Systematic revision of the pantropical whip spider family Charinidae Quintero, 1986 (Arachnida, Amblypygi)
Abstract
The whip spider family Charinidae Quintero, 1986 is the most speciose and widely distributed in the arachnid order Amblypygi Thorell, 1883. It comprises three genera and 95 species distributed across all tropical continents and the eastern Mediterranean. Despite recent advances in the taxonomy of the family, a global revision of all its species, necessary to advance understanding of its systematics, biogeography and evolution, has never been conducted. In the present contribution, the family is revised in its entirety for the first time, including all previous names and 33 new species, 24 in the genus Charinus Simon, 1892: C. alagoanus sp. nov., C. apiaca sp. nov., C. carinae sp. nov., C. carioca sp. nov., C. carvalhoi sp. nov., C. cearensis sp. nov., C. diamantinus sp. nov., C. euclidesi sp. nov., C. goitaca sp. nov., C. guayaquil sp. nov., C. imperialis sp. nov., C. loko sp. nov., C. magalhaesi sp. nov., C. miskito sp. nov., C. mocoa sp. nov., C. monasticus sp. nov., C. palikur sp. nov., C. perquerens sp. nov., C. puri sp. nov., C. renneri sp. nov., C. sooretama sp. nov., C. souzai sp. nov., C. susuwa sp. nov., C. una sp. nov.; eight in the genus Sarax Simon, 1892: S. bilua sp. nov., S. dunni sp. nov., S. gravelyi sp. nov., S. indochinensis sp. nov., S. lembeh sp. nov., S. palau sp. nov., S. rahmadii sp. nov., S. tiomanensis sp. nov.; and one in the genus Weygoldtia Miranda et al., 2018: W. consonensis sp. nov. Taxonomic keys to the 132 species (excluding four nomina dubia) are presented and several taxonomic rearrangements implemented. Four subspecies are elevated to species level: Charinus cavernicolus Weygoldt, 2006, C. elegans Weygoldt, 2006, C. longipes Weygoldt, 2006, and Sarax bispinosus (Nair, 1934). Sarax batuensis Roewer, 1962 is removed from synonymy with Sarax buxtoni (Gravely, 1915) and S. buxtoni newly synonymized with Sarax rimosus (Simon, 1901). Stygophrynus moultoni Gravely, 1915 is transferred to Sarax, resulting in Sarax moultoni (Gravely, 1915) comb. nov. Ten species are transferred from Charinus to Sarax, resulting in new combinations: S. abbatei (Delle Cave, 1986) comb. nov., S. bengalensis (Gravely, 1911) comb. nov., S. dhofarensis (Weygoldt, Pohl & Polak, 2002) comb. nov., S. ioanniticus (Kritscher, 1959) comb. nov., S. israelensis (Miranda et al., 2016) comb. nov., S. omanensis (Delle Cave, Gardner & Weygoldt, 2009) comb. nov., S. pakistanus (Weygoldt, 2005) comb. nov., S. seychellarum (Kraepelin, 1898) comb. nov., S. socotranus (Weygoldt, Pohl & Polak, 2002) comb. nov. and S. stygochthobius (Weygoldt & Van Damme, 2004) comb. nov.
References
Agapakis G. & Miranda G.S. 2019. First record of Charinus ioanniticus (Arachnida, Amblypygi, Charinidae) from continental Europe. Arachnologische Mitteilungen/Arachnology Letters 58: 13–15. https://doi.org/10.30963/aramit5805
Arldt T. 1908. Die Ausbreitung einiger Arachnidenordnung (Mygalomorphen, Skorpione, Pedipalpen, Solifugen, Palpigraden). Archiv für Naturgeschichte 74: 389–458.
Armas L.F. 2000a. La artropodofauna cavernicola de las Antillas Mayores. Boletín de la Sociedad Entomológica Aragonesa 27: 134–138.
Armas L.F. 2000b. Parthenogenesis in Amblypygi (Arachnida). Avicennia 12/13: 133–134.
Armas L.F. 2004. Arácnidos de República Dominicana. Palpigradi, Schizomida, Solifugae y Thely-phonida (Chelicerata: Arachnida). Revista Ibérica de Aracnología 2: 1–64.
Armas L.F. 2005. Notas sobre la biología reproductiva del amblipígido partenogenético Charinus acosta (Quintero, 1983) (Amblypygi: Charinidae). Boletín de la Sociedad Entomológica Aragonesa 36: 271–273.
Armas L.F. 2006a. Nueva especie de Charinus Simon, 1892 (Amblypygi: Charinidae) de Cuba central. Revista Ibérica de Aracnología 13: 167–170.
Armas L.F. 2006b. Sinopsis de los amblipígidos antillanos (Arachnida: Amblypygi). Boletín Sociedad Entomológica Aragonesa 38: 223–245.
Armas L.F. 2010. Nuevos arácnidos de Puerto Rico (Arachnida: Amblypygi, Araneae, Opiliones, Parasitiformes, Schizomida, Scorpiones). Boletín de la Sociedad Entomológica Aragonesa 47: 55–64.
Armas L.F. 2013a. Los amblipigios cavernícolas de Cuba (Arachnida: Amblypygi). Revista Ibérica de Aracnología 22: 15–19.
Armas L.F. 2013b. Pedipalpi (Arachnida: Amblypygi, Schizomida, Thelyphonida) de Cuba Occidental. Revista Ibérica de Aracnología 22: 91–94.
Armas L.F. 2014. Los amblipigios de Cuba (Arachnida: Amblypygi). Revista Ibérica de Aracnología 24: 29–51.
Armas L.F. 2017. Charinus victori Armas, 2010 is a synonym of Charinus aguayoi Moyá-Guzman, 2009 (Amblypygi: Charinidae). Revista Ibérica de Aracnología 31: 113–114.
Armas L.F. & Abud Antun A.J. 2000. El alacrán en la cultura de República Dominicana. Revista Ibérica de Aracnología 1: 77–79.
Armas L.F. & Alayón García G. 1984. Sinopsis de los arácnidos cavernícolas de Cuba (excepto ácaros). Poeyana 276: 1–25.
Armas L.F. & Alayón García G. 2014. Aracnofauna del Archipiélago de los Canarreos. Revista Cubana de Ciencias Biológicas 3:41–52.
Armas L.F. & Ávila Calvo A. 2000. Dos nuevos amblipígidos de Cuba, con nuevos sinónimos y registros (Arachnida: Amblypygi). Anales de la Escuela Nacional de Ciencias Biologicas, Mexico 46: 289–303.
Armas L.F. & Maes J.-M. 2000. Los amblipígidos de Nicaragua (Arachnida: Amblypygi). Revista Ibérica de Aracnología 1: 39–44.
Armas L.F. & Pérez A. 2001. Los amblipígidos de República Dominicana (Arachnida: Amblypygi). Revista Ibérica de Aracnología 3: 47–66.
Armas L.F. & Teruel R. 1997. A new Charinus (Amblypygi: Charontidae) from St. John, U.S. Virgin Islands. Avicennia 6/7: 43–46.
Armas L.F., García G.A. & Hernández J.M.R. 2009. Aracnofauna (excepto Acari) del Macizo Guamu-haya, Cuba Central. Primera Aproximación. Boletín Sociedad Entomológica Aragonesa 45: 135−146.
Armas L.F., Chiriví D.J., Botero-Trujillo R., Cortés G.P.C. & García-Hernández S. 2012. Presencia en Colombia de la familia Charinidae (Arachnida: Amblypygi). Boletín de la Sociedad Entomológica Aragonesa 50: 321–322.
Armas L.F., Palomino-Cárdenas A.C. & Castillo-Espinoza M.d. 2016. Amblipigios de los Departamentos Cusco y Madre de Dios, Perú, con la descripción de un nuevo Charinus (Amblypygi: Charinidae, Phrynidae). Revista Ibérica de Aracnología 28: 45–50.
Armas L.F., Trujillo R.E. & Agreda E.O. 2018. Sinopsis de los amblipigios de Guatemala (Arachnida: Amblypygi). Revista Ibérica de Aracnología 32: 55–72.
Ávila Calvo A. & Armas L.F. 1997. Lista de los amblipigidos (Arachnida: Amblypygi) de México, Centroamérica y las Antillas. Cocuyo 6: 31–32.
Ax P. 2001. Multicellular Animals: The Phylogenetic System of the Metazoa. Springer-Verlag, Berlin.
Baehr M. 1974. Wiederfund der einzigen bekannten australischen Geisselspinne Charinus pescotti Dunn 1949 (Chelicerata, Arachnida, Amblypygi). Entomologische Zeitschrift 84: 101–105.
Baert L. & Mahnert V. 2015. The distribution of the non-Araneae and non-Acari arachnids of Galápagos. Belgian Journal of Entomology 28: 1–76.
Baert L., Maelfait J.-P. & Desender K. 1996. Distribution of the arachnid species of the orders Scorpiones, Solifugae, Amblypygi, Schizomida, Opiliones and Pseudoscorpiones in Galápagos. Bulletin de l’Institut royal des Sciences naturelles de Belgique, Entomologie 65: 5–19.
Ballón-Estacio R.J. & Armas L.F. 2019. Una especie nueva de Charinus (Amblypygi: Charinidae) del Departamento de Huánuco, Perú. Revista Ibérica de Aracnología 35: 15–19.
Banks N. 1902. Papers from the Hopkins Stanford Galapagos Expedition, 1898–1899. VII. Entomological results (6), Arachnida, part 1. Proceedings of the Washington Academy of Sciences 4: 49–70.
Baptista R.L.C. & Giupponi A.P.L. 2002. A new troglomorphic Charinus from Brazil (Arachnida: Amblypygi: Charinidae). Revista Ibérica de Aracnología 6: 105–110.
Baptista R.L.C. & Giupponi A.P.L. 2003. A new troglomorphic Charinus from Minas Gerais State, Brazil (Arachnida: Amblypygi: Charinidae). Revista Ibérica de Aracnología 7: 79–84.
Barnes R.D. 1987. Invertebrate Zoology. Saunders College Publishing, Philadelphia.
Barrows W.M. 1925. Modification and development of the arachnid palpal claw, with special reference to spiders. Annals of the Entomological Society of America XVIII: 483–516. https://doi.org/10.1093/aesa/18.4.483
Benoit P.L.G. 1979. Contributions à l’étude de la faune terrestre des îles granitiques de l’archipel des Séchelles. (Mission P.L.G. Benoit – J.J. van Mol 1972. Amblypygi et Scorpiones). Revue de Zoologie africaine 93: 458–460.
Blick T. & Seiter M. 2016. Whip spiders (Amblypygi, Arachnida) of the Western Palaearctic – a review. Zootaxa 4161 (4): 586–592. https://doi.org/10.11646/zootaxa.4161.4.11
Börner C. 1904. Beiträge zur Morphologie der Arthropoden. I. Ein Beitrag zur Kenntnis der Pedipalpen. Zoologie 42: 1–174. https://doi.org/10.5962/bhl.title.59544
Bristowe W.S. 1952. The arachnid fauna of the Batu Caves in Malaya. Annals and Magazine of Natural History 12: 697–707. https://doi.org/10.1080/00222935208654340
Butler A.G. 1873. A monographic revision of the genus Phrynus, with descriptions of four remarkable new species. Annals and Magazine of Natural History, Serie 4 12: 117–125. https://doi.org/10.1080/00222937308680723
Buxton B.H. 1917. Notes on the anatomy of arachnids. Journal of Morphology 29: 1–31. https://doi.org/10.1002/jmor.1050290102
Caporiacco L. di. 1947. Diagnosi preliminari di specie nuove di aracnidi della Guiana Britannica raccolte dai Professori Beccari e Romiti. Monitore Zoologico Italiano 56: 20–34.
Caporiacco L. di. 1948. Arachnida of British Guiana collected in 1931 & 1936 by Professors Beccari and Romiti. Proceedings of the Zoological Society of London 118: 607–747. https://doi.org/10.1111/j.1096-3642.1948.tb00402.x
Carnaval A.C., Waltari E., Rodrigues M.T., Rosauer D., VanDerWal J., Damasceno R., Prates I., Strangas M., Spanos Z., Rivera D., Pie M.R., Firkowski C.R., Bornschein M.R., Ribeiro L.F. & Moritz C. 2014. Prediction of phylogeographic endemism in an environmentally complex biome. Proceedings of the Royal Society B 281 (1792): 20141461. https://doi.org/10.1098/rspb.2014.1461
Chapin K.J. & Hebets E.A. 2016. The behavioral ecology of amblypygids. Journal of Arachnology 44: 1–14. https://doi.org/10.1636/v15-62.1
Cloudsley-Thompson J.L. 1968. Spiders, Scorpions, Centipedes & Mites, Revised Edition. Pergamon Press, New York. https://doi.org/10.1016/B978-0-08-012322-6.50015-8
Colla A., Legittimo C.M., Castellucci F., Simeon E. & Miranda G.S. 2020. First record of Amblypygi from Italy: Charinus ioanniticus (Charinidae). Arachnologische Mitteilungen/Arachnology Letters 18: 642–648. https://doi.org/10.13156/arac.2020.18.6.642
Cope E.D. 1886a. An analytical table of the genera of snakes. Proceedings of the American Philosophical Society 23: 479–499.
Cope E.D. 1886b. Three problematical genera of Mexican boaeform snakes. American Naturalist 20: 293–294.
Delle Cave L. 1986. Biospeleology of the Somaliland Amblypygi (Arachnida, Chelicerata) of the caves of the Showli Berdi and Mugdile (Bardera, Somaliland). Redia 69: 143–170.
Delle Cave L., Gardner A.S. & Weygoldt P. 2009. A new troglomorphic whip spider of the genus Charinus from the Sultanate of Oman (Amblypygi: Charinidae). Fauna of Arabia 24: 129–134.
Dittmar K., Porter M.L., Price L., Svenson G. & Whitling M.F. 2005. A brief survey of invertebrates in caves of Peninsular Malaysia. Malayan Nature Journal 57: 221–233.
Dunlop J.A. 1994. An Upper Carboniferous amblypygid from the Writhlington Geological Nature Reserve. Proceedings of the Geologists’ Association 105: 245–250. https://doi.org/10.1016/S0016-7878(08)80177-0
Dunlop J.A. & Martill D.M. 2002. The first whipspider (Arachnida: Amblypygi) and three new whipscorpions (Arachnida: Thelyphonida) from the Lower Cretaceous Crato Formation of Brazil. Transactions of the Royal Society of Edinburgh, Earth Sciences 92: 325–334. https://doi.org/10.1017/S0263593300000262
Dunn R.A. 1949. New Pedipalpi from Australia and the Solomon Islands. Memoirs of the National Museum of Victoria 16: 7–15. https://doi.org/10.24199/j.mmv.1949.16.01
El-Hennawy H.K. 2002. The first record of Amblypygi from Egypt. Journal of Arachnology 30: 452–453. https://doi.org/cpm8fj
Engel M.S. & Grimaldi D.A. 2014. Whipspiders (Arachnida: Amblypygi) in amber from the Early Eocene and mid-Cretaceous, including maternal care. Novitates Paleoentomologicae 9: 1–17. https://doi.org/10.17161/np.v0i9.4765
Fage L. 1929. Fauna of the Batu Caves, Selangor. X. Arachnida: Pedipalpi (part) and Araneae. Journal of the Federated Malay States Museums 14: 356–364.
Fage L. 1939. Les Pédipalpes africains du genre Charinus à propos d’une espèce nouvelle du Fouta-Djalon: Charinus milloti, n. sp. Bulletin de la Société entomologique de France 44: 153–160.
Fage L. 1946a. Complément à la faune des arachnides de Madagascar. Bulletin du Muséum national d’histoire naturelle, Paris 2: 256–267.
Fage L. 1946b. Scorpions et pédipalpes de l’Indochine Française. Annales de la Société entomologique de France 113: 71–81.
Fage L. 1954. Remarques sur la distribution géographique des pédipalpes amblypyges africains, accompagnées de la description d’une espèce nouvelle de Madagascar: Charinus madagascariensis, nov. sp. Annales du Musée du Congo belge, Sciences zoologiques 1: 180–184.
Fage L. & Simon E. 1936. Mission scientifique de l’Omo, Tome III. – Fascicule 30. Arachnida III. Pedipalpi, Scorpiones, Solifuga et Araneae (1re partie). Mémoires du Muséum national d’histoire naturelle, Paris 4: 293–340.
Gainett G., Ballesteros J.A., Kanzler C.R., Zehms J.T., Zern J.M., Aharon S., Gavish-Regev E. & Sharma P.P. 2020. Systemic paralogy and function of retinal determination network homologs in arachnids. BMC Genomics 21: 811. https://doi.org/10.1186/s12864-020-07149-x
Galán C. & Herrera F.F. 2006. Fauna cavernícola de Venezuela: Una Revisión. Boletín de la Sociedad Venezolana de Espeleología 40: 39–57.
Garwood R.J., Dunlop J.A., Knecht B.J. & Hegna T.A. 2017. The phylogeny of fossil whip spiders. BMC Evolutionary Biology 17: 105. https://doi.org/10.1186/s12862-017-0931-1
Gaubert P. 1892. Recherches sur les organes des sens et sur les systemes tegumentaires, glandulaires et musculaires des appendices des Arachnides. Annales des Sciences naturelles, Zoologie et Paléontologie 7: 31–184.
Gibbons A.T., Idnurm A., Seiter M., Paul S. Dyer M.K., Goodacre S.L., Gorb S.N. & Wolff J.O. 2019. Amblypygid-fungal interactions: The whip spider exoskeleton as a substrate for fungal growth. Fungal Biology 123: 497–506. https://doi.org/10.1016/j.funbio.2019.05.003
Giltay L. 1931. Scorpions et Pédipalpes. In: Résultats scientifiques du voyage aux Indes orientales néerlandaises de LL. AA. RR. le Prince et la Princesse Léopold de Belgique. Mémoires du Musée royal d’Histoire naturelle de Belgique 3: 1–28.
Giupponi A.P.L. & Baptista R.L.C. 2003. A case of homonymy in Charinus Simon, 1892 (Amblypygi: Charinidae). Revista Ibérica de Aracnología 7: 173–174.
Giupponi A.P.L. & Kury A.B. 2002. A new species of Charinus Simon, 1892 from northeastern Brazil (Arachnida, Amblypygi, Charinidae). Boletim do Museu Nacional 477: 1–7.
Giupponi A.P.L. & Kury A.B. 2013. Two new species of Heterophrynus Pocock, 1894 from Colombia with distribution notes and a new synonymy (Arachnida: Amblypygi: Phrynidae). Zootaxa 3647: 329–342. https://doi.org/10.11646/zootaxa.3647.2.5
Giupponi A.P.L. & Miranda G.S. 2012. A new species of Sarax Simon, 1892 from the Philippines (Arachnida: Amblypygi: Charinidae). Anais da Academia Brasileira de Ciências 84: 165–173. https://doi.org/10.1590/S0001-37652012000100017
Giupponi A.P.L. & Miranda G.S. 2016. Eight new species of Charinus Simon, 1892 (Arachnida: Amblypygi: Charinidae) endemic for the Brazilian Amazon, with notes on their conservational status. PLoS One 11: e0148277. https://doi.org/10.1371/journal.pone.0148277
Gonçalves-Souza T., Giupponi A.P.L. & Hernandes F.A. 2014. A rare finding of mites (Arachnida: Acari: Leeuwenhoekiidae) parasitising a whip spider (Arachnida: Amblypygi: Charinidae). Folia Parasitologica 61:182–184.
González-Sponga M.A. 1998. Arachnida of Venezuela. Two new species of the genus Charinides Gravely, 1911 (Amblypygi: Charontidae). Acta Biologica Venezuelica 18: 1–8.
Goodnight C.J. & Goodnight M.L. 1946. A new species of pedipalp from South America. Transactions of the American Microscopical Society 65: 323–327. https://doi.org/10.2307/3223544
Gravely F.H. 1911. Notes on Pedipalpi in the collection of the Indian Museum. I. New Pedipalpi from Calcutta. II. A preliminary note on a new Sarax from Singapore. Records of the Indian Museum 6: 33–38. Available from https://www.biodiversitylibrary.org/page/11135481 [accessed 2 Jul. 2021].
Gravely F.H. 1912. Exhibition of living Pedipalpi, with remarks on the distribution of the order. Proceedings of the Asiatic Society of Bengal: cxxiii–cxxv.
Gravely F.H. 1915a. Notes on the habits of Indian insects, myriapods & arachnids. Records of the Indian Museum 11: 483–539.
Gravely F.H. 1915b. A revision of the Oriental subfamilies of Tarantulidae (order Pedipalpi). Records of the Indian Museum 11: 433–455.
Gray M.R. & Robinson M.L. 1986. Observations on the behaviour and taxonomy of the Australian tailless whipscorpion Charinus pescotti Dunn (Amblypygi: Charontidae). Proceedings of the Linnean Society of New South Wales 108: 217–224.
Hansen H.J. 1893. Organs and characters in different orders of arachnids. Entomologiske Meddelelser 4: 135–251.
Hansen H.J. 1921. The Pedipalpi, Ricinulei, and Opiliones (exc. Op. Laniatores) collected by Mr. Leonardo Fea in tropical West Africa and adjacent islands. Studies on Arthropoda 1: 1–55. Gyldendalske Boghandel, Copenhagen.
Hansen H.J. 1930. On the comparative morphology of the appendages in the Arthropoda. B. Crustacea (supplement), Insecta, Myriopoda, and Arachnida. Studies on Arthropoda 3: 1–376. Gyldendalske Boghandel, Copenhagen.
Harms D. 2018. A new species of Charinus (Amblypygi: Charinidae) from Ghana, with notes on West African whip spiders. Evolutionary Systematics 2: 45–53. https://doi.org/10.3897/evolsyst.2.24505
Harvey M.S. 1985. Amblypygi. In: Walton D.W. (ed.) Zoological Catalogue of Australia 3: 156–157.
Harvey M.S. 1992. The phylogeny and classification of the Pseudoscorpionida (Chelicerata: Arachnida). Invertebrate Taxonomy 6 (6): 1373–1435. https://doi.org/10.1071/IT9921373
Harvey M.S. 2001. The Schizomida (Arachnida) of the Seychelle Islands. Invertebrate Taxonomy 15: 681–693. https://doi.org/10.1071/IT01006
Harvey M.S. 2002a. The neglected cousins: What do we know about the smaller arachnid orders? Journal of Arachnology 30: 357–372. https://doi.org/d6443q
Harvey M.S. 2002b. Nomenclatural notes on Solifugae, Amblypygi, Uropygi and Araneae (Arachnida). Records of the Western Australian Museum 20: 449–459.
Harvey M.S. 2002c. The first Old World species of Phrynidae (Amblypygi): Phrynus exsul from Indonesia. Journal of Arachnology 30: 470–474.
Harvey M.S. 2003. Catalogue of the Smaller Arachnid Orders of the World. Amblypygi, Uropygi, Schizomida, Palpigradi, Ricinulei and Solifugae. CSIRO Publishing, Collingwood, Australia. https://doi.org/10.1071/9780643090071
Harvey M.S. 2013. Whip Spiders of the World, version 1.0. Western Australian Museum, Perth. Available from http://museum.wa.gov.au/catalogues-beta/whip-spiders [accessed 2 Jul. 2021].
Harvey M.S. & West P.L.J. 1998. New species of Charon (Amblypygi, Charontidae) from northern Australia and Christmas Island. Journal of Arachnology 26: 273–284. Available from https://www.jstor.org/stable/3706233 [accessed 26 Mar. 2021].
Harvey M.S., Rahmadi C. & Aland K. 2012. The first record of whip spiders (Amblypygi) in Western Australia. Australasian Arachnology 84: 8–9.
Haupt J. 2009. Chelicerae of arachnids: Facts and fiction. Arthropoda Selecta 18: 17–19.
Hernández Pacheco J.J., Izquierdo Zamora I. & Oromi Masoliver P. 1992. Catálogo espeleológico de las Islas Galápagos. Resultados Científicos del Proyecto Galápagos: Patrimonio de la Humanidad TMFC 2: 71–180.
Hirst S. 1913. Second report on the Arachnida – The scorpions, Pedipalpi, and supplementary notes on the Opiliones and pseudoscorpions. Transactions of the Linnean Society of London, Zoology 2: 31–37. https://doi.org/10.1111/j.1096-3642.1914.tb00122.x
Igelmund P. 1987. Morphology, sense organs, and regeneration of the forelegs (whips) of the whip spider Heterophrynus elaphus (Arachnida, Amblypygi). Journal of Morphology 193: 75–89. https://doi.org/10.1002/jmor.1051930108
Janetschek H. 1957. Das seltsamste Tier Tirols. Palpenläufer (Arachn., Palpigradida): Stellung, Verbreitung, Arten, Bibliographie. Schlern-Schriften 158: 190–214.
Jiménez M.-L. & Llinas-Gutiérrez J. 2005. Description of male Phrynus asperatipes (Amblypygi, Phrynidae) from Baja California Sur, Mexico. Journal of Arachnology 33: 862–865. https://doi.org/10.1636/h03-42.1
Jocqué M. & Giupponi A.P.L. 2012. Charinus bromeliaea sp. n. (Amblypygi: Charinidae); a new species of bromeliad inhabiting whip spider from French Guyana. Zootaxa 3158 (2): 53–59. https://doi.org/10.11646/zootaxa.3158.1.4
Kästner A. 1932. Ordnung der Arachnida: Pedipalpi Latreille = Geissel-Scorpione. In: Kükenthal W. (ed.) Handbuch der Zoologie: 1–75. Walter de Gruyter und Co., Berlin.
Kästner A. 1956. Wirbellose. Lehrbuch der Speziellen Zoologie. Gustav Fischer, Jena.
Kaestner A. 1968. Arthropod Relatives, Chelicerata, Myriapoda. Invertebrate Zoology Vol. II. John Wiley and Sons, New York.
Kaestner A. 1969. Wirbellose. Lehrbuch der Speziellen Zoologie. Gustav Fischer, Stuttgart.
Klingel H. 1963. Paarungsverhalten bei Pedipalpen (Telyphonus [sic] caudatus L., Holopeltidia, Uropygi und Sarax sarawakiensis Simon, Charontiinae, Amblypygi. Verhandlungen der Deutschen Zoologischen Gesellschaft 26: 452–459.
Kovařík F. & Vlasta D. 1996. First report of Amblypygi (Charinidae: Charinus ioanniticus) from Turkey. Klapalekiana 32: 57–58.
Kraepelin K. 1895. Revision der Tarantuliden Fabr. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg 13: 3–53.
Kraepelin K. 1898. Neue Pedipalpen und Scorpione des Hamburger Museums. Mitteilungen aus dem Naturhistorischen Museum in Hamburg 15: 1–6.
Kraepelin K. 1899. Scorpiones und Pedipalpi. In: Dahl F. (ed.) Das Tierreich 8: i–xviii, 1–265.
Kraepelin K. 1901. Catalogue des Pédipalpes des collections du Muséum d’Histoire Naturelle de Paris. Bulletin du Muséum national d’histoire naturelle 7: 263–264.
Kraepelin K. 1914. Die Skorpione und Pedipalpen von Neu-Caledonien und den benachbarten Inselgruppen. In: Sarasin F. & Roux J. (eds) Nova Caledonia, Zoologie 1: 327–337. C.W. Kreidals Verlag, Wiesbaden.
Kraus O. 1961. Charontidae aus Israel, ein zoogeographisch bemerkenswertes Vorkommen (Arach., Pedipalpi – Amblypygi). Senckenbergiana Biologica 42: 491–493.
Kraus O. 1970. Genitalmorphologie und Systematik der Amblypygi (Arachnida). Bulletin du Muséum national d’histoire naturelle 2: 176–180.
Kritscher E. 1959. Ergebnisse der von Dr. O. Paget und Dr. E. Kritscher auf Rhodos durchgeführten zoologischen Exkursionen. II. Pedipalpi (Amblypygi). Annalen des Naturhistorischen Museums in Wien 63: 453–457.
Kritscher E. 1966. Die Pedipalpen- und Araneenausbeute der Österreichischen Neukaledonien-Expedition 1965. Annalen des Naturhistorischen Museums in Wien 69: 373–391.
Koch L. 1867. Beschreibungen neuer Arachniden und Myriapoden. Verhandlungen der Kaiserlich-Königlichen Zoologischen-Botanischen Gesellschaft in Wien 17: 173–250. Available from https://www.biodiversitylibrary.org/page/26724654 [accessed 30 May 2021].
Koch L. & Keyserling E. 1885. Die Arachniden Australiens nach der Natur beschrieben und abgebildet. Bauer und Raspe, Nürnberg. Available from https://www.biodiversitylibrary.org/page/51908653 [accessed 30 May 2021].
Lawrence R.F. 1968. The structure of the cleaning brush on the pedipalps of some African Amblypygi (Arachnida). Journal of Zoology, London 153: 1–8.
Lawrence R.F. 1969. A collection of African Amblypygi with keys to the subfamilies, genera and species of the Ethiopian fauna. Revue de Zoologie et de Botanique africaine 80: 80–87.
Legendre R. 1972. Les arachnides de Madagascar. In: Battistini R. & Richard-Vindard G. (eds) Biogeography and Ecology in Madagascar: 427–457. W. Junk, The Hague.
McArthur I.W., Miranda G.S., Seiter M. & Chapin K.J. 2018. Global patterns of sexual dimorphism in Amblypygi. Zoologischer Anzeiger 273: 56–64. https://doi.org/10.1016/j.jcz.2018.02.005
McClure H.E., Lim B.-L. & Winn S.E. 1967. Fauna of the Dark Cave, Batu Caves, Kuala Lumpur, Malaysia. Pacific Insects 9: 399–428.
Mello-Leitão C. 1931. Pedipalpos do Brasil e algumas notas sobre a ordem. Archivos do Museu Nacional 33: 7–72.
Messana G., Chelazzi L. & Baccetti N. 1985. Biospeleology of Somalia. Mugdile and Showli Berdi Caves. Monitore Zoologico Italiano, n.s., Supplemento 20: 325–340. https://doi.org/10.1080/03749444.1985.10736702
Millot J. 1949a. Classe de Arachnides (Arachnida). I. – Morphologie générale et anatomie interne.
In: Grassé P.-P. (ed.) Traité de zoologie, Vol. 6: 263–319. Masson et Cie, Paris.
Millot J. 1949b. Ordre des Amblypyges. In: Grassé P.-P. (ed.) Traité de zoologie, Volume 6: 563–558. Masson et Cie, Paris.
Miranda G.S. & Giupponi A.P.L. 2011. A new synanthropic species of Charinus Simon, 1892 from Brazilian Amazonia and notes on the genus (Arachnida: Amblypygi: Charinidae). Zootaxa 2980: 61–68. https://doi.org/10.11646/zootaxa.2980.1.5
Miranda G.S. & Reboleira A.S.P.S. 2019. Amblypygids of Timor-Leste: First records of the order from the country with the description of a remarkable new species of Sarax (Arachnida, Amblypygi, Charinidae). ZooKeys 820: 1–12. https://doi.org/10.3897/zookeys.820.30139
Miranda G.S., Aharon S., Gavish-Regev E., Giupponi A.P.L. & Wizen G. 2016a. A new species of Charinus Simon, 1892 (Arachnida: Amblypygi: Charinidae) from Israel and new records of C. ioanniticus (Kritscher, 1959). European Journal of Taxonomy 234: 1–17. https://doi.org/10.5852/ejt.2016.234
Miranda G.S., Giupponi A.P.L. & Wizen G. 2016b. Two new species of whip spider (Amblypygi): an epigean and a cave dwelling Charinus Simon, 1892 from Belize. Zootaxa 4098 (3): 545–559. https://doi.org/10.11646/zootaxa.4098.3.7
Miranda G.S., Milleri-Pinto M., Gonçalves-Souza T., Giupponi A.P.L. & Scharff N. 2016c. A new species of Charinus Simon, 1892 (Amblypygi, Charinidae) from Brazil, with notes on behavior. ZooKeys 621: 15–36. https://doi.org/10.3897/zookeys.621.9980
Miranda G.S., Giupponi A.P., Prendini L. & Scharff N. 2018a. Weygoldtia, a new genus of Charinidae Quintero, 1986 (Arachnida, Amblypygi) with a reappraisal of the genera in the family. Zoologischer Anzeiger 273: 23–32. https://doi.org/10.1016/j.jcz.2018.02.003
Miranda G.S., Kury A.B. & Giupponi A.P.L. 2018b. Review of Trichodamon Mello-Leitão 1935 and phylogenetic placement of the genus in Phrynichidae (Arachnida, Amblypygi). Zoologischer Anzeiger 273: 33–55. https://doi.org/10.1016/j.jcz.2018.02.006
Miranda G.S., Giupponi A.P., Scharff N. & Prendini L. 2021. Phylogeny and biogeography of the pantropical whip spider family Charinidae (Arachnida: Amblypygi). Zoological Journal of the Linnean Society. https://doi.org/10.1093/zoolinnean/zlaa101
Monte B.G.O. do, Gallão J.E., Schimonsky D.M. von & Bichuette M.E. 2015. New records of two endemic troglobitic and threatened arachnids (Amblypygi and Opiliones) from limestone caves of Minas Gerais state, southeast Brazil. Biodiversity Data Journal 3: e5260. https://doi.org/10.3897/BDJ.3.e5260
Monteith G.M. 1965. Notes on the order Amblypygi (Arachnida) in Australia. Journal of the Entomological Society of Queensland 4: 87. https://doi.org/10.1111/j.1440-6055.1965.tb00663.x
Moritz M. & Fischer S.-C. 1980. Die Typen der Arachniden-Sammlung des Zoologischen Museums Berlin. Mitteilungen aus dem Zoologischen Museum in Berlin 56: 137–154.
Morrone J.J. 2006. Biogeographic areas and transition zones of Latin America and the Caribbean Islands based on panbiogeographic and cladistic analysis of the entomofauna. Annual Review of Entomology 51: 467–494. https://doi.org/10.1146/annurev.ento.50.071803.130447
Moseley M., Lim T.W. & Lim T.T. 2012. Fauna reported from Batu caves, Selangor, Malaysia: Annotated checklist and bibliography. Cave and Karst Science 39: 77–92.
Moyá-Guzmán S. 2009. Taxonomía de los guabás (Arachnida: Amblypygi) en Puerto Rico y la descripción de una nueva especie de Charinus Gravely. Ceiba 8: 68–78.
Muchmore W.B. 1993. List of terrestrial invertebrates of St. John, U.S. Virgin Islands (exclusive of Acarina and Insecta), with some records of freshwater species. Caribbean Journal of Science 29: 30–38.
Nair K.B. 1934. On a new sub-species of Phrynichosarax cochinensis Gravely (Pedipalpi: Tarantulidae). Records of the Indian Museum 36: 475–476.
Peck S.B. & Kukalova-Peck J. 1986. Preliminary summary of the subterranean fauna of the Galapagos Islands, Ecuador. Part I. Introduction and non-insect invertebrates. Proceedings of the 9th International Congress of Speleology, Barcelona 2: 164–166.
Peck S.B. & Peck J. 1986. The Galapagos Islands. Volcanic caves and cave fauna of the Galapagos Islands. Canadian Cavernicolous 18: 42–49.
Pérez Y. & Teruel R. 2004. La fauna de arácnidos de dos localidades de Cuba oriental (Arachnida: Scorpiones, Amblypygi, Schizomida, Ricinulei). Revista Ibérica de Aracnología 10: 167–178.
Pike N. 1872. A visit to the Seychelles Islands. Transactions of the Royal Society of Arts and Sciences of Mauritius series B 6: 83–142.
Pinto-da-Rocha R., Machado G. & Weygoldt P. 2002. Two new species of Charinus Simon, 1892 from Brazil with biological notes (Arachnida; Amblypygi; Charinidae). Journal of Natural History 36: 107–118. https://doi.org/10.1080/00222930110110152
Pocock R.I. 1893. On some points in the morphology of the Arachnida (s.s.), with notes on the classification of the group. Annals and Magazine of Natural History, Ser. 6 11: 1–19. https://doi.org/10.1080/00222939308677457
Pocock R.I. 1898. Scorpions, Pedipalpi, and spiders from the Solomon Islands. Annals and Magazine of Natural History, Ser. 7 1: 457–475. https://doi.org/10.1080/00222939808678002
Pocock R.I. 1900. The Fauna of British India, Including Ceylon and Burma. Arachnida. Taylor & Francis, London.
Proctor H.C. 1998. Indirect sperm transfer in arthropods: Behavioral and evolutionary trends. Annual Review of Entomology 43: 153–174. https://doi.org/10.1146/annurev.ento.43.1.153
Pyron R.A., Reynolds R.G. & Burbrink F.T. 2014. A taxonomic revision of boas (Serpentes: Boidae). Zootaxa 3846: 249–260. https://doi.org/10.11646/zootaxa.3846.2.5
Quintero D., Jr. 1975. Scanning electron microscope observations on the tarsi of the legs of the amblypygids (Arachnida, Amblypygi). In: Proceedings of the 6th International Arachnological Congress: 161–163. Free University of Amsterdam, The Netherlands.
Quintero D.J. 1981. The amblypygid genus Phrynus in the Americas (Amblypygi, Phrynidae). Journal of Arachnology 9: 117–166.
Quintero D.J. 1983. Revision of the amblypygid spiders of Cuba and their relationships with the Caribbean and continental American amblypygid fauna. Studies on the Fauna of Curaçao & other Caribbean Islands 65: 1–54.
Quintero D., Jr. 1986. Revision de la clasification de Amblypygidos pulvinados: Creacion de subordenes, una neuva familia y un nuevo genero con tres nuevas especies (Arachnida: Amblypygi).
In: Eberhard, W.G, Lubin, Y.D. & Robinson B.C. (eds) Proceedings of the 9th International Congress of Arachnology, Panama 1983: 203–212. Smithsonian Institution Press, Washington D.C.
Quintero D.J. & Shear W.A. 2016. Charinidae Gray 1849 (Reptilia, Squamata, Serpentes): Proposed suppression. Bulletin of Zoological Nomenclature 73: 25–29. https://doi.org/10.21805/bzn.v73i1.a21
Rahmadi C. & Harvey M.S. 2008. A first epigean species of Stygophrynus Kraepelin (Amblypygi: Charontidae) from Java and adjacent island, Indonesia with notes on S. dammermani Roewer, 1928. Raffles Bulletin of Zoology 56: 281–288.
Rahmadi C. & Kojima J.-i. 2010. Whip spiders of the genus Sarax in the Papuan region, with description of two new species (Amblypygi: Charinidae). Journal of Arachnology 38: 475–484. https://doi.org/10.1636/Ha09-95.1
Rahmadi C., Harvey M.S. & Kojima J.-i. 2010. Whip spiders of the genus Sarax Simon 1892 (Amblypygi: Charinidae) from Borneo Island. Zootaxa 2612: 1–21. https://doi.org/10.11646/zootaxa.2612.1.1
Rahmadi C., Harvey M.S. & Kojima J.-i. 2011. The status of the whip spider subgenus Neocharon (Amblypygi: Charontidae) and the distribution of the genera Charon and Stygophrynus. Journal of Arachnology 39: 223–229. https://doi.org/10.1636/ca10-77.1
Ravelo P.O. 1975. Speleophrynus tronchonii, nuevo género y especie de amblipigios de la familia Charontidae, en una cueva de Venezuela. Boletín de la Sociedad Venezolana de Espeleología 6: 77–85.
Ravelo P.O. 1977. Speleophrynus bordoni nuevo especie de amblipigios de la familia Charontidae, en una cueva de Venezuela (Arachnida: Amblypygy). Boletín de la Sociedad Venezolana de Espeleología 8: 17–25.
Ray J.M. & Ruback P. 2015. Updated checklists of snakes for the provinces of Panamá and Panamá Oeste, Republic of Panama. Mesoamerican Herpetology 2: 168–188.
Remillet M. 1973. Aperçu de la faune souterraine à Madagascar. In: Livre du cinquantenaire de L’Institut de Spéologie “Émile Racovitza”: 135–160. Editura Academiei Republicii Socialiste România, Bucharest.
Réveillion F. & Maquart P.O. 2015. A new species of Charinus Simon, 1892 (Amblypygi, Charinidae) from termite nests in French Guiana. Zootaxa 4032: 190–196. https://doi.org/10.11646/zootaxa.4032.2.3
Rodríguez-Cabrera T.M. & Teruel R. 2016. Distribution, conservation status and taxonomic update of the Cuban endemic whip-spider Charinus tomasmicheli Armas, 2006 (Amblypygi: Charinidae). Revista Ibérica de Aracnología 29: 1–8.
Roewer C.F. 1962. Einige Arachniden aus den Batu Caves in Malaya. Pacific Insects 4: 517–520.
Rosin R. & Shulov A. 1960. Representatives of the order Amblypygi (Arachnida) found in Israel. Bulletin of the Research Council of Israel 9B: 167–168.
Roth V.D. & Craig P.R. 1970. Arachnida of the Galápagos Islands (excluding Acari). Résultats scientifiques de la Mission zoologique belge aux îles Galapagos et en Ecuador (N. et J. Leleup, 1964–1965) 2: 107–124. Royal Museum for Central Africa, Tervuren.
Ruppert E.E. & Barnes R.D. 1994. Invertebrate Zoology. Saunders College Publishers, Fort Worth, Texas.
Schaller F. 1971. Indirect sperm transfer by soil arthropods. Annual Review of Entomology 16: 407–446. https://doi.org/10.1146/annurev.en.16.010171.002203
Segovia J.M.G., Neco L.C. & Willemart R.H. 2018. On the habitat use of the Neotropical whip spider Charinus asturius (Arachnida: Amblypygi). Zoologia 35: e12874. https://doi.org/10.3897/zoologia.35.e12874
Segovia J.M.G., Gainett G. & Willemart R.H. 2020. Predatory behavior and sensory morphology of the whip spider Charinus asturius (Arachnida: Amblypygi). Journal of Ethology 38: 273–280. https://doi.org/10.1007/s10164-020-00648-0
Seiter M. & Hörweg C. 2013. The whip spider collection (Arachnida, Amblypygi) held in the Natural History Museum Vienna, Austria. Arachnologische Mitteilungen 46: 47–53. https://doi.org/10.5431/aramit4606
Seiter M. & Wolff J. 2014. Description of Sarax buxtoni (Gravely 1915) (Arachnida: Amblypygi: Charinidae) and a new case of parthenogenesis in Amblypygi from Singapore. Journal of Arachnology 42: 233–239. https://doi.org/10.1636/Ha14-13.1
Seiter M., Wolff J. & Hörweg C. 2015. A new species of the South East Asian genus Sarax Simon, 1892 (Arachnida: Amblypygi: Charinidae) and synonymization of Sarax mediterraneus Delle Cave, 1986. Zootaxa 4012 (3): 542–552. https://doi.org/10.11646/zootaxa.4012.3.8
Seiter M., Lanner J. & Karolyi F. 2017. Mating behaviour and spermatophore morphology of four Cuban whip spiders (Arachnida, Amblypygi, Phrynidae) – Taxonomic relevance and evolutionary trends. Zoologischer Anzeiger: 117–126. https://doi.org/10.1016/j.jcz.2017.08.006
Seiter M., Schramm F.D. & Schwaha T. 2018. Description of a new Charinus species (Amblypygi: Charinidae) from the Monseñor Nouel province, Dominican Republic. Zootaxa 4438: 349–361. https://doi.org/10.11646/zootaxa.4438.2.9
Selden P.A. & Dunlop J.A. 1998. Fossil taxa and relationships of Chelicerata. In: Edgecombe G. (ed.) Arthropod Fossils and Phylogeny: 303–331. Columbia University Press, New York.
Seyyar O. & Demir H. 2007. A new locality for Charinus ioanniticus (Kritscher, 1959) (Amblypygi: Charinidae) in Turkey. Serket 10: 109–111.
Shakhatreh M.R., Miranda G.S., Bader-Katbeh A., Baker M.A.A. & Amr Z.S. 2020. Charinus ioanni-ticus (Amblypygi: Charinidae), first record of a whip spider from Jordan. Arachnologische Mitteilungen/Arachnology Letters 59: 35–37. https://doi.org/10.30963/aramit5906
Shear W.A., Selden P.A., Rolfe W.D.I., Bonamo P.M. & Grierson J.D. 1987. New terrestrial arachnids from the Devonian of Gilboa, New York (Arachnida, Trigonotarbida). American Museum Novitates 2901: 1–74.
Shultz J.W. 1989. Morphology of locomotor appendages in Arachnida: Evolutionary trends and phylogenetic implications. Zoological Journal of the Linnean Society 97: 1–56. https://doi.org/10.1111/j.1096-3642.1989.tb00552.x
Simon E. 1892. Arachnides. In: Raffray A., Bolivar I., Simon E. (eds) Etude sur les Arthropodes cavernicoles de l’île de Luzon, Voyage de M. E. Simon aux îles Philippines (mars et avril 1890). 4è mémoire. Annales de la Société entomologique de France 61: 35–52.
Simon E. 1901. On the Arachnida collected during the “Skeat Expedition” to the Malay Peninsula, 1899–1900. Proceedings of the Zoological Society of London 71: 45–84. https://doi.org/10.1111/j.1469-7998.1901.tb08164.x
Snodgrass R.E. 1902. Papers from the Hopkins Stanford Galapagos Expedition, 1898–1899. VII. Entomological results (6). Arachnida, part 2. Field notes on species described in part I. Proceedings of the Washington Academy of Sciences 4: 71–80. Available from https://www.biodiversitylibrary.org/page/8666034#page/87/mode/1up [ accessed 2 Jul. 2021].
Snodgrass R.E. 1948. The feeding organs of Arachnida, including mites and ticks. Smithsonian Miscellaneous Collection 110: 1–93.
Speijer E.A.M. 1937. A collection of pedipalps from the Raffles Museum. Bulletin of the Raffles Museum 13: 171–175.
Stockton W.D. 1976. Amblypygi. In: Leleup N. (ed.) Mission zoologique belge aux îles Galapagos et en Ecuador (N. et J. Leleup, 1964–1965): 55–65. Royal Museum for Central Africa, Brussels.
Strinati P. & Aellen V. 1983. Voyage biospéologique autour du monde. Memoires du Speléo-Club de Paris 9: 1–29.
Tagliatela J., Gallao J.E. & Bichuette M.E. 2019. Behavioral repertoire of Charinus mysticus Giupponi & Kury 2002. Espeleo-Tema 29: 57–63.
Taiti S. & Ferrara F. 1985. Biospeleology of Somalia. Two new species of terrestrial Isopoda (Crustacea Oniscidea) from Mugdile and Showli Berdi Caves. Monitore Zoologico Italiano 20: 341–349. https://doi.org/10.1080/03749444.1985.10736703
Teruel R. 2011. Nuevos registros de Charinus acosta (Quintero, 1983) en Cuba (Amblypygi: Charinidae). Boletín de la Sociedad Entomológica Aragonesa 49: 345–346.
Teruel R. 2016. The genus Charinus Simon, 1892 (Amblypygi: Charinidae) on the Island of Hispaniola, Greater Antilles. Revista Ibérica de Aracnología 28: 3–12.
Teruel R. & Coulis M. 2017. First record of the genus Charinus Simon, 1892 from Martinique, Lesser Antilles, with description of a new species (Amblypygi: Charinidae). Ecologica Montenegrina 13: 30–36. https://doi.org/10.37828/em.2017.13.3
Teruel R. & Friedrich S. 2016. Unusual observation of a whip spider in Germany (Arachnida, Amblypygi). Spixiana 39: 22.
Teruel R. & Questel K. 2011. A new species of Charinus Simon 1892 (Amblypygi: Charinidae) from the Leeward Islands, Lesser Antilles. Boletín de la Sociedad Entomológica Aragonesa 49: 15–18.
Teruel R. & Questel K. 2015. A new species of Charinus Simon, 1892 (Amblypygi: Charinidae) from Guadeloupe, Lesser Antilles. Revista Ibérica de Aracnología 26: 43–47.
Teruel R. & Rodríguez-Cabrera T.M. 2016. Actualización de la distribución conocida de Charinus acosta (Quintero, 1983) en Cuba (Amblypygi: Charinidae). Revista Ibérica de Aracnología 28: 135–136.
Teruel R., Armas L.F. & Rodríguez T.M. 2009. Nuevos datos sobre la distribución geográfica y ecología de los amblipígidos de Cuba (Arachnida: Amblypygi). Boletín de la Sociedad Entomológica Aragonesa 44: 201–211.
Thomé M.T.C., Lyra M.L., Lemes P., Teixeira L.S., Carnaval A.C., Haddad C.F.B. & Canedo C. 2020. Outstanding diversity and microendemism in a clade of rare Atlantic Forest montane frogs. Molecular Phylogenetics and Evolution 149: 106813. https://doi.org/10.1016/j.ympev.2020.106813
Thorell T. 1888. Pedipalpi e scorpioni dell’Arcipelago Malese conservati nel Museo Civico di Storia Naturale di Genova. Annali del Museo Civico di Storia Naturale di Genova 26: 327–428.
Torres-Contreras R., García D.M.Á. & Armas L.F. 2015. Una especie nueva de Charinus Simon, 1892 (Amblypygi: Charinidae) del Caribe Colombiano. Revista Ibérica de Aracnología 27: 145–148.
Torres-Contreras R.A., Arroyo-Pérez W. & Alvarez-Garcia D.M. 2019. Charinus rocamadre (Amblypygi, Charinidae): Description of the female, ultrastructure of the male and female gonopods, and mitochondrial DNA sequences. Zootaxa 4664: 129–141. https://doi.org/10.11646/zootaxa.4664.1.6
Trajano E. & Bichuette M.E. 2010. Diversity of Brazilian subterranean invertebrates, with a list of troglomorphic taxa. Subterranean Biology 7: 1–16.
Vandel A. 1965. Biospeleology: The Biology of Cavernicolous Animals. Pergamon Press, Edinburgh.
Van der Hammen L. 1986. Comparative studies in Chelicerata. IV. Apatellata, Arachnida, Scorpionida, Xiphosura. Zoölogische Verhandelingen (Leiden) 226: 1–52.
Vanni S., Bartolozzi L. & Whitman-Mascherini S. 1988. Cataloghi del Museo Zoologico ‘La Specola’. 6. Arachnida, Solifugae e Amblypygi: Tipi. Atti della Società Toscana di Scienze Naturali, Memorie B 94: 373–377.
Vasconcelos A.C.O. & Ferreira R.L. 2016. Description of two new species of Charinus Simon, 1892 from Brazilian caves with remarks on conservation (Arachnida: Amblypygi: Charinidae). Zootaxa 4072 (2): 185–202. https://doi.org/10.11646/zootaxa.4072.2.3
Vasconcelos A.C.O. & Ferreira R.L. 2017. Two new species of cave-dwelling Charinus Simon, 1892 from Brazil (Arachnida: Amblypygi: Charinidae). Zootaxa 4312: 277–292. https://doi.org/10.11646/zootaxa.4312.2.4
Vasconcelos A.C., Giupponi A.P.L. & Ferreira R.L. 2013 . A new species of Charinus Simon, 1892 from northeastern Brazil with comments on the potential distribution of the genus in Central and South Americas (Arachnida: Amblypygi: Charinidae). Zootaxa 3737: 488–500. https://doi.org/10.11646/zootaxa.3737.4.9
Vasconcelos A.C.O., Giupponi A.P.L. & Ferreira R.L. 2014. A new species of Charinus from Minas Gerais State, Brazil, with comments on its sexual dimorphism (Arachnida: Amblypygi: Charinidae). Journal of Arachnology 42: 155–162. https://doi.org/10.1636/H14-01.1
Vasconcelos A.C.O., Giupponi A.P.L. & Ferreira R.L. 2016. Description of a new troglomorphic species of Charinus Simon, 1892 from Brazil (Arachnida, Amblypygi, Charinidae). ZooKeys 600: 35–52. https://doi.org/10.3897/zookeys.600.8580
Víquez C., Miranda R. & Armas L.F 2012. First record of the genus Charinus (Amblypygi: Charinidae) from Panama. Revista Iberica de Aracnologia 21: 56–58.
Víquez C., Chiriví D.J., Moreno-González J.A. & Christensen J.A. 2014. Heterophrynus armiger Pocock, 1902 (Amblypygi: Phrynidae): First record from Colombia, with notes on its historic distribution records and natural history. Check List 10: 457–460.
Weidner H. 1959. Die Entomologischen Sammlungen des Zoologischen Staatsinstituts und Zoologischen Museums Hamburg. I. Teil. Pararthropoda und Chelicerata I. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 57: 89–142.
Werner F. 1935. Klasse: Arachnoidea, Spinnentiere. Pedipalpen. In: Bronn H.G. (ed.) Klassen und Ordnungen des Tierreichs: 317–490. Akademische Verlagsgesellschaft, Leipzig.
Weygoldt P. 1972a. Charinus koepckei n. sp. aus Peru (Amblypygi: Charontidae). Senckenbergiana Biologica 53: 281–286.
Weygoldt P. 1972b. Charontidae (Amblypygi) aus Brasilien. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere 99: 107–132.
Weygoldt P. 1972c. Geisselskorpione und Geisselspinnen (Uropygi und Amblypygi). Zeitschrift des Kölner Zoo 15: 95–107.
Weygoldt P. 1972d. Spermatophorenbau und Samenübertragung bei Uropygen (Mastigoproctus brasilianus C.L. Koch) und Amblypygen (Charinus brasilianus Weygoldt und Admetus pumilio C.L. Koch) (Chelicerata, Arachnida). Zeitschrift für Morphologie und Ökologie der Tiere 71: 23–51. https://doi.org/10.1007/BF00282139
Weygoldt P. 1974. Kampf und Paarung bei Geisselspinne Charinus montanus Weygoldt (Arachnida, Amblypygi, Charontidae). Zeitschrift für Tierpsychologie 34: 217–223.
Weygoldt P. 1975. Die indirekte Spermatophorenübertragung bei Arachniden. Verhandlungen der Deutschen Zoologischen Gesellschaft: 308–313.
Weygoldt P. 1994. Amblypygi. In: Juberthie C. & Decu V. (eds) Encyclopaedia Biospeologica: 241–247. Société de Biospéologie, Moulis and Bucarest.
Weygoldt P. 1996a. Evolutionary morphology of whip spiders: towards a phylogenetic system (Chelicerata: Arachnida: Amblypygi). Journal of Zoological Systematics and Evolution Research 34: 185–202. https://doi.org/10.1111/j.1439-0469.1996.tb00825.x
Weygoldt P. 1996b. The relationships of the south east African whip spiders Hemiphrynus machadoi Fage, 1951 and Phrynichus scullyi Purcell, 1901: Introduction of the new generic names Xerophrynus and Phrynichodamon (Chelicerata: Amblypygi). Zoologischer Anzeiger 235: 117–130.
Weygoldt P. 1998a. The identity of Oligacanthophrynus guianensis di Caporiacco, 1947 (Arachnida: Amblypygi: Charinidae). Bulletin of the British Arachnological Society 11: 72.
Weygoldt P. 1998b. Revision of the species of Phrynichus Karsch, 1879 and Euphrynichus Weygoldt, 1995 (Chelicerata, Amblypygi). Zoologica, Stuttgart 147: 1–65.
Weygoldt P. 1999a. Revision of the genus Damon C. L Koch, 1850 (Chelicerata: Amblypygi: Phrynichidae). Zoologica, Stuttgart 150: 1–45.
Weygoldt P. 1999b. Sperm transfer, spermatophore morphology, and female genitalia of three species of whip spiders: Charinus seychellarum Kraepelin, 1898, Damon medius (Herbst, 1797), and Phrynichus scaber (Gervais, 1844) (Chelicerata, Amblypygi). Zoologica, Stuttgart 150: 47–64.
Weygoldt P. 1999c. Spermatophores and the evolution of female genitalia in whip spiders (Chelicerata, Amblypygi). Journal of Arachnology 27: 103–116.
Weygoldt P. 2000a. Whip spiders (Chelicerata: Amblypygi). Their Biology, Morphology and Systematics. Apollo Books, Stenstrup.
Weygoldt P. 2000b. African whip spiders. Synopsis of the Amblypygi (Chelicerata: Arachnida) reported from Africa. Memorie della Società Entomologica Italiana 78: 339–359.
Weygoldt P. 2002a. Amblypygi. In: Adis J. (ed.) Amazonian Arachnida and Myriapoda: 293–302. Pensoft Publishers, Moscow.
Weygoldt P. 2002b. Chelicerata, Spinnentiere. In: Rieger R. (ed.) Spezielle Zoologie: 449–497. Heidelberg, Berlin.
Weygoldt P. 2002c. Sperm transfer and spermatophore morphology of the whip spiders Sarax buxtoni, S. brachydactylus (Charinidae), Charon cf. grayi, and Stygophrynus brevispina nov. spec. (Charontidae) (Chelicerata, Amblypygi). Zoologischer Anzeiger 241: 131–148. https://doi.org/10.1078/s0044-5231(04)70069-8
Weygoldt P. 2005. Biogeography, systematic position, and reproduction of Charinus ioanniticus (Kritscher, 1959) with the description of a new species from Pakistan (Chelicerata, Amblypygi, Charinidae). Senckenbergiana Biologica 85: 1–14.
Weygoldt P. 2006a. Courtship and sperm transfer in Charinus neocaledonicus Kraepelin, 1895 and Charinus australianus (L. Koch, 1867) (Arachnida, Amblypygi, Charinidae). Zoologischer Anzeiger 244: 239–247. https://doi.org/10.1016/j.jcz.2006.01.002
Weygoldt P. 2006b. New Caledonian whip spiders: Notes on Charinus australianus, Charinus neocaledonicus and other south-western Pacific species of the Charinus australianus species group (Chelicerata, Amblypygi, Charinidae). Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg 42: 5–37.
Weygoldt P. 2007. Parthenogenesis and reproduction in Charinus ioanniticus (Kritscher, 1959) (Chelicerata, Amblypygi, Charinidae). Bulletin of the British Arachnological Society 14: 81–82. https://doi.org/10.13156/arac.2007.14.2.81
Weygoldt P. 2008. Spermatophores, female genitalia, and courtship behaviour of two whip spider species, Charinus africanus and Damon tibialis (Chelicerata: Amblypygi). Zoologischer Anzeiger 247: 223–232. https://doi.org/10.1016/j.jcz.2007.12.002
Weygoldt P. & Van Damme K. 2004. A new troglomorphic whip spider of the genus Charinus (Amblypygi: Charinidae) from Socotra Island. Fauna of Arabia 20: 327–334.
Weygoldt P., Weisemann A. & Weisemann K. 1972. Morphologisch-histologische Untersuchungen an den Geschlechtsorganen der Amblypygi unter besonderer Berücksichtigung von Tarantula marginemaculata C.L. Koch (Arachnida). Zeitschrift für Morphologie und Ökologie der Tiere 73: 209–247. https://doi.org/10.1007/BF00297206
Weygoldt P., Hans P. & Polak S. 2002. Arabian whip spiders: four new species of the genera Charinus and Phrynichus (Chelicerata: Amblypygi) from Oman and Socotra. Fauna of Arabia 19: 289–309.
Wolf B. 1938. Animalium cavernarum Catalogus. W. Junk, The Hague.
Wolfe J.M., Daley A.C., Legg D.A. & Edgecombe G.D. 2016. Fossil calibrations for the arthropod Tree of Life. Earth-Science Reviews 160: 43–110. https://doi.org/10.1016/j.earscirev.2016.06.008
Wolff J.O., Seiter M. & Gorb S.N. 2015. Functional anatomy of the pretarsus in whip spiders (Arachnida, Amblypygi). Arthropod Structure & Development 44: 524–540. https://doi.org/10.1016/j.asd.2015.08.014
Wolff J.O., Schwaha T., Seiter M. & Gorb S.N. 2016. Whip spiders (Amblypygi) become water-repellent by a colloidal secretion that self-assembles into hierarchical microstructures. Zoological Letters 2: 23. https://doi.org/10.1186/s40851-016-0059-y
Wolff J.O., Seiter M. & Gorb S.N. 2017. The water-repellent cerotegument of whip-spiders (Arachnida: Amblypygi). Arthropod Structure & Development 46: 116–129. https://doi.org/10.1016/j.asd.2016.10.010
Yussof S. 1997. The Natural and Other Histories of Batu Caves. Malaysian Nature Society, Kuala Lumpur.
Copyright (c) 2021 Gustavo Silva de Miranda, Alessandro P.L. Giupponi, Lorenzo Prendini, Nikolaj Scharff
This work is licensed under a Creative Commons Attribution 4.0 International License.
Creative Commons Copyright Notices
Authors who publish with this journal agree to the following terms:
- Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a Creative Commons Attribution License (CC BY 4.0) that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.
- Authors are able to enter into separate, additional contractual arrangements for the non-exclusive distribution of the journal's published version of the work (e.g., post it to an institutional repository or publish it in a book), with an acknowledgement of its initial publication in this journal.
- Authors are NOT ALLOWED TO post their work online (e.g., in institutional repositories or on their website) prior to and during the submission process, as it can lead to taxonomic issues.
