workers have suggested that P. pectinata may comprise multiple species. A. cuspidata taxonomic status is uncertain since different morphotypes, perhaps due to sexual dimorphism, have been seen. Perhaps the major problem in sawfish taxonomy concerns the largetooth group. This group is the most 2 conserved in terms of morphology. The historical taxonomy behind the origin of most available nominal species is also confusing. All these factors combined have resulted in disagreement of use of names and species distribution range that makes very little or no sense from a biogeographical standpoint. Dissertation organization The main purpose of this dissertation is to review the taxonomy of sawfishes based on critical evaluation of their historical taxonomic literature supplemented with empirical observations on morphology, genetics, and geographic distribution (Chapter 2). (This chapter is co-authored with Anthropologist and independent researcher Matthew McDavitt.) Once the taxonomy was resolved a molecular phylogenetics study based on both mitochondrial and nuclear genes was carried out to estimate the evolutionary interrelationships among the different species of sawfishes (Chapter 3). Relationships within cohesive species were subsequently investigated for their level of structuring within each major ocean basin (species distribution range). This was based on small fragments of mitochondrial DNA and rostral tooth counts obtained from museum specimens (Chapter 4). A summary of findings of the three preceding chapters is provided in the general conclusions chapter (Chapter 5). This chapter also outlines some general conclusions about the importance of taxonomy as a vital pre-requisite to any study of a given taxa. In addition, this chapter outlines some potential areas of future research that would further our understanding of the sawfishes. 3 CHAPTER 2. TAXONOMIC REVIEW OF THE SAWFISHES (CHONDRICHTHYES, PRISTIDAE) A paper to be submitted to the Zoological Journal of the Linnean Society Vicente Faria and Matthew McDavitt Abstract Sawfishes (Pristidae) are considered endangered or critically endangered species worldwide, but conservation and management measures have been undermined due to poor understanding of the group’s taxonomy. In the present study seven living species were identified in the family Pristidae. Pristis pristis (Linneaus, 1758) cannot be assigned to a single species. Pristis pristis has historically been associated with features from several different species and as such is a chimaeric taxon that does not exist in nature. The commonly asserted association between this nominal species and the largetooth sawfishes is the product of taxonomic misinterpretation. Sawfish species are assigned to Pristis clavata Garman 1906, Pristis pectinata Latham 1794, Pristis zijsron Bleeker 1851, Pristis microdon Latham 1794, Pristis perotteti Müller & Henle 1841, Pristis zephyreus Jordan & Starks 1895, and Anoxypristis cuspidata (Latham 1794). A thorough chronological revision of sawfish historical taxonomy is provided, as is an identification key to the sawfish species.
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