Skip to main content Skip to main navigation menu Skip to site footer
Published: 2022-08-04

Bowie gen. nov., a diverse lineage of ground-dwelling spiders occurring from the Himalayas to Papua New Guinea and northern Australia (Araneae: Ctenidae: Cteninae)

Arachnology, Senckenberg Research Institute, Mertonstraße 17–21, 60325 Frankfurt am Main, Germany
Araneae Taxonomy systematics revision first records new species unknown sexes transfers South Asia Southeast Asia Queensland


A new genus in the spider family Ctenidae Keyserling, 1877 is described from Asia: Bowie gen. nov. belongs to the subfamily Cteninae and all members formerly described were placed so far in the genus Ctenus Walckenaer, 1805. After morphological as well as preliminary molecular characters were checked, it was clear that a new genus had to be erected to accommodate this predominantly Asian lineage of ground-dwelling spiders. As is the case in most Ctenidae generally, it was not easy to find apomorphic characters diagnosing this new taxon. Therefore, a combination of morphological characters is used to define all congeners. An important and newly introduced character in this respect is –among others– the fused patellar crack of the male palp. Forty-nine valid species are transferred to the new genus (first country/province records of particular species underlined): Bowie martensi (Jäger, 2012) comb. nov. (Nepal), B. bomdilaensis (Tikader & Malhotra, 1981) comb. nov. (India), B. indicus (Gravely, 1931) comb. nov. (India), B. cladarus (Jäger, 2012) comb. nov. (Myanmar), B. pingu (Jäger & Minn, 2015) comb. nov. (Myanmar), B. natmataung (Jäger & Minn, 2015) comb. nov. (Myanmar), B. sikkimensis (Gravely, 1931) comb. nov. (India), B. ramosus (Thorell, 1887) comb. nov. (Myanmar), B. goaensis (Bastawade & Borkar, 2008) comb. nov. (India), B. himalayensis (Gravely, 1931) comb. nov. (India), B. meghalayaensis (Tikader, 1976) comb. nov. (India), B. narashinhai (Patel & Reddy, 1988) comb. nov. (India), B. ceylonensis (F.O. Pickard-Cambridge, 1897) comb. nov. (Sri Lanka), B. andamanensis (Gravely, 1931) comb. nov. (Andaman Islands), B. kapuri (Tikader, 1973) comb. nov. (Andaman Islands), B. cochinensis (Gravely, 1931) comb. nov. (India), B. thorelli (F.O. Pickard-Cambridge, 1897) comb. nov. (Sri Lanka), B. lishuqiang (Jäger, 2012) comb. nov. (China: Sichuan), B. banna (Yao & Li in Chu et al. 2022) comb. nov. (China: Yunnan), B. theodorianum (Jäger, 2012) comb. nov. (Thailand, Laos, Vietnam), B. robustus (Thorell, 1897) comb. nov. (Myanmar, Thailand, Laos), B. yaeyamensis (Yoshida, 1998) comb. nov. (Taiwan), B. yulin (Yao & Li in Chu et al. 2022) comb. nov. (China: Yunnan), B. simplex (Thorell, 1897) comb. nov. (Myanmar, Thailand, Laos), B. bayeri (Jäger 2012) comb. nov. (Laos), B. holthoffi (Jäger, 2012) comb. nov. (Laos), B. saci (Ono, 2010) comb. nov. (Vietnam), B. floweri (F.O. Pickard-Cambridge, 1897) comb. nov. (Malaysia Peninsula), B. argentipes (Hasselt, 1893) comb. nov. (Malaysia Peninsula, Singapore, Indonesia: Sumatra), B. palembangensis (Strand, 1906) comb. nov. (Indonesia: Sumatra), B. angigitanus (Roewer, 1938) comb. nov. (Papua New Guinea), B. pulvinatus (Thorell, 1890) comb. nov. (Malaysia: Sarawak), B. hosei (F.O. Pickard-Cambridge, 1897) comb. nov. (Malaysia: Sarawak, Brunei), B. monaghani (Jäger, 2013) comb. nov. (Laos), B. javanus (Pocock, 1897) comb. nov. (Indonesia: Sumatra, Java, Bali), B. fungifer (Thorell, 1890) comb. nov. (Malaysia Peninsula), B. valvularis (Hasselt, 1882) comb. nov. (Indonesia: Sumatra), B. bicostatus (Thorell, 1890) comb. nov. (Malaysia: Sarawak), B. bantaengi (Merian, 1911) comb. nov. (Indonesia: Sulawesi), B. bowonglangi (Merian, 1911) comb. nov. (Indonesia: Sulawesi), B. celebensis (Pocock, 1897) comb. nov. (Indonesia: Sulawesi), B. sagittatus (Giltay, 1935) comb. nov. (Indonesia: Sulawesi), B. kochi (Simon, 1897b) comb. nov. (Indonesia: West Papua), B. sarawakensis (F.O. Pickard-Cambridge, 1897) comb. nov. (Malaysia: Sarawak), B. philippinensis (F.O. Pickard-Cambridge, 1897) comb. nov. (Philippines), B. aruanus (Strand, 1911) comb. nov. (Indonesia: Maluku), B. angularis (Roewer, 1938) comb. nov. (Indonesia: Maluku), B. rufisternis (Pocock, 1898) comb. nov. (Papua New Guinea: New Britain), and B. corniger (F.O. Pickard-Cambridge, 1898) comb. nov. (South Africa). For thirty-two species, illustrations of their respective copulatory organs, as well as habitus photos, are provided. Fifty-five new species are described, these are listed, together with the already described species, according to their geographic occurrence and to their affiliation to species groups as far as the latter could be recognised (type species indicated by an asterisk): Bowie hunkydory spec. nov. (Nepal), B. ziggystardust spec. nov. (Nepal), B. ladystardust spec. nov. (Nepal), B. aladdinsane spec. nov. (India), B. majortom spec. nov. (Nepal), B. jeangenie spec. nov. (India), B. heroes spec. nov. (India), B. fascination spec. nov. (Vietnam), B. low spec. nov. (Thailand), B. dodo spec. nov. (Vietnam), B. stationtostation spec. nov. (Myanmar), B. candidate spec. nov. (Vietnam), B. diamonddogs spec. nov. (Vietnam), B. yassassin spec. nov. (Taiwan), B. bemywife spec. nov. (Thailand), B. subterraneans spec. nov. (Thailand), B. afterall spec. nov. (Thailand), B. warszawa spec. nov. (Thailand), B. artdecade spec. nov. (Cambodia), B. bigbrother spec. nov. (Vietnam), *B. rebelrebel spec. nov. (Vietnam), B. youngamericans spec. nov. (Vietnam), B. right spec. nov. (Vietnam), B. stay spec. nov. (Vietnam), B. fame spec. nov. (Vietnam), B. win spec. nov. (Vietnam), B. joethelion spec. nov. (Malaysia Peninsula), B. mossgarden spec. nov. (Malaysia Peninsula), B. neukoeln spec. nov. (Malaysia Peninsula), B. scarymonsters spec. nov. (Indonesia: Sumatra), B. teenagewildlife spec. nov. (Indonesia: Sumatra), B. letsdance spec. nov. (Indonesia: Java), B. crystaljapan spec. nov. (Indonesia: Sumatra), B. tonight spec. nov. (Malaysia: Sarawak), B. catpeople spec. nov. (Malaysia: Sabah), B. ricochet spec. nov. (Indonesia: Kalimantan), B. fashion spec. nov. (Malaysia Peninsula), B. withinyou spec. nov. (Malaysia: Sarawak), B. abdulmajid spec. nov. (Singapore), B. blackout spec. nov. (Malaysia Peninsula), B. modernlove spec. nov. (Malaysia: Sabah), B. chinagirl spec. nov. (Malaysia: Sabah), B. withoutyou spec. nov. (Malaysia: Sabah), B. magicdance spec. nov. (Indonesia: Sulawesi), B. bluejean spec. nov. (Malaysia: Sabah), B. criminalworld spec. nov. (Malaysia: Sabah), B. shakeit spec. nov. (Malaysia: Sabah), B. ashestoashes spec. nov. (Indonesia: Kalimantan), B. underground spec. nov. (Indonesia: Kalimantan), B. lodger spec. nov. (Philippines), B. redsails spec. nov. (Philippines), B. thenextday spec. nov. (Indonesia: Papua), B. lazarus spec. nov. (Papua New Guinea), B. thiesi spec. nov. (Papua New Guinea) and B. blackstar spec. nov. (Papua New Guinea). Formerly unknown sexes are described for the first time for the following species: Bowie martensi comb. nov. (female), B. indicus comb. nov. (only male RTA tip), B. narashinhai comb. nov. (male), B. argentipes comb. nov. (female) and B. celebensis comb. nov. (male). Fourteen species groups are proposed on the basis of morphological characters: cladarus-species group (17 species), robustus-species group (14 species), bemywife-species group (2 species), rebelrebel-species group (10 species), youngamericans-species group (3 species), floweri-species group (3 species), scarymonsters-species group (2 species), teenagewildlife-species group (2 species), argentipes-species group (10 species), javanus-species group (5 species), chinagirl-species group (11 species), shakeit-species group (5 species), lodger-species group (3 species) and blackstar-species group (7 species). Thus, 93 species are grouped, leaving 11 species without a current assignment to any of these groups. Ctenus kandyensis Kim & Ye, 2014 syn. nov. is recognised as a junior synonym of Bowie thorelli comb. nov., and C. calcarifer F.O. Pickard-Cambridge, 1902 syn. nov. as junior synonym of B. sarawakensis comb. nov. Five species previously placed in Ctenus are considered nomina dubia, four of them are transferred to Bowie gen. nov., one to Nilus: Bowie barbatus (Thorell, 1895) comb. nov., B. tumidulus (Simon, 1887b) comb. nov., B. flavidus (Hogg, 1922) comb. nov., B. pollii (Hasselt, 1893) comb. nov. and Nilus marginatus (Walckenaer, 1847) comb. nov. Beside the latter species, three species are transferred from Ctenus, all of them to Anahita: A. periculosa (Bristowe, 1931) comb. nov., A. dangsa (Reddy & Patel, 1994) comb. nov. and A. tuniensis (Patel & Reddy, 1988) comb. nov. All in all, Bowie gen. nov. is the second largest genus within the family Ctenidae, with 108 species in total including nomina dubia. It occurs from Nepal in the Himalayas and South India across large parts of South and South East Asia to Papua New Guinea. One undescribed species is known from northern Australia (Queensland). Representatives are known to live in the leaf litter of forests, with most species having small distribution ranges, usually occurring within a 100 km radius. With this revision, the family Ctenidae contains now 586 species and 48 genera, and the number of species assigned to the genus Ctenus, so far used as nomenclatural “waste bin”, is reduced to 164.



  1. Alayón, G. (2002a) Notas sobre la familia Ctenidae en Cuba con la descripción de una nueva especie de Ctenus de una caverna y la hembra de C. coxanus Bryant (Arachnida: Araneae). Revista Ibérica de Aracnología, 6, 135–139.
    Alayón, G. (2002b) Dos especies nuevas de Ctenus (Araneae: Ctenidae) de Quintana Roo, México. Solenodon, 2, 11–16.
    Alayón, G. (2004) Notas sobre la familia Ctenidae (Arachnida: Araneae) en La Hispaniola, con la descripción de tres nuevas especies. Revista Ibérica de Aracnología, 9, 277–283.
    Arizala, S., Labarque, F.M. & Polotow, D. (2021) Revision of the Neotropical spider genus Acanthoctenus (Araneae: Ctenidae: Acanthocteninae). Zootaxa, 4920 (1), 1–55.
    Arts, L.des. (1912) Zusammenstellung der afrikanischen Arten der Gattung Ctenus. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 29 (2), 183–218.
    Badcock, H.D. (1932) Reports of an expedition to Paraguay and Brazil in 1926–7, supported by the Trustees of the Percy Sladen Memorial Fund and the Executive Committee of the Carnegie Trust for the Universities of Scotland. Arachnida from the Paraguayan Chaco. Journal of the Linnean Society of London, Zoology, 38 (257), 1–48.
    Banks, N. (1892) A classification of North American spiders. The Canadian Entomologist, 24 (4), 88–97.
    Banks, N. (1902) Papers from the Hopkins Stanford Galapagos Expedition; 1898–1899. VII. Entomological Results (6). Arachnida. With field notes by Robert E. Snodgrass. Proceedings of the Washington Academy of Sciences, 4, 49–86.
    Bastawade, D.B. & Borkar, M. (2008) Arachnida (orders Scorpiones, Uropygi, Amblypygi, Araneae and Phalangida). In: Fauna of Goa, State Fauna Series. Zoological Survey of India, Kolkata, 16, 211–242.
    Bayer, S. (2011). Revision of the pseudo-orbweavers of the genus Fecenia Simon, 1887 (Araneae, Psechridae), with emphasis on their pre-epigyne. ZooKeys, 153, 1–56.
    Bayer, S. (2012) The lace-sheet-weavers – a long story (Araneae: Psechridae: Psechrus). Zootaxa, 3379 (1), 1–170.
    Bayer, S. & Jäger, P. (2009) Heteropoda species from limestone caves in Laos (Araneae: Sparassidae: Heteropodinae). Zootaxa, 2143 (1), 1–23.
    Bayer, S. & Jäger, P. (2010) Expected species richness in the genus Psechrus in Laos (Araneae: Psechridae). Revue Suisse de Zoologie, 117, 57–75.
    Benoit, P.L.G. (1977) Etudes sur les Ctenidae africains (Araneae). III. Le remembrement du genre Anahita Karsch 1879. Revue Zoologique africaine, 91, 368–380.
    Benoit, P.L.G. (1978) Espèces est-africaines du genre Ctenus Walckenaer (Araneae—Ctenidae). Revue Zoologique Africaine, 92 (2), 525–532.
    Benoit, P.L.G. (1979). Etudes sur les Ctenidae africains (Araneae) VIII. Gen. Ctenus Walck.-groupe abditus. Revue Zoologique africaine, 93, 425–444.
    Benoit, P.L.G. (1981) Etudes sur les Ctenidae africains (Araneae) XI. Etude des mâles isolés. Revue Zoologique Africaine, 95, 29–44.
    Bertkau, P. (1872) Über die Respirationsorgane der Araneen. Archiv für Naturgeschichte, 38, 208–233.
    Bertkau, P. (1880) Verzeichniss der von Prof. Ed. van Beneden auf seiner im Auftrage der Belgischen Regierung unternommenen wissenschaftlichen Reise nach Brasilien und La Plata I. J. 1872–73 gesammelten Arachniden. Mémoires Couronnés et Mémoires des Savants Étrangers de l’Académie Royale des Sciences, des Lettres et des Beaux-Arts de Belgique, 43, 1–120.
    Bertkau, P. (1882) Über das Cribellum und Calamistrum. Ein Beitrag zur Histologie, Biologie und Systematik der Spinnen. Archiv für Naturgeschichte, 48, 316–362.
    Blackwall, J. (1841) The difference in the number of eyes with which spiders are provided proposed as the basis of their distribution into tribes; with descriptions of newly discovered species and the characters of a new family and three new genera of spiders. Transactions of the Linnean Society of London, 18 (4), 601–670.
    Bloom, T., Binford, G., Esposito, L. A., Alayón G., G., Peterson, I., Nishida, A., Loubet-Senear, K. & Agnarsson, I. (2014) Discovery of two new species of eyeless spiders within a single Hispaniola cave. Journal of Arachnology, 42, 148–154.
    Bonnet, P. (1956) Bibliographia araneorum. Douladoure, Toulouse, 2 (2), 919–1926.
    Brescovit, A.D. (1996) Revisão do gênero Centroctenus Mello-Leitão (Araneae, Ctenidae, Cteninae). Revista Brasileira de Entomologia, 40, 301–313.
    Brescovit, A.D. & Simó, M. (2007) On the Brazilian Atlantic Forest species of the spider genus Ctenus Walckenaer, with the description of a neotype for C. dubius Walckenaer (Araneae, Ctenidae, Cteninae). Bulletin of the British arachnological Society, 14, 1–17.
    Brignoli, P.M. (1983) A catalogue of the Araneae described between 1940 and 1981. Manchester University Press, 755 pp.
    Bristowe, W.S. (1931) A preliminary note on the spiders of Krakatau. Proceedings of the Zoological Society of London, 101 (4), 1387–1400., pl. 1.
    Chu, C., Lu, Y., Yao, Z. & Li, S. (2022) A survey of ctenid spiders (Araneae, Ctenidae) from Xishuangbanna Tropical Botanical Garden, Yunnan, China. Biodiversity Data Journal, 10, e87597.
    Daudin, F.M. (1802) Histoire naturelle, générale et particulière, des reptiles. F. Dufart, Paris, 3, 1–452., pls. 29–45.
    Davies, V.T. (1994) The huntsman spiders Heteropoda Latreille and Yiinthi gen. nov. (Araneae: Heteropodidae) in Australia. Memoirs of the Queensland Museum, 35, 75–122.
    Döbler, H. (1982) Katalog der in den Sammlungen der Abteilung Taxonomie der Insekten des Institutes für Pflanzenschutzforschung, Bereich Eberswalde (ehemals Deutsches Entomologisches Institut), auf bewahrten Typen—XX (Coleóptera: Lycidae, Lampyridae, Cantharidae, Phengodidae, Drilidae, Malachiidae, Karumiidae, Dasytidae, Melyridae, Cleridae, Lymexylonidae, Derodontidae)]. Beiträge zur Entomologie, 32 (2), 393–435.
    Dufour, L. (1820) Descriptions de cinq arachnides nouvelles. Annales Générales des Sciences Physiques, 5, 198–209.
    Dupérré, N. (2015) Description of a new genus and thirteen new species of Ctenidae (Araneae, Ctenidae) from the Chocó region of Ecuador. Zootaxa, 4028 (4), 451–484.
    Foelix, R.F. & Erb, B. (2011) Microscopical studies on exuviae of the jumping spider Phidippus regius. Peckhamia, 90.1, 1–15.
    Forcart, L. (1961) Katalog der Typusexemplare in der Arachnida- Sammlung des Naturhistorischen Museums zu Basel: Scorpionidea, Pseudoscorpionidea, Solifugae, Opilionidea and Araneida. Verhandlungen der Naturforschenden Gesellschaft Basel, 72 (1), 47–87.
    Giltay, L. (1935) Liste des arachnides d’Extrême-Orient et des Indes orientales recueillis, en 1932, par S. A. R. le Prince Léopold de Belgique. Bulletin du Musée Royal d’Histoire Naturelle de Belgique, 11 (20), 1–15.
    Gravely, F.H. (1931) Some Indian spiders of the families Ctenidae, Sparassidae, Selenopidae and Clubionidae. Records of the Indian Museum Calcutta, 33, 211–282.
    Hasselt, A.W.M.van. (1882) Araneae. In: Veth, P.J. (Ed.), Midden-Sumatra 4A(11). Reizen en onderzoekingen der Sumatra-expeditie, uitgerust door het aardrijkskundig genootschap, 1877–1879. Brill, Leiden, pp. 1–56., pls. 1–5.
    Hasselt, A.W.M.van. (1893) Spinnen van Java, Sumatra en Ceylon voor den Heer J. R. H. Neervoort van de Poll door den Heer J. Z. Kannegieter aldaar verzameld. Tijdschrift voor Entomologie, 36, 129–158.
    Heimer, S. & Nentwig, W. (1991) Spinnen Mitteleuropas: Ein Bestimmungsbuch. Paul Parey, Berlin, 543 pp.
    Helversen, O.V. & Martens, J. (1972) Unrichtige Fundort-Angaben in der Arachniden-Sammlung Roewer. Senckenbergiana biologica, 53, 109–123.
    Henrard, A. & Jocqué, R. (2017) Morphological and molecular evidence for new genera in the Afrotropical Cteninae (Araneae, Ctenidae) complex. Zoological Journal of the Linnean Society, 180 (1), 82–154 & Suppl.
    Hogg, H.R. (1922) Some spiders from south Annam. Proceedings of the Zoological Society of London, 92 (2), 285–312.
    Huber, K.C., Haider, T.S., Müller, M.W., Huber, B.A., Schweyen, R.J. & Barth, F.G. (1993) DNA sequence data indicates the polyphyly of the family Ctenidae (Araneae). Journal of Arachnology, 21, 194–201.
    Hyatt, K.H. (1954) The African spiders of the family Ctenidae in the collections of the British Museum (Natural History). Annals and Magazine of Natural History, (12) 7, 877–894.
    Jäger, P. (1998) First results of a taxonomic revision of the SE Asian Sparassidae (Araneae). In: Selden, P.A. (Ed.), Proceedings of the 17th European Colloquium of Arachnology. British Arach. Soc., Burnham Beeches, Bucks, Edinburgh, 1997, pp. 53–59.
    Jäger, P. (2001) Diversität der Riesenkrabbenspinnen im Himalaya. Über eine Radiation zweier Gattungen in den Schneetropen (Araneae: Sparassidae: Heteropodinae). Courier Forschungsinstitut Senckenberg, 232, 1–136.
    Jäger, P. (2006) Lengthening of embolus and intromittent duct: a review of an evolutionary trend in the spider family Sparassidae (Arachnida: Araneae). In: Deltshev, C. & Stoev, P. (Eds.), European Arachnology 2005. Acta zoologica bulgarica, Suppl. No 1, 49–62.
    Jäger, P. (2007) Spiders (Araneae) from Laos with descriptions of new species. Acta Arachnologica, 56 (1), 29–58.
    Jäger, P. (2008a) Revision of the huntsman spider genus Heteropoda Latreille 1804: species with exceptional male palpal conformations (Araneae: Sparassidae: Heteropodinae). Senckenbergiana biologica, 88 (2), 239–310.
    Jäger, P. (2008b) Barylestis Simon, 1910 -- first record in Asia with comments on its zoogeography (Araneae: Sparassidae: Heteropodinae). Arthropoda Selecta, 17, 105–110.
    Jäger, P. (2012) Asian species of the genera Anahita Karsch 1879, Ctenus Walckenaer 1805 and Amauropelma Raven, Stumkat & Gray 2001 (Arachnida: Araneae: Ctenidae). Zootaxa, 3429 (1), 1–63.
    Jäger, P. (2013) Ctenus monaghani spec. nov., a nocturnal hunter from the forest floor in Laos (Araneae: Ctenidae). Zootaxa, 3670 (1), 91–93.
    Jäger, P. (2014) Heteropoda Latreille 1804: new species, synonymies, transfers and records (Araneae: Sparassidae: Heteropodinae). Arthropoda Selecta, 23 (2) (Special Issue Pekka Lehtinen 80th Anniversary), 145–188.
    Jäger, P. (2020) The spider genus Olios Walckenaer, 1837 (Araneae: Sparassidae)—Part 1: species groups, diagnoses, identification keys, distribution maps and revision of the argelasius-, coenobitus- and auricomis-groups. Zootaxa, 4866 (1), 1–119.
    Jäger, P. & Kulkarni, S. (2016) An unexpected new species of the genus Pseudopoda (Araneae: Sparassidae: Heteropodinae) from the Western Ghats in India. Zookeys, 577, 55–62.
    Jäger, P. & Minn, M.Z. (2015) New species in the family Ctenidae Keyserling 1877 from high altitude habitats in Myanmar, with the first case of penetration of the female’s cuticle by a male in the RTA-clade (Arachnida: Araneae: Ctenidae). Zootaxa, 3994 (2), 235–252.
    Jäger, P. & Ono, H. (2000) Sparassidae of Japan. I. New species of Olios, Heteropoda, and Sinopoda, with notes on some known species (Araneae: Sparassidae: Sparassinae and Heteropodinae). Acta Arachnologica, 49, 41–60.
    Jocqué, R. & Steyn, T. (1997) Petaloctenus, a new genus of Ctenidae from West-Africa (Arachnida, Araneae). Bulletin de l’Institut Royal des Sciences Naturelles de Belgique (Entomologie), 67, 107–117.
    Karsch, F. (1879a) Baustoffe zu einer Spinnenfauna von Japan. Verhandlungen des naturhistorischen Vereins der preussischen Rheinlande und Westfalens, 36, 57–105.
    Karsch, F. (1879b) Sieben neue Spinnen von Sta Martha. Entomologische Zeitschrift, Stettin, 40, 106–109.
    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, pls. 10–12.
    Karthika, K. & Sunil Jose, K. (2020) New distributional record of Ctenus Indicus (Gravely 1931) from Western Ghats, Kerala. International Journal of Entomology Research, 5 (6), 54–56.
    Keyserling, E. (1877) Ueber amerikanische Spinnenarten der Unterordnung Citigradae. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien, 26, 609–708.
    Keyserling, E. (1891) Die Spinnen Amerikas. Brasilianische Spinnen. Bauer & Raspe, Nürnberg, 3, 1–278.
    Kim, J.P. & Ye, S.H. (2014) A new species of the genus Ctenus Walckenaer, 1805 (Araneae: Ctenidae) from Sri Lanka. Korean Arachnology, 30 (1), 61–70.
    Koch, C.L. (1847) Die Arachniden. J.L. Lotzbeck, Nürnberg, Vierzehnter Band, pp. 89–210., pls. 481–504., Fünfzehnter Band, pp. 1–136, pls. 505–540., Sechszehnter und letzter Band, pp. 1–80, pls. 541–563., Index 64 pp.
    Koch, C.L. (1850) Übersicht des Arachnidensystems. Heft 5. J. L. Lotzbeck, Nürnberg, 77 pp.
    Koch, L. (1878a) Die Arachniden Australiens, nach der Natur beschrieben und abgebildet. Bauer & Raspe, Nürnberg, 969–1044., pl. 85–91.
    Koch, L. (1878b) Japanesische Arachniden und Myriapoden. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien, 27, 735–798., pls. 15–16.
    Kraus, O. (1955) Spinnen aus El Salvador (Arachnoidea, Araneae). Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 493, 1–112.
    Latreille, P.A. (1804) Tableau methodique des Insectes. Nouveau Dictionnaire d’Histoire Naturelle, Paris, 24, 129–295.
    Lehtinen, P.T. (1967) Classification of the cribellate spiders and some allied families, with notes on the evolution of the suborder Araneomorpha. Annales Zoologici Fennici, 4, 199–468.
    Lenz, H. (1886) Beiträge zur Kenntniss der Spinnenfauna Madagascars. Zoologische Jahrbücher, Zeitschrift für Systematik, Geographie und Biologie der Thiere, 1 (2), 379–408., pl. 10.
    Lessert, R. de (1935) Description de deux araignées cavernicoles du Congo belge. Revue de Zoologie et de Botanique Africaines, 27, 326–332.
    Lin, S.-M., Chen, C.A. & Lue, K.-Y. (2002) Molecular phylogeny and biogeography of the grass lizards genus Takydromus (Reptilia: Lacertidae) of East Asia. Molecular Phylogenetics and Evolution, 22, 276–288.
    Linnaeus, C. (1767) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus differentiis, synonymis, locis. Editio duodecima, reformata. Laurentius Salvius, Holmiae, 533–1327.
    Lo, Y.-Y., Cheng, R.-C. & Lin, C.-P. (2021) Species delimitation and taxonomic revision of Oxyopes (Araneae: Oxyopidae) of Taiwan, with description of two new species. Zootaxa, 4927 (1), 58–86.
    Martins, R. & Bertani, R. (2007) The non-Amazonian species of the Brazilian wandering spiders of the genus Phoneutria Perty, 1833 (Araneae: Ctenidae), with the description of a new species. Zootaxa, 1526 (1), 1–36.
    Marusik, Y.M. & Omelko, M.M. (2016) Redescription of two Ctenidae (Araneae) from Taiwan. Zootaxa, 4173 (2), 155–162.
    Marusik, Y.M., Zhang, F. & Omelko, M.M. (2012) A new genus and species of ctenid spiders (Aranei: Ctenidae) from south China. Arthropoda Selecta, 21 (1), 61–66.
    Mello-Leitão, C. (1929) Aranhas do Pernambuco, colhidas por D. Bento Pickel. Anais da Academia Brasileira de Ciências, 1, 91–112.
    Mello-Leitão, (1930) Aranhas do Cuminá. Arquivos do Museu Nacional do Rio de Janeiro, 32, 51–75.
    Mello-Leitão, (1941) Las arañas de la provincia de Santa Fe colectadas por el Profesor Birabén. Revista del Museo de La Plata (New Series, Zoology), 2, 199–225.
    Merian, P. (1911) Die Spinnenfauna von Celebes. Beiträge zur Tiergeographie im Indoaustralischen Archipel. Zoologische Jahrbücher, Abteilung für Systematik, Geographie und Biologie der Tiere, 31, 165–354.
    Miller, J.A. & Rahmadi, C. (2012) A troglomorphic spider from Java (Araneae, Ctenidae, Amauropelma). ZooKeys, 163, 1–11.
    Ono, H. (2010) Four new spiders (Arachnida, Araneae) of the families Liphistiidae, Ctenizidae, Araneidae and Ctenidae from Vietnam. Memoirs of the National Museum for Nature and Science, 46, 1–12.
    Patel, B.H. & Reddy, T.S. (1988) Two new species of genus Ctenus Walckenaer (Araneae: Ctenidae) from coastal Andhra Pradesh, India. Entomon, 13, 103–107.
    Pereira, M.P., Labarque, F.M. & Polotow, D. (2020) Description of a new Neotropical spider genus in the family Ctenidae Keyserling, 1877 (Araneae: Lycosoidea) from the Brazilian Amazon rainforest. Zootaxa, 4890 (3), 375–396.
    Perty, M. (1833) Arachnides Brasilienses. In: de Spix, J.B. & Martius, F.P. (eds) Delectus animalium articulatorum quae in itinere per Braziliam ann. 1817 et 1820 colligerunt. Monachii [=Munich], pp. 191–209., pls. 38–39.
    Petrunkevitch, A. (1910) Some new or little known American Spiders. Annals of the New York Academy of Science, 19, 205–224.
    Petrunkevitch, A. (1925). Arachnida from Panama. Transactions of the Connecticut Academy of Arts and Sciences, 27, 51–248.
    Pickard-Cambridge, O. (1876) Catalogue of a collection of spiders made in Egypt, with descriptions of new species and characters of a new genus. Proceedings of the Zoological Society of London, 44 (3), 541–630, pl. 58–60.
    Pickard-Cambridge, F.O. (1897) On cteniform spiders from the lower Amazons and other regions of North and South America, with a list of all known species of these groups hitherto recorded from the New World. The Annals and Magazin of Natural History, (6) 19, 52–106.
    Pickard-Cambridge, F.O. (1898) On the cteniform spiders of Africa, Arabia and Syria. Proceedings of the Zoological Society of London, 66 (1), 13–32, pls III–IV.
    Pickard-Cambridge, F.O. (1902) New species of spiders belonging to the genus Ctenus, with supplementary notes. The Annals and Magazin of Natural History, (7) 9, 401–415.
    Platnick, N.I. (1989) Advances in spider taxonomy 1981-1987: a supplement to Brignoli’s A catalogue of the Araneae described between 1940 and 1981. Manchester University Press, Manchester, 673 pp.
    Platnick, N.I. (1993) Advances in spider taxonomy 1988-1991, with synonymies and transfers 1940-1980. The New York Entomological Society, New York, 846 pp.
    Platnick, N.I. (1998) Advances in spider taxonomy 1992-1995 with redescriptions 1940-1980. New York Entomological Society, New York, 976 pp.
    Platnick, N.I. (2001) The world spider catalog, version 2.0. American Museum of Natural History. Available from: (accessed 12 May 2022)
    Pocock, R.I. (1897) Spinnen (Araneae). In: Kükenthal, W. (Ed.), Ergebnisse einer zoologische Forschungsreise in dem Molukken und Borneo. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 23, 591–629.
    Pocock, R.I. (1898) Scorpions, pedipalpi and spiders collected by Dr Willey in New Britain, the Solomon Islands, Loyalty Islands, etc. In: Willey, A. (ed.) Zoological results based on material from New Britain, New Guinea, Loyalty Islands and elsewhere, collected during the years 1895, 1896 and 1897. Part I. University Press, Cambridge, pp. 95–120., pls. 10–11.
    Polotow, D. & Brescovit, A.D. (2009a) Revision and cladistics analysis of Isoctenus and description of a new neotropical genus (Araneae, Ctenidae, Cteninae). Zoological Journal of the Linnean Society, 155, 583–614.
    Polotow, D. & Brescovit, A.D. (2009b) Description of Toca, a new neotropical spider genus (Araneae, Ctenidae, Calocteninae). Journal of Arachnology, 37, 243–245.
    Polotow, D. & Brescovit, A.D. (2009c) Revision of the new wandering spider genus Ohvida and taxonomic remarks on Celaetycheus Simon, 1897 (Araneae: Ctenidae). Zootaxa, 2115 (1), 1–20.
    Polotow, D. & Brescovit, A.D. (2012) Redescription of five Amazonian species of Ctenus Walckenaer (Araneae, Ctenidae, Cteninae). Zootaxa, 3338 (1), 49–59.
    Polotow, D. & Brescovit, A.D. (2013) New species of the Neotropical spider genus Celaetycheus Simon, 1897 (Araneae: Ctenidae). Zootaxa, 3637 (2), 139–c157.
    Polotow, D. & Brescovit, A.D. (2014) Phylogenetic analysis of the tropical wolf spider subfamily Cteninae (Arachnida, Araneae, Ctenidae). Zoological Journal of the Linnean Society, 170 (2), 333–361.
    Polotow, D. & Brescovit, A.D. (2018) Kiekie, a new Neotropical spider genus of Ctenidae (Cteninae, Araneae). Zootaxa, 4531 (3), 353–373.
    Polotow, D. & Brescovit, A.D. (2019) Guasuctenus gen. nov., a new Neotropical spider genus of Ctenidae (Araneae). Zootaxa, 4624 (4), 539–550.
    Polotow, D. & Jocqué, R. (2015) Afroneutria, a new spider genus of Afrotropical Ctenidae (Arachnida, Araneae). European Journal of Taxonomy, 121, 1–27.
    Raven, R.J., Stumkat, K. & Gray, M.R. (2001) Revisions of Australian ground-hunting spiders: I. Amauropelma gen. nov. (Araneomorphae: Ctenidae). Records of the Western Australian Museum, Supplement, 64, 187–227.
    Raychaudhuri, D. & Saha, S. (2015) Spiders (Araneae: Arachnida) of reserve forests of Dooars: Gorumara National Park, Chapramari Wildlife Sanctuary and Mahananda Wildlife Sanctuary. World Scientific News, 20, 1–336.
    Raychaudhuri, D. & Saha, S. (2017) Litter and ground dwelling spiders (Araneae: Arachnida) of reserve forests of Dooars, West Bengal. World Scientific News, 63, 1–240.
    Reddy, T.S. & Patel, B.H. (1994) A new species of genus Ctenus Walckenaer (Araneae: Ctenidae) from India. Entomon, 19, 131–133.
    Rheims, C.A. (2020) Revision of the spider genus Sparianthis Simon, 1880 (Araneae, Sparassidae, Sparianthinae). Zootaxa, 4890 (2), 151–191.
    Roewer, C.F. (1938) Araneae. In: Résultats scientifiques du Voyage aux indes orientales néerlandaises de la SS. AA. RR. le Prince et la Princesse Leopold de Belgique. Mémoires du Musée Royal d’Histoire Naturelle de Belgique, 3 (19), 1–94.
    Roewer, C.F. (1955) Katalog der Araneae von 1758 bis 1940, bzw. 1954. 2. Band, Abt. a (Lycosaeformia, Dionycha [excl. Salticiformia]). 2. Band, Abt. b (Salticiformia, Cribellata) (Synonyma-Verzeichnis, Gesamtindex). Institut royal des Sciences naturelles de Belgique, Bruxelles, 1751 pp.
    Sankaran, P.M. & Sebastian, P.A. (2018) First record of Africactenus Hyatt, 1954 and redescriptions of two poorly known species of Ctenus Walckenaer, 1805 (Araneae, Ctenidae, Cteninae) from India. Zootaxa, 4388 (3), 395–406.
    Schenkel, E. (1953) Bericht über einige Spinnentiere aus Venezuela. Verhandlungen der Naturforschenden Gesellschaft in Basel, 64, 1–57.
    Schmidt, G. (1956) Genus- und Speziesdiagnosen neuer, mit Bananen eingeschleppter Spinnen nebst Mitteilung über das Auffinden der Männchen zweier Spinnenarten. Zoologischer Anzeiger, 157, 24–31.
    Silva-Davila, D. (2003) Higher-level relationships of the spider family Ctenidae (Araneae: Ctenoidea). Bulletin of the American Museum of Natural History, 274, 1–86.<0001:HLROTS>2.0.CO;2
    Simon, E. (1876) Les arachnides de France. Tome troisième. Roret, Paris, 364 pp., pls. IX–XIII.
    Simon, E. (1884) Arachnides recueillis en Birmanie par M. le chevalier J. B. Comotto et appartenant au Musée civique d’histoire naturelle de Gènes. Annali del Museo Civico di Storia Naturale di Genova, 20, 325–372.
    Simon, E. (1885) Matériaux pour servir à la faune arachnologiques de l’Asie méridionale. I. Arachnides recueillis à Wagra-Karoor près Gundacul, district de Bellary par M. M. Chaper. II. Arachnides recueillis à Ramnad, district de Madura par M. l’abbé Fabre. Bulletin de la Société Zoologique de France, 10, 1–39, pl. 10.
    Simon, E. (1886) Etudes arachnologiques. 18e Mémoire. XXVI. Matériaux pour servir à la faune des Arachnides du Sénégal. (Suivi d’une appendice intitulé: Descriptions de plusieurs espèces africaines nouvelles). Annales de la Société Entomologique de France, (6) 5, 345–396.
    Simon, E. (1887a) Observation sur divers arachnides: synonymies et descriptions. Annales de la Société Entomologique de France, (6) 7 (Bull.), 158–159, 167, 175–176, 186–187, 193–195.
    Simon, E. (1887b) Etude sur les arachnides de l’Asie raignéese faisant partie des collections de l’Indian Museum (Calcutta). I. Arachnides recueillis a Tavoy (Tenasserim) par Moti Ram. Journal of the Asiatic Society Bengal, 56, 101–117.
    Simon, E. (1888) Etudes arachnologiques. 21e Mémoire. XXIX. Descriptions d’espèces et de genres nouveaux de l’Amérique centrale et des Antilles. Annales de la Société Entomologique de France, (6) 8, 203–216.
    Simon, E. (1890) Etudes arachnologiques. 22e Mémoire. XXXIV. Etude sur les arachnides de l’Yemen. Annales de la Société Entomologique de France, (6) 10, 77–124.
    Simon, E. (1891) Descriptions de quelques arachnides du Costa Rica communiqués pa M. A. Getaz (de Genève). Bulletin de la Société Zoologique de France, 16, 109–112.
    Simon, E. (1897a) Histoire naturelle des araignées. Deuxième édition, tome second. Roret, Paris, pp. 1–192.
    Simon, E. (1897b) Etudes arachnologiques. 27e Mémoire. XLII. Descriptions d’espèces nouvelles de l’ordre des Araneae. Annales de la Société Entomologique de France, 65, 465–510.
    Simon, E. (1898) Histoire naturelle des araignées. Deuxième édition, tome second. Roret, Paris, pp. 193–380.
    Simon, E. (1905) Arachnides de Java, recueillis par le Prof. K. Kraepelin en 1904. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 22, 49–73.
    Simon, E. (1909) Arachnides recueillis par L. Fea sur la côte occidentale d’Afrique. 2e partie. Annali del Museo Civico di Storia Naturale di Genova, 44, 335–449.
    Strand, E. (1906) Sumatra und Neu-Guinea Spinnen des naturhistorischen Museums zu Wiesbaden. Jahrbücher des nassauischen Vereins für Naturkunde, 59, 257–278.
    Strand, E. (1908) Neue außereuropäische Spinnen. Zoologischer Anzeiger, 33, 5–7.
    Strand, E. (1909) Neue oder wenig bekannte neotropische cteniforme Spinnen des Berliner Museums. Zoologische Jahrbücher, Abteilung für Systematik, Geographie und Biologie der Tiere, 28 (4), 401–428.
    Strand, E. (1911) Araneae von den Aru- und Kei-Inseln. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 34, 127–199.
    Strand, E. (1916) Systematische-faunistische Studien über paläarktische, afrikanische und amerikanische Spinnen des Senckenbergischen Museums. Archiv für Naturgeschichte, 81 (A9), 1–153.
    Sundevall, C.J. (1833) Conspectus Arachnidum. C. F. Berling, Londini Gothorum, pp. 1–39.
    Taczanowski, L. (1874) Les aranéides de la Guyane française. Horae Societatis Entomologicae Rossicae, 10, 56–115.
    Thorell, T. (1873) Remarks on synonyms of European spiders. Part IV. C. J. Lundström, Uppsala, pp. 375-645.
    Thorell, T. (1878) Studi sui ragni Malesi e Papuani. II. Ragni di Amboina raccolti Prof. O. Beccari. Annali del Museo Civico di Storia Naturale di Genova, 13, 5–317.
    Thorell, T. (1887) Viaggio di L. Fea in Birmania e regioni vicine. II. Primo saggio sui ragni birmani. Annali des Museo civico di storia naturale Genova, 25, 5–417.
    Thorell, T. (1890) Diagnoses aranearum aliquot novarum in Indo-Malesia inventarum. Annali del Museo Civico di Storia Naturale di Genova, 30, 132–172.
    Thorell, T. (1892) Studi sui ragni Malesi e Papuani. IV, 2. Annali del Museo Civico di Storia Naturale di Genova, 31, 1–490.
    Thorell, T. (1895) Descriptive catalogue of the spiders of Burma. London, 406 pp.
    Thorell, T. (1897) Araneae paucae Asiae australis. Bihang till Kongliga Svenska Vetenskaps-Akademiens Handlingar, 22 (IV, 6), 1–36.
    Tikader, B.K. (1973) A new species of rare spider of the genus Ctenus (family-Ctenidae) from Andaman Islands, India. Current Science, 42, 862–863.
    Tikader, B.K. (1976) A new species of spider of the genus Ctenus (family: Ctenidae) from Meghalaya, India. Journal of the Bombay natural History Society, 72, 791–793.
    Tikader, B.K. & Malhotra, M.S. (1981) Revision of spiders of the genus Ctenus Walckenaer from India (Araneae: Ctenidae). Records of the Zoological Survey of India, 79, 105–124.
    Töpfer-Hofmann, G., Cordes, D. & Helversen, O. von (2000) Cryptic species and behavioural isolation in the Pardosa lugubris group (Araneae, Lycosidae), with description of two new species. Bulletin of the British arachnological Society, 11 (7), 257–274.
    Wagner, W.A. (1887) Copulationsorgane des Männchens als Criterium für die Systematik der Spinnen. Horae Societatis Entomologicae Rossicae, 22, 3–132., pl. 1–10.
    Walckenaer, C.A. (1805) Tableau des aranéides ou caractères essentiels des tribus, genres, familles et races que renferme le genre Aranea de Linné, avec la désignation des espèces comprises dans chacune de ces divisions. Paris, 88 pp.
    Walckenaer, C.A. (1837) Histoire naturelle des insectes. Aptères. Tome premier. Roret, Paris, 682 pp., pls. 1–15.
    Walckenaer, C.A. (1847) Dernier Supplément. In: Walckenaer, C.A. & Gervais, P. (Eds.), Histoire naturelles des Insects. Aptères. Tome quatrième. Roret, Paris, pp. 365–596.
    Wheeler, W.C., Coddington, J.A., Crowley, L.M., Dimitrov, D., Goloboff, P.A., Griswold, C.E., Hormiga, G., Prendini, L., Ramírez, M.J., Sierwald, P., Almeida-Silva, L.M., Álvarez-Padilla, F., Arnedo, M.A., Benavides, L.R., Benjamin, S.P., Bond, J.E., Grismado, C.J., Hasan, E., Hedin, M., Izquierdo, M.A., Labarque, F.M., Ledford, J., Lopardo, L., Maddison, W.P., Miller, J.A., Piacentini, L.N., Platnick, N.I., Polotow, D., Silva-Dávila, D., Scharff, N., Szűts, T., Ubick, D., Vink, C., Wood, H.M. & Zhang, J.X. (2017) The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling. Cladistics, 33 (6), 576–616.
    World Spider Catalog (2022) World Spider Catalog. Version 23.5. Natural History Museum Bern. Available from: (accessed 28 June 2022)
    Yeh, W.-B, Chang, Y.-L., Lin, C.-H., Wu, F.S. & Yang, J.T. (2004) Genetic differentiation of Loxoblemmus appendicularis complex (Orthoptera: Gryllidae): speciation through vicariant and glaciation events. Annals of the Entomological Society of America, 97 (4), 613–623.[0613:GDOLAC]2.0.CO;2
    Yoshida, H. (1998) A new species of the genus Ctenus (Araneae: Ctenidae) from the Yaeyama Islands, Japan and Taiwan. Acta arachnologica, 47, 117–120.
    Yoshida, H. (2009) Ctenidae. In: Ono, H. (Ed.), The Spiders of Japan with keys to the families and genera and illustrations of the species. Tokai Univ. Press, Kanagawa, pp. 467–468.
    Zhang, F., Hu, D.S. & Han, G.X. (2011) A new species of the genus Anahita Karsch, 1879 (Araneae: Ctenidae) from Hainan Island, China. Zootaxa, 2839 (1), 85–88.
    Zhu, M.S., Chen, H.M. & Song, D.X. (1999) A new species of the genus Anahita (Araneae: Ctenidae) from China. In: The China Zoological Society: Zoological Studies in China. Chinese Forestry Publishing House, Beijing, pp. 210–212.