The Spider Cytogenetic Database

Current version: 14 (Feb 28, 2026)

Douglas Araujo, Marielle Cristina Schneider, Emygdio Paula-Neto and Doralice Maria Cella​
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Lycosidae  Sundevall, 1833

Cited as = species name adopted by the researcher that performed the cytogenetic analysis; 2n = diploid number of males and in parenthesis, diploid number of females; n = haploid number; 3n = triploid number; 4n = tetraploid number; SCS = sex chromosome system; T = telocentric; A = acrocentric; St = subtelocentric; Sm = submetacentric; M = metacentric; H = holocentric; S = supernumerary chromosome; (?) = dubious description; (*) = designation of the species name not recorded in the taxonomic database.

Valid name Cited as 2n SCS Chromosomal morphology Collection site Bibliography
Aglaoctenus castaneus (Mello-Leitão, 1942) ---- 14/16 X1X2 14/16T+X1X2 Brazil Sanches et al., 2025
Aglaoctenus lagotis (Holmberg, 1876) ---- 14/16 X1X2 14/16T+X1X2 Brazil Sanches et al., 2025
Aglaoctenus oblongus (C. L. Koch, 1847) ---- 28/30 X1X2 28/30T+X1X2 Brazil Sanches et al., 2025
Alopecosa aculeata (Clerck, 1757) Tarentula aculeata 28 X1X2 26A+X1X2A Finland Hackman, 1948
Alopecosa albofasciata (Brullé, 1832) ---- 28 X1X2 26A+X1X2A Israel Gorlova et al., 1997
Alopecosa pulverulenta (Clerck, 1757) Tarentula pulverulenta 28 X1X2 26A+X1X2A Finland Hackman, 1948
Anomalomma harishi Dyal, 1935 ---- 28 X1X2 ---- India Mittal, 1961
Anomalomma harishi Dyal, 1935 ---- 28 X1X2 26A+X1X2A India Mittal, 1963
Anomalomma sp. ---- 28 X1X2 26A+X1X2A India Sharma et al., 1958
Arctosa alpigena (Doleschall, 1852) ---- 26 X1X2 24A+X1X2A Finland Hackman, 1948
Arctosa alpigena (Doleschall, 1852) ---- (30) ---- 28A+2St Austria Dolejš et al., 2011
Arctosa alpigena lamperti Dahl, 1908 ---- 28 X1X2 24A+2St+X1X2A Czech Republic Dolejš et al., 2011
Arctosa cinerea (Fabricius, 1777) ---- 28 X1X2 26A+X1X2A Czech Republic Dolejš et al., 2011
Arctosa cinerea (Fabricius, 1777) ---- 28 X1X2 26A+X1X2 Turkey Kumbiçak et al., 2011
Arctosa figurata (Simon, 1876) ---- 28 X1X2 26A+X1X2A Czech Republic Dolejš et al., 2011
Arctosa leopardus (Sundevall, 1833) ---- 26 X1X2 24A+X1X2A Finland Hackman, 1948
Arctosa leopardus (Sundevall, 1833) ---- 28 X1X2 26A+X1X2A Czech Republic Dolejš et al., 2011
Arctosa lutetiana (Simon, 1876) Tricca lutetiana 28 X1X2 26A+X1X2A Czech Republic Dolejš et al., 2011
Arctosa maculata (Hahn, 1822) ---- 28 X`1X2 26A+X1X2A Czech Republic Dolejš et al., 2011
Arctosa mulani (Dyal, 1935) Pardosa mulani 28 X1X2 26A+X1X2A India Sharma et al., 1958
Arctosa perita (Latreille, 1799) ---- (12) ---- ---- Turkey Akan et al., 2005
Arctosa perita (Latreille, 1799) ---- 28 X1X2 26A+X1X2A Czech Republic Dolejš et al., 2011
Arctosa renidescens Buchar & Thaler, 1995 ---- 28 X1X2 26A+X1X2A Austria Dolejš et al., 2011
Arctosa sp. ---- 28 X1X2 ---- India Mittal, 1960
Arctosa sp. ---- 28 X1X2 26A+X1X2A India Mittal, 1963
Crocodilosa leucostigma (Simon, 1885) Lycosa leucostigma 28 X1X2 ---- India Srivastava & Shukla, 1986
Diapontia uruguayensis Keyserling, 1877 ---- 28/30 X1X2 28/30T+X1X2 Brazil Sanches et al., 2025
Draposa lyrivulva (Bösenberg & Strand, 1906) Pardosa leucopalpis 28 X1X2 26A+X1X2A India Bole-Gowda, 1958
Draposa lyrivulva (Bösenberg & Strand, 1906) Pardosa leucopalpis 28 X1X2 26A+X1X2A India Mittal, 1963
Draposa lyrivulva (Bösenberg & Strand, 1906) Pardosa leucopalpis 24 X1X2 ---- India Srivastava & Shukla, 1986
Draposa oakleyi (Gravely, 1924) Pardosa lahorensis 28 X1X2 26A+X1X2A India Sharma et al., 1958
Draposa oakleyi (Gravely, 1924) Pardosa oakleyi 26 X1X2 ---- India Srivastava & Shukla, 1986
Evippa praelongipes (O.Pickard-Cambridge, 1871) ---- 26 X1X2 24A+X1X2A India Sharma et al., 1958
Evippa praelongipes (O.Pickard-Cambridge, 1871) ---- 26 X1X2 23A+1M+X1X2A Israel Gorlova et al., 1997
Gladicosa pulchra (Keyserling, 1877) Lycosa insopita 26+2S X1X2 ---- USA Montgomery, 1905
Hippasa agelenoides (Simon, 1884) ---- 28 X1X2 26A+X1X2A India Bole-Gowda, 1958
Hippasa agelenoides (Simon, 1884) ---- 24 X1X2 22A+X1X2A India Sharma et al., 1958
Hippasa agelenoides (Simon, 1884) ---- 28 X1X2 26A+X1X2A India Sharma & Ramakrishna, 2019
Hippasa deserticola Simon, 1889 Hippasa pisaurina 26 X1X2 ---- India Mittal, 1960
Hippasa deserticola Simon, 1889 Hippasa pisaurina 26 X1X2 24A+X1X2A India Mittal, 1963
Hippasa deserticola Simon, 1889 Hippasa madhuae 28 X1X2 26A+X1X2A India Parida et al., 1986
Hippasa deserticola Simon, 1889 Hippasa pisaurina 28 X1X2 ---- India Srivastava & Shukla, 1986
Hippasa holmerae Thorell, 1895 ---- 26 ---- ---- ---- Bugayong et al., 1999
Hippasa lycosina Pocock, 1900 ---- 24 X1X2O 10M+14Sm China Zhang et al., 2025
Hippasa olivacea (Thorell, 1887) Hippasa oliracea 28 X1X2 26A+X1X2A India Parida & Sharma, 1987b
Hippasa olivacea (Thorell, 1887) ---- 28 X1X2 ---- India Parida & Sharma, 1987a
Hippasa olivacea (Thorell, 1887) ---- 28 X1X2 ---- India Sharma & Parida, 1987
Hippasa sp. ---- 22 X1X2 ---- India Parida & Sharma, 1987a
Hippasa sp. ---- 22 X1X2 ---- India Sharma & Parida, 1987
Hippasosa kumari (Dyal, 1935) Ocyale kumari 22 X1X2 20A+X1X2A India Sharma et al., 1958
Hogna sternalis (Bertkau, 1880) ---- 19 X 18T+XT Brazil Araujo et al., 2015a
Hygrolycosa rubrofasciata (Ohlert, 1865) ---- 20 X1X2 18A+X1X2A Russia Gorlov et al., 1995
Lycosa barnesi Gravely, 1924 ---- 27 X ---- India Srivastava & Shukla, 1986
Lycosa bistriata Gravely, 1924 ---- 28 X1X2 26A+X1X2A India Bole-Gowda, 1958
Lycosa carmichaeli Gravely, 1924 ---- 28 X1X2 ---- India Mittal, 1961
Lycosa carmichaeli Gravely, 1924 ---- 28 X1X2 26A+X1X2A India Mittal, 1963
Lycosa carmichaeli Gravely, 1924 ---- 22 X1X2 ---- India Srivastava & Shukla, 1986
Lycosa chaperi Simon, 1885 ---- 22 X1X2 20A+X1X2A India Mittal, 1963
Lycosa coelestis L. Koch, 1878 ---- 26 X1X2 24A+X1X2A Japan Suzuki, 1954
Lycosa erythrognatha Lucas, 1836 Lycosa erythrognata 22 X1X2 20A+X1X2A ---- Diaz & Saez, 1966
Lycosa erythrognatha Lucas, 1836 ---- 22 X1X2 20T+X1X2T Argentina Chemisquy et al., 2008
Lycosa erythrognatha Lucas, 1836 ---- 22 X1X2 20T+X1X2T Brazil Araujo et al., 2015a
Lycosa erythrognatha Lucas, 1836 ---- 22 X1X2 20A+X1X2A Brazil Cavenagh et al., 2022
Lycosa grahami Fox, 1935 ---- 18 X1X20 12St+6T China Zhang et al., 2025
Lycosa madani Pocock, 1901 ---- 24 X1X2 22A+X1X2A India Mittal, 1963
Lycosa nigrotibialis Simon, 1884 ---- 28 X1X2 ---- India Mittal, 1961
Lycosa nigrotibialis Simon, 1884 ---- 28 X1X2 26A+X1X2A India Mittal, 1963
Lycosa nigrotibialis Simon, 1884 ---- 24 X1X2 ---- India Srivastava & Shukla, 1986
Lycosa nordenskjoldi Tullgren, 1905 Lycosa nordenskölii 19 X 18A+XA ---- Diaz & Saez, 1966
Lycosa nordenskjoldi Tullgren, 1905 ---- 19 X 18T+XT Brazil Araujo et al., 2015a
Lycosa gr. nordenskjoldi ---- 19 X 18A+XA Brazil Cavenagh et al., 2022
Lycosa pampeana Holmberg, 1876 ---- 22 X1X2 20T+X1X2T Argentina Chemisquy et al., 2008
Lycosa phipsoni Pocock, 1899 ---- ---- X1X2 ---- India Mittal, 1961
Lycosa praegrandis C.L. Koch, 1836 Lycosa cf. nordmanni 22 X1X2 20A+X1X2A Israel Gorlova et al., 1997
Lycosa sericovittata Mello-Leitão, 1939 ---- 22 X1X2 20T+X1X2T Brazil Araujo et al., 2015a
Lycosa sericovittata Mello-Leitão, 1939 ---- (26) ---- 26T Brazil Araujo et al., 2015a
Lycosa sericovittata Mello-Leitão, 1939 ---- 22 X1X2 20A+X1X2A Brazil Cavenagh et al., 2022
Lycosa tarantula (Linnaeus, 1758) Lycosa narborensis 18 ---- ---- Turkey Akan et al., 2005
Lycosa sp. ---- 20-24? ---- ---- France Carnoy, 1885
Lycosa sp. ---- ---- X ---- ---- Chickering & Hard, 1935
Lycosa sp. ---- 28 X1X2 26A+X1X2A Japan Suzuki, 1954
Lycosa sp. ---- 28 X1X2 26A+X1X2A India Bole-Gowda, 1958
Lycosa sp. ---- 28 X1X2 26A+X1X2A India Sharma et al., 1958
Lycosa sp. ---- 28 X1X2 ---- ---- Sokolov, 1960
Lycosa sp. ---- 28 X1X2 26A+X1X2A India Mittal, 1963
Lycosa sp. ---- 22 X1X2 20T+X1X2T Uruguay Brum-Zorrilla & Postiglioni, 1980
Lycosa sp. (grupo thorelli) ---- 23 X1X2X3 20T+X1X2X3T Uruguay Postiglioni & Brum-Zorrilla, 1981
Lycosa sp. ---- 22 X1X2 20T+X1X2T Uruguay Postiglioni & Brum-Zorrilla, 1981
Lycosa sp. ---- 18 X1X2 ---- India Srivastava & Shukla, 1986
Lycosa sp. ---- 28 X1X2 ---- India Srivastava & Shukla, 1986
Lycosa sp. ---- 22 X1X2 20A+X1X2A India Parida & Sharma, 1987b
Lycosa sp. ---- 22 X1X2 ---- India Parida & Sharma, 1987a
Lycosa sp. ---- 28 X1X2 ---- India Parida & Sharma, 1987a
Lycosa sp. ---- 22 X1X2 ---- India Sharma & Parida, 1987
Lycosa sp. ---- 28 X1X2 ---- India Sharma & Parida, 1987
Lycosa sp. ---- 21 X1X2Y 16T+2St+X1X2M/Sm+YT Peru Navia et al., 2006
Margonia himalayensis (Gravely, 1924) Venonia himalayensis 28 X1X2 ---- India Mittal, 1961
Margonia himalayensis (Gravely, 1924) Venonia himalayensis 28 X1X2 26A+X1X2A India Mittal, 1963
Pardosa agrestis (Westring, 1861) ---- 28 X1X2 26A+X1X2A Russia Gorlov et al., 1995
Pardosa agricola (Thorell, 1856) Lycosa fluviatilis 28 X1X2 26A+X1X2A Finland Hackman, 1948
Pardosa alacris (C.L. Koch, 1833) ---- 28 X1X2 26A+X1X2A Czech Republic Kumbiçak et al., 2009
Pardosa amentata (Clerck, 1757) Lycosa saccata 28 X1X2 26A+X1X2A Finland Hackman, 1948
Pardosa amentata (Clerck, 1757) Lycosa paludicola 28 X1X2 26A+X1X2A Finland Hackman, 1948
Pardosa amentata (Clerck, 1757) Lycosa pullata 28 X1X2 26A+X1X2A Finland Hackman, 1948
Pardosa amentata (Clerck, 1757) ---- 28 X1X2 ---- ---- Sokolov, 1960
Pardosa astrigera L. Koch, 1878 Lycosa T-insignita 28 X1X2 ---- ---- Suzuki, 1950a
Pardosa astrigera L. Koch, 1878 Lycosa T-insignita 28 X1X2 26A+X1X2A Japan Suzuki, 1954
Pardosa astrigera L. Koch, 1878 Lycosa cinereofusca 28 X1X2 26A+X1X2A Japan Suzuki, 1954
Pardosa astrigera L. Koch, 1878 Pardosa T-insignita 28 X1X2 ---- Japan Matsumoto, 1977
Pardosa astrigera L. Koch, 1878 ---- 28 X1X2 26T+X1X2T Japan Kageyama et al., 1978
Pardosa astrigera L. Koch, 1878 ---- 28 X1X2 26T+X1X2T China Zhenling et al., 1996
Pardosa bifasciata (C.L. Koch, 1834) ---- 28 X1X2 26A+X1X2A Turkey Kumbiçak et al., 2011
Pardosa birmanica Simon, 1884 Lycosa birmanica 28 X1X2 26A+X1X2A India Bole-Gowda, 1958
Pardosa birmanica Simon, 1884 ---- 28 X1X2 ---- India Datta & Chatterjee, 1983
Pardosa birmanica Simon, 1884 Lycosa birmanica 28 X1X2 ---- India Srivastava & Shukla, 1986
Pardosa birmanica Simon, 1884 ---- 28 X1X2 26A+X1X2A India Parida & Sharma, 1987b
Pardosa birmanica Simon, 1884 ---- 28 X1X2 ---- India Parida & Sharma, 1987a
Pardosa birmanica Simon, 1884 ---- 28 X1X2 ---- India Sharma & Parida, 1987
Pardosa birmanica Simon, 1884 ---- 28 X1X2 26T+X1X2T India Datta & Chatterjee, 1989
Pardosa fletcheri (Gravely, 1924) Lycosa fletcheri 28 X1X2 ---- India Srivastava & Shukla, 1986
Pardosa laura Karsch, 1879 ---- 28 X1X2 26T+X1X2T Japan Kageyama et al., 1978
Pardosa lugubris (Walckenaer, 1802) ---- 28 X1X2 26A+X1X2A Russia Gorlov et al., 1995
Pardosa morosa (L. Koch, 1870) ---- 26 X1X2 24A+X1X2A Czech Republic/Slovakia Král et al., 2011
Pardosa morosa (L. Koch, 1870) ---- 25? X1X2? 24A+1M? Czech Republic Král et al., 2011
Pardosa palustris (Linnaeus, 1758) Lycosa monticola 28 X1X2 26A+X1X2A Finland Hackman, 1948
Pardosa palustris (Linnaeus, 1758) Lycosa tarsalis 28 X1X2 24A+2M+X1X2A Finland Hackman, 1948
Pardosa palustris (Linnaeus, 1758) ---- 28 X1X2 26A+X1X2A Russia Gorlov et al., 1995
Pardosa plumipes (Thorell, 1875) ---- 28 X1X2 26A+X1X2A Russia Gorlov et al., 1995
Pardosa pseudoannulata (Bösenberg & Strand, 1906) Lycosa pseudoannulatus 28 X1X2 26A+X1X2A Japan Suzuki, 1954
Pardosa pseudoannulata (Bösenberg & Strand, 1906) Lycosa annandalei 28 X1X2 26A+X1X2A India Bole-Gowda, 1958
Pardosa pseudoannulata (Bösenberg & Strand, 1906) Lycosa basiri 22 X1X2 ---- India Mittal, 1960
Pardosa pseudoannulata (Bösenberg & Strand, 1906) Lycosa basiri 22 X1X2 20A+X1X2A India Mittal, 1963
Pardosa pseudoannulata (Bösenberg & Strand, 1906) Lycosa pseudoannulata 28 X1X2 26T+X1X2T Japan Kageyama et al., 1978
Pardosa pseudoannulata (Bösenberg & Strand, 1906) Lycosa annandalei 28 X1X2 ---- India Srivastava & Shukla, 1986
Pardosa pseudoannulata (Bösenberg & Strand, 1906) ---- n=15 ---- ---- China Yu et al., 2024
Pardosa saltans Töpfer-Hofmann, 2000 ---- 28 X1X2 26A+X1X2A Czech Republic Kumbiçak et al., 2009
Pardosa sumatrana (Thorell, 1890) Lycosa sumatrana 24 X1X2 22A+X1X2A India Sharma, 1961
Pardosa sumatrana (Thorell, 1890) Lycosa sumatrana 28 X1X2 ---- India Srivastava & Shukla, 1986
Pardosa wagleri (Hahn, 1822) ---- 28 X1X2 26A+X1X2A ---- Kořínková & Král, 2013
Pardosa sp. ---- ---- X ---- ---- Chickering & Hard, 1935
Pardosa sp. ---- 28 X1X2 26T+X1X2T India Sharma & Gupta, 1956
Pardosa sp. ---- 28 X1X2 26A+X1X2A India Bole-Gowda, 1958
Pardosa sp. ---- 28 X1X2 ---- India Mittal, 1960
Pardosa sp. ---- 28 X1X2 26A+X1X2A India Mittal, 1963
Pirata piraticus (Clerck, 1757) ---- 26 X1X2 24A+X1X2A Finland Hackman, 1948
Pirata piraticus (Clerck, 1757) ---- 26 X1X2 24T+X1X2T ---- Yu et al., 1998a
Pirata punctipes (Gravely, 1924) Lycosa punctipes 28 X1X2 26A+X1X2A India Sharma, 1961
Pirata subpiraticus (Bösenberg & Strand, 1906) ---- 26 X1X2 24T+X1X2T Japan Kageyama et al., 1978
Pirata subpiraticus (Bösenberg & Strand, 1906) ---- 26 X1X2 24T+X1X2T ---- Yu et al., 1998b
Pirata sp. ---- 26 X1X2 ---- ---- Sokolov, 1960
Piratula latitans (Blackwall, 1841) Pirata latitans 24 X1X2 22A+X1X2A India Mittal, 1963
Piratula piratoides (Bösenberg & Strand, 1906) Pirata piratoides 26 X1X2 24T+X1X2T ---- Zhenling et al., 1997a
Piratula procurva (Bösenberg & Strand, 1906) Pirata procurvus 26 X1X2 24T+X1X2T Japan Kageyama et al., 1978
Piratula uliginosa (Thorell, 1856) Pirata uliginosus 24 X1X2 22A+X1X2A Finland Hackman, 1948
Rabidosa rabida (Walckenaer, 1837) Lycosa rabida 28 X1X2 26T+X1X2T USA Wise, 1983
Rabidosa rabida (Walckenaer, 1837) Lycosa rabida 28 X1X2 ---- USA Tugmon et al., 1990
Rabidosa rabida (Walckenaer, 1837) Lycosa rabida 28 X1X2 26T+X1X2T USA Wise & Taylor, 1995
Schizocosa bilineata (Emerton, 1885) ---- ---- X ---- ---- Chickering & Hard, 1935
Schizocosa communis (Emerton, 1885) Lycosa communis 22 X1X2 ---- ---- Painter, 1914
Schizocosa crassipes (Walckenaer, 1837) ---- 22 X1X2 ---- USA Hard, 1939
Schizocosa malitiosa (Tullgren, 1905) Lycosa malitiosa 22 X1X2 20T+X1X2T Uruguay Brum-Zorrilla & Cazenave, 1974
Schizocosa malitiosa (Tullgren, 1905) Lycosa malitiosa 22 X1X2 ---- Uruguay Benavente & Wettstein, 1977
Schizocosa malitiosa (Tullgren, 1905) Lycosa malitiosa 22 X1X2 20T+X1X2T Uruguay Brum-Zorrilla & Postiglioni, 1980
Schizocosa malitiosa (Tullgren, 1905) ---- 22 X1X2 20T+X1X2T Argentina Chemisquy et al., 2008
Schizocosa ocreata (Hentz, 1844) ---- 22 X1X2 ---- USA Stratton, 1997
Schizocosa rovneri Uetz & Dondale, 1979 ---- 22 X1X2 ---- USA Stratton, 1997
Schizocosa stridulans Stratton, 1984 ---- 22 X1X2 ---- USA Stratton, 1997
Schizocosa sp. ---- 28 X1X2 ---- India Mittal, 1960
Schizocosa sp. ---- 28 X1X2 26A+X1X2A India Mittal, 1963
Schizocosa sp. (grupo malitiosa) Lycosa sp. (grupo malitiosa) 23 X 22T+XM Uruguay Postiglioni & Brum-Zorrilla, 1981
Serratacosa himalayensis (Gravely, 1924) Lycosa himalayensis 28 X1X2 26A+X1X2A India Mittal, 1963
Tigrosa georgicola (Walckenaer, 1837) Lycosa georgicola 28 X1X2 24T+2A+X1X2T USA Wise, 1983
Tigrosa helluo (Walckenaer, 1837) Lycosa helluo 28 X1X2 26T+X1X2T USA Wise & Taylor, 1995
Trochosa ruricola (De Geer, 1778) ---- 26 X1X2 24A+X1X2A Finland Hackman, 1948
Trochosa ruricola (De Geer, 1778) Trachosa ruricula 20 X1X2 ---- Turkey Taşdemir et al., 2012
Trochosa spinipalpis (F.O.Pickard-Cambridge, 1895) ---- 26 X1X2 24A+X1X2A Finland Hackman, 1948
Trochosa spinipalpis (F.O.Pickard-Cambridge, 1895) ---- 26 X1X2 24A-T+X1X2A-T ---- Zhenling et al., 1997b
Tropicosa thorelli (Keyserling, 1877) Lycosa thorelli 22 X1X2 20T+X1X2T Uruguay Brum-Zorrilla & Postiglioni, 1980
Tropicosa thorelli (Keyserling, 1877) Lycosa thorelli 22 X1X2 20T+X1X2T Uruguay Postiglioni & Brum-Zorrilla, 1981
Venonia sp. ---- 26 X1X2 ---- India Mittal, 1961
Venonia sp. ---- 26 X1X2 ---- India Mittal, 1961
Venonia sp. ---- 26 X1X2 24A+X1X2A India Mittal, 1963
Wadicosa fidelis (O.Pickard-Cambridge, 1872) ---- 28 X1X2 26A+X1X2A Canary Island, Israel Forman et al., 2013
Wadicosa quadrifer (Gravely, 1924) Lycosa quadrifer 27 X ---- India Srivastava & Shukla, 1986
Xerolycosa miniata (C.L. Koch, 1834) ---- 22 X1X2 20A+X1X2A Finland Hackman, 1948
Xerolycosa miniata (C.L. Koch, 1834) ---- 22 X1X2 20A+X1X2A Russia Gorlov et al., 1995
Xerolycosa miniata (C.L. Koch, 1834) ---- 22 X1X2 20A+X1X2A Czech Republic Dolejš et al., 2011
Xerolycosa nemoralis (Westring, 1861) ---- 26 X1X2 24A+X1X2A Finland Hackman, 1948
Xerolycosa nemoralis (Westring, 1861) ---- 22 X1X2 20A+X1X2A Russia Gorlov et al., 1995
Xerolycosa nemoralis (Westring, 1861) ---- 22 X1X2 20A+X1X2A Czech Republic Dolejš et al., 2011