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Systematics, phylogeny and biogeography

Systematics and biogeography of the sheet-web building wolf spider genus Venonia (Araneae : Lycosidae)

Jung-Sun Yoo A B C D and Volker W. Framenau A
+ Author Affiliations
- Author Affiliations

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

B School of Animal Biology, University of Western Australia, Crawley, Western Australia 6009, Australia.

C Research Institute for Natural Science, Dongguk University, Seoul 100-715, Korea.

D Corresponding author. Email:

Invertebrate Systematics 20(6) 675-712
Submitted: 19 April 2006  Accepted: 21 September 2006   Published: 15 December 2006


The Australian/Oriental wolf spider genus Venonia Thorell, 1894 (type species V. coruscans Thorell, 1894) belongs to one of the few true web-building genera within the Lycosidae. Their small sheet-webs with funnel-like retreats are generally found in the ground layer of vegetation, such as on lawns and meadows, but also in depressions of soil and under roots of trees. Members of the genus Venonia are easily identified within the Lycosidae due to a unique combination of somatic and genitalic characters. Most conspicuous is a posterodorsal white spot on the abdomen just above the elongated posterior spinnerets on an otherwise uniformly coloured, small, and relatively slender spider. The cymbium of the male pedipalp is highly asymmetrical appearing retrolaterally truncated. Its tegular apophysis is membranous. The female epigyne is generally not sclerotised and has a posterior central incision. Our revision recognises fifteen species of which seven are new to science: V. chaiwooi, sp. nov.; V. choiae, sp. nov.; V. cinctipes (Simon, 1898); V. coruscans Thorell, 1894; V. infundibulum, sp. nov.; V. joejim, sp. nov.; V. kimjoopili, sp. nov.; V. kokoda Lehtinen & Hippa, 1979; V. micans (Simon, 1898) (= Venonia gabrielae Barrion & Litsinger, 1995, new synonymy); V. micarioides (L. Koch, 1877); V. milla Lehtinen & Hippa, 1979; V. muju (Chrysanthus, 1967); V. nata, sp. nov.; V. sungahae, sp. nov.; and V. vilkkii Lehtinen & Hippa, 1979. A phylogenetic analysis including representatives of the venoniine genera Anomalosa Roewer, 1960 and Allotrochosina Roewer, 1960 with Pirata subpiraticus (Bösenberg & Strand, 1906) as outgroup suggests a Gondwanan origin of the Venoniinae and one dispersal event within Venonia from the Australian region into Wallacea and only one dispersal event by V. coruscans into the Oriental region. Venonia spirocysta Chai, 1991 from China is not a true Venonia and is here considered incerta sedis. We reject the inclusion of the genus Zoica Simon, 1898 in the subfamily Venoniinae Lethinen & Hippa, 1979 due to considerable morphological differences in representatives of this genus (in particular in the male pedipalp), and therefore consider the subfamily Zoicinae Lehtinen & Hippa, 1979 as valid.


We are grateful to the following people for the loan of specimens used in this study: Mike Gray and Graham Milledge (AM); Bruce Halliday and Sandy Roy (ANIC); Janet Beccaloni (BMNH); Yin Chang-Min (HNU); Gary Jahn and Josie Catindig (IRRI); Anne-Elise Leguin and Christine Rollard (MNHN); Peter Lillywhite, Ken Walker and Richard Marchant (MV); Nigel Monaghan (NMI); Erik Nieukerken and B. Van Bekkum-Ansari (RMNH); Gavin Dally and Tracy Churchill (NTM); Rob Raven and Owen Seeman (QM); David Hirst (SAM); Seppo Koponen (ZMT). In particular, we want to acknowledge the help of Jim Berry (Butler University, Indianapolis) and Joe Beatty (Southern Illinois University at Carbondale) who made their Pacific islands wolf spider material available for our study. Young-Bo Lee and An-Ja Ko (National Institute of Agricultural Science and Technology, RDA, Suwon, Korea) took the SEM photographs. Bill Humphreys made one of his stereomicroscopes available for the use of the senior author. We thank Sung-Ah Kim for compiling some tables and parts of this study. The senior author is particularly grateful to his father and mother for their lifelong and continuing support without which this study would not have been possible. The contribution of Joo-Pil Kim (Dongguk University, Seoul) and Yung-Chul Park (Ewha Womans University, Seoul) to the senior author’s study of Australasian wolf spiders is acknowledged. The Australian Biological Resources Study (ABRS) (Mark Harvey, WAM, and Andy Austin, The University of Adelaide) provided funding for VWF. This work was supported by the Korea Research Foundation Grant funded by the Korean Government (MOEHRD). (KRF-2005- 214-C00226) to JSY.


Barrion A. T., and Litsinger J. A. (1995). ‘Riceland Spiders of South and Southeast Asia.’ (CAB International: Wallingford, UK.)

Bonnet P. (1955). ‘Bibliographia Araneorum. Tome II (1me partie: A–B).’ (Douladoure: Toulouse, France.)

Bonnet P. (1959). ‘Bibliographia Araneorum. Tome II (5me partie: T–Z).’ (Douladoure: Toulouse, France.)

Bösenberg W., Strand E. (1906) Japanische Spinnen. Abhandlungen der Senckenbergischen naturforschenden Gesellschaft 30, 93–422.

Bremer K. (1988) The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution 42, 795–803.
CrossRef |

Bremer K. (1994) Branch support and tree stability. Cladistics 10, 295–304.
CrossRef |

Brignoli P. M. (1983). ‘A Catalogue of the Araneae described between 1940 and 1981.’ (British Arachnological Society: Manchester, UK.)

Brown J. H., and Lomolino M. V. (1998). ‘Biogeography. 2nd edn.’ (Sinauer Associates: Sunderland, MA, USA.)

Chai B. Q. (1991). In ‘Fauna of Zhejiang: Araneida’. (Eds Z. F. Chen and Z. H. Zhang.) p. 208. (Zhejiang Science and Technology Publishing House: Zhejiang, China.)

Chrysanthus F. (1967) Spiders from South New Guinea VIII. Nova Guinea Zoology 37, 401–426.

Dondale C. D. (1986). The subfamilies of wolf spiders (Araneae: Lycosidae). Actas X Congreso Internacional de Aracnología, Jaca, España 1, 327–332.

Dondale C. D., and Redner J. H. (1990). ‘The Insects and Arachnids of Canada, Part 17. The Wolf Spiders, Nursery Web Spiders, and Lynx Spiders of Canada and Alaska, Araneae: Lycosidae, Pisauridae, and Oxyopidae.’ (Biosystematics Research Centre, Research Branch, Agriculture Canada: Ottawa, Canada.)

Foelix R. F. (1970) Structure and function of tarsal sensilla in the spider Araneus diadematus. The Journal of Experimental Zoology 175, 99–108.
CrossRef |

Framenau V. W. (2002) Review of the wolf spider genus Artoria Thorell (Araneae: Lycosidae). Invertebrate Systematics 16, 209–235.
CrossRef |

Framenau V. W. (2005) The wolf spider genus Artoria Thorell in Australia: new synonyms and generic transfers (Araneae, Lycosidae). Records of the Western Australian Museum 22, 265–292.

Framenau V. W. (2006a) Review of the Australian wolf spider genus Anomalosa Roewer 1960. Zootaxa 1304, 1–20.

Framenau V. W. (2006b) Review of the wolf spider genus Diahogna Roewer, 1960 (Araneae, Lycosidae). Journal of Natural History 40, 273–292.
CrossRef |

Framenau V. W., Vink C. J. (2001) Revision of the wolf spider genus Venatrix Roewer (Araneae: Lycosidae). Invertebrate Taxonomy 15, 927–970.
CrossRef |

Framenau V. W., Yoo J.-S. (2006) Systematics of the new Australian wolf spider genus Tuberculosa (Araneae: Lycosidae). Invertebrate Systematics 20, 185–202.
CrossRef |

Framenau V. W., Gotch T. B., Austin A. D. (2006) The wolf spiders of artesian springs in arid South Australia, with a revalidation of Tetralycosa (Araneae, Lycosidae). The Journal of Arachnology 34, 1–36.
CrossRef |

Goloboff P. (1993). ‘NONA, version 2.0.’ (Instituto Miguel Lillo: Sierra Madre de Tucuman, Argentina.) Available online at: [verified 20 November 2006].

Gravely F. H. (1924) Some Indian spiders of the family Lycosidae. Records of the Indian Museum Calcutta 26, 587–613.

Hippa H., Lehtinen P. T. (1983) The Zantheres group of Zoicinae (Araneae, Lycosidae) and a relimitation of the subfamily. Annales Zoologici Fennici 20, 151–156.

ICZN  (1999). ‘International Code of Zoological Nomenclature. Fourth Edition.’ (The International Trust for Zoological Nomenclature: London, UK.)

Job W. (1968) Das Röhrengewebe von Aulonia albimana Walckenaer (Araneida: Lycosidae) und seine systematische Bedeutung. Zoologischer Anzeiger 180, 403–409.

Job W. (1974) Beiträge zur Biologie der fangnetzbauenden Wolfsspinne Aulonia albimana (Walckenaer 1805) (Arachnida, Araneae, Lycosidae, Hippasinae). Zoologische Jahrbücher für Systematik 101, 560–608.

Koch C. L. (1847). ‘Die Arachniden. 14. Band.’ (Zeh’sche Buchhandlung: Nürnberg, Germany.)

Koch L. (1877). ‘Die Arachniden Australiens, nach der Natur beschrieben und abgebildet.’ pp. 889–968. (Bauer and Raspe: Nürnberg, Germany.)

Lehtinen P. T., Hippa H. (1979) Spiders of the Oriental–Australian region I. Lycosidae: Venoniinae and Zoicinae. Annales Zoologici Fennici 16, 1–22.

Lundgren I. (2002). ‘Palau Nature Facts.’ (The Nature Conservancy: Koror, Palau.) Available online at: [verified 12 April 2006].

McKay R. J. (1973) The wolf spiders of Australia (Araneae: Lycosidae): 1. The bicolor group. Memoirs of the Queensland Museum 16, 375–398.

McKay R. J. (1985). Lycosidae. In ‘Zoological Catalogue of Australia, Vol. 3. Arachnida, Mygalomorphae, Araneomorphae in Part, Pseudoscorpionida, Amblypygida, Palpigradi’. (Ed. D. E. Walton.) pp. 73–88. (Australian Government Publishing Service: Canberra, Australia.)

Murphy N. P., Framenau V. W., Donellan S. C., Harvey M. S., Park Y.-C., Austin A. D. (2006) Phylogenetic reconstruction of the wolf spiders (Araneae, Lycosidae) using sequences from 12S rRNA, 28S rRNA and NADH1 genes: implications for classification, biogeography and the evolution of web-building behavior. Molecular Phylogenetics and Evolution 38, 583–602.
CrossRef | PubMed |

Nixon K. C. (2002). ‘WinClada, version 1.00.08.’ (Published by the author: Ithaca, NY, USA.) Available online at: http:/ [verified 20 November 2006].

Page R. D. M. (2001). ‘Nexus Data Editor for Windows, version 0.5.0.’ (Published by the author: Glasgow, UK.) Available at: [verified 20 November 2006].

Petrunkevitch A. (1928) Systema Aranearum. Transactions of the Connecticut Academy of Arts and Science 29, 1–270.

Platnick N. I. (1993). ‘Advances in Spider Taxonomy, 1988–1991. With Synonymies and Transfers 1940–1980.’ (New York Entomological Society in association with the American Museum of Natural History: New York, USA.)

Platnick N. I. (1998). [imprint date 1997]. ‘Advances in Spider Taxonomy, 1992–1995. With Redescriptions 1940–1980.’ (New York Entomological Society in association with the American Museum of Natural History: New York, USA.)

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

Rainbow W. J. (1911) A census of Australian Araneidae. Records of the Australian Museum 9, 107–319.

Roewer C. F. (1955). [imprint date 1954]. ‘Katalog der Araneae von 1758 bis 1940. Vol. 2a.’ (Institut Royal des Sciences Naturelles de Belgique: Bruxelles, Belgium.)

Roewer C. F. (1959) Araneae Lycosiformae II (Lycosidae). Exploration du Parc National de l’Upemba – Mission GF de Witte 55, 1–518.

Sheehan D. C., and Hrapchak B. B. (1980). ‘Theory and Practice of Histotechnology.’ 2nd edn. (Battelle Press: Columbus, OH, USA.)

Sierwald P. (2000) Description of the male of Sosippus placidus, with notes on the subfamily Sosippinae (Araneae, Lycosidae). The Journal of Arachnology 28, 133–140.
CrossRef |

Simon E. (1885) Matériaux pour servir à la faune arachnologiques de l’Asie méridionale. I. Arachnides recuellis à Wagra-Karoor près Gundacul, district de Bellary par M. M. Chaper. II. Arachnides recuellis à Ramnad, district de Madura par M. l’abbé Fabre. Bulletin de la Société Zoologique de France 10, 1–39.

Simon E. (1887) Observation sur divers arachnides: synonymies et descriptions. Annales de la Société Entomologique de France 7, 158–195.

Simon E. (1890). (in Hasselt, A. W. M. van) Araneae ex Archipelago Malayano. In ‘Zoologische Ergebnisse einer Reise in Niederländisch Ost-Indien 1’. (Ed. M. Weber.) pp. 193–210. (Verlag von E. J. Brill : Leiden, The Netherlands.)

Simon E. (1898a). ‘Histoire naturelle des araignées. 2.’ (Roret: Paris, France.)

Simon E. (1898b) Descriptions d’arachnides nouveaux des familles des Agelenidae, Pisauridae, Lycosidae et Oxyopidae. Annales de la Société Entomologique de Belgique 42, 1–34.

Storey B. C. (1995) The role of mantle plumes in continental breakup: case histories from Gondwanaland. Nature 377, 301–308.
CrossRef |

Sundevall J. C. (1833). ‘Conspectus Arachnidum.’ (C. F. Berling: Lund, Sweden.)

Tanaka H. (1988) Lycosid spiders of Japan I. The genus Pirata Sundevall. Acta arachnologica. Tokyo 36, 33–77.

Thorell T. (1894) Decas aranearum in ins. Singapore a Cel. Th. Workman inventarum. Bolletino della Società Entomologica Italiana 26, 321–355.

Vink C. J. (2001) A revision of the genus Allotrochosina Roewer (Lycosidae: Araneae). Invertebrate Taxonomy 15, 461–466.
CrossRef |

Vink C. J., Paterson A. M. (2003) Combined molecular and morphological phylogenetic analyses of the New Zealand wolf spider genus Anoteropsis (Araneae: Lycosidae). Molecular Phylogenetics and Evolution 28, 576–587.
CrossRef | PubMed |

Workman T. (1896). ‘Malaysian Spiders.’ pp. 25–104. (Workman: Belfast, Ireland.)

Zyuzin A. A. (1993) Studies on the wolf spiders (Araneae: Lycosidae). I. A new genus and species from Kazakhstan, with comments on the Lycosinae. Memoirs of the Queensland Museum 33, 693–700.

Appendix 1.  Characters and character states for the cladistic analysis of Venonia and allied general
All character states are considered unordered for the cladistic analysis in NONA
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Appendix 2.  Character states for phylogenetic analysis of Venonia and allied genera

Appendix 3.  Specimens used in phylogenetic analysis
Australian states: Qld, Queensland; NT, Northern Territory; Vic., Victoria; WA, Western Australia. Types: AT, allotype; HT, holotype; LT, lectotype; PT, paratype
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