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THE SYSTEMATICS AND BIOLOGY OF THE 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. 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 ...... 186 5.5.2. Analysis ...... 186 5.6. Results ...... 187 5.6.1. COI data ...... 187 5.6.2. ITS2 data ...... 189 5.6.3. Morphological data ...... 191 5.6.4. Combined data ...... 191 5.7. Discussion ...... 191

CHAPTER 6. FIELD STUDIES ...... 195

6.1. Introduction and aims ...... 195 6.2. Web-site tenacity ...... 196 6.2.1. Background ...... 196 6.2.2. Methods ...... 201 6.2.2.1. Pilot study ...... 201 6.2.2.2. The regular long-term transect ...... 203 6.2.2.3. Data analyses ...... 208 6.2.3. Results ...... 212 6.2.4. Discussion ...... 219 6.3. orientation with respect to camouflage and predators ...... 222 6.3.1. Background ...... 222 6.3.2. Aims ...... 227 6.3.3. Methods ...... 227 6.3.4. Results ...... 229 6.3.5. Discussion ...... 231

CHAPTER 7. GROWTH AND DEVELOPMENT DURING CAPTIVE REARING ...... 237

7.1. Introduction and aims ...... 237 7.2. Rearing in captivity ...... 238 7.2.1. Methods for captive rearing of Poltys ...... 238 7.2.1.1. Procedure ...... 238 7.2.1.2. The food mix ...... 241 7.3. Abdominal shapes and species status ...... 242 7.3.1. Methods ...... 243 7.3.2. Results ...... 243 7.3.2.1. Abdominal shapes of spiders from the same egg sac ...... 243 7.3.2.2. Confirmed pairings of NE Queensland males and females ...... 245 7.3.2.3. Epigyne variation in P. laciniosus ...... 246 7.3.3. Discussion ...... 247 7.4. An examination of life history through rearing data ...... 248 7.4.1. Methods ...... 251

v 7.4.2. Results ...... 253 7.4.2.1. Egg sac statistics ...... 253 7.4.2.2. Measuring growth ...... 257 7.4.2.3. The development of abdominal shape ...... 265 7.4.3. Discussion ...... 267

CHAPTER 8. GENERAL BIOLOGY & BEHAVIOUR ...... 269

8.1. Introduction and aims ...... 269 8.2. A review of the published information concerning Australian Poltys ...... 269 8.3. Observations arising from the present studies...... 271 8.3.1. Habitat ...... 271 8.3.2. Webs ...... 273 8.3.3. Diurnal concealment ...... 280 8.3.4. Prey and prey capture ...... 284 8.3.5. Dimorphism, courtship and mating ...... 289 8.3.6. Egg sacs ...... 294 8.3.7. Nocturnal predators and parasites ...... 296 8.3.8. Competition and other interactions ...... 298 8.3.9. Brief comments on the use of light and insect repellent during field studies ...... 300

CONCLUDING REMARKS ...... 302

REFERENCES ...... 306

APPENDICES ...... 332

Appendix 1. Generic characters ...... 332 Appendix 2. Details of specimens examined for generic analysis ...... 335 Appendix 3. Character attribute codings for newly added Poltyini taxa ...... 338 Appendix 4. Specimens sequenced and used for phylogenies...... 339 Appendix 5. COI sequences...... 341 Appendix 6. ITS2 sequences ...... 343 Appendix 7. Pairwise frequencies ...... 344 Appendix 8. Model parameters for ML...... 351 Appendix 9. Character codings for morphological characters...... 352 Appendix 10. Complete codings for combined analysis ...... 353 Appendix 11. Base data for Waitara Creek transect (pictorial) ...... 355 Appendix 12. Extracted tabular data for Waitara Creek transect...... 363 Appendix 13. Rest position survey—full data ...... 368 Appendix 14. Egg sac data...... 378 Appendix 15. Reared spiderlings: measurement data...... 380 Appendix 16. Reared spiderlings: basic data for other specimens...... 386 Appendix 17. Reared spiderlings: P. grayi moult dates ...... 388

vi SUMMARY

The genus Poltys C.L. Koch is revised for Australia and partly examined for the wider Australasian region. Five of the ten species originally described from Australia are found to be synonymous with South East Asian species, a further three are synonymous with an Australian species and four which were previously overlooked are newly described, resulting in a total of eight current species recorded from Australia.

Poltys coronatus Keyserling , P. keyserlingi Keyserling, P. multituberculatus Rainbow and P. penicillatus Rainbow are synonymised with P. illepidus C.L. Koch; P. microtuberculatus Rainbow is synonymised with P. stygius Thorell; P. bimaculatus

Keyserling, P. mammeatus Keyserling and P. salebrosus Rainbow are synonymised with P. laciniosus Keyserling; P. sigillatus Chrysanthus from New Guinea is synonymised with P. frenchi Hogg. Five new species are described, four from

Australia, P. grayi sp.nov., P. jujorum sp.nov. , P. milledgei sp.nov. and P. noblei sp.nov. , and P. timmeh sp.nov. from New Caledonia. A checklist of all Poltys types described from the region, including illustrations, is included.

The delimitation of the Australian species is aided and confirmed by DNA sequencing. Sequences from two genes and morphological characters are used to reconstruct a phylogeny of the Australian species.

The generic relationships of Poltys are examined in the context of the putative tribe Poltyini (Simon, 1895). No firm conclusions about the relationships of Poltys can be made, however the results indicate that the Poltyini is polyphyletic.

The results of field studies are presented; these indicate that P. noblei is less likely to move between web sites than diurnal taxa referenced from other studies.

Specimens were shown to sometimes occupy the same, or a closely adjacent web site,

vii for over eight months. Spiders most often move during spring and summer but often remain in the same site throughout winter. Specimens of Poltys noblei are also shown to be unevenly distributed on trees and bushes in respect to aspect and position. It is suggested that these observations indicate the importance of camouflage to deter wasp and bird predators.

Specimens of several Poltys species were reared from egg sacs, confirming male–female identification and showing the variation in abdominal shape between siblings. Growth data indicate that sibling males and females cannot normally interbreed; males mature after 2–4 moults, females after 8–11 moults.

Field and cage observations of general aspects of Poltys biology are presented including preferred habitat, prey capture and handling, courtship and mating, competition and web construction.

viii ACKNOWLEDGMENTS

A ‘cast of thousands’ has been involved at some point or other over the extended period of this study, some people in several different ways; to everyone involved, I extend the most sincere thanks. I wish to emphasize that any mistakes, omissions and other problems which may have crept into this work are all my own doing—everyone who has provided me with advice or assistance has done so in good faith and I take full responsibility for any misapplication of this help. I hope I have not overlooked anyone in the list below; sincere apologies for any omissions.

Firstly, none of this would have been possible without the extended and wonderfully knowledgeable support of my supervisors, Mike Gray at the Australian

Museum and Harley Rose, at the University of Sydney; both have earned a well deserved rest from the barrage of manuscripts, questions and occasional crises.

The heart of any taxonomic project is the specimens and their data. This goes hand in hand with a wealth of knowledge about the collections and their history that is held by Curators, Collection Managers and Research Scientists. I am indebted to all these people who have been so helpful to me whilst I have been searching for specimens and information. So, for hosting visits, arranging loans of specimens and for information on collections, I thank: Graham Milledge and Mike Gray (AM); Bruce

Halliday (ANIC); A. Barrion; Janet Beccaloni and Paul Hillyard (BMNH); Tino

Gonsalves and Alistair Ramsdale (BPBM); Christa Deeleman-Reinhold; Manfred

Grasshoff (Frankfurt); Tamás Szûts (HNHM); Joseph Koh; Herbert Levi (MCZ),

Margaret Humphrey (MMUS); Christine Rollard (MNHNP); Rudy Jocqué (MRAC);

Giuliano Doria (MSNG); John Murphy; Ansie Dippenaar-Schoeman & Elizabeth

ix Kassimatis (NCAP); Jürgen Gruber (NHMW); Torbjorn Kronestedt (NHRM); Peter

Lillywhite and Ken Walker (NMV); Graham Brown and Gavin Dally (NTM); James

Hogan and G. McGavin (OUM); Val Davies, Phil Lawless, Robert Raven and Owen

Seeman (QM); Tim Kingston (QVM Launceston), N. Sivasothi (Raffles Mus.); Domir

De Bakker (RBIN); Kees van den Berg and Erik van Nieukerken (RMNH); David Hirst

(SAMA); Fernando Alvarez and César Durán-Barrón (UNAM), Mark Harvey and

Julianne Waldock (WAM); Wanda Wesolowska (Wroclaw); Jason Dunlop (ZMB); H.

Dastych (ZMH); Nikolaj Scharff (ZMUC); Marek Zabka.

There are many other facets to what turned into quite a broad study and I wish to thank the following for many and various things: Kate Dempsey and Alberto Daglio for helping to bridge a communications divide; Gerry Cassis, Sally Cowan, Greta Gray,

Mike Gray, Graham Milledge, Tim Moulds, John Noble, John Olive, Judy Thompson,

Doug Wallace and Jenni Webber for assistance in collecting and rearing specimens;

Herbert Levi & Matjaz Kuntner for making available various drawings and notes; Akio

Tanikawa for supplying Japanese reference material; Marek Zabka for sending me coquille board; Shane Ahyong, Max Moulds and Greg Edgecomb for assistance with phylogenetics; Carl Bento, Mike Gray, David Hain and Joseph Koh for photographs and information; Bruce Henderson for information on N.C. Coleman; Trevor

Reddicliffe for access to N.C. Coleman’s old property at Edmonton; David McAlpine for identifying parasitic flies; Sue Gibbs, Emma Gray, Klaus Heil and Pam Mudford for translations; Sam Fenton, Anina Hainsworth, Leone Lemmer, Fiona Simpson and other library staff for assistance with references, and Pam Mudford for photocopying and other library assistance; Miquel Arnedo for advice on sequences for analysis; all the past and present EBU staff who have helped with different parts of the project: Sarah

x Brown, Don Colgan, Rebecca Johnson, Denis O’Meally and Deidre Sharkey, and Sue

Lindsay for help preparing specimens and taking SEM images; Todd Blackledge, Roz and Peter Cusack, Greg Edgecomb,Volker Framenau, Mariella Herberstein, Rebecca

Johnson, Matjaz Kuntner, Nikolaj Scharff and my supervisors for advice, proof reading and much other assistance with various manuscripts and thesis chapters; Margaret

Humphrey for showing me how to prepare chromosome preps and providing an introduction to electrophoresis (neither of which I used subsequently!).

In personal matters, I sincerely thank my long neglected family & friends in

Europe and Australia for putting up with infrequent visits fitted around conferences, specimens and museum visits; my former partner for his encouragement and support at the commencement of the project; my colleague & present partner Graham Milledge who was always prepared to help in the field, provided home support and constant encouragement, and without (much) complaint put up with the disruptive schedule necessitated by keeping odd hours and raising spiders at home (which sometimes made the ensuite bathroom “out of bounds” at night for weeks on end).

This project was generously supported by provision of funding and equipment by The Australian Museum and The University of Sydney, a Norman Scott Noble

Scholarship administered by The University of Sydney, and two awards from the Joyce

W. Vickery Scientific Research Fund administered through the Linnean Society of

New South Wales. Attendance at the 16 th International Congress of Arachnology in

Gent, Belgium was supported by assistance from the Congress Organising Committee.

I would also like to thank the conservation and forestry bodies of New South Wales,

Northern Territory, Queensland, South Australia and Western Australia for permission to collect in protected areas.

xi ABBREVIATIONS AND NOMENCLATURE

Abbreviations

The following abbreviations are used for repository institutions and private

collections:

AM: Australian Museum, Sydney, MSNG: Museo Civico di Storia

Australia; Naturale “G. Doria”, Genoa,

BMNH: The Natural History Museum, Italy;

London, UK; NCAP: National Collection of

BPBM: Bernice P. Bishop Museum, Arachnida, Pretoria, South

Honolulu, USA; Africa;

CLD: C.L. Deeleman-Reinhold, The NHMW: Naturhistorisches Museum,

Netherlands; Wien, Austria;

HNHM: Hungarian Natural History NHRM: Swedish Museum of Natural

Museum, Budapest, Hungary; History, Stockholm, Sweden;

JAM: J. Murphy, UK; NMV: Museum Victoria, Melbourne,

JK: J. Koh, Singapore; Australia;

MMUS: Macleay Museum, University NTM: Museums and Art Galleries of

of Sydney, Sydney, Australia; the Northern Territory, Darwin,

MNHNP: Muséum National d’Histoire Australia;

Naturelle, Paris, France; OUM: Oxford University Museum,

MRAC: Koninklijk Museum voor Oxford, UK;

Midden Afrika, Tervuren, QM: Queensland Museum, Brisbane,

Belgium; Australia;

xii RBIN: Royal Belgian Institute of WAM: Western Australian Museum,

Natural Sciences, Brussels, Perth, Australia;

Belgium; ZMB: Museum für Naturkunde,

RMNH: National Museum of Natural Zentralinstitut der Humboldt-

History, Leiden, The Universität, Berlin, Germany;

Netherlands; ZMH: Zoologisches Institut und

SAMA: South Australian Museum, Zoologisches Museum der

Adelaide, Australia; Universität Hamburg, Hamburg,

UNAM, Instituto de Biología, Germany;

Universidad Nacional ZMUC: Universitets København,

Autónoma de México, Mexico Zoologisk Museum,

D.F., Mexico; Copenhagen, Denmark.

Within Australian museums, specimen numbers prefixed KS are AM, N or

NN are SAMA, S or W are QM, WA are WAM, others are indicated.

The following abbreviations are used in collection records:

G,M&S: Gray, Milledge and Smith; MRG: M.R. Gray;

HMS: H.M. Smith; NP: National Park;

M&S: Milledge and Smith; SF: State Forest.

Collector’s names which only appear once or twice are not abbreviated.

xiii The nomenclature and most abbreviations of male palpal sclerites follow

Coddington (1990). The following abbreviations for morphological features, molecular terms and miscellaneous items are used in the tables, text and figures

(molecular reagents are not included) ac: aciniform spigot(s); fl: flagelliform spigot; agg: aggregate spigot(s); I, II, III, IV: Roman numerals used in

ALE: anterior lateral eye(s); reference to legs 1–4;

ALS: anterior lateral spinneret(s); ITS2: internal transcribed spacer 2;

AME: anterior median eye(s); LLs and LsLS: large-large-small and

BH: basal haematodocha; large-small-large-small bp: base pair; (prolateral cheliceral teeth);

CD: copulation duct; M0–Mx: moults 0 (normally pre-

CL: carapace length; emergence from egg sac)

Co: conductor; onwards;

COI: cytochrome oxidase c subunit I; MA: median apophysis;

CY: cymbium; mAP: minor ampullate spigot; cyl: cylindrical spigot(s); MAP: major ampullate spigot; df: degrees of freedom; MEA: metine embolic apophysis;

DH: distal haematodocha; ML: maximum likelihood;

DNA: deoxyribonucleic acid (bases A, MP: maximum parsimony

adenine; C, cytosine; G, mtDNA: mitochondrial DNA;

guanine; T, thymine); N: number of samples;

E: embolus; NSW: New South Wales;

EBU: Evolutionary Biology Unit (AM) NT: Northern Territory;

FD: fertilisation duct; nu: nubbin;

xiv P: pocket; S: stipes;

P+TL: patella and tibia overall SA: South Australia;

combined length, dorsal; SD: standard deviation;

P+TI: as P+TL specifically for leg I SDu: sperm duct;

(Ch. 7); SEM: scanning electron microscopy or

Pc: paracymbium; microscope; pir: piriform spigot(s); Sp: spermatheca;

PCR: polymerase chain reaction s/syn: senior synonym;

PLE: posterior lateral eye(s); ST: sub tegulum

PLS: posterior lateral spinneret(s); STA: sub-terminal apophysis;

PM: paramedian apophysis; Syn.: synonymised;

PME: posterior median eye(s); T: tegulum;

PMS: posterior median spinneret(s); TA: terminal apophysis;

PNG: Papua New Guinea; TL: tegular lobe;

Qld: Queensland; UV: ultra-violet (radiation);

R: radix; WA: Western Australia

Nomenclature

For spiders, current scientific names and their authors have been sourced from

Platnick (2006). For all Poltys species discussed (the primary taxon dealt with), citation of scientific names and their authors follows Recommendation 51A of the

International Code of Zoological Nomenclature (International Commission on

Zoological Nomenclature (ICZN), 1999). For these taxa the full author citation of name and date are given at least once; for non-Australian species this is in the

xv Checklist (Table 2.1). Authors of taxa other than Poltys are cited only by name on the first usage of that taxon name. Authors of plant names are not included.

ADDITIONAL ACKNOWLEDGEMENT FOR ON-LINE PDF VERSION

I am grateful to The Bulletin of the British Arachnological Society, The

Journal of Arachnology and The Records of the Australian Museum for permission to include work published in the following:

Smith, H.M. (2003). Shape variation in Australian Poltys species (Araneae:

Araneidae). Bulletin of the British Arachnological Society 12: 355–

360.

Smith, H.M. (2005). A preliminary study of the relationships of taxa included

in the tribe Poltyini (Araneae, Araneidae). Journal of Arachnology 33:

468–481.

Smith, H.M. (2006). A revision of the genus Poltys in Australasia (Araneae:

Araneidae). Records of the Australian Museum 58: 43–96.

Links to these publications, where available, are given in the bibliography.

xvi