Liocheles australasiae. Photo by Alex Tietz.

Belongs within: Iurida.
Contains: Urodacus.

The Scorpionoidea are a clade of scorpions characterised by katoikogenic development, with females producing small eggs with no yolk or embryonic membranes whose embryos are nourished via attachment of the embryonic chelicerae to modified outgrowths of the ovariuterus (Sharma et al. 2015). Members include the pantropical Hormuridae which have a low median ocular tubercle on the carapace, not raised above the carapace surface, and distal denticles on the cheliceral movable finger that are subequal in length. The Urodacidae of Australia and Madagascar have multiple median denticle rows on the chelal fingers together with a single row of four to twenty trichobothria on the ventral surface of the pedipalp patella. The Diplocentridae have a disjunct distribution with species found in the Americas and the Middle East.

Scorpionoidea [Pandinidae, Pandinini, Pandinoidae]
    |  i. s.: Heteroscorpion Birula 1903 VHP12
    |         Protoischnuridae CL01
    |           |--Araripescorpius Campos 1986 CL01, S-BF04
    |           |    `--A. ligabuei Campos 1986 SF01
    |           `--Protoischnurus Carvalho & Lourenço 2001 CL01
    |                `--*P. axelrodorum Carvalho & Lourenço 2001 CL01
    |--Hormuridae [Ischnuridae, Liochelidae] SF15
    |    |--Iomachus Pocock 1893 K77
    |    `--Liocheles Sundevall 1833 K77 (see below for synonymy)
    |         |--*L. australasiae (Fabricius 1775) FB01 (see below for synonymy)
    |         |--‘Hormurus’ asiaticus Koch & Keyserling 1884-1885 KK90
    |         |--L. karschii (Keyserling 1885) (see below for synonymy) K77
    |         |--L. longimanus Locket 1995 L95
    |         |--L. nigripes (Pocock 1897) K77 [=Hormurus nigripes K14]
    |         `--L. waigiensis (Gervais 1844) K77 (see below for synonymy)
    `--+--Urodacidae [Urodacinae] SF15
       |    |--Urodacus K77
       |    `--Aops Volschenk & Prendini 2008 T13
       |         `--A. oncodactylus Volschenk & Prendini 2008 T13
       `--+--Diplocentrus Peters 1861 SF15, S-LFP13 [Diplocentridae]
          |    |  i. s.: D. anophthalmus L95
          |    |         D. cueva L95
          |    |         D. diablo SF15
          |    |         D. lesueuri B06
          |    |         D. mitchelli L95
          |    |         D. ochoterenai SF01
          |    |         D. peloncillensis Francke 1975 AS-N08
          |    |--D. mexicanus group S-LFP13
          |    |    |--*D. mexicanus Peters 1861 S-LFP13
          |    |    |--D. tehuano Francke 1977 S-LFP13
          |    |    `--D. whitei S-LFP13
          |    `--D. keyserlingii group S-LFP13
          |         |--D. coylei Fritts & Sissom 1996 [incl. D. malinalco Armas & Martín-Frías 2000] S-LFP13
          |         |--D. formosus Armas & Martín-Frías 2003 S-LFP13
          |         |--D. hoffmanni Francke 1977 S-LFP13
          |         |--D. keyserlingii Karsch 1880 [incl. D. reticulatus Francke 1977] S-LFP13
          |         |--D. kraepelini Santibáñez-López, Francke & Prendini 2013 S-LFP13
          |         |--D. mitlae Francke 1977 S-LFP13
          |         |--D. rectimanus Pocock 1898 [incl. D. moritzi Stahnke 1981] S-LFP13
          |         |--D. sagittipalpus Santibáñez-López, Francke & Prendini 2013 S-LFP13
          |         |--D. sissomi Santibáñez-López, Francke & Prendini 2013 S-LFP13
          |         `--D. tenango Santibáñez-López & Francke 2008 S-LFP13
          `--Scorpionidae SF15
               |--Lipsominae K77
               |--Hemiscorpioninae K77
               `--Scorpioninae K77
                    |  i. s.: Opisthophthalmus carinatus K77, AM07
                    |         Palamnaeus Thorell 1876 K77, T89
                    |           |--P. laevigatus Thorell 1876 KK90
                    |           |--P. longimanus [incl. P. angustimanus] T89
                    |           |--P. petersii Thorell 1876 [incl. P. bengalensis Sim. 1884] T89
                    |           `--P. silenus T89
                    |--+--Heterometrus SF15
                    |  |    |--H. cyanaeus (Koch 1836) K77
                    |  |    |--H. megacephalus [=Pandinus megacephalus, Buthus megacephalus] T89
                    |  |    |--H. petersii SF01
                    |  |    `--H. spinifer S07
                    |  `--Pandinus Thorell 1876 SF15, K92
                    |       |--P. ceylonicus K92
                    |       |--P. imperator Koch 1842 S00
                    |       |--P. indicus (Linnaeus 1754) [=Scorpio indicus] T94
                    |       |--P. kochi K92
                    |       `--P. platycheles Werner 1916 [=Pandinus (Pandinops) platycheles] J98
                    `--+--Hadogenes Kraepelin 1894 SF15, K77
                       |    `--H. troglodytes AM07
                       `--+--Opisthacanthus Peters 1861 SF15, K77
                          |    |--O. asper SF15
                          |    |--O. davydovi Birula 1904 K77
                          |    `--O. madagascariensis SF15
                          `--Scorpio Linnaeus 1758 SF15, L58
                               |--S. afer Linnaeus 1758 L58
                               |--S. americus Linnaeus 1758 L58
                               |--S. australis Linnaeus 1758 L58
                               |--S. europaeus Linnaeus 1758 L58
                               |--S. maurus Linnaeus 1758 L58
                               |    |--S. m. maurus SS08
                               |    |--S. m. fuscus SS08
                               |    `--S. m. palmatus SF15
                               `--S. schweiggeri Holl 1829 S93

Liocheles Sundevall 1833 K77 [incl. Hormurus Thorell 1876 FB01, Ischnurus Koch 1837 FB01; Hormurini, Ischnurini]

*Liocheles australasiae (Fabricius 1775) FB01 [=Scorpio australasiae FB01, Hormurus australasiae K77, Ischnurus australasiae FB01; incl. H. boholiensis Kraepelin 1914 K77, H. caudicula boholiensis K77, Buthus brevicaudatus Rainbow 1897 K77, *Ischnurus complanatus Koch 1838 K77, Hormurus australasiae var. complanatus T89, Sisyphus complanatus K77, Scorpio cumingii Walckenaer & Gervais 1844 K77, Sc. gracilicauda Guérin-Meneville 1843 K77, Hormurus australasiae suspectus Thorell 1888 K77, Ischnurus pistaceus Simon 1877 K77]

Liocheles karschii (Keyserling 1885) [=Hormurus karschii, H. caudicula karschii; incl. H. karschii keyensis Kraepelin 1914, H. papuanus Kraepelin 1914, H. caudicula papuanus] K77

Liocheles waigiensis (Gervais 1844) K77 [=Scorpio (Ischnurus) waigiensis K77, Hormurus waigiensis K14; incl. Ischnurus caudicula Koch 1867 K77, *H. caudicula K77, Liocheles caudicula K77, Ischnurus dechangei Becker 1880 K77, Hormurus dechangei K14, H. insculptus Thorell 1888 K77, H. caudicula insculptus K77, Ischnurus neocalendonicus Simon 1877 K77, H. neocaledonicus K14, H. caudicula novaeguineae Giltay 1931 K77, H. sarasini Kraepelin 1914 K77, H. caudicula sarasini K77, H. weberi Pocock 1893 K77, H. caudicula weberi K77]

*Type species of generic name indicated


[AM07] Acosta, L. E., & G. Machado. 2007. Diet and foraging. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 309–338. Harvard University Press: Cambridge (Massachusetts).

[AS-N08] Aguiar, A. P. N., P. L. Santana-Neto, J. R. B. Souza & C. M. R. de Albuquerque. 2008. Relationship between litter characteristics and female size in Tityus stigmurus (Scorpiones, Buthidae). Journal of Arachnology 36 (2): 464–467.

[B06] Banks, N. 1906. Arachnida from the Bahamas. Bulletin of the American Museum of Natural History 22: 185–189.

[CL01] Carvalho, M. da G. P. de, & W. R. Lourenço. 2001. A new family of fossil scorpions from the Early Cretaceous of Brazil. Comptes Rendus de L’Academie des Sciences Serie II Sciences de la Terre et des Planètes 332: 711–716.

[FB01] Fet, V., & G. Bechly. 2001. Liochelidae, fam. nov. (Scorpiones): proposed introduction as a substitute name for Ischnuridae Simon, 1879, as an alternative to the suggested amendment of Ischnurinae Fraser, 1957 (Insecta, Odonata) to Ischnurainae in order to remove homonymy. Bulletin of Zoological Nomenclature 58 (4): 280–281.

[J98] Jäger, P. 1998. Das Typenmaterial der Spinnentiere (Arachnida: Acari, Amblypygi, Araneae, Opiliones, Pseudoscorpiones, Scorpiones, Uropygi) aus dem Museum Wiesbaden. Jahrbuecher des Nassauischen Vereins fuer Naturkunde 119: 81–91.

[K92] Karsch, F. 1892. Arachniden von Ceylon und von Minikoy gesammelt von den Herren Doctoren P. und F. Sarasin. Berliner Entomologische Zeitschrift 36 (2): 267–310.

[K77] Koch, L. E. 1977. The taxonomy, geographic distribution and evolutionary radiation of Australo-Papuan scorpions. Records of the Western Australian Museum 5 (2): 83–367.

[KK90] Koch, L., & E. Keyserling. 1884–1890. Die Arachniden Australiens nach der Natur beschrieben und abgebildet vol. 2. Bauer & Raspe: Nürnberg.

[K14] Kraepelin, K. 1914. Die Skorpione und Pedipalpen von Neu-Caledonien und den benachbarten Inselgruppen. In: Sarasin, F., & J. Roux (eds) Nova Caledonia: Forschungen in Neu-Caledonian und auf den Loyalty-Inseln. A. Zoologie vol. 1 pt 4 pp. 327–337. C. W. Kreidels Verlag: Wiesbaden.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[L95] Locket, N. A. 1995. A new ischnurid scorpion from the Northern Territory, Australia. Records of the Western Australian Museum Supplement 52: 191–198.

[S-BF04] Santiago-Blay, J. A., V. Fet, M. E. Soleglad & P. R. Craig. 2004. The second Cretaceous scorpion specimen from Burmese amber (Arachnida: Scorpiones). Journal of Systematic Palaeontology 2 (2): 147–152.

[S-LFP13] Santibáñez-López, C. E., O. F. Francke & L. Prendini. 2013. Systematics of the keyserlingii group of Diplocentrus Peters, 1861 (Scorpiones: Diplocentridae), with descriptions of three new species from Oaxaca, Mexico. American Museum Novitates 3777: 1–47.

[SS08] Sarmento, S. M. N., A. M. de Souza, M. V. Meiado & C. M. R. de Albuquerque. 2008. Notes on the life history traits of Rhopalurus rochai (Scorpiones, Buthidae) under different feeding regimes. Journal of Arachnology 36 (2): 476–479.

[S93] Selden, P. A. 1993. Arthropoda (Aglaspidida, Pycnogonida and Chelicerata). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 297–320. Chapman & Hall: London.

[SF15] Sharma, P. P., R. Fernández, L. A. Esposito, E. González-Santillán & L. Monod. 2015. Phylogenomic resolution of scorpions reveals multilevel discordance with morphological phylogenetic signal. Proceedings of the Royal Society of London Series B—Biological Sciences 282: 20142953.

[S00] Shultz, J. W. 2000. Skeletomuscular anatomy of the harvestman Leiobunum aldrichi (Weed, 1893) (Arachnida: Opiliones: Palpatores) and its evolutionary significance. Zoological Journal of the Linnean Society 128: 401–438.

[S07] 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.

[SF01] Soleglad, M. E., & V. Fet. 2001. Evolution of scorpion orthobothriotaxy: a cladistic approach. Euscorpius 1: 1–38.

[T13] Taylor, C. K. 2013. Annotated bibliography for Barrow Island terrestrial invertebrates. Records of the Western Australian Museum Supplement 83: 135–144.

[T89] Thorell, T. 1889. Viaggio di Leonardo Fea in Birmania e regioni vicine. XXI.—Aracnidi Artrogastri Birmani raccolti da L. Fea nel 1885–1887. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 521–729.

[T94] Thorell, T. 1894. Förteckning öfver Arachnider från Java och närgränsande öar, insamlade af docenten D:r Carl Aurivillius; jemte beskrifingar å några sydasiatiska och sydamerikanska Spindlar. Bihang till K. Svenska Vetenskaps-Akademiens Handlingar 20 pt 4 (4): 1–63.

[VHP12] Volschenk, E. S., M. S. Harvey & L. Prendini. 2012. A new species of Urodacus (Scorpiones: Urodacidae) from Western Australia. American Museum Novitates 3748: 1–18.

Last updated: 16 June 2020.

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