Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 18 No. 2: 31 Dec. 2025
Type: Article
Published: 2025-12-31
DOI: 10.11646/megataxa.18.2.2
Page range: 392-401
Abstract views: 26
PDF downloaded: 17

All genera of the world: Order Amblypygi (Animalia: Arthropoda: Arachnida)

Arachnology Lab and Scorpion Systematics Research Group, Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, U.S.A.
Euamblypygi Paleoamblypygi Charinoidea Charontoidea Phrynoidea Charinidae Charontidae Paracharontidae Phrynichidae Phrynidae †Weygoldtinidae Damoninae Heterophryninae Phrynichinae Phryninae generic names synonyms type species amblypygids whip spiders tailless whip scorpions

Abstract

Forty-one genus-group names are available in the arachnid Order Amblypygi Thorell, 1883 by the end of 2025. Twenty-five of these refer to currently accepted genera (18 extant and seven extinct) and two are nomina dubia. Fourteen genus-group names are in synonymy. Eleven (44%) of the currently accepted genera were described since 1950. Amblypygi includes 299 currently accepted species-group names (284 extant species, including two extant subspecies elevated to the rank of species herein, and 15 extinct species) and 13 nomina dubia. The suprageneric classification of the order is well structured due to extensive phylogenetic analyses.

References

  1. Alexander, A.J. (1962) Courtship and mating in amblypygids (Pedipalpi, Arachnida). Proceedings of the Zoological Society of London, 138 (3), 379–383. https://doi.org/10.1111/j.1469-7998.1962.tb05705.x
  2. Beccaloni, J. (2009) Arachnids. Natural History Museum, London, U.K., 320 pp.
  3. Blanchard, E. (1852) Arachnides. In: L’organisation du règne animal. Vol. 2 [2nd ed.]. E. Blanchard, Paris, 232 pp.
  4. 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
  5. 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.
  6. Chapin, K.J. & Hebets, E.A. (2016) The behavioral ecology of amblypygids. Journal of Arachnology, 44 (1), 1–14. https://doi.org/10.1636/V15-62.1
  7. Coddington, J.A., Giribet, G., Harvey, M.S., Prendini, L. & Walter, D.E. (2004) Arachnida. In: Cracraft, J. & Donoghue, M. (Eds), Assembling the Tree of Life. Oxford University Press, Oxford, U.K., pp. 296–318. https://doi.org/10.1093/oso/9780195172348.003.0019
  8. 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.
  9. Delle Cave, L. & Simonetta, A.M. (1975) Taxonomic notes on the Amblypygi (Arachnida Chelicerata) from Ethiopia and Somalia. Monitore Zoologico Italiano, N. S., Supplemento, 6, 141–166. https://doi.org/10.1080/03749444.1975.10736812
  10. Dumortier, B. (1963) Morphology of sound emission apparatus in Arthropoda. In: Busnel, R.-G. (Ed.), Acoustic behaviour of animals. Elsevier, Amsterdam, pp. 277–345.
  11. Dunlop, J.A. (2018) Systematics of the Coal Measures whip spiders (Arachnida: Amblypygi). Zoologischer Anzeiger, 273, 14–22. https://doi.org/10.1016/j.jcz.2017.11.004
  12. 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
  13. Dunlop, J.A. & Mrugalla, B. (2015) Redescription of the Chiapas amber whip spider Electrophrynus mirus (Amblypygi). Journal of Arachnology, 43(2), 220–223. https://doi.org/10.1636/H15-07
  14. Dunlop, J.A. & Penney, D. (2012) Fossil arachnids. Monograph Series Vol. 2. Siri Scientific Press, Rochdale, U.K., 192 pp.
  15. Dunlop, J.A., Penney, D. & Jekel, D. (2023) A summary list of fossil spiders and their relatives. In: World Spider Catalog. Natural History Museum Bern. Available from: http://wsc.nmbe.ch (last accessed on 26 Nov. 2025).
  16. 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
  17. 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
  18. Esposito, L.A., Bloom, T., Caicedo-Quiroga, L., Alicea-Serrano, A.M., Sánchez-Ruíz, J.A., May-Collado, L.J., Binford, G.J. & Agnarsson, I. (2015) Islands within islands: Diversification of tailless whip spiders (Amblypygi, Phrynus) in Caribbean caves. Molecular Phylogenetics and Evolution, 93, 107–117. https://doi.org/10.1016/j.ympev.2015.07.005
  19. Fabricius, J.C. (1793) Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species adjectis synonimis, locis, obervationibus, descriptionibus Hafniae 2. C.G. Proft, Copenhagen, 519 pp. https://doi.org/10.5962/bhl.title.125869
  20. Fage, L. (1939) Sur une phryne du sud Marocain pourvue d’un appareil stridulent Musicodamon atlanteus, n. gen., nov. sp. Bulletin de la Société Zoologique de France, 64, 100–114.
  21. Fage, L. (1946) Scorpions et pédipalpes de l’Indochine Française. Annales de la Société Entomologique de France, 113, 71–81. https://doi.org/10.1080/21686351.1944.12279016
  22. Fage, L. (1951) Pedipalpes (Amblypyges) récoltés en Angola par M. A. de Barros Machado. Companhia de Diamantes de Angola, Publiçacões Culturais, 13, 9–17.
  23. 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.
  24. Fitzinger, L. (1843) Systema Reptilium. Fasciculus primus: Amblyglossae. Braumüller & Seidel, Vindobonae [Vienna],106 pp. https://doi.org/10.5962/bhl.title.4694
  25. Foelix, R.F. & Hebets, E. (2001) Sensory biology of whip spiders (Amblypygi). Andrias, 15, 129–140.
  26. Fowler-Finn, K.D. & Hebets, E.A. (2006) An examination of agonistic interactions in the whip spider Phrynus marginemaculatus (Arachnida; Amblypygi). Journal of Arachnology, 34(1), 62–76. https://doi.org/10.1636/S04-104.1
  27. Fritsch [Frič], A. (1901) Fauna der Gaskohle und der Kalksteine der Permformation Böhmens. Myriopoda pars II. Arachnoidea. Vol. IV(2). Selbstverlag, Prag [Prague], pp. 56–63, pls 153, 154.
  28. Garwood, R.J. & Dunlop, J.A. (2014) Three-dimensional reconstruction and the phylogeny of extinct chelicerate orders. PeerJ, 2, e641. https://doi.org/10.7717/peerj.641
  29. 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
  30. Gerstaecker, A. (1873) Gliedertierfauna des Sansibargebietes. Arachnoidea. In: Decken, C.C. von der (Ed.), Reisen in Ost-afrika in den Jahren 1859–1865, 3. C.F. Winter, Leipzig and Heidelberg, pp. 461–503.
  31. Gervais, P. (1842) Entomologie. L’Institut, Journal Universal des Sciences et des Sociétés Savantes en France et a l’Étranger, 1ère Section, 10, 76.
  32. Gervais, P. (1844) Histoire Naturelle des Insectes. Aptères. Vol. 3. Librairie Encyclopédique de Rôret, Paris, 494 pp.
  33. Giupponi, A.P. de L. & Miranda, G.S. de (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
  34. Gravely, F.H. (1911) Notes on Pedipalpi in the collection of the Indian Museum. I. New Pedipalpi from Calcutta. Records of the Indian Museum, 6, 33–36. https://doi.org/10.26515/rzsi/v6/i1/1911/163717
  35. Gravely, F.H. (1915) A revision of the Oriental subfamilies of Tarantulidae (order Pedipalpi). Records of the Indian Museum, 11, 433–455. https://doi.org/10.26515/rzsi/v11/i6/1915/163049
  36. Hansen, H.J. (1921) The Pedipalpi, Ricinulei, and Opiliones (exc. Op. Laniatores) collected by Mr. Leonardo Fea in tropical West Africa and adjacent islands. In: Studies on Arthropoda, 1. Gyldendalske Boghandel, Kjøbenhavn, pp. 1–55. https://doi.org/10.5962/bhl.title.30252
  37. Harvey, M.S. (2002) Nomenclatural notes on Solifugae, Amblypygi, Uropygi and Araneae (Arachnida). Records of the Western Australian Museum, 20, 449–459.
  38. Harvey, M.S. (2003) Catalogue of the smaller arachnid orders of the world: Amblypygi, Uropygi, Schizomida, Palpigradi, Ricinulei and Solifugae. CSIRO Publishing, Melbourne, 400 pp. https://doi.org/10.1071/9780643090071
  39. Harvey, M.S. (2013) Whip spiders of the World, version 1.0. Western Australian Museum, Perth. Available from: http://www.museum.wa.gov.au/catalogues/whip-spiders (last accessed on 26 Nov. 2025).
  40. Hebets, E.A. (2002) Relating the unique sensory system of amblypygids to the ecology and behavior of Phrynus parvulus from Costa Rica (Arachnida, Amblypygi). Canadian Journal of Zoology, 80 (2), 286–295. https://doi.org/10.1139/z02-006
  41. Herzig, V., Hauke, T.J. & Lüddecke, T. (2023) Unmasking trends and drivers of the international arachnid trade. Frontiers in Arachnid Science, 2, 1161383. https://doi.org/10.3389/frchs.2023.1161383
  42. Horn, G.H. (1889) A synopsis of the Halticinae of boreal America. Transactions of the American Entomological Society, 16, 163–320. https://doi.org/10.2307/25076525
  43. Hu, X., Lei, X., Luo, C., Jarzembowski, E.A., Wang, B. & Xiao, C. (2020) A new whip spider (Arachnida: Amblypygi) in mid-Cretaceous Kachin amber. Cretaceous Research, 116, 104596. https://doi.org/10.1016/j.cretres.2020.104596
  44. ICZN (International Commission on Zoological Nomenclature) (2017) Opinion 2396 (Case 2169) – Phrynus Lamarck, 1801 (Arachnida, Amblypygi): Conserved. Bulletin of Zoological Nomenclature, 74, 108–110. https://doi.org/10.21805/bzn.v74.a026
  45. Karsch, F. (1879) Ueber eine neue Eintheilung der Tarantuliden (Phrynidae aut.). Archiv für Naturgeschichte, 45, 189–197.
  46. Koch, C.L. (1840) Die Arachniden, 8(1–2) C.H. Zeh’sche Buchhandlung, Nürnberg, pp. 1–40, pl. CCLIII–CCLXVIII.
  47. Koch, C.L. (1850) Übersicht des Arachnidensystems. Vol. 5. J.L. Lotzbeck, Nürnberg, 77 pp.
  48. Koch, L. (1867) Beschreibungen neuer Arachniden und Myriapoden. Verhandlungen der Kaiserlich-Königlichen Zoologischen-Botanischen Gesellschaft in Wien, 17, 173–250.
  49. Kraepelin, K. (1895) Revision der Tarantuliden Fabr. Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg, 13 (3), 3–53.
  50. Kraepelin, K. (1899) Scorpiones und Pedipalpi. In: Dahl, F. (Ed.), Das Tierreich. Herausgegeben von der Deutschen Zoologischen Gesellschaft, 8. R. Friedländer und Sohn Verlag, Berlin, pp. i–xviii, 1–265. https://doi.org/10.5962/bhl.title.2028
  51. 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.
  52. Lamarck, J.B.P.A. de (1801) Systême des animaux sans vertèbres. Lamarck & Deterville, Paris, 432 pp. + 8 tables.
  53. Latreille, P.A. (1804) Histoire naturelle, générale et particulière, des Crustacés et des Insectes. Ouvrage faisant suite aux oeuvres de Leclerc de Buffon, et partie de cours complet d’histoire naturelle redigée par C.S. Sonnini, 7. De l’Imprimerie de F. Dufart, Paris, 413 pp. https://doi.org/10.5962/bhl.title.15764
  54. Latreille, P.A. (1810) Considérations générales sur l’ordre naturel des animaux composant les classes des Crustacés, des Arachnides et des Insectes. F. Schoell, Paris, 444 pp. https://doi.org/10.5962/bhl.title.39620
  55. Lawrence, R.F. (1967) Additions to the fauna of South West Africa: Solifuges, scorpions and Pedipalpi. Scientific Papers of the Namib Desert Research Station, 34, 1–19.
  56. Lichtenstein, A.A.H. & Herbst, J.F.W. (1797) Naturgeschichte der Insekten-Gattungen Solpuga und Phalangium. In: Herbst, J.F.W. (Ed.), Natursystem der Ungeflügelten Insekten, 1(8). Gottl. August. Lange, Berolini [Berlin], pp. 1–88.
  57. Linnaeus, C. [C. von Linné]. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, I. Ed. 10. Laurentii Salvii, Holmiae [Stockholm], 821 pp. https://doi.org/10.5962/bhl.title.542
  58. Marshall, B.M., Strine, C.T., Fukushima, C.S., Cardoso, P., Orr, M.C. & Hughes, A.C. (2022) Searching the web builds fuller picture of arachnid trade. Communications Biology, 5, 448. https://doi.org/10.1038/s42003-022-03374-0
  59. McLean, C.J., Garwood, R.J. & Brassey, C.A. (2018) Sexual dimorphism in the arachnid orders. PeerJ, 6, e5751. https://doi.org/10.7717/peerj.5751
  60. McLean, C.J., Garwood, R.J. & Brassey, C.A. (2020) Sexual dimorphism in the size and shape of the raptorial pedipalps of giant whip spiders (Arachnida: Amblypygi). Journal of Zoology, 310 (1), 45–54. https://doi.org/10.1111/jzo.12726
  61. Mello-Leitão, C. (1931) Pedipalpos do Brasil e algumas notas sobre a ordem. Archivos do Museu Nacional do Rio de Janeiro, 33, 8–72.
  62. Mello-Leitão, C. (1935) Novo pedipalpo do Brasil. Annaes da Academia Brasileira de Sciencias, 7, 183–184.
  63. Millot, J. (1949) Ordre des Amblypyges. In: Grassé, P.-P. (Ed.), Traité de zoologie, anatomie, systématique, biologie, 6. Masson et Cie, Paris, pp. 563–588.
  64. Miranda, G.S. de, Giupponi, A.P. de L., 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
  65. Miranda, G.S. de, Kulkarni, S.S., Tagliatela, J., Baker, C.M., Giupponi, A.P. de L., Labarque, F.M., Gavish-Regev, E., Rix, M.G., Carvalho, L.S., Fusari, L.M., Harvey, M.S., Wood, H.M. & Sharma, P.P. (2024) The rediscovery of a relict unlocks the first global phylogeny of whip spiders (Amblypygi). Systematic Biology, 73 (3), 495–505. https://doi.org/10.1093/sysbio/syae021
  66. Miranda, G.S. de, Kury, A.B. & Giupponi, A.P. de 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
  67. Miranda, G.S. de, Scharff, N., Giupponi, A.P. de L. & Prendini, L. (2021) Phylogeny and biogeography of the pantropical whip spider family Charinidae (Arachnida, Amblypygi). Zoological Journal of the Linnean Society, 194 (1), 136–180. https://doi.org/10.1093/zoolinnean/zlaa101
  68. Moreno, A. (1940) Pedipalpida emmendata. Memorias de la Sociedad Cubana de Historia Natural “Felipe Poey”, 14, 167–168.
  69. Moreno-González, J.A., Guiterrez-Estrada, M. & Prendini, L. (2023) Systematic revision of the whip spider family Paracharontidae (Arachnida: Amblypygi) with description of a new troglobitic genus and species from Colombia. American Museum Novitates, 4000, 1–36. https://doi.org/10.1206/4000.1
  70. Pallas, P.S. (1772) Spicilegia Zoologica. Vol. I(9). Gottl. August. Lange, Berolini [Berlin], 86 pp.
  71. Perty, M. (1833) [1834] Arachnides Brasilienses. In: Spix, J.B. de & Martius, C.F.P. de (eds), Delectus Animalium Articulatorum quae in itinere per Brasiliam annis MDCCCXVII-MDCCCXX jussu et auspiciis Maximiliani Josephi I. Friedrich Fleischer, Monachii [München], 191–209. https://doi.org/10.5962/bhl.title.102991
  72. Petrunkevitch, A. (1913) A monograph of the terrestrial Palaeozoic Arachnida of North America. Transactions of the Connecticut Academy of Arts and Sciences, 18, 1–137.
  73. Petrunkevitch, A. (1945) Calcitro fisheri. A new fossil arachnid. American Journal of Science, 243, 320–329 (erratum: p. 466). https://doi.org/10.2475/ajs.243.6.320
  74. Petrunkevitch, A. (1971) Chiapas amber spiders, II. University of California Publications in Entomology, 63, 1–44.
  75. Pfeffer, G. (1893) Ostafrikanische Reptilien und Amphibien, gesammelt von Herrn Dr. F. Stuhlmann im Jahre 1888 und 1889. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten, 10, 69–105.
  76. Pocock, R.I. (1894a) Notes on the Pedipalpi of the family Tarantulidae contained in the collection of the British Museum. Annals and Magazine of Natural History, (6), 14 (82), 273–298. https://doi.org/10.1080/00222939408677804
  77. Pocock, R.I. (1894b) Contributions to our knowledge of the arthropod fauna of the West Indies. III. Supplement on the Pedipalpi of the West Indies. Journal of the Linnean Society of London, Zoology, 24, 527–542. https://doi.org/10.1111/j.1096-3642.1894.tb02494.x
  78. Pocock, R.I. (1897) Report upon the Scorpiones and Pedipalpi obtained on the Lower Amazons by Messrs. E. E. Austen and F. Pickard-Cambridge during the trip of Mr. Siemens’s Steamship ‘Faraday’. Annals and Magazine of Natural History, (6), 19 (112), 357–368. https://doi.org/10.1080/00222939708680549
  79. Pocock, R.I. (1902a) Arachnida. Scorpiones, Pedipalpi, and Solifugae. In: Godman, F.D. & Salvin, O. (Eds), Biologia Centrali-Americana, 3. Taylor & Francis, London, pp. 1–71.
  80. Pocock, R.I. (1902b) A contribution to the systematics of the Pedipalpi. Annals and Magazine of Natural History, (7), 9 (51), 157–165. https://doi.org/10.1080/00222930208678562
  81. Pocock, R.I. (1911) A monograph of the terrestrial Carboniferous Arachnida of Great Britain. Monographs of the Palaeontographical Society, London, 64, 1–84. https://doi.org/10.1080/02693445.1911.12035553
  82. Pomel, M. (1853) Catalogue méthodique et descriptif des vertébrés fossiles découverts dans le bassin hydrographique supérieur de la Loire et surtout dans la vallée de son affluent principal, l’Allier. J.-B. Baillière, Paris, 193 pp.
  83. Prendini, L. (2011) Order Amblypygi Thorell, 1883. In: Zhang, Z.-Q. (ed.), Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa, 3148, 154. https://doi.org/10.11646/zootaxa.3148.1.28
  84. Prendini, L., Weygoldt, P. & Wheeler, W.C. (2005) Systematics of the Damon variegatus group of African whip spiders (Chelicerata: Amblypygi): Evidence from behaviour, morphology and DNA. Organisms, Diversity & Evolution, 5, 203–236. https://doi.org/10.1016/j.ode.2004.12.004
  85. Purcell, W.F. (1902) On some South African Arachnida belonging to the orders Scorpiones, Pedipalpi, and Solifugae. Annals of the South African Museum, 2, 137–225.
  86. Quintero, D. (1982) Phrynus Lamarck, 1801 (Arachnida, Amblypygi): Proposed conservation. Bulletin of Zoological Nomenclature, 39, 40–44.
  87. Quintero, D. (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 and other Caribbean Islands, 65, 1–54.
  88. Quintero, D. (1986) Revision de la clasification de amblypygidos pulvinados: Creacion de subordenes, una nueva 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 Ninth International Congress of Arachnology, Panama 1983. Smithsonian Institution Press, Washington D.C., pp. 203–212.
  89. Rahmadi, C., Harvey, M.S. & Kojima, J. (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
  90. Ravelo, P.O. (1975) Speleophrynus tronchonii nuevo genero y especie de amblipigios de la familia Charontidae, en una cueva de Venezuela (Arachnida: Amblipygy). Boletin de la Sociedad Venezolana de Espeleologia, 6, 77–85.
  91. Rayor, L.S. & Taylor, L.A. (2006) Social behavior in amblypygids, and a reassessment of arachnid social patterns. Journal of Arachnology, 34 (2), 399–421. https://doi.org/10.1636/S04-23.1
  92. 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, 67–74.
  93. Santer, R.D. & Hebets, E. (2009) Prey capture by the whip spider Phrynus marginemaculatus C.L. Koch. Journal of Arachnology, 37 (1), 109–112. https://doi.org/10.1636/ST07-139.1
  94. Scudder, S.H. (1876) New and interesting insects from the Carboniferous of Cape Breton. Proceedings of the American Association for the Advancement of Science, 24B, 110–111.
  95. Scudder, S.H. (1890) Illustrations of the Carboniferous Arachnida of North America, of the orders Anthracomarti and Pedipalpi. Memoirs of the Boston Society of Natural History, 4, 443–456.
  96. Shultz, J.W. (1990) Evolutionary morphology and phylogeny of Arachnida. Cladistics, 6 (1), 1–38. https://doi.org/10.1111/j.1096-0031.1990.tb00523.x
  97. Shultz, J.W. (2007) A phylogenetic analysis of the arachnid orders based on morphological characters. Zoological Journal of the Linnean Society, 150 (2), 221–265. https://doi.org/10.1111/j.1096-3642.2007.00284.x
  98. Simon, E. (1892) Arachnides. In: Raffray, A., Bolívar, I. & Simon, E. (eds), Voyage de M. E. Simon aux îles Philippines (Mars et Avril 1890) 4e Mémoire (1): Étude sur les arthropodes cavernicoles de l’île de Luzon, Planches 1 et 2. Annales de la Société Entomologique de France, 61, 35–52. https://doi.org/10.1080/21686351.1892.12279283
  99. Thorell, T. (1883) Descrizione di alcuni aracnidi inferiori dell’Arcipelago Malese. Annali del Museo Civico di Storia Naturale di Genova, 18, 21–69.
  100. 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.
  101. Thorell, T. (1889) Aracnidi artrogastri Birmani raccolti da L. Fea nel 1885–1887. Annali del Museo Civico di Storia Naturale di Genova (2), 7, 521–729. https://doi.org/10.5962/bhl.title.10400
  102. Uhen, M.D., Allen, B., Behboudi, N., Clapham, M.E., Dunne, E., Hendy, A., Holroyd, P.A., Hopkins, M., Mannion, P., Novack-Gottshall, P., Pimiento, C. & Wagner, P. (2023) Paleobiology Database User Guide Version 1.0. PaleoBios, 40 (11), 1–56. https://doi.org/10.5070/P9401160531
  103. Werner, F. (1934) Scorpiones, Pedipalpi. In: Bronn, H.G. (Ed.), Klassen und Ordnungen des Tierreichs. 5: Arthropoda. IV: Arachnoidea, 5(IV)(8)(1–2). Akademische Verlagsgesellschaft m.b.H., Leipzig, pp. 1–316.
  104. Weygoldt, P. (1970) Lebenszyklus und postembryonale Entwicklung der Geisselspinne Tarantula marginemaculata C.L. Koch (Chelicerata, Amblypygi) im Laboratorium. Zeitschrift für Morphologie und Ökologie der Tiere, 67, 58–85. https://doi.org/10.1007/BF00280716
  105. Weygoldt, P. (1972) Charontidae (Amblypygi) aus Brasilien. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere, 99, 107–132.
  106. Weygoldt, P. (1975) Die indirekte Spermatophorenübertragung bei Arachniden. Verhandlungen der Deutschen Zoologischen Gesellschaft, 67, 308–313.
  107. Weygoldt, P. (1990) Arthropoda-Chelicerata: Sperm transfer. In: Adiyodi, K.G. & Adiyodi, R.G. (Eds), Reproductive biology of invertebrates, Vol. 4(B). Oxford & IBH Publishing Co., New Delhi, India, pp. 77–119.
  108. Weygoldt, P. (1995) The development of the phrynichid “hand”: Notes on allometric growth and introduction of the new generic name Euphrynichus (Arachnida, Amblypygi). Zoologischer Anzeiger, 234 (1), 75–84.
  109. 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
  110. 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.
  111. Weygoldt, P. (1998a) Mating and spermatophore morphology in whip spiders (Phrynichodamon scullyi [Purcell, 1901], Damon gracilis nov. spec., Damon variegatus [Perty, 1834], and Euphrynichus bacillifer [Gerstaecker, 1873]) (Arachnida: Amblypygi: Phrynichidae). Zoologischer Anzeiger 236, 259–276.
  112. Weygoldt, P. (1998b) The identity of Oligacanthophrynus guianensis di Caporiacco, 1947 (Arachnida: Amblypygi: Charinidae). Bulletin of the British Arachnological Society, 11, 72.
  113. Weygoldt, P. (1999) Revision of the genus Damon C.L. Koch, 1850 (Chelicerata: Amblypygi: Phrynichidae). Zoologica, Stuttgart, 150, 1–45.
  114. Weygoldt, P. (2000) Whip spiders. Their biology, morphology and systematics. Apollo Books, Stenstrup, Denmark, 163 pp. https://doi.org/10.1163/9789004473539
  115. Weygoldt, P. (2002a) Fighting, courtship, and spermatophore morphology of the whip spider Musicodamon atlanteus Fage, 1939 (Phrynichidae) (Chelicerata, Amblypygi). Zoologischer Anzeiger, 241 (3), 245–254. https://doi.org/10.1078/0044-5231-00072
  116. Weygoldt, P. (2002b) Amblypygi. In: Adis, J. (Ed.), Amazonian Arachnida and Myriapoda. Pensoft Publishers, Sofia, pp. 293–302.
  117. Weygoldt, P. (2003) Reproductive biology of two species of Phrynichus, P. exophthalmus Whittick, 1940 and P. delfersi arabicus nov. ssp. (Chelicerata: Amblypygi). Zoologischer Anzeiger, 242 (3), 193–208. https://doi.org/10.1078/0044-5231-00097
  118. Weygoldt, P. (2008) Spermatophores, female genitalia, and courtship behaviour of two whip spider species, Charinus africanus and Damon tibialis (Chelicerata: Amblypygi). Zoologischer Anzeiger 247 (3), 223–232. https://doi.org/10.1016/j.jcz.2007.12.002
  119. Weygoldt, P. (2009) Courtship and mating in the whip spider Phrynichus orientalis Weygoldt, 1998 (Chelicerata: Amblypygi). Zoologischer Anzeiger 248 (3), 177–181. https://doi.org/10.1016/j.jcz.2009.09.001
  120. Weygoldt, P. & Paulus, H.F. (1979) Untersuchungen zur Morphologie, Taxonomie und Phylogenie der Chelicerata. II. Cladogramme und die Entfaltung der Chelicerata. Zeitschrift für die Zoologische Systematik und Evolutionforschung, 17, 177–200. https://doi.org/10.1111/j.1439-0469.1979.tb00699.x
  121. 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
  122. Wood, H.C. (1869) On the Phalangia and Pedipalpi collected by Professor Orton in western South America, with the description of new African species, accompanied with a plate. Transactions of the American Philosophical Society, new series, 13, 435–442. https://doi.org/10.2307/1005373
  123. World Amblypygi Catalog (2025) Natural History Museum Bern. Available from: https://wac.nmbe.ch/order/amblypygi/1 (last accessed on 20 Dec. 2025).