Lycosidae

Female Xerolycosa nemoralis, from Spiders of the Netherlands.

Belongs within: Amaurobioidea.
Contains: Evippa, Tarentula, Lycosinae, Arctosa, Artoriinae, Piratinae, Venonia.

The Lycosidae, wolf spiders, are a cosmopolitan group of mostly ground-hunting spiders, characterised by having the anterior eyes smaller than the posterior eyes and forming a single row near the front margin of the carapace. Subgroups of the Lycosidae include the Venoniinae, a group of small, slender spiders with glistening hairs on the abdomen, found in the Indo-Australian region, that construct simple sheet webs. The Zoicinae, another Indo-Australian group, are characterised by a strong lateral apophysis on the male palp. Xerolycosa is a genus of medium-sized wolf spiders found in dry litter in the northern Palaearctic, females of which may dig shallow depressions in soil.

Big bad wolfies
Published 27 April 2010
Female wolf spider with an abdomen-load of young. Photo by J. Centavo.

The wolf spiders (I usually call them simply ‘wolfies’) of the family Lycosidae are one of the more easily recognisable groups of ground-dwelling spiders. Their eyes are placed in three rows clustered together at the front of the cephalothorax with the median posterior eyes large and sitting above a straight row of the small anterior eyes. Some wolf spiders reach relatively large sizes and large wolfies tend to usually be some variant of brown or grey with longitudinal stripes. Most members of the family, particularly the larger species, tend to be morphologically quite conservative and despite the recognition of well over 2000 species in the family (a number that is increasing with no sign of slowing down) distinguishing those species is not usually easy without close examination. Wolf spiders are not often inclined to bite humans and their bites are not usually regarded as dangerous (though a bite from a large species could be painful).

A slightly more distinctively coloured member of the family—Geolycosa archboldi from central Florida. Photo by H. K. Wallace.

Wolf spiders get their name because most members of the family are active hunters rather than snare builders though a smaller number of genera build distinctive sheet webs with a silk retreat tunnel. Most authors have regarded the sheet web builders as retaining the ancestral behaviour for the family but phylogenetic analysis has not determined this conclusively (Murphy et al. 2006); if web building is ancestral then it has been convergently lost on numerous occasions. Those lycosids that do not build sheet webs may be permanently vagrant or they may dig themselves a home burrow into which they retreat when not hunting. All wolf spiders wrap their eggs in a silken egg-sac which the female carries on the underside of her spinnerets; after the eggs hatch she carries her young around clinging to her abdomen.

A more typical lycosid photographed by Sander van der Molen.

Recent studies have shown the need for a fair amount of revision of lycosid systematics; the main genus Lycosa in particular had been shown to be a polyphyletic assemblage of conservative large lycosids. Researchers are slowly chipping away at the necessary revisions; a great deal of progress has been made (see, for instance, Volker Framenau’s webpage on Australian lycosids), but a great deal remains to be done. Matters have not been helped by the fact that wolf spiders were another group of arachnids to be subjected to the loving care and attention of Carl-Friedrich Roewer, demonstrating his usual talent for producing extensive revisions based on the most superficial and inconsequential of characters. Also, until recently there was debate over the identity of Lycosa‘s type species, the Mediterranean L. tarantula originally named by Linnaeus. This species, it should be noted, was the original tarantula; it was only later that the name became associated with South American mygalomorph spiders.

Systematics of Lycosidae

Characters (from Jocqué & Dippenaar-Schoeman 2007): Small to very large; eight eyes present in three rows, with anterior row of four small eyes, second row with two large (posterior median) eyes and third row of two intermediate-sized eyes on anterolateral surface of carapace; legs with three tarsal claws; ecribellate; male palp usually without retrolateral tibial apophysis; entelegyne; female carries egg-sac attached to spinnerets.

<==Lycosidae [Hippaseae, Hippasinae, Lycoseae, Lycosoidae]
    |--+--‘Arctosa’ ebichaPY07
    |  `--‘Arctosa’ kwangreungensisPY07
    `--+--Xerolycosa Dahl 1908PY07, MGK03
       |    |--X. albofasciata (Brullé 1832)K55
       |    |--X. miniata (Koch 1834)MGK03
       |    `--X. nemoralis (Westring 1861)MGK03
       `--+--+--LycosinaePY07
          |  `--+--‘Trochosa’ oraria (Koch 1876)PY07, VMP02
          |     `--+--ArctosaPY07
          |        `--ArtoriinaePY07
          `--+--PiratinaePY07
             `--VenoniinaeYF06
                  |  i. s.: *Satta cannibalorum Lehtinen & Hippa 1979YF06
                  |         Agalenocosa Mello-Leitão 1944FL13
                  |--‘Venonia’ spirocysta Chai 1991YF06
                  `--+--Allotrochosina Roewer 1960PY07, YF06
                     |    |--*A. schauinslandi (Simon 1899)PVD10
                     |    `--A. karriYF06
                     `--+--VenoniaYF06
                        `--Anomalosa Roewer 1960YF06
                             |--A. kochi Simon 1898YF06
                             `--A. oz Framenau 2006YF06
Lycosidae incertae sedis:
  Tricca alpigena (Doleschall 1852)K02
  Brevilabus gillonorum Cornic 1980J02
  Acantholycosa lignaria (Clerck 1757)RKD02
  EvippaMGK03
  Wadicosa Zyuzin 1985 [Wadicosinae]MGK03
    |--W. fidelis (Pickard-Cambridge 1872)MGK03
    `--W. quadrifera (Gravely 1924)MG03
  Allohogna Roewer 1955MGK03
    `--A. singoriensis (Laxmann 1770) [=Lycosa singoriensis]MGK03
  Mustelicosa Roewer 1960MGK03
    `--M. dimidiata (Thorell 1875) [=Trochosa dimidiata]MGK03
  Tarentuloides Doleschall 1859K92, D59
    `--*T. boiei (Doleschall 1859) [=Lycosa (Tarentuloides) boiei]D59
  Megarctosa leopardus (Sundevall 1832)K55
  SosippinaeF07
    |--Sosippus Simon 1888YF06
    |    `--S. nitidus (Mello-Leitão 1944) [=Hippasella nitida]C90
    `--PorrimosaC90
         |--P. callipoda (n. d.)C90
         |--P. castaneaC90
         |--P. diversa (n. d.)C90
         |--P. glieschi (n. d.)C90
         |--P. harknessiC90
         |--P. lagotisC90
         `--P. securifera (n. d.)C90
  Schizocosa Chamberlin 1904RU14
    |--S. crassipesPR01
    |--S. ocreata (Hentz 1844)RU14
    |--S. rovneri (Uetz & Dondale 1979)RU14
    |--S. saltatrixE95
    |--S. stridulans Stratton 1984RU14
    |--S. uetzi Stratton 1997RU14
    `--S. yurae (Strand 1908) [=Tarentula yurae, Avicosa yurae]J98
  TarentulaS06
  ZoicinaeFL13
    |--Zoica Simon 1898YF06
    |--Zantheres Thorell 1887YF06
    |--Lysania Thorell 1890YF06
    |--Shapna Hippa & Lehtinen 1983YF06
    `--Margonia Hippa & Lehtinen 1983YF06
         `--M. himalayensis (Gravely 1924) [=Venonia himalayensis]YF06
  Aglaoctenus Tullgren 1905 [incl. Porrima]YF06
    `--A. lagotis (Holmberg 1876)FM11
  Anomalomma Simon 1890YF06
    `--*A. lycosinum Simon 1890YF06
  TricassinaeF07
  Passiena torbjoerni Lehtinen 2005FY06
  TrabeopsD90
  Piratula minuta (Emerton 1885)SB14
  GladicosaSB14
    |--G. gulosaCM07
    `--G. pulchra (Keyserling 1877)SB14
  Donacosa merliniJD-S07
  LinoptesP92

*Type species of generic name indicated

References

[C90] Capocasale, R. M. 1990. Las especies de la subfamilia Hippasinae de America del Sur (Araneae, Lycosidae). Journal of Arachnology 18: 131–142.

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).

[D59] Doleschall, C. L. 1859. Tweede Bijdrage tot de kennis der Arachniden van den Indischen Archipel. Verhandelingen der Natuurkundige Vereeniging in Nederlandsch Indie [Acta Societatis Scientiarum Indo-Neêrlandicae] 5 (5): 1–60, pls 1–18.

[D90] Dondale, C. D. 1990. Litter Araneae (Araneida). In: Dindal, D. L. (ed.) Soil Biology Guide pp. 477–502. John Wiley & Sones: New York.

[E95] Elgar, M. A. 1995. The duration of copulation in spiders: comparative patterns. Records of the Western Australian Museum Supplement 52: 1–11.

[F07] Framenau, V. W. 2007. Revision of the new Australian genus Artoriopsis in a new subfamily of wolf spiders, Artoriinae (Araneae: Lycosidae). Zootaxa 1391: 1–34.

[FL13] Framenau, V. W., & A. E. Leung. 2013. Costacosa, a new genus of wolf spider (Araneae, Lycosidae) from coastal north-west Western Australia. Records of the Western Australian Museum Supplement 83: 173–184.

[FY06] Framenau, V. W., & J.-S. Yoo. 2006. Systematics of the new Australian wolf spider genus Tuberculosa (Araneae: Lycosidae). Invertebrate Systematics 20: 185–202.

[FM11] Freire-Jr, G. de B., & P. C. Motta. 2011. Effects of experimental fire regimes on the abundance and diversity of cursorial arachnids of Brazilian savannah (cerrado biome). Journal of Arachnology 39 (2): 263–272.

[J98] Jäger, P. 1998. Das Typenmaterial der Spinnentiere (Arachnida: Acari, Amblypygi, Araneae, Opiliones, Pseudoscorpiones, Scorpiones, Uropygi) aus dem Museum Wiesbaden. Jahrbuecher des Nassauischen Vereins fuer Naturkunde 119: 81–91.

[J02] Jocqué, R. 2002. Genitalic polymorphism—a challenge for taxonomy. Journal of Arachnology 30 (2): 298–306.

[JD-S07] Jocqué, R., & A. S. Dippenaar-Schoeman. 2007. Spider Families of the World. Royal Museum for Central Africa: Tervuren (Belgium).

[K92] Karsch, F. 1892. Arachniden von Ceylon und von Minikoy gesammelt von den Herren Doctoren P. und F. Sarasin. Berliner Entomologische Zeitschrift 36 (2): 267–310.

[K02] Koponen, S. 2002. Ground-living spiders in bogs in northern Europe. Journal of Arachnology 30 (2): 262–267.

[K55] Kraus, O. 1955. Spinnen von Korsika, Sardinien und Elba (Arach., Araneae). Senckenbergiana Biologica 36: 371–394.

[MG03] Marusik, Yu. M., & E. F. Guseinov. 2003. Spiders (Arachnida: Aranei) of Azerbaijan. 1. New family and genus records. Arthropoda Selecta 12 (1): 29–46.

[MGK03] Marusik, Yu. M., E. F. Guseinov & S. Koponen. 2003. Spiders (Arachnida: Aranei) of Azerbaijan. 2. Critical survey of wolf spiders (Lycosidae) found in the country with description of three new species and brief review of Palaearctic Evippa Simon, 1885. Arthropoda Selecta 12 (1): 47–65.

Murphy, N. P., V. W. Framenau, S. C. Donnellan, M. S. Harvey, Y.-C. Park & A. D. Austin. 2006. Phylogenetic reconstruction of the wolf spiders (Araneae: Lycosidae) using sequences from the 12S rRNA, 28S rRNA, and NADH1 genes: implications for classification, biogeography, and the evolution of web building behavior. Molecular Phylogenetics and Evolution 38 (3): 583–602.

[PVD10] Paquin, P., C. J. Vink & N. Dupérré. 2010. Spiders of New Zealand: annotated family key and species list. Manaaki Whenua Press: Lincoln (New Zealand).

[PY07] Park, Y. C., J.-S. Yoo, M. P. Schwarz, N. Murphy & J.-P. Kim. 2007. Molecular phylogeny of east Asian wolf spiders (Araneae: Lycosidae) inferred from mitochondrial 12S ribosomal DNA. Annals of the Entomological Society of America 100 (1): 1–8.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[PR01] Punzo, F., & C. Reeves. 2001. Geographical variation in male courtship behaviour of the giant whipscorpion Mastigoproctus giganteus (Lucas) (Arachnida, Uropygi). Bulletin of the British Arachnological Society 12 (2): 93–96.

[RKD02] Relys, V., S. Koponen & D. Dapkus. 2002. Annual differences and species turnover in peat bog spider communities. Journal of Arachnology 30 (2): 416–424.

[RU14] Rutledge, J. M., & G. W. Uetz. 2014. Juvenile experience and adult female mating preferences in two closely related Schizocosa species. Journal of Arachnology 42 (2): 170–177.

[S06] Strand, E. 1906. Die arktischen Araneae, Opiliones und Chernetes. In: Römer, F., & F. Schaudinn (eds) Fauna Arctica. Eine Zusammenstellun der arktischen Tierformen, mit besonder Berücksichtigung des Spitzbergen-Gebietes auf Grund der Ergebnisse der Deutschen Expedition in das Nördliche Eismeer im Jahre 1898 vol. 4 pp. 431–478. Gustav Fischer: Jena.

[SB14] Suter, R. B., & K. Benson. 2014. Nocturnal, diurnal, crepuscular: activity assessments of Pisauridae and Lycosidae. Journal of Arachnology 42 (2): 178–191.

[VMP02] Vink, C. J., A. D. Mitchell & A. M. Paterson. 2002. A preliminary molecular analysis of phylogenetic relationships of Australasian wolf spider genera (Araneae, Lycosidae). Journal of Arachnology 30 (2): 227–237.

[YF06] Yoo, J.-S., & V. W. Framenau. 2006. Systematics and biogeography of the sheet-web building wolf spider genus Venonia (Araneae: Lycosidae). Invertebrate Systematics 20: 675–712.

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