Female Chaerilus celebensis, photographed by A. Tietz.

Belongs within: Scorpiones.
Contains: Iurida, Buthoidea.

The Orthosterni is a clade including the living scorpions and some fossil relatives, characterised by the presence of trichobothria on the pedipalp (Soleglad & Fet 2001). Molecular phylogenetic analysis supports the division of living scorpions between two basal lineages, the Iurida and Buthida (Sharma et al. 2015).

<==Orthosterni [Dionychopodes, Neoscorpii, Orthosternina]
    |  i. s.: Corniops mapesiiSF01
    |         Palaeoburmesebuthus Lourenço 2002S-BF04
    |           `--P. grimaldii Lourenço 2002S-BF04
    |--+--Pulmonoscorpius kirktonensisLSE13, GD14
    |  `--AnthracoscorpiiF04
    |       |--Cyclophthalmus senior Corda 1835F04
    |       |--Microlabis sternbergii Corda 1839F04
    |       |--Anthracoscorpio juvenis Kušta 1888 [=Anthracomartus juvenis]F04
    |       |--Mazonia woodianaF04
    |       |--Eobuthus Fritsch 1904F04
    |       |    `--*E. rakovnicensis Fritsch 1904F04
    |       |--Feistmantelia Fritsch 1904F04
    |       |    `--*F. ornata Fritsch 1904F04
    |       |--Glyptoscorpius [incl. Centromachus]F04
    |       |    `--G. caledonicus Peach 1881–1882F04
    |       |--Isobuthus Fritsch 1904F04
    |       |    |--I. kralupensis [=Cyclophthalmus kralupensis]F04
    |       |    `--I. nyranensis Fritsch 1904F04
    |       `--Eoscorpius Meek & Worth 1868LSE13, F04
    |            |--E. anglicus Woodward 1876F04
    |            `--E. carbonariusLSE13
       |    |--Cryptoscorpius americanusSF01
       |    |--CompsoscorpiusLSE13
       |    |    |--C. buthiformisGD14
       |    |    `--C. elegans Petrunkevitch 1949S93
       |    `--PalaeopisthacanthusSF01
       |         |--P. schuchertiSF01
       |         `--P. vogelandurdeniSF01
       `--+--Archaeobuthus Lourenço 2001SF01, S-BF04 [Archaeobuthidae]
          |    `--*A. estephani Lourenço 2001S-BF04
             `--Buthida [Chaerilida, Pseudochactida]SF15
                  |  `--Troglokhammouanus steineriBSL19
                  `--+--Chaerilus Sim. 1877T94 [ChaerilidaeSF15, Chaerilini, Chaeriloidea]
                     |    |--C. cavernicola [incl. Timogenes sumatranus Simon 1880]K77
                     |    |--C. celebensis Pocock 1893T94
                     |    `--C. chapmani (Vachon & Lourenço 1985)L95
                     `--Pseudochactidae [Pseudochactoidea]SF15
                          |--Vietbocap laoSF15
                          |--Toglokhammouanus steineriSF15
                          `--Pseudochactas Gromov 1998SF01
                               `--P. ovchinnikovi Gromov 1998SF01

*Type species of generic name indicated


[BSL19] Ballesteros, J. A., C. E. Santibáñez López, Ľ. Kováč, E. Gavish-Regev & P. P. Sharma. 2019. Ordered phylogenomic subsampling enables diagnosis of systematic errors in the placement of the enigmatic arachnid order Palpigradi. Proceedings of the Royal Society of London Series B—Biological Sciences 286: 20192426.

[F04] Fritsch, A. 1904. Palaeozoische Arachniden. Selestverlag: Prague.

[GD14] Garwood, R. J., & J. Dunlop. 2014. Three-dimensional reconstruction and the phylogeny of extinct chelicerate orders. PeerJ 2: e641.

[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.

[LSE13] Legg, D. A., M. D. Sutton & G. D. Edgecombe. 2013. Arthropod fossil data increase congruence of morphological and molecular phylogenies. Nature Communications 4 (2485): 1–7.

[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.

[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.

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

[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.

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