Lycosa

Female Lycosa godeffroyi carrying an egg-sac, copyright Fir0002.

Belongs within: Lycosinae.

The genus Lycosa has been used in the past for a diverse array of relatively large wolf spiders that construct silken burrows. It is now known to be polyphyletic in its earlier circumscriptions; the genus as currently recognised is most diverse in the western Palaearctic, but remains poorly defined and many species remain to be reclassified. The type species, Lycosa tarantula, is found in southern Europe and was the original spider referred to as ‘tarantula’ before the name became associated with members of the Theraphosidae. Other members of the genus include L. coelestis, found in eastern Asia.

Wolf underground
Published 17 April 2024

The wolf spiders of the Lycosidae can be a conservative bunch, and their classification has long proven a challenge. Perhaps no subgroup of wolfies epitomises this issue more than the type genus Lycosa.

Lycosa tarantula alongside its burrow, copyright Nicolas Helitas.

Over two hundred species have been assigned to Lycosa from subtropical climates around the world. However, the genus has historically been poorly defined and long suspected of being a dumping ground for species not readily assignable to other genera. Lycosa species have an elevated cephalothorax, relatively large posterior eyes, and relatively short, strong legs with black markings ventrally. Most construct burrows from which they ambush passing prey. A core group of species around the type, L. tarantula, has also been characterised by shared genital characters: females with a tongue-shaped or rhomboid epigyne septum confined to the posterior half of the epigyne and a reduced or absent septal pedicel, and males with a wide and flat tegular apophysis ending with a distal process directed retrolaterally to posteriorly (Armiach Steinpress et al. 2022). However, the vast majority of Lycosa species found outside the Mediterranean region are yet to have their relationships with this core group confirmed.

Lycosa oculata rapidly retreating, copyright Mike Steinmann.

Investigation of the genetic relationships between Lycosa sensu stricto species in the western Mediterranean by Planas et al. (2013) identified four distinct species groups. Armiach Steinpress et al. (2022), reviewed species found in Israel and found that they could also be divided between Planas et al.’s groups. As well as molecular data, members of the groups could also be separated by their living habits. Lycosa tarantula and its two most immediate relatives are relatively large species constructing permanent burrows whose entrances they surround with a high turret of leaves and other debris held together with silk. The four members of the L. baulnyi group also constructed entrance turrets but their burrows are not as deep as those of the L. tarantula group. The four or more species of the L. oculata group forego the turrets, instead camouflaging their burrows with sandy trapdoors. Finally, two species in the L. fasciiventris group do not construct permanent burrows, instead sheltering under rocks. Relationships between the groups (and to related genera) suggest that the burrowing habit is ancestral for Lycosa. Did the L. fasciiventris group lose their burrowing skills because they moved into microhabitats too rocky for digging?

Systematics of Lycosa
<==Lycosa Latreille 1804F08
    |--L. coelestis Koch 1878PY07, VMP02
    `--+--L. godeffroyi Koch 1865PY07, MGK03
       `--L. suzukiiPY07
Lycosa incertae sedis:
  *L. tarantula (Linnaeus 1758) [=Aranea tarantula]F08
  L. accentuataK01
  L. aerescensS99
  L. affinis Lucas 1846E12
  L. agricola Thorell 1856S06
  L. albopatellaS06
  L. albosparsa (n. d.)F06
  L. algidaH09
  L. annulataP-C06
  L. arapensis (Strand 1908) [=Tarentula arapensis, Mimohogna arapensis]J98
  L. arenivagaFF99
  L. argenteomarginata Lycas 1846E12
  L. atalanta Koch 1880S06
  L. atrata Thorell 1872S06
  L. auroguttata (Keyserling 1891)FM11
  L. avida Walckenaer 1837N37
  L. baltimoriana (Keyserling 1876) [=Tarentula baltimoriana; incl. L. benedicta Chamberlin 1925]F37
  L. bellicosaFF99
  L. biimpressa Lucas 1846E12
  L. borealis Sund. 1833S06
  L. camtschadalica Kulcz. 1885S06
  L. canescensS99
  L. carolinensisC90
  L. cecilii Pocock 1901P01
  L. ceratiolaMC05
  L. corallina McKay 1974FL13
  L. cuneataK01
  L. cursorK01
  L. darlingiP01
  L. dimidiata [incl. L. dimidiata var. virgata Kulczyński 1901]K01
  L. eiseni Thorell 1875S06
  L. erratica Lucas 1846E12
  L. erythrognatha Lucas 1836FM11
  L. exilipes Lucas 1846E12
  L. felina Koch 1878MGK03
  L. ferruginea (Koch 1867)S06
  L. florissanti Petrunkevitch 1922S93
  L. fulvipes Collett 1875S06
  L. fulvolineata Lucas 1846E12
  L. fumigata (L. 1758)S06
  L. furcifera Thorell 1875S06
  L. fuscula Thorell 1875S06
  L. giebeli Pavesi 1873S06
  L. gilberta Hogg 1905 [=Venonia gilberta]YF06
  L. glacialis Thorell 1872 [incl. L. aquilonaris]S06
  L. gracilenta Lucas 1846E12
  L. grisea Risso 1826R26
  L. groenlandica Thorell 1872 [incl. L. dromaea]S06
  L. guernei Simon 1887S06
  L. hawaiiensis Simon 1899S99
  L. herbigrada Blackw. 1857S06
  L. hilarisH09
  L. hyperborea Thorell 1871S06
  L. indagatrix Walckenaer 1837K92
  L. indecora (Koch 1880)S06
  L. innocua Doleschall 1859D59
  L. inornata Blackwall 1862FM11
  L. inquilinaR13
  L. ishikariana (Saito 1934)STK06
  L. labradoriensis Thorell 1875S06
  L. laeta Koch 1877E07
  L. lanca Karsch 1879K92
  L. lapponica Thorell 1872S06
  L. lasciva (Koch 1880)S06
  L. latefasciataS95
  L. latisepta Kulcz. 1885S06
  L. lebakensis Doleschall 1859D59
  L. leuckartii (Thorell 1870)LF10
  L. lignaria Sund. 1833S06
  L. mackenziana Keys. 1876S06
  L. malayana Doleschall 1859D59
  L. malitiosaE95
  L. marshalli Pocock 1901P01
  L. mauraS99
  ‘Salticus’ melvillensis [=Erigone melvillensis]S06
  L. miniataK01
  L. monscalvana Risso 1826R26
  L. montanaS06
  L. monticola Sund. 1833S06
  L. musgraveiB96
  L. narbonensis Walckenaer 1806MGK03 [=L. (Tarentula) narbonensisD59]
  L. nauticaS99
  L. nemoralisK01
  L. norvegica Thorell 1870S06
  L. numida Lucas 1846E12
  L. oahuensis Koch & Keyserling 1890KK90
  L. palustris (L. 1758) [incl. L. palustris var. islandica Strand 1906]S06
  L. paramushirensis Nakatsudi 1937N37
  L. pastoralisS95
  L. pernix Thorell 1871S06
  L. pilipes Lucas 1846E12
  L. praegrandis Koch 1836MGK03
  L. prativaga (Koch 1870)S06
  L. pulverulentaK01
  L. quadripunctata Lucas 1846E12
  L. rabidaFF99
  L. raboti Simon 1887S06
  L. ralphiF64
  L. retirugaH09
  L. riparia (Koch 1848)S06
  L. rufimanoides (Strand 1908) [=Tarentula rufimanoides]J98
  L. salifodina McKay 1976FL13
  L. schauinslandi Simon 1899S99
  L. septentrionalis Westring 1861S06
  L. subantarctica Forster 1964F64
  L. subaustralis (Strand 1908) [=Tarentula subaustralis, Hogna subaustralis]J98
  L. sylvicola Lucas 1846E12
  L. tachypoda Thorell 1878S06
  L. tayloriS99
  L. thorelli (Keyserling 1877)FM11
  L. timida Lucas 1846E12
  L. transvaalicaP01
  L. tremensS99
  L. tremula Simon 1899S99
  L. turbidaH09
  L. uliginosaS99
  L. umbrataS99
  L. uncata Thorell 1877S06
  L. vagabunda Lucas 1846E12
  L. valida Lucas 1846E12
  L. venatrix Lucas 1846E12
  L. villica Lucas 1846E12
Nomina nuda: Lycosa capitataS06
             Lycosa feroxS06
             Lycosa insularisS06
             Lycosa luteolaS06
             Lycosa naeviaS06
             Lycosa nivalisS06
             Lycosa pellitaS06
             Lycosa pilosaS06
             Lycosa simmoS06
             Lycosa turneriS06
             Lycosa undataS06
             Lycosa ungavensisS06

*Type species of generic name indicated

References

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[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.

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[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.

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[LF10] Langlands, P. R., & V. W. Framenau. 2010. Systematic revision of Hoggicosa Roewer, 1960, the Australian ‘bicolor‘ group of wolf spiders (Araneae: Lycosidae). Zoological Journal of the Linnean Society 158: 83–123.

[MC05] Machado, G., P. C. Carrera, A. M. Pomini & A. J. Marsaioli. 2005. Chemical defense in harvestmen (Arachnida, Opiliones): do benzoquinone secretions deter invertebrate and vertebrate predators? Journal of Chemical Ecology 31 (11): 2519–2539.

[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.

[N37] Nakatsudi, K. 1937. Notes on a new genus and two new species of Arachnida from the island of Paramushir, northern Kuriles, Japan. Tokyo Nogyo Daigaku Nogaku Shuho 1 (1): 22–27.

[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.

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Planas, E., C. Fernández-Montraveta & C. Ribera. 2013. Molecular systematics of the wolf spider genus Lycosa (Araneae: Lycosidae) in the Western Mediterranean Basin. Molecular Phylogenetics and Evolution 67: 414–428.

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[S99] Simon, E. 1899. Ergebnisse einer Reise nach dem Pacific (Schauinsland 1896–1897). Arachnoideen. Zoologische Jahrbücher, Abteilung für Systematik, Geographie un Biologie der Thiere 12 (4): 411–437.

[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.

[STK06] Suzuki, S., N. Tsurusaki & Y. Kodama. 2006. Distribution of an endangered burrowing spider Lycosa ishikariana in the San’in Coast of Honshu, Japan (Araneae: Lycosidae). Acta Arachnologica 55 (2): 79–86.

[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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