Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography

Systematics of the new Australian wolf spider genus Tuberculosa (Araneae : Lycosidae)

Volker W. Framenau A D and Jung-Sun Yoo A B C
+ 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(2) 185-202
Submitted: 18 August 2005  Accepted: 17 January 2006   Published: 26 April 2006


The new Australian wolf spider genus Tuberculosa is revised, with T. harveyi, sp. nov. from the Northern Territory as type species. The genus includes a further three species from northern Queensland: T. austini, sp. nov., T. hoggi (Framenau & Vink, 2001), comb. nov. and T. monteithi, sp. nov. The genus is defined by a unique sexual dimorphism: males carry modified tubercular setae on the ventral side of the third coxae, which are here compared to the knobbed setae that can be found on the ventral surface of the abdomen in Hygrolycosa rubrofasciata Ohlert, 1865 and Passiena torbjoerni Lehtinen, 2005 from two different lycosid subfamilies. Based on the structure of the male pedipalp, Tuberculosa belongs to the subfamily Lycosinae sensu Dondale (1986) with close affinities to Venatrix Roewer, 1960, because males of both genera have a tubercle on the outer edges of their fangs and the cymbium tip carries claw-like macrosetae. A cladistic analysis of all four Tuberculosa species, based on 12 morphological characters and with Venatrix konei (Berland, 1924) as outgroup, revealed a topology with T. austini and T. harveyi as sister-species in the most derived clade (V. konei (T. monteithi (T. hoggi (T. harveyi, T. austini))). The distribution of Tuberculosa in the tropical north of Australia supports an origin of the genus, as well as its putative sister-genus Venatrix, in the Palaearctic region, in contrast to all other Australian Lycosinae, which appear to be of Gondwanan origin.


We are grateful to Mike Gray and Graham Milledge (AM), Robert Raven and Owen Seeman (QM) and Gavin Dally (NTMAG) for the loan of specimens for this study. We are particularly grateful to Barbara Baehr (QM) and Tracey Churchill (NTMAG) and their respective families for their hospitality during visits of the senior author to Brisbane and Darwin. Young-Bo Lee and An-Ja Ko (National Institute of Agricultural Science and Technology, RDA, Suwon, Korea) took the SEM photographs. Melissa Thomas, Mark Harvey and Pekka Lehtinen gave helpful comments on earlier drafts of this manuscript. The Australian Biological Resource Study (ABRS) (to 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).


Barthel J., Helversen O. (1990) Pardosa wagleri (Hahn 1822) and Pardosa saturatior Simon 1937, a pair of sibling species (Araneae, Lycosidae). Bulletin de la Société Europeenne d’Arachnologie 1, 17–23.

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

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

Cracraft J. (1982) Geographic differentiation, cladistics, and vicariance biogeography: reconstructing the tempo and mode of evolution. American Zoologist 22, 411–424.

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

Doutch H. F. (1972). The palaeogeography of northern Australia and New Guinea and its relevance to the Torres Strait area. In ‘Bridge and Barrier: The Natural and Cultural History of Torres Strait’. (Ed. D. Walker.) pp. 1–10. (Reseach School of Pacific Studies, Department of Biogeography and Geomorphology, Australian National University: Canberra, Australia.)

Elgar M. A. (1991) Sexual cannibalism, size dimorphism, and courtship behaviour in orb-weaving spiders (Araneidae). Evolution 45, 444–448.

Engelhardt W. (1964) Die mitteleuropäischen Arten der Gattung Trochosa C. L. Koch, 1848 (Araneae, Lycosidae). Morphologie, Chemotaxonomie, Biologie, Autökologie. Zeitschrift für Morphologie und Ökologie der Tiere 54, 219–392.
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) Gender specific differences in activity and home range reflect morphological dimorphism in wolf spiders (Araneae, Lycosidae). Journal of Arachnology 33, 334–346.

Framenau V. W. (in press a) Mainosa, a new genus for the Australian ‘Shuttlecock Wolf Spider’ (Araneae, Lycosidae). Journal of Arachnology. ,

Framenau V. W. (in press b) The wolf spider genus Venatrix: new species, synonyms and generic transfers. Records of the Western Australian Museum ,

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., Gotch T. B., Austin A. D. (In press) The wolf spiders of artesian springs in arid South Australia, with a revalidation of Tetralycosa (Araneae, Lycosidae). Journal of Arachnology ,

Fuhn I. E., Niculescu-Burlacu F. (1971) Lycosidae. Fauna Republicii Socialiste România. Arachnida 5, 1–256.

Gasnier T. R., de Azevedo C. S., Torres-Sanchez M. P., Höfer H. (2002) Adult size of eight hunting spider species in central Amazonia: temporal variations and sexual dimorphism. Journal of Arachnology 30, 146–154.

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

Greenstone M. H. (1982) Ballooning frequency and habitat predictability in two wolf spider species (Lycosidae: Pardosa). The Florida Entomologist 65, 83–89.

Head G. (1995) Selection on fecundity and variation in degree of sexual size dimorphism among species (class: Araneae). Evolution 49, 776–781.

Hebets E. A., Uetz G. W. (1999) Female responses to isolated signals from multimodal male courtship display in the wolf spider genus Schizocosa (Araneae: Lycosidae). Animal Behaviour 57, 865–872.
CrossRef | PubMed |

Hebets E. A., Uetz G. W. (2000) Leg ornamentation and the efficacy of courtship display in four species of wolf spider (Araneae: Lycosidae). Behavioral Ecology and Sociobiology 47, 280–286.
CrossRef |

Kikkawa J., Monteith G. B., and Ingram G. (1981). Cape York Peninsula: the major region of faunal interchange. In ‘Ecological Biogeography in Australia’. (Ed. A. Keast.) pp. 1697–1742 (Junk: The Hague, The Netherlands.)

Köhler D., Tembrock G. (1987) Akustische Signale bei der Wolfsspinne Hygrolycosa rubrofasciata (Arachnida: Lycosidae). Zoologischer Anzeiger 219, 147–153.

Kotiaho J., Alatalo R. V., Mappes J., Parri S. (1996) Sexual selection in a wolf spider: male drumming activity, body size, and viability. Evolution 50, 1977–1981.

Kronestedt T. (1996) Vibratory communication in the wolf spider Hygrolycosa rubro-fasciata (Araneae, Lycosidae). Revue Suisse de Zoologie hors série I , 341–354.

Lehtinen P. T. (2005) Review of the Oriental wolf spider genus Passiena (Lycosidae, Pardosinae). Journal of Arachnology 33, 398–407.

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

Maddison W. P., Donague M. J., Maddison D. R. (1984) Outgroup analysis and parsimony. Systematic Zoology 33, 83–103.

Mappes J., Alatalo R. V., Kotiaho J., Parri S. (1996) Viability costs of condition-dependent sexual male display in a drumming wolf spider. Proceedings of the Royal Society of London. Series B. Biological Sciences 263, 785–789.

Miller G. L., Stratton G. E., Miller P. R., Hebets E. A. (1998) Geographic variation in male courtship behaviour and sexual isolation in wolf spiders of the genus Schizocosa. Animal Behaviour 56, 937–951.
CrossRef | PubMed |

Monteith G. B. (1997) Revision of the Australian flat bugs of the subfamily Mezirinae (Insecta: Hemiptera: Aradidae). Memoirs of the Queensland Museum 41, 1–169.

Moya-Laraño J., Halaj J., Wise D. H. (2002) Climbing to reach females: Romeo should be small. Evolution 56, 420–425.
PubMed |

Moya-Laraño J., Taylor P. W., Fernández-Montraveta C. (2003) Body patterns as potential amplifiers of size and condition in a territorial spider. Biological Journal of the Linnean Society 78, 355–364.
CrossRef |

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: [verified March 2006].

Nixon K. C., Carpenter J. M. (1993) On outgroups. Cladistics 9, 413–426.
CrossRef |

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

Prenter J., Elwood R. W., Montgomery W. I. (1999) Sexual size dimorphism and reproductive investment by female spiders: a comparative analysis. Evolution 53, 1987–1994.

Richter C. J. J. (1970) Aerial dispersal in relation to habitat in eight wolf spider species (Pardosa, Araneae, Lycosidae). Oecologia 5, 200–214.
CrossRef |

Sheehan D. C., and Hrapchack 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). Journal of Arachnology 28, 133–140.

Tietjen W. J., and Rovner J. S. (1982). Chemical communication in lycosids and other spiders. In ‘Spider Communication: Mechanisms and Ecological Significance’. (Eds P. N. Witt and J. S. Rovner.) pp. 249–279 (Princeton University Press: Princeton, NJ, USA.)

Uetz G. W., and Stratton G. E. (1982). Acoustic communication and reproductive isolation in spiders. In ‘Spider Communication: Mechanisms and Ecological Significance’. (Ed. P. N. Witt and J. S. Rovner.) pp. 123–159 (Princeton University Press: Princeton, NJ, USA.)

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 |

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

Walker S. E., Rypstra A. L. (2001) Sexual dimorphism in functional response and troph morphology in Rabidosa rabida (Araneae: Lycosidae). American Midland Naturalist 146, 161–170.

Walker S. E., Rypstra A. L. (2002) Sexual dimorphism in trophic morphology and feeding behavior of wolf spiders (Araneae: Lycosidae) as a result of differences in reproductive roles. Canadian Journal of Zoology 80, 679–688.
CrossRef |

Watrous L. E., Wheeler Q. D. (1981) The out-group comparison method of character analysis. Systematic Zoology 30, 1–11.

Wiebes J. T. (1959) The Lycosidae and Pisauridae (Araneae) of the Netherlands. Zoologische Verhandelingen 42, 1–79.

Yan C. Y., Kroenke L. W. (1993) A plate tectonic reconstruction of the southwest Pacific, 0–100 Ma. Proceedings of the Ocean Drilling Program, Scientific Results 130, 697–709.

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.

Rent Article (via Deepdyve) Export Citation Cited By (6)