Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography
RESEARCH ARTICLE

The systematics and biology of the spider genus Nephila (Araneae : Nephilidae) in the Australasian region

Mark S. Harvey A D , Andrew D. Austin B and Mark Adams C
+ Author Affiliations
- Author Affiliations

A Department of Terrestrial Invertebrates, Western Australian Museum, Locked Bag 49, Welshpool DC, WA 6986, Australia.

B Australian Centre for Evolutionary Biology and Biodiversity, School of Earth & Environmental Sciences, The University of Adelaide, SA 5005, Australia.

C Evolutionary Biology Unit, South Australian Museum, North Terrace, Adelaide, SA 5000, Australia.

D Corresponding author. Email: mark.harvey@museum.wa.gov.au

Invertebrate Systematics 21(5) 407-451 https://doi.org/10.1071/IS05016
Submitted: 20 April 2005  Accepted: 10 July 2007   Published: 7 November 2007

Abstract

Five species of the nephilid genus Nephila Leach are found in the Australasian region, which for the purposes of this study was defined as Australia and its dependencies (including Lord Howe I., Norfolk I., Christmas I., Cocos (Keeling) Is), New Guinea (including Papua New Guinea and the Indonesian province of West Papua), Solomon Is, Vanuatu, New Caledonia, Fiji, Tonga, Niue, New Zealand and other parts of the south-west Pacific region. All species are redescribed and illustrated. Nephila pilipes (Fabricius) occurs in the closed forests of eastern and northern Australia, New Guinea, Solomon Is and Vanuatu (through to South-East Asia); N. plumipes (Latreille) is found in Australia (including Lord Howe I. and Norfolk I.), New Guinea, Vanuatu, Solomon Is and New Caledonia; N. tetragnathoides (Walckenaer) inhabits Fiji, Tonga and Niue; N. antipodiana (Walckenaer) occurs in northern Australia (as well as Christmas I.), New Guinea and Solomon Is (through to South-East Asia); and N. edulis (Labillardière) is found in Australia (including Cocos (Keeling) Is), New Guinea, New Zealand and New Caledonia. Epeira (Nephila) walckenaeri Doleschall, E. (N.) hasseltii Doleschall, N. maculata var. annulipes Thorell, N. maculata jalorensis Simon, N. maculata var. novae-guineae Strand, N. pictithorax Kulczyński, N. maculata var. flavornata Merian, N. pictithorax Kulczyński, N. maculata var. flavornata Merian, N. maculata piscatorum de Vis, and N. (N.) maculata var. lauterbachi Dahl are proposed as new synonyms of N. pilipes. Nephila imperialis var. novaemecklenburgiae Strand, N. ambigua Kulczyński, N. sarasinorum Merian and N. celebesiana Strand are proposed as new synonyms of N. antipodiana. Meta aerea Hogg, N. meridionalis Hogg, N. adelaidensis Hogg and N. meridionalis hermitis Hogg are proposed as new synonyms of N. edulis. Nephila picta Rainbow is removed from the synonymy of N. plumipes and treated as a synonym of N. edulis, and N. nigritarsis insulicola Pocock is removed from the synonymy of N. plumipes and treated as a synonym of N. antipodiana. Allozyme data demonstrate that N. pilipes is distinct at the 80% FD level from N. edulis, N. plumipes and N. tetragnathoides. Nephila plumipes and N. tetragnathoides, deemed to represent sister-taxa owing to the shared presence of a triangular protrusion of the male pedipalpal conductor, were found to differ at 15% FD in the genetic study. No genetic differentiation was found between 10 populations of N. edulis sampled across mainland Australia. Species of the genus Nephila have been extensively used in ecological and behavioural studies, and the biology of Nephila species in the Australasian region is extensively reviewed and compared with studies on Nephila species from other regions of the world.


Acknowledgements

Numerous people have generously supplied much of the material utilised in this study. In particular, we wish to thank those people who kindly obtained at our request the live material utilised in the allozyme analysis: Anthony Desmond, Mark Elgar, Gordon Graham, Bill Humphreys, Jane Rienks, Erich Volschenk, Julianne Waldock and Doug Wallace. Funds were made available for the collection of material by the following organisations: the Australian Biological Resources Study; National Geographic Society, the American Museum of Natural History and the Queensland Museum, facilitated by Dr N. I. Platnick and Dr R. J. Raven, respectively; Adelaide University and the Australian Entomological Society for a research grant that enabled the collection of live material from Uluru.


We wish to thank the many curators who allowed access to the collections in their care: G. Brown (NTMAG), C. E. Griswold (CAS), H. Dastych (ZMH), J. W. Early (AMNZ), M. R. Gray, R. Harris and G. Milledge (AM), G. Hall (NZAC), R. B. Halliday (ANIC), A. Hänggi and J. Ineichen-Reidi (NMB), P. D. Hillyard (BMNH), D. Hirst (SAM), T. Huflejt (MZPW), T. Kronestedt (SMNH), S. Larcher (USNM), L. Leibensperger and H. W. Levi (MCZ), G. Milledge (MV), M. Moritz (ZMB), E. J. Nieukerken (RMNH), N. I. Platnick (AMNH), R. J. Raven (QM), C. Rollard (MNHN), U. Schreiber (SMF), P. Sirvid and L. J. Boutin (MNZ), S. Swift (BPBM) and E. Turner (TMAG). Permission to export specimens from Fiji was graciously granted by Mr J. Vakabua (Ministry of Agriculture, Fisheries and Forests, Fiji). Volker Framenau and Michael Rix kindly allowed us to publish their photographs of Nephila. We also thank Carolyn Horne, Malcolm Krieg and Julianne Waldock for technical assistance, Bradley Durrant for the habitus drawings, and Matjaž Kuntner, Volker Framenau, Gustavo Hormiga and Mark Elgar for their extremely helpful comments on a draft of the manuscript.


References


Adams A. (1847) Notes on the habits of certain exotic spiders. Annals and Magazine of Natural History 20, 289–297.

Adams M., Baverstock P. R., Watts C. H. S., Reardon T. (1987) Electrophoretic resolution of species boundaries in Australian Microchiroptera: I. Eptesicus (Chiroptera: Vespertilionidae). Australian Journal of Biological Sciences 40, 143–162.

Alvarez-Padilla F. (2007) Systematics of the spider genus Metabus O. P-Cambridge, 1899 (Araneoidea: Tetragnathidae) with additions to the tetragnathid fauna of Chile and comments on the phylogeny of Tetragnathidae. Zoological Journal of the Linnean Society 151, 285–335.
CrossRef |

Archer A. F. (1958) Studies in the orbweaving spiders (Argiopidae). 4. American Museum Novitates 1922, 1–21.

Austin A. D. (1977) A note on the life-history of Anatrachyntis terminella (Walker) (Lepidoptera: Cosmopterigidae), whose larvae are predaceous on the eggs of Nephila edulis (Koch) (Araneae: Araneidae). Journal of the Australian Entomological Society 16, 427–428.
CrossRef |

Austin A. D. (1985) The function of spider egg sacs in relation to parasitoids and predators, with special reference to the Australian fauna. Journal of Natural History 19, 359–376.
CrossRef |

Austin A. D., Anderson D. T. (1978) Reproduction and development of the spider Nephila edulis (Koch) (Araneidae: Araneae). Australian Journal of Zoology 26, 501–518.
CrossRef |

Benoit P. L. G. (1962) Les Araneidae-Nephilinae africains. Revue de Zoologie et de Botanique Africaines 65, 217–231.

Berland L. (1931) Sur quelques Araignées envoyées de Nouvelle-Calédonie par M. Risbec. Bulletin du Muséum National d’Histoire Naturelle, Série 2 3, 666–672.

Bjorkman-Chiswell B., Kulinski M. M., Muscat R. L., Nguyen K. A., Norton B. A., Symonds M. R. E., Westhorpe G. E., Elgar M. A. (2004) Web-building spiders attract prey by storing decay matter. Die Naturwissenschaften 91, 245–248.
CrossRef | PubMed |

Blackwall J. (1833) Characters of some undescribed genera and species of Araneidae. London, Edinburgh and Dublin Philosophical Magazine and Journal of Science (3) 3, 104–443.

Blackwall J. (1864) Description of seven new species of East-Indian spiders received from the Rev. O. P. Cambridge. Annals and Magazine of Natural History, Series 3 14, 36–45.

Blackwall J. (1865) Descriptions of recently discovered species, and characters of a new genus, of Araneidea from the east of central Afric. Annals and Magazine of Natural History, Series 3 16, 336–352.

Blackwall J. (1877) A list of spiders captured in the Seychelle Islands by Professor E. Percival Wright, M.D., F.L.S.; with descriptions of species supposed to be new to arachnologists. Proceedings of the Royal Irish Academy. Section B: Biological, Geological, and Chemical Science, Series 2 3, 1–22.

Bonnet P. (1958). ‘Bibliographia Araneorum, Vol. 2, Number 4.’ (Douladoure: Toulouse, France.)

Bristowe W. S. (1932) Insects and other invertebrates for human consumption in Siam. Transactions of the Entomological Society of London 80, 387–404.

Brown S. G., Christenson T. E. (1983) The relationships between web parameters and spiderling predatory behavior in the orb-weaver, Nephila clavipes. Zeitschrift fur Tierpsychologie 63, 241–250.

Bruce M. J. (2006) Silk decorations: controversy and consensus. Journal of Zoology 269, 89–97.
CrossRef |

Brunet B. (1994). ‘The Silken Web: A Natural History of Australian Spiders.’ (Reed Books: Sydney Australia.)

Butler A. G. (1879) On Arachnida from the Mascarene Islands and Madagascar. Proceedings of the Zoological Society of London 1879, 729–734.

Champion de Crespigny F. E., Herbstein M. E., Elgar M. A. (2001) Food caching in orb-web spiders (Araneae: Araneoidea). Die Naturwissenschaften 88, 42–45.
CrossRef | PubMed |

Christenson T. E., Goist K. C. (1979) Cost and benefits of male-male competition in the orb weaving spider Nephila clavipes. Behavioral Ecology and Sociobiology 5, 87–92.
CrossRef |

Christenson T. E., Wenzl P. A. (1980) Egg-laying of the golden silk spider, Nephila clavipes L. (Araneae, Araneidae): functional analysis of the egg sac. Animal Behaviour 28, 1110–1118.
CrossRef |

Christenson T. E., Wenzl P. A., Legum P. (1979) Seasonal variation in egg hatching and certain egg parameters of the golden silk spider Nephila clavipes (Araneidae). Psyche 86, 137–147.

Christenson T. E., Brown S. G., Wenzl P. A., Hill E. M., Goist K. C. (1985) Mating behavior of the golden orb-weaving spider Nephila clavipes 1. Female receptivity and male courtship. Journal of Comparative Psychology (Washington, D.C.: 1983) 99, 160–166.
CrossRef |

Chrysanthus F. (1959) Spiders from South New Guinea II. Nova Guinea (n.s.) 10, 197–206.

Chrysanthus F. (1960) Spiders from South New Guinea III. Nova Guinea (n.s.) 3, 23–42.

Chrysanthus F. (1971) Further notes on the spiders of New Guinea I (Argyopidae). Zoologische Verhandelingen 113, 1–52.

Clyne D. (1969). ‘A Guide to Australian Spiders.’ (Nelson: Melbourne, Australia.)

Coddington J. A. (1990) Ontogeny and homology in the male palpus of orb-weaving spiders and their relatives, with comments on phylogeny (Araneoclada: Araneoidea, Deinopoidea). Smithsonian Contributions to Zoology 496, 1–52.

Coddington J. A., Hormiga G., Scharff N. (1997) Giant female or dwarf male spiders? Nature 385, 687–688.
CrossRef |

Craig C. L. (2003). ‘Spiderwebs and Silk: Tracing Evolution from Molecules to Genes to Phenotypes.’ (Oxford University Press: New York, USA.)

Dahl H. (1911) Die Verbreitung der Spinnen spricht gegen ein frühere Landverbindung der Südspitzen unsrer Kontinente. Zoologischer Anzeiger 37, 270–282.

Dahl H. (1912) Seidenspinne und Spinnenseide. Mitteilung aus dem Zoologischen Museum in Berlin 6, 1–90.

de Vis C. W. (1911) A fisherman’s spider. Annals of the Queensland Museum 10, 167–168.

Doleschall C. L. (1857) Bijdrage tot de Kennis der Arachniden van den Indischen Archipel. Natuurkundig Tijdschrift voor Nederlandsch-Indië 13, 399–434.

Doleschall C. L. (1859) Tweede Bijdrage tot de Kennis der Arachniden van den Indischen Archipel. Acta Societatis Scientarium Indo-Neerlandicae 5, 1–60.

Dondale C. D. (1966) The spider fauna (Araneida) of deciduous orchards in the Australian Capital Territory. Australian Journal of Zoology 14, 1157–1192.
CrossRef |

Eisner T., Nowicki S. (1983) Spider web protection through visual advertisement: role of the stabilimenta. Science 219, 185–187.
CrossRef | PubMed |

Elgar M. A. (1989) Kleptoparasitism: a cost of aggregating for an orb-weaving spider. Animal Behaviour 37, 1052–1055.
CrossRef |

Elgar M. A. (1993) Inter-specific associations involving spiders: kleptoparasitism, mimicry and mutualism. Memoirs of the Queensland Museum 33, 411–430.

Elgar M. A. (1998). Sexual selection and sperm competition in arachnids.In ‘Sperm Competition and Sexual Selection’. (Eds T. R. Birkhead andA. P. Møller.) pp. 307–337. (Academic Press: San Diego, CA, USA.)

Elgar M. A., Fahey B. F. (1996) Sexual cannibalism, competition, and size dimorphism in the orb-weaving spider Nephila plumipes Latreille (Araneae: Araneoidea). Behavioral Ecology 7, 195–198.
CrossRef |

Elgar M. A., Schneider M. J. (2004) Kleptoparasitism: a cost of aggregating for an orb-weaving spider. Advances in the Study of Behavior 34, 135–163.

Elgar M. A., Bruce M. J., Champion de Crespigny F. E., Cutler A. R., Cutler C. L., Gaskett A. C., Herberstein M. E., Ramamurthy S., Schneider J. M. (2003a) Male mate choice and pattern of paternity in the polyandrous, sexually cannibalistic orb-web spider Nephila plumipes. Australian Journal of Zoology 51, 357–365.
CrossRef |

Elgar M. E., Champion de Crespigny F., Ramamurthy S. (2003b) Male copulation behaviour and the risk of sperm competition. Animal Behaviour 66, 211–216.
CrossRef |

Fabricius J. C. (1781). ‘Species Insectorum Exhibentes Eorum Differentias Specificas, Synonyma Auctorum, Loca Natalia, Metamorphosin Adiectis Observationibus, Descriptionibus.’ Vol. 1. (C. E. Bohnii: Hamburgi, Germany.)

Fabricius J. C. (1793). ‘Entomologia Systematica Emendata et Aucta. Secundum Classes, Ordines, Genera, Species Adjectis Synonimis, Locis, Obervationibus, Descriptionibus.’ Vol. 2. (C. G. Proft: Hafniae (Copenhagen, Denmark).)

Farr J. A. (1976) Social behavior of the golden silk spider, Nephila clavipes (Linnaeus) (Araneae, Araneidae). The Journal of Arachnology 4, 137–144.

Felsenstein J. (1993). ‘PHYLIP (Phylogeny Inference Package. Version 3.5c).’ (Distributed by the author: University of Washington, Seattle, WA, USA.)

Fincke O. M., Higgins L., Rojas E. (1990) Parasitism of Nephila clavipes (Araneae, Tetragnathidae) by an ichneumonid (Hymenoptera, Polysphinctini) in Panama. The Journal of Arachnology 18, 321–330.

Fromhage L., Schneider J. M. (2004) Safer sex with feeding females: sexual conflict in a cannibalistic spider. Behavioral Ecology 16, 377–382.
CrossRef |

Gerstäcker A. (1873). Gliedertierfauna des Sansibargebietes. Arachnoidea. In ‘Reisen in Ost-afrika in den Jahren 1859–1865.’ Vol. 3. (Ed. C. C. von der Decken.) pp. 461–503. (C. F. Winter: Leipzig, Germany.)

Green A. (1981) Golden orb weaver spiders (Nephila sp.) in Tasmania. Tasmanian Naturalist 67, 12–13.

Griffiths B. V., Holwell G. I., Herberstein M. E., Elgar M. A. (2003) Frequency, composition and variation in external food stores constructed by orb-web spiders: Nephila edulis and Nephila plumipes (Araneae: Araneoidea). Australian Journal of Zoology 51, 119–128.
CrossRef |

Griswold C. E., Coddington J. A., Hormiga G., Scharff N. (1998) Phylogeny of the orb-web building spiders (Araneae, Orbiculariae: Deinopoidea, Araneoidea). Zoological Journal of the Linnean Society 123, 1–99.
CrossRef |

Grostal P., Walter D. E. (1997) Kleptoparasites or commensals? Effects of Argyrodes antipodianus (Araneae: Theridiidae) on Nephila plumipes (Araneae: Tetragnathidae). Oecologia 111, 570–574.
CrossRef |

Herberstein M. E., Elgar M. A. (1994) Foraging strategies of Eriophora transmarina and Nephila plumipes (Araneae: Araneoidea): Nocturnal and diurnal orb-weaving spiders. Australian Journal of Ecology 19, 451–457.
CrossRef |

Herberstein M. E., Craig C. L., Coddington J. A., Elgar M. A. (2000) The functional significance of silk decorations of orb-web spiders: a critical review of the imperical evidence. Biological Reviews 75, 649–669.
PubMed |


Herberstein M. E., Schneider J. M., Elgar M. A. (2002) Costs of courtship and mating in a sexually cannibalistic orb-web spider: female mating strategies and their consequences for males. Behavioral Ecology and Sociobiology 51, 440–446.
CrossRef |

Higgins L. E. (1987) Time budget and prey of Nephila clavipes Linnaeus (Araneae, Araneidae) in southern Texas, USA. The Journal of Arachnology 15, 401–418.

Higgins L. E. (1992a) Developmental changes in barrier web structure under different levels of predation risk in Nephila clavipes (Araneae, Tetragnathidae). Journal of Insect Behavior 5, 635–655.
CrossRef |

Higgins L. E. (1992b) Developmental plasticity and fecundity in the orb-weaving spider Nephila clavipes. The Journal of Arachnology 20, 94–106.

Hill E. M., Christenson T. (1981) Effects of prey characteristics and web structure on feeding and predatory responses of Nephila clavipes spiderlings. Behavioral Ecology and Sociobiology 8, 1–5.
CrossRef |

Hill E. M., Christenson T. (1988) Male residency on juvenile female Nephila clavipes (Araneae, Araneidae) webs. The Journal of Arachnology 16, 257–259.

Hingston R. W. G. (1922a) The snare of the Giant Wood Spider (Nephila maculata), part I. Journal of the Bombay Natural History Society 28, 642–649.

Hingston R. W. G. (1922b) The snare of the giant wood spider (Nephila maculata), part II: the continuation of the snare. Journal of the Bombay Natural History Society 28, 911–917.

Hingston R. W. G. (1922c) The snare of the giant wood spider (Nephila maculata), part III: further lessons of the Nephila. Journal of the Bombay Natural History Society 28, 917–923.

Hingston R. W. G. (1923) The giant wood spider (Nephila maculata). Journal of the Bombay Natural History Society 29, 70–76.

Hirst A. S. (1909) Arachnida and Myriapoda. Proceedings of the Zoological Society of London 1909, 132–160.

Hodge M. A., Uetz G. W. (1992) Antipredator benefits of single and mixed-species grouping by Nephila clavipes L. (Araneae, Tetragnathidae). The Journal of Arachnology 20, 212–216.

Hogg H. R. (1896). Araneidae. In ‘Report on the Work of the Horn Scientific Expedition to Central Australia, Part II. Zoology’. (Ed. B. Spencer.) pp. 309–356. (Dulau and Co.: London, UK.)

Hogg H. R. (1910) Two new Nephilae from South Australia. Transactions and Proceedings of the Royal Society of South Australia 34, 59–62.

Hogg H. R. (1914) Spiders from the Montebello Islands. Proceedings of the Zoological Society of London 1914(1), 69–92.

Hormiga G., Eberhard W. G., Coddington J. A. (1995) Web construction behaviour in Australian Phonognatha and the phylogeny of nephiline and tetragnathid spiders (Araneae: Tetragnathidae). Australian Journal of Zoology 43, 313–364.
CrossRef |

Hormiga G., Scharff N., Coddington J. A. (2000) The phylogenetic basis of sexual size dimorphism in orb-weaving spiders (Araneae, Orbiculariae). Systematic Biology 49, 435–462.
CrossRef | PubMed |

Humphreys W. F. (1993) Criteria for identifying thermal behaviour in spiders: a low technology approach. Memoirs of the Queensland Museum 33, 543–550.

Karsch F. (1879) West-afrikanische Myriapoden und Arachniden. Zeitschrift für die Gesammten Naturwissenschaften 52, 825–837.

Keyserling E. (1887). Nachtrage. In ‘Die Arachniden Australiens’. Vol. 2. (Eds L. Koch and E. Keyserling.) pp. 153–232. (Bauer und Raspe: Nürnberg, Germany.)

King P. P. (1827). ‘Narrative of a Survey of the Intertropical and Western coasts of Australia. Performed Between the Years 1818 and 1822.’ Vol. 1–2. (J. Murray: London, UK.)

Klärner D., Barth F. G. (1982) Vibratory signals and prey capture in orb weaving spiders (Zygiella x-notata, Nephila clavipes; Araneidae). Journal of Comparative Physiology. A, Sensory, Neural, and Behavioral Physiology 148, 445–456.
CrossRef |

Koch C. L. (1839). ‘Die Arachniden. Getreu nach der Natur Abgebildet und Beschrieben.’ Vol. 6. (C. H. Zeh’schen: Nürnberg, Germany.)

Koch L. (1867) Beschreibungen neuer Arachniden und Myriapoden. Verhandlungen der K. K. Zoologischen-Botanischen Gesellschaft in Wien 17, 173–250.

Koch L. (1872). ‘Die Arachniden Australiens, Vol. 1.’ pp. 105–368. (Bauer und Raspe: Nürnberg, Germany.)

Koch L. (1878) Japanesische Arachniden und Myriapoden. Verhandlungen der K. K. Zoologischen-Botanischen Gesellschaft in Wien 27, 735–798.

Koh J. (1989). ‘Guide to Common Singapore Spiders.’ (Singapore Science Centre: Singapore.)

Krakauer T. (1972) Thermal responses of the orb-weaving spider Nephila clavipes (Araneae: Argiopidae). American Midland Naturalist 88, 245–250.
CrossRef |

Kulczyński W. (1911). Spinnen aus Nord-Neu-Guinea. In ‘Nova Guinea. Résultats de l’expedition scientifique Néerlandaise à la Nouvelle Guinée en 1903 sous les auspices d’Arthur Wichmann. Vol. 3’. pp. 423–518. (E. J. Brill: Leiden, The Netherlands.)

Kuntner M. (2002) The placement of Perilla (Araneae, Araneidae) with comments on araneid phylogeny. The Journal of Arachnology 30, 281–287.
CrossRef |

Kuntner M. (2005) A revision of Herennia (Araneae: Nephilidae: Nephilinae), the Australian ‘coin’ spiders. Invertebrate Systematics 19, 391–436.
CrossRef |

Kuntner M. (2006) Phylogenetic systematics of the Gondwanan nephilid spider lineage Clitaetrinae (Araneae, Nephilidae). Zoologica Scripta 35, 19–62.
CrossRef |

Kuntner M. (2007) A monograph of Nephilengys, the pantropical ‘hermit spiders’ (Araneae, Nephilidae, Nephilinae). Systematic Entomology 32, 95–135.
CrossRef |

Kuntner M., Hormiga G. (2002) The African spider genus Singafrotypa (Araneae, Araneidae). The Journal of Arachnology 30, 129–139.
CrossRef |

Kuntner M., Coddington J. A., Hormiga G. (in press) Phylogeny ofextant nephilid orb-weaving spiders (Araneae, Nephilidae): testingmorphological and ethological homologies. Cladistics ,

Labillardière J. (1799). ‘Relation du Voyage à la Recherche de la Pérouse, fait par Ordre de l’Assemblée Constituante. Vol. 2.’ (H. J. Jansen: Paris, France.)

Latreille P. A. (1804). ‘Histoire Naturelle, Generale et Particulière, des Crustacés et des Insectes. Vol. 7.’ (F. Dufart: Paris, France.)

Leach W. E. (1815). ‘The Zoological Miscellany; being Descriptions of New or Interesting Animals. Vol. 2.’ (Nodder: London, UK.)

Levi H. W. (1980) The orb-weaver genus Mecynogea, the subfamily Metinae and the genera Pachygnatha, Glenognatha and Azilia of the subfamily Tetragnathinae north of Mexico (Araneae: Araneidae). Bulletin of the Museum of Comparative Zoology 149, 1–75.

Levi H. W., von Eickstedt V. R. D. (1989) The Nephilinae spiders of the neotropics (Araneae: Tetragnathidae). Memorias do Instituto Butantan 51, 43–56.

Lucas H. (1858). Aptères. In ‘Archives Entomologiques. Vol. 2’. (Ed. M. J. Thomson.) pp. 377–445. (Bureau du Trésorier de la Société Entomologique de France: Paris, France.)

Macleay W. S. (1827). Annulosa. Catalogue of insects, collected by Captain King, R. N. In ‘Narrative of a Survey of the Intertropical and Western Coasts of Australia. Performed between the Years 1818 and 1822. Vol. 2’. (Ed. P. P. King.) pp. 438–469. (J. Murray: London, UK.)

Main B. Y. (1976). ‘Spiders.’ (Collins: Sydney, Australia.)

Main B. Y. (1980). ‘Spiders of Australia.’ (Axiom Books: Stepney, SA, Australia.)

Mascord R. E. (1970). ‘Australian Spiders in Colour.’ (Reed: Sydney, Australia.)

Merian P. (1911) Die Spinnenfauna von Celebes. Beiträge zur Tiergeographie im Indoaustralischen Archipel. Zoologische Jahrbücher, Abteilung für Systematik. Geographie und Biologie der Tiere 31, 165–354.

Miyashita T. (1986) Growth, egg production, and population density of the spider, Nephila clavata in relation to food conditions in the field. Researches on Population Ecology 28, 135–149.
CrossRef |

Miyashita T. (1992) Feeding rate may affect dispersal in the orb-web spider Nephila clavata. Oecologia 92, 339–342.
CrossRef |

Moore C. W. (1977) The life cycle, habitat and variation in selected web parameters in the spider, Nephila clavipes (Koch) (Araneidae). American Midland Naturalist 98, 95–108.
CrossRef |

Moritz M., Fischer S.-C. (1986) Die Typen der Arachniden-Sammlung des Zoologischen Museums Berlin. VII. Araneae: Araneidae und Tetragnathidae. Mitteilung aus dem Zoologischen Museum in Berlin 62, 187–214.
CrossRef |

Nei M. (1978) Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics 89, 583–590.
PubMed |


Olivier A. G. (1789). Araignée, In ‘Aranea. Encyclopédie méthodique. Histoire naturelle. Insectes. Vol. 4’. (Ed. C.-L. Panckoucke.) pp. 173–240. (Panckoucke: Paris, France.)

Pallas P. S. (1772). ‘Spicilegia Zoologic. Vol. 1(9).’ (Gottl. August. Lange: (Berlin, Germany.)

Pan H., Zhou K. Y., Song D., Yang Q. (2004) Phylogenetic placement of the spider genus Nephila (Araneae: Araneoidea) inferred from rRNA and MaSp1 gene sequences. Zoological Science 21, 343–351.
CrossRef | PubMed |

Platnick N. I. (2006). ‘The World Spider Catalog, Version 7.0.’ (American Museum of Natural History: New York, USA.) Available at http://research.amnh.org/entomology/spiders/catalog/index.html [Verified July 2007]

Pocock R. I. (1900). Chilopoda, Diplopoda, and Arachnida. In ‘A monograph of Christmas Island (Indian Ocean)’. (Ed. C. W. Andrews.) pp. 153–162. (British Museum (Natural History): London, UK.)

Rack G. (1961) Die Entomologischen Sammlungen des Zoologischen Staatsinstituts und Zoologischen Museums Hamburg. II. Teil. Chelicerata II: Araneae. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 59, 1–60.

Rainbow W. J. (1895) Descriptions of some new Araneidae of New South Wales. No. 5. Proceedings of the Linnean Society of New South Wales, Series 2 10, 347–360.

Rainbow W. J. (1896) Descriptions of some new Araneidae of New South Wales. No. 6. Proceedings of the Linnean Society of New South Wales 21, 320–344.

Richardson B. J., Baverstock P. R., and Adams M. (1986). ‘Allozyme Electrophoresis: a Handbook for Animal Systematics and Population Biology.’ (Academic Press: Sydney, Australia.)

Robinson M. H., Mirick H. (1971) The predatory behavior of the golden-web spider Nephila clavipes (Araneae: Araneidae). Psyche 78, 123–139.

Robinson M. H., Robinson B. (1973) Ecology and behavior of the giant wood spider Nephila maculata (Fabr.) in New Guinea. Smithsonian Contributions to Zoology 149, 1–73.

Robinson M. H., Robinson B. (1974a) The stabilimentum of Nephila clavipes and the origin of stabilimentum-building in araneids. Psyche 80, 277–288.

Robinson M. H., Robinson B. (1974b) Adaptive complexity: the thermoregulation postures of the golden-web spider, Nephila clavipes, at low latitudes. American Midland Naturalist 92, 386–396.
CrossRef |

Robinson M. H., Robinson B. (1976) The ecology and behavior of Nephila maculata: a supplement. Smithsonian Contributions to Zoology 218, 1–22.

Robinson M. H., Robinson B. (1978a) The stabilamentum of Nephila clavipes and the origin of stabilamentum-building in araneids. Zoological Journal of the Linnean Society 64, 87–102.

Robinson M. H., Robinson B. (1978b) Thermoregulation in orb-web spiders: new descriptions of thermoregulatory postures and experiments on the effects of posture and coloration. Zoological Journal of the Linnean Society 64, 87–102.

Robinson M. H., Robinson B. (1980) Comparative studies of the courtship and mating behaviour of tropical araneid spiders. Pacific Insects Monographs 36, 1–218.

Robinson M. H., Robinson B. C., Murphy F. M., and Corley W. S. (1986). Egg-sac burying behavior in Nephila maculata: a specialized adaptation. In ‘Proceedings of the Ninth International Congress of Arachnology’. (Eds W. G. Eberhard, Y. D. Lubin, and B. C. Robinson.) pp. 245–254. (Smithsonian Institution Press: Washington, DC, USA.)

Roewer C. F. (1942). ‘Katalog der Araneae. Vol. 1.’ (R. Friedländer und Söhne: Bremen, Germany.)

Rypstra A. L. (1985) Aggregations of Nephila clavipes (Araneae, Araneidae) in relation to prey availability. The Journal of Arachnology 13, 71–78.

Schneider J. M., Elgar M. A. (2002) Sexual cannibalism in Nephila plumipes as a consequence of female life history strategies. Journal of Evolutionary Biology 15, 84–91.
CrossRef |

Schneider J. M., Herberstein M. E., de Crespigny F. C., Ramamurthy S., Elgar M. A. (2000) Sperm competition and small size advantage for males of the golden-orb spider Nephila edulis. Journal of Evolutionary Biology 13, 939–946.
CrossRef |

Schneider J. M., Thomas M. L., Elgar M. A. (2001) Ectomised conductors in the golden orb-spider Nephila plumipes (Araneoidea): male adaptation to sexual conflict. Behavioral Ecology and Sociobiology 49, 410–415.
CrossRef |

Schneider J. M., Fromhage L., Uhl G. (2005) Copulation patterns in the golden orb-web spider Nephila madagascariensis. Journal of Ethology 23, 51–55.
CrossRef |

Simon E. (1877) Arachnides recueillis aus îles Philippines par MM. G.-A. Baer et Laglaise. Annales de la Société Entomologique de France, séries 5 7, 53–96.

Simon E. (1894). ‘Histoire Naturelle des Araignées. 2nd edn, Vol. 1(3).’ (Encyclopédie Roret: Paris, France.)

Simon E. (1901) On the Arachnida collected during the “Skeat Expedition” to the Malay Peninsula, 1899–1900. Proceedings of the Zoological Society of London 71, 45–84.

Strand E. (1906a) Über einige Vogelspinnen und afrikanische Spinnen des Naturhistorischen Museums zu Wiesbaden. Jahrbuch des Nassauischen Vereins für Naturkunde 59, 1–45.

Strand E. (1906b) Sumatra- und Neu-Guinea-Spinnen des Naturhistorischen Museums zu Wiesbaden. Jahrbuch des Nassauischen Vereins für Naturkunde 59, 257–278.

Strand E. (1907) Vorläufige Diagnosen afrikanischer und südamerikanischer Spinnen. Zoologischer Anzeiger 31, 525–558.

Strand E. (1911a) Vorläufige Diagnosen neuer Spinnen, insbesondere aus der Südsee, des Senckenbergischen Museums. Archiv für Naturgeschichte 77, 202–207.

Strand E. (1911b) Araneae von den Aru- und Kei-Inseln. Abhandlungen herausgegeben von der Senckenbergischen Naturforschenden Gesellschaft 34, 129–199.

Strand E. (1915) Indoaustralische, papuanische und polynesische Spinnen des Senckenbergischen Museums, gesammelt von Dr. E. Wolf, Dr. J. Elbert u.a. Abhandlungen herausgegeben von der Senckenbergischen Naturforschenden Gesellschaft 36, 181–274.

Su Y.-C., Chang Y.-H., Lee S.-C., Tso I.-M. (2007) Phylogeography of the giant wood spider (Nephila pilipes, Araneae) from Asian-Australian regions. Journal of Biogeography 34, 177–191.
CrossRef |

Swofford D. L., Olsen G. J., Waddell P. J., and Hillis D. M. (1996). Phylogenetic inference. In ‘Molecular Systematics’. (Eds D. M. Hillis, C. Moritz and B. K. Mabel.) pp. 407–514. (Sinauer Associates: Sunderland, MA, USA.)

Thorell T. (1859) Nya exotiska Epeirider. Öfversigt af Kongl. Vetenskaps-Akademiens Förhandlingar 16, 299–304.

Thorell T. (1875) On some spiders from New Caledonia, Madagascar, and Réunion. Proceedings of the Zoological Society of London 1875, 130–149.

Thorell T. (1877) Studi sui Ragni Malesi e Papuani. I. Ragni di Selebes raccolti nel 1874 dal Dott. O. Beccari. Annali del Museo Civico di Storia Naturale di Genova 10, 341–634.

Thorell T. (1881) Studi sui Ragni Malesi e Papuani. Part III. Ragni dell’Austro Malesia e del Capo York, conservati nel Museo Civico di Storia Naturale di Genova. Annali del Museo Civico di Storia Naturale di Genova 17, 1–720.

Tso I.-M., Tai P.-L., Ku T.-H., Kuo C.-H., Yang E.-C. (2002) Colour associated foraging success and population genetic structure in a sit-and-wait predator Nephila maculata (Araneae: Tetragnathidae). Animal Behaviour 63, 175–182.
CrossRef |

Tikader B. K. (1962) Studies on some Indian spiders (Araneae: Arachnida). Journal of the Linnean Society of London. Zoology 44, 561–584.

Uhl G., Vollrath F. (1998a) Genital morphology of Nephila edulis: implications for sperm competition in spiders. Canadian Journal of Zoology 76, 39–47.
CrossRef |

Uhl G., Vollrath F. (1998b) Little evidence of size-selective sexual cannibalism in two species of Nephila (Araneae). Zoology 101, 101–106.

Uhl G., Vollrath F. (2000) Extreme size variability does not extend to genitalia in Nephila edulis. Journal of Zoology 251, 7–14.
CrossRef |

Vasconcellos-Neto J., Lewinsohn T. M. (1984) Discrimination and release of unpalatable butterflies by Nephila clavipes a Neotropical orb weaving spider. Ecological Entomology 9, 337–344.

Vinson A. (1863). ‘Aranéides des îles de la Réunion, Maurice et Madagascar.’ (Roret: Paris, France.)

Vollrath F. (1985) Web spider’s dilemma: a risky move or site dependent growth. Oecologia 68, 69–72.
CrossRef |

Vollrath F., Houston A. (1986) Previous experience and site tenacity in the orb spider Nephila (Araneae: Araneidae). Oecologia 70, 305–308.
CrossRef |

Vollrath F., Parker G. A. (1992) Sexual dimorphism and distorted sex ratios in spiders. Nature 360, 156–159.
CrossRef |

Vollrath F., Parker G. A. (1997) Giant female or dwarf male spiders? Nature 385, 688.
CrossRef | PubMed |

Walckenaer C. A. (1805). ‘Tableau des Aranéides ou Caractères Essentiels des Tribus, Genres, Families et Races que Renferme le Genre Aranea de Linné, avec la Désignation des Espèces Comprises dans Chacune de ces Divisions.’ (Published by the author: Paris, France.)

Walckenaer C. A. (1842). ‘Histoire Naturelle Des Insectes. Aptères. Vol. 2.’ (Librairie Encyclopédique de Roret: Paris, France.)

Walckenaer C. A. (1847). ‘Histoire Naturelle des Insectes. Aptères. Vol. 4.’ (Librairie Encyclopédique de Roret: Paris, France.)

Zarr J. H. (1984). ‘Biostatistical Analysis.’ (Prentice-Hall: Upper Saddle River, NJ, USA.)










Appendix 1.  Index to species names
Valid names in italics, synonyms in Roman. [Note that numbers refer to page numbers in the PDF version of this paper]
A1



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