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I the SYSTEMATICS and BIOLOGY of the GENUS POLTYS CORE Metadata, citation and similar papers at core.ac.uk Provided by Sydney eScholarship THE SYSTEMATICS AND BIOLOGY OF THE GENUS POLTYS (ARANEAE: ARANEIDAE) IN AUSTRALASIA HELEN MOTUM SMITH Thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy Faculty of Agriculture, Food and Natural Resources, The University of Sydney, New South Wales 2006 Submitted August 2006 i DECLARATION All work presented in this thesis is the original work of the author, except where otherwise stated. This thesis contains no material that has been submitted for the award of a higher degree to any other university or institution. Helen M. Smith Note All new scientific names used in this thesis have been accepted for publication in accordance with the requirements of Article 8.1 of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature (ICZN), 1999). Use of these names in this thesis does not imply prior publication or otherwise affect the availability of these names (Articles 8.2 and 8.3). ii TABLE OF CONTENTS SUMMARY............................................................................................................vii ACKNOWLEDGMENTS .......................................................................................ix ABBREVIATIONS AND NOMENCLATURE ......................................................xii ADDITIONAL ACKNOWLEDGEMENT FOR ON-LINE PDF VERSION .......... xvi CHAPTER 1. INTRODUCTION .............................................................................1 1.1. Historical and contemporary review .............................................................1 1.1.1. The genus Poltys .................................................................................1 1.1.2. Generic affiliations .............................................................................. 4 1.1.3. General biology ................................................................................... 6 1.1.4. Growth and development .....................................................................7 1.1.5. Araneid behavioural studies .................................................................7 1.2. Aims, rationale and general methodology .....................................................8 1.2.1. Aims ................................................................................................... 8 1.2.2. Rationale ............................................................................................. 9 1.2.3. General methodology ..........................................................................9 1.3. A comment on scope and fieldwork ...........................................................10 CHAPTER 2. TAXONOMY OF AUSTRALASIAN POLTYS ...............................12 2.1. Introduction, aims and a comment on methodology ....................................12 2.2. Material and methods ................................................................................. 13 2.2.1. General .............................................................................................. 13 2.2.2. Order of treatment of species .............................................................15 2.2.3. Measurements and descriptions .........................................................15 2.2.4. Distribution maps and records ...........................................................16 2.2.5. Abbreviations .................................................................................... 16 2.3. Taxonomy .................................................................................................. 17 2.3.1. Genus Poltys C.L. Koch ....................................................................17 2.3.2. Key to Australian Poltys species ........................................................ 28 2.3.3. The Poltys illepidus -group .................................................................34 2.3.3.1. Poltys illepidus C.L. Koch ........................................................ 35 2.3.3.2. Poltys stygius Thorell ................................................................49 2.3.4. The Poltys columnaris -group ............................................................. 54 2.3.4.1. Poltys jujorum sp. nov. ............................................................. 56 2.3.4.2. Poltys milledgei sp. nov. ........................................................... 67 2.3.5. The Poltys frenchi -group ...................................................................72 2.3.5.1. Poltys frenchi Hogg ..................................................................73 2.3.5.2. Poltys timmeh sp. nov. ..............................................................82 2.3.6. The Poltys laciniosus -group .............................................................. 86 2.3.6.1. Poltys laciniosus Keyserling...................................................... 87 iii 2.3.6.2. Poltys grayi sp. nov. .................................................................98 2.3.6.3. Poltys noblei sp. nov. .............................................................. 108 2.3.7. Australasian species currently considered nomina dubia .................. 114 2.4. Notes on Southeast Asian Poltys species .................................................. 115 CHAPTER 3. GENERIC RELATIONSHIPS ....................................................... 123 3.1. Introduction and aims ............................................................................... 123 3.2. Cladistics.................................................................................................. 126 3.2.1. A brief background to cladistics ..................................................... 126 3.2.2. Character polarity and outgroups .................................................... 127 3.2.3. Tree-building algorithms ................................................................. 128 3.2.4. Consensus trees .............................................................................. 132 3.2.5. Polytomies and zero length branches .............................................. 133 3.2.6. Tree support indices ........................................................................ 135 3.3. Methods ................................................................................................... 137 3.3.1. Taxa ................................................................................................ 137 3.3.2. Characters ....................................................................................... 140 3.3.3. Analysis .......................................................................................... 145 3.4. Results ..................................................................................................... 148 3.5. Discussion and outgroup selection for phylogenetic studies ...................... 151 CHAPTER 4. DNA SEQUENCING AND THE APPLICATION OF DNA DATA TO SPECIES SEPARATION PROBLEMS .............................................. 153 4.1. Introduction and aims ............................................................................... 153 4.2. Perceived problems in Poltys and the relevance of sequence differences in this context ............................................................................................... 156 4.3. Choice of sequences used in this study ..................................................... 158 4.4. Methods of DNA extraction and sequencing ............................................ 161 4.4.1. DNA extraction ............................................................................... 161 4.4.2. Primers and PCR ............................................................................. 162 4.4.3. Cycle sequencing ............................................................................ 163 4.4.4. Sequence editing and alignment ...................................................... 163 4.5. Methods for species separations ............................................................... 164 4.6. Results .................................................................................................... 165 4.6.1. The sequences ................................................................................. 165 4.6.2. Species separations .......................................................................... 166 4.7. Discussion and conclusions ...................................................................... 168 CHAPTER 5. CLADISTIC ANALYSIS—RELATIONSHIPS OF THE POLTYS LACINIOSUS -GROUP AND P. FRENCHI ............................................... 172 5.1. Introduction and aims ............................................................................... 172 5.2. Outgroups ............................................................................................... 172 5.3. The treatment of data from different sources ............................................ 173 5.4. Data and analyses—separate data sets ...................................................... 175 5.4.1. DNA methodology .......................................................................... 175 iv 5.4.2. Morphological data ......................................................................... 178 5.4.2.1. Methods .................................................................................. 178 5.4.2.2. The characters ......................................................................... 178 5.5. Data and analyses—combined data sets ................................................... 186 5.5.1. Preparation
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