Liocheles australasiae, photographed by Alex Tietz.

Belongs within: Chactoidea.
Contains: Urodacus.

The Scorpionoidea are a clade of scorpions characterised by katoikogenic development. Females produce 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. Diplocentrids are distinguished from Scorpionidae by the presence of a subaculear spine on the telson.

The Protoischnuridae are a fossil group from the Early Cretaceous of Brazil resembling Hormuridae in the presence of two carinae on the ventral face of the basal metasomal segments (Carvalho & Lourenço 2001).

Scorpionoidea [Pandinidae, Pandinini, Pandinoidae]
| i. s.: Heteroscorpion Birula 1903VHP12
| ProtoischnuridaeCL01
| |--Araripescorpius Campos 1986CL01, S-BF04
| | `--A. ligabuei Campos 1986SF01
| `--Protoischnurus Carvalho & Lourenço 2001CL01
| `--*P. axelrodorum Carvalho & Lourenço 2001CL01
|--Hormuridae [Ischnuridae, Liochelidae]SF15
| |--Iomachus Pocock 1893K77
| `--Liocheles Sundevall 1833K77 [incl. Hormurus Thorell 1876FB01, Ischnurus Koch 1837FB01; Hormurini, Ischnurini]
| |--*L. australasiae (Fabricius 1775)FB01 (see below for synonymy)
| |--‘Hormurus’ asiaticus Koch & Keyserling 1884-1885KK90
| |--L. karschii (Keyserling 1885) (see below for synonymy)K77
| |--L. longimanus Locket 1995L95
| |--L. nigripes (Pocock 1897)L95 [=Hormurus nigripesK14]
| `--L. waigiensis (Gervais 1844)K77 (see below for synonymy)
`--+--Urodacidae [Urodacinae]SF15
| |--UrodacusSF15
| `--Aops Volschenk & Prendini 2008T13
| `--A. oncodactylus Volschenk & Prendini 2008T13
`--+--Diplocentrus Peters 1861SF15, S-LFP13 [Diplocentridae]
| | i. s.: D. anophthalmusL95
| | D. cuevaL95
| | D. diabloSF15
| | D. lesueuriB06
| | D. mitchelliL95
| | D. ochoterenaiSF01
| | D. peloncillensis Francke 1975AS-N08
| |--D. mexicanus groupS-LFP13
| | |--*D. mexicanus Peters 1861S-LFP13
| | |--D. tehuano Francke 1977S-LFP13
| | `--D. whiteiS-LFP13
| `--D. keyserlingii groupS-LFP13
| |--D. coylei Fritts & Sissom 1996 [incl. D. malinalco Armas & Martín-Frías 2000]S-LFP13
| |--D. formosus Armas & Martín-Frías 2003S-LFP13
| |--D. hoffmanni Francke 1977S-LFP13
| |--D. keyserlingii Karsch 1880 [incl. D. reticulatus Francke 1977]S-LFP13
| |--D. kraepelini Santibáñez-López, Francke & Prendini 2013S-LFP13
| |--D. mitlae Francke 1977S-LFP13
| |--D. rectimanus Pocock 1898 [incl. D. moritzi Stahnke 1981]S-LFP13
| |--D. sagittipalpus Santibáñez-López, Francke & Prendini 2013S-LFP13
| |--D. sissomi Santibáñez-López, Francke & Prendini 2013S-LFP13
| `--D. tenango Santibáñez-López & Francke 2008S-LFP13
| i. s.: Opisthophthalmus carinatusK77, AM07
|--+--Pandinus Thorell 1876SF15, K92
| | |--P. ceylonicusK92
| | |--P. imperator Koch 1842S00
| | |--P. indicus (Linnaeus 1754) [=Scorpio indicus]T94
| | |--P. kochiK92
| | `--P. platycheles Werner 1916 [=Pandinus (Pandinops) platycheles]J98
| `--Heterometrus Ehrenberg & Hemprich 1828SF15, S72 [incl. Palamnaeus Thorell 1876S72]
| |--*H. spinifer (Ehrenberg & Hemprich 1828) (see below for synonymy)S72
| |--H. cyanaeus (Koch 1836)K77
| |--‘Palamnaeus’ laevigatus Thorell 1876KK90
| |--‘Scorpio’ longimanus Herbst 1800S72 [=Palamnaeus longimanusT89; incl. P. angustimanusT89]
| |--H. megacephalus [=Pandinus megacephalus, Buthus megacephalus]T89
| |--H. petersii (Thorell 1876)SF15, S72 [=*Palamnaeus petersiiS72; incl. P. bengalensis Sim. 1884T89]
| |--‘Palamnaeus’ silenusT89
| `--‘Palamnaeus’ thorelliS72
`--+--Hadogenes Kraepelin 1894SF15, K77
| `--H. troglodytesSF15
`--+--Opisthacanthus Peters 1861SF15, K77
| |--O. asperSF15
| |--O. davydovi Birula 1904K77
| `--O. madagascariensisSF15
`--Scorpio Linnaeus 1758SF15, L58
|--S. afer Linnaeus 1758L58
|--S. americus Linnaeus 1758L58
|--S. australis Linnaeus 1758L58
|--S. europaeus Linnaeus 1758L58
|--S. maurus Linnaeus 1758 [=Heterometrus maurus]S72
| |--S. m. maurusSF15
| |--S. m. fuscusSS08
| `--S. m. palmatus (Ehrenberg & Hemprich 1828)SF15, S72 [=Buthus (Heterometrus) palmatusS72]
|--S. pallipes Risso 1826R26
`--S. schweiggeri Holl 1829S93

*Heterometrus spinifer (Ehrenberg & Hemprich 1828) [=Buthus (Heterometrus) spinifer, Palamnaeus spinifer]S72

*Liocheles australasiae (Fabricius 1775)FB01 [=Scorpio australasiaeFB01, Hormurus australasiaeK77, Ischnurus australasiaeFB01; incl. H. boholiensis Kraepelin 1914K77, H. caudicula boholiensisK77, Buthus brevicaudatus Rainbow 1897K77, *Ischnurus complanatus Koch 1838K77, Hormurus australasiae var. complanatusT89, Sisyphus complanatusK77, Scorpio cumingii Walckenaer & Gervais 1844K77, Sc. gracilicauda Guérin-Meneville 1843K77, Hormurus australasiae suspectus Thorell 1888K77, Ischnurus pistaceus Simon 1877K77]

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) waigiensisK77, Hormurus waigiensisK14; incl. Ischnurus caudicula Koch 1867K77, *H. caudiculaK77, Liocheles caudiculaK77, Ischnurus dechangei Becker 1880K77, Hormurus dechangeiK14, H. insculptus Thorell 1888K77, H. caudicula insculptusK77, Ischnurus neocalendonicus Simon 1877K77, H. neocaledonicusK14, H. caudicula novaeguineae Giltay 1931K77, H. sarasini Kraepelin 1914K77, H. caudicula sarasiniK77, H. weberi Pocock 1893K77, H. caudicula weberiK77]

*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.

[R26] Risso, A. 1826. Histoire naturelle des principales productions de l’Europe méridionale et particulièrement de celles des environs de Nice et des Alpes maritimes vol. 5. F.-G. Levrault: Paris.

[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.

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

[S72] Stahnke, H. L. 1972. The nomenclatural status of Heterometrus and Palamnaeus (Scorpionidae). Journal of the Bombay Natural History Society 69 (1): 205–207.

[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.

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