Bothriuridae

Black scorpion Bothriurus bonariensis, photographed by Gustavo Schmidt.

Belongs within: Iurida.

The Bothriuridae is a group of scorpions found in Australia and South America characterised by legs with two pedal spurs, a vertically compressed sternum, and pedipalps with chelal trichobothrium Et₂ on the ventral surface and ib at the extreme base of the fixed finger or on the palm.

Bothriuridae [Bothriuroidae, Bothriuroidea]
    |--Vachonianinae K77
    |--Brachistosternus Pocock 1893 OA02b [Brachistosterninae K77]
    |    |  i. s.: B. andinus Chamberlin 1916 OA02b
    |    |         B. ehrenbergii (Gervais 1841) OA02b [incl. B. ehrenbergi var. politus AO02]
    |    |         B. ferrugineus (Thorell 1877) AO02
    |    |         B. holmbergi Carbonell 1923 OA02b
    |    |         B. peruvianus Piza 1974 OA02b
    |    `--B. (Leptosternus Maury 1973) OA02b
    |         |--B. (L.) castroi Mello-Leitão 1940 OA02b
    |         |--B. (L.) intermedius Lönnberg 1902 OA02b
    |         |--B. (L.) montanus Roig Alsina 1977 OA02b
    |         |--B. (L.) quiscapata Ochoa & Acosta 2002 OA02b
    |         |--B. (L.) titicaca Ochoa & Acosta 2002 OA02b
    |         `--B. (L.) weijenberghii (Thorell 1876) OA02b
    `--Bothriurinae [Cercophoninae] K77
         |--Cercophonius Peters 1861 [incl. Acanthochirus Peters 1861] K77
         |    `--*C. squama (Gervais 1844) (see below for synonymy) K77
         |--Urophonius Pocock 1893 K77
         |    |--U. brachycentrus (Thorell 1877) K77
         |    `--U. jeheringi Pocock 1893 K77
         |--Timogenes Simon 1880 K77
         |    |--T. dorbignyi (Guérin-Méneville 1843) [=Scorpio dorbignyi] AO02
         |    |--T. elegans (Mello-Leitão 1931) AO02
         |    `--T. haplochirus Maury & Roig Alsina 1977 CFS14
         `--Bothriurus Peters 1861 CFS14
              |--B. araguayae Vellard 1934 FM11
              |--B. asper Pocock 1893 [=B. bonariensis asper] AO02
              |--B. bocki Kraepelin 1911 AO02
              |--B. bonariensis (Koch 1843) AO02
              |--B. bucherli (San Martin 1965) K77
              |--B. burmeisteri Kraepelin 1894 CFS14
              |--B. chacoensis Maury & Acosta 1993 AO02
              |--B. dumayi AO02
              |--B. flavidus (Maury 1971) K77
              |--B. inermis Maury 1981 AO02
              |--B. maculatus Kraepelin 1911 [=B. bonariensis var. maculata] AO02
              |--B. olaen Acosta 1997 AO02
              |--B. prospicuus A02
              |--B. rochensis K77
              |--B. trivittatus Werner 1939 AO02
              `--B. vittatus Guérin-Méneville 1838 AO02

Bothriuridae incertae sedis:
  Orobothriurus Maury 1976 OA02a
    |--O. alticola (Pocock 1899) OA02a
    |--O. atiquipa Ochoa & Acosta 2002 OA02a
    |--O. calchaqui Ochoa et al. 2011 CFS14
    |--O. compagnucci Ochoa et al. 2011 CFS14
    |--O. curvidigitus (Kraepelin 1911) OA02a [incl. Bothriurus lampei Werner 1916 J98]
    |--O. famatina Acosta & Ochoa 2001 CFS14
    |--O. grismadoi Ojanguren-Affilastro et al. 2009 CSF14
    |--O. iskay Acosta & Ochoa 2001 AO02
    |--O. paessleri (Kraepelin 1911) OA02a
    `--O. wawita Acosta & Ochoa 2000 AO02
  Tehuankea Cekalovic 1973 K77

*Cercophonius squama (Gervais 1844) [=Scorpio (Telegonus) squama; incl. C. granulosus Kraepelin 1908, C. kershawi Glauert 1930, C. michaelseni Kraepelin 1908, C. sulcatus Kraepelin 1908, *Acanthochirus testudinarius Peters 1861] K77

*Type species of generic name indicated

REFERENCES

[A02] Acosta, L. E. 2002. Patrones zoogeográficos de los Opiliones Argentinos (Arachnida: Opiliones). Revista Ibérica de Aracnología 6: 69–84.

[AO02] Acosta, L. E., & J. A. Ochoa. 2002. Lista de los escorpiones bolivianos (Chelicerata: Scorpiones), con notas sobre su distribución. Revista de la Sociedad Entomológica Argentina 61 (3–4): 15–23.

[CSF14] Campón, F. F., S. L. Silnik & L. A. Fedeli. 2014. Scorpion diversity of the Central Andes in Argentina. Journal of Arachnology 42 (2): 163–169.

[FM11] Freire-Jr, G. de B., & P. C. Motta. 2011. Effects of experimental fire regimes on the abundance and diversity of cursorial arachnids of Brazilian savannah (cerrado biome). Journal of Arachnology 39 (2): 263–272.

[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.

[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.

[OA02a] Ochoa, J. A., & L. E. Acosta. 2002a. Orobothriurus atiquipa, a new bothriurid species (Scorpiones) from Lomas in southern Peru. Journal of Arachnology 30: 98–103.

[OA02b] Ochoa, J. A., & L. E. Acosta. 2002b. Two new Andean species of Brachistosternus Pocock (Scorpiones: Bothriuridae). Euscorpius 2: 1–13.

Buthinae

Marbled scorpion Lychas marmoreus, copyright Reiner Richter.

Belongs within: Buthoidea.

The Buthinae are a pantropical group of scorpions with oblique, non-overlapping rows of tooth-like granules on the movable finger of the pedipalp.

<==Buthinae K77
    |--Isometroides Keyserling 1885 K77
    |    `--*I. vescus (Karsch 1880) [=Isometrus vescus; incl. Isometroides angusticaudus Keyserling 1885] K77
    |--Buthus F04
    |    |--B. cyaneus D57
    |    |--B. eupeus [incl. B. eupeus var. cognatus] K01
    |    |--B. gibbosus F04
    |    |--B. grammurus Thorell 1889 T89
    |    |--B. groutii Wood 1869 W69
    |    |--B. hottentotae T89
    |    |--B. martensi Karsch 1879 [incl. B. confucius Sim. 1880] K01
    |    |--B. occitanus GK04
    |    |--B. reticulatus D57
    |    `--B. tamulus K77
    |--Isometrus Hemprich & Ehrenberg 1828 [=Atreus Walckenaer & Gervais 1844] K77
    |    |--I. maculatus (De Geer 1778) SP10 (see below for synonymy)
    |    |--I. acanthurus Pocock 1899 K77
    |    |--I. antillanus T89
    |    |--I. assamensis Oates 1888 K77
    |    |--I. atomarius T89
    |    |--I. basilicus Karsch 1879 K77
    |    |--I. brachycentrus Pocock 1899 K77
    |    |--I. feae Thorell 1889 T89
    |    |--I. formosus Pocock 1893 K77
    |    |--I. fuscus T89
    |    |--I. madagassus Roewer 1943 K77
    |    |--I. melanodactylus (Koch 1867) (see below for synonymy) K77
    |    |--I. perfidus Koch & Keyserling 1884-1885 KK90
    |    |--I. phipsoni T89
    |    |--I. rigidulus Pocock 1897 K77
    |    |--I. shoplandii Oates 1888 T89
    |    |--I. thurstoni Pocock 1893 K77
    |    |--I. thwaitesii Pocock 1897 K77
    |    `--I. vittatus Pocock 1900 K77
    `--Lychas Koch 1850 [incl. Archisometrus Kraepelin 1891, Hemilychas Hirst 1911] K77
         |--*L. scutilus Koch 1845 K77 [=Isometrus scutilus T89]
         |--L. albimanus Henderson 1919 K77
         |--L. alexandrinus Hirst 1911 (see below for synonymy) K77
         |--L. feae (Thorell 1889) K77
         |--L. flavimanus (Thorell 1888) K77
         |--L. gravelyi Henderson 1913 K77
         |--L. guineensis Lucas 1858 SP10 [=Scorpio (Lychas) guineensis K77]
         |--L. hendersoni (Pocock 1897) K77
         |--L. infuscatus (Pocock 1890) K77
         |--L. laevifrons (Pocock 1897) K77
         |--L. marmoreus (Koch 1845) (see below for synonymy) K77
         |--L. mucronatus (Fabricius 1798) K77 (see below for synonymy)
         |--L. nigrimanus (Kraepelin 1898) K77
         |--L. nigristernis (Pocock 1899) K77
         |--L. perfidus (Keyserling 1887) K77
         |--L. rugosus (Pocock 1897) K77
         |--L. scaber (Pocock 1893) K77
         |--L. shelfordi (Borelli 1904) K77
         |--L. shoplandi Oates 1888 K77
         |--L. tweediei Kopstein 1937 K77
         `--L. variatus (Thorell 1877) (see below for synonymy) K77

Isometrus maculatus (De Geer 1778) SP10 [=Scorpio maculatus K77, Lychas maculatus K77; incl. S. americanus Herbst 1800 K77, Centrurus (Isometrus) americanus K77, Lychas americanus K77, Scorpio dentatus Herbst 1800 K77, Buthus (*Isometrus) filum Hemprich & Ehrenberg 1828 K77, S. (*Atreus) filum KK90, K77, S. (Lychas) gabonensis Lucas 1858 K77]

Isometrus melanodactylus (Koch 1867) [=Lychas melanodactylus; incl. I. gracilis Thorell 1877, I. melanodactylus inflatus Glauert 1925, I. papuensis Werner 1916] K77

Lychas alexandrinus Hirst 1911 [=L. (*Hemilychas) alexandrinus; incl. L. annulatus Glauert 1925, L. mjobergi Kraepelin 1916, L. truncatus Glauert 1925] K77

Lychas marmoreus (Koch 1845) [=Tityus marmoreus, *Archisometrus marmoreus, Lychas marmoreus typicus; incl. Isometrus bituberculatus Pocock 1891, Archisometrus bituberculatus, Lychas jonesae Glauert 1925, L. marmoreus nigrescens Kraepelin 1916, L. marmoreus obscurus Kraepelin 1916, L. marmoreus splendens Kraepelin 1916] K77

Lychas mucronatus (Fabricius 1798) K77 [=Scorpio mucronatus T94, Archisometrus mucronatus T94, Isometrus mucronatus T89; incl. Tityus varius Koch 1845 T89, I. varius T89]

Lychas variatus (Thorell 1877) [=Isometrus variatus, Archisometrus variatus, Lychas marmoreus variatus; incl. Isometrus armatus Pocock 1890, Archisometrus armatus, Lychas armatus, L. spinatus besti Glauert 1925, L. marmoreus kimberleyanus Kraepelin 1916, L. lappa Glauert 1954, L. spinatus pallidus Glauert 1925, Isometrus variatus papuanus Thorell 1888, L. papuanus, L. spinatus Kraepelin 1916, Isometrus thorelli Keyserling 1885] K77

*Type species of generic name indicated

REFERENCES

[D57] Doleschall, C. L. 1857. Bijdrage tot de kennis der Arachniden van den Indischen Archipel. Natuurkundig Tijdschrift voor Nederlandsch Indië, series 3, 3 (5–6): 399–434, pls 1–2.

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

[GK04] Gantenbein, B., & P. D. Keightley. 2004. Rates of molecular evolution in nuclear genes of east Mediterranean scorpions. Evolution 58 (11): 2486–2497.

[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.

[K01] Kulczyński, V. 1901. Arachnoideák [Arachnoidea]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 311–369. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[SP10] Santos, R., & C. E. Prieto. 2010. Los Assamiidae (Opiliones: Assamiidae) de Río Muni (Guinea Ecuatorial), con la descripción de ocho nuevas especies. Revista de Biologia Tropical 58 (1): 203–243.

[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 2^a, 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.

[W69] Wood, H. C., Jr. 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 13: 435-442, pl. 17.

Buthoidea

Durango bark scorpion Centruroides suffusus, copyright Tomascastelazo.

Belongs within: Orthosterni.
Contains: Microtityus, Buthinae, Tityus.

The Buthoidea are a group of scorpions that lack ventral trichobothria on the patella of the chela (Soleglad & Fet 2001). Members are currently divided between the cosmopolitan Buthidae and the Madagascan Microcharmidae though it has been suggested that the latter represents a subgroup of the former.

<==Buthoidea
    |--Microcharmidae SF01
    `--Buthidae [Androctonini, Androctonoidae, Buthoidae, Centruridae, Centrurini] B-TN08
         |  i. s.: Charmus laneus K92
         |         Uroplectes J98
         |           |--U. emiliae (Werner 1916) [=Lychas emiliae] J98
         |           `--U. occidentalis L49
         |         Microtityus B-TN08
         |         Lissothus SF01
         |         Karasbergia S-BF04
         |         Microbuthus S-BF04
         |         Orthochirus SF01
         |         Zabius SF01
         |         Pakistanorthochirus SF01
         |         Baloorthochirus SF01
         |         Orthochiroides SF01
         |         Apistobuthus SF01
         |         Razianus SF01
         |         Palaeolychas balticus Lourenço & Weitschat 1996 S-BF04
         |         Palaeotityobuthus longiaculeus Lourenço & Weitschat 2000 S-BF04
         |         Palaeoprotobuthus pusillus Lourenço & Weitschat 2000 S-BF04
         |         Palaeoakentrobuthus knodeli Lourenço & Weitschat 2000 S-BF04
         |         Palaeoananteris ribnitiodamgartensis Lourenço & Weitschart 2001 S-BF04
         |         Grosphus hirtus Kraepelin 1901 AS-N08
         |         Stenochirus Karsch 1892 K92
         |           `--*S. sarasinorum Karsch 1892 K92
         |         Rhopalurus Thorell 1876 B-TN08
         |           `--R. rochai (Borelli 1910) SS08
         |         Centrurus B06 [Centrurinae K77]
         |           |--C. degeeri T89
         |           |--C. nitidus T89
         |           |--C. tenuis T89
         |           `--C. vittatus B06
         |         Ananteris Thorell 1891 B-TN08 [Ananterinae K77]
         |           |--*A. balzanii Thorell 1891 B-T08
         |           |--A. arcadioi Botero-Trujillo 2008 B-T08
         |           |--A. ashmolei Lourenço 1981 B-T08
         |           |--A. charlescorfieldi Lourenço 2001 AO02
         |           |--A. columbiana Lourenço 1991 B-TN08
         |           |--A. cussinii Borelli 1910 B-T08
         |           |--A. dorae Botero-Trujillo 2008 B-T08
         |           |--A. ehrlichi Lourenço 1994 B-T08
         |           |--A. gorgonae Lourenço & Flórez 1989 B-T08
         |           |--A. leilae Lourenço 1999 B-T08
         |           |--A. myriamae Botero-Trujillo 2007 B-T08
         |           |--A. pydanieli Lourenço 1982 L-M-HG08
         |           `--A. tolimana Teruel & García 2007 B-T08
         |--+--Hottentotta trilineatus SF15
         |  `--+--Androctonus Ehrenberg 1828 SF15, SO18
         |     |    |--A. australis (Linnaeus 1758) GE02
         |     |    `--A. bicolor SF01
         |     `--Mesobuthus Vachon 1950 SF15, GK04
         |          |  i. s.: M. martensii SF15
         |          |--M. caucasicus (Nordmann 1840) GK04
         |          |--M. eupeus (Koch 1839) GK04
         |          `--+--M. cyprius Gantenbein & Kropf 2000 GK04
         |             `--M. gibbosus (Brullé 1832) GK04
         `--+--Buthinae SF15
            `--+--Parabuthus SF15
               |    |--P. brevimanus [=Buthus brevimanus] P01
               |    |--P. cristatus Pocock 1901 P01
               |    |--P. granulatus (Ehrenberg 1831) [incl. P. granulatus bergeri Werner 1916] J98
               |    |    |--P. g. granulatus P01
               |    |    `--P. g. fuscus Pocock 1901 P01
               |    |--P. laevifrons (Simon 1888) [incl. P. ibelli Werner 1916] J98
               |    `--P. transvaalicus SF15
               `--+--Tityus SF15
                  `--Centruroides Marx 1890 SF15, B-TN08
                       |--C. exilicauda (Wood 1863) SS08
                       |--C. gracilis (Latreille 1804) SS08
                       |--C. hentzi CS90
                       |--C. hoffmanni Armas 1996 S-LFP13
                       |--C. limpidus WS10
                       |--C. margaritatus (Gervais 1841) AO02
                       |--C. nigrimanus (Pocock 1898) C-MF13
                       |--C. nigrovariatus (Pocock 1898) S-LFP13
                       |--C. nitidus [incl. C. beynai] SF01
                       |--C. sculpturatus M-RL88
                       |--C. suffusus Pocock 1902 C-MF13
                       `--C. vittatus (Say 1821) S00

*Type species of generic name indicated

REFERENCES

[AO02] Acosta, L. E., & J. A. Ochoa. 2002. Lista de los escorpiones bolivianos (Chelicerata: Scorpiones), con notas sobre su distribución. Revista de la Sociedad Entomológica Argentina 61 (3–4): 15–23.

[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.

[B-T08] Botero-Trujillo, R. 2008. The scorpion genus Ananteris in Colombia: comments on the taxonomy and description of two new species (Scorpiones, Buthidae). Journal of Arachnology 36 (2): 287–299.

[B-TN08] Botero-Trujillo, R., & J. A. Noriega. 2008. First record of the scorpion genus Microtityus from Colombia, with the description of a new species (Scorpiones, Buthidae). Journal of Arachnology 36 (2): 259–266.

[CS90] Corey, D. T., & I. J. Stout. 1990. Ground surface arachnids in sandhill communities of Florida. Journal of Arachnology 18: 167–172.

[C-MF13] Cupul-Magaña, F. G., & O. F. Francke. 2013. Tres especies de alacranes (Scorpiones: Buthidae y Vaejovidae) embotellados en mezcal mexicano. Revista Ibérica de Aracnología 22: 123–124.

[GK04] Gantenbein, B., & P. D. Keightley. 2004. Rates of molecular evolution in nuclear genes of east Mediterranean scorpions. Evolution 58 (11): 2486–2497.

[GE02] Giribet, G., G. D. Edgecombe, W. C. Wheeler & C. Babbitt. 2002. Phylogeny and systematic position of Opiliones: a combined analysis of chelicerate relationships using morphological and molecular data. Cladistics 18: 5–70.

[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.

[L49] Lawrence, R. F. 1949. A collection of Opiliones and scorpions from north-east Angola made by Dr. A. de Barros Machados in 1948. Diamang Publicações Culturais 1949: 1–20.

[L-M-HG08] Lo-Man-Hung, N. F., T. A. Gardner, M. A. Ribeiro-Júnior, J. Barlow & A. B. Bonaldo. 2008. The value of primary, secondary, and plantation forests for Neotropical epigeic arachnids. Journal of Arachnology 36 (2): 394–401.

[M-RL88] Meyer-Rochow, V. B., & A. R. Liddle. 1988. Structure and function of the eyes of two species of opilionid from New Zealand glow-worm caves (Megalopsalis tumida: Palpatores, and Hendea myersi cavernicola: Laniatores). Proceedings of the Royal Society of London Series B—Biological Sciences 233: 293–319.

[P01] Pocock, R. I. 1901. Descriptions of some new African Arachnida. Annals and Magazine of Natural History, series 7, 7: 284–288.

[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.

[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.

[SO18] Šťáhlavský, F., V. Opatova, P. Just, L. N. Lotz & C. R. Haddad. 2018. Molecular technique reveals high variability of 18S rDNA distribution in harvestmen (Opiliones, Phalangiidae) from South Africa. Comparative Cytogenetics 12 (1): 41–59.

[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 2^a, 7: 521–729.

[WS10] Wang, M., S. Sun, C. Li & X. Shen. 2010. Distinctive mitochondrial genome of the calanoid copepod Calanus sinicus, a useful molecular marker for phylogenetic and population studies. In: China-Russia Bilateral Symposium: Proceedings of the China-Russia Bilateral Symposium of "Comparison on Marine Biodiversity in the Northwest Pacific Ocean", 10–11 October 2010, Qingdao (China) pp. 187–192. Institute of Oceanology, Chinese Academy of Sciences; A. V. Zhirmunsky Institute of Marine Biology, Far East Branch of the Russian Academy of Sciences.

Last updated: 16 June 2020.

Scorpiones

Palaeophonus hunteri, from Fritsch (1904).

Belongs within: Euchelicerata.
Contains: Orthosterni.

The Scorpiones, scorpions, are a highly distinctive group of arachnids in which the pedipalps form large chelae and the rear section of the opisthosoma is modified to form a mobile 'tail' bearing a terminal sting. Modern scorpions are invariably terrestrial but aquatic habits have been suggested for the earliest known Silurian–Devonian scorpions such as Palaeoscorpius, Proscorpius and the Protoscorpiones. Members of the extinct Palaeoscorpiones and modern Neoscorpionina possess an enlarged prosomal sternum separating the leg coxae; in Palaeoscorpius and the Palaeophonidae, the coxae are positioned anterior to the sternum. Palaeoscorpius devonicus has been suggested to have been an entirely gnathobasic feeder, in contrast to the extraintestinal feeding of modern scorpions (Giribet et al. 2002).

<==Scorpiones [Apoxypodes, Branchioscorpionina, Protoscorpiones, Scorpiniata, Scorpionida]
    |--Palaeoscorpius devonicus Lehmann 1944 GE02, S93
    `--+--Palaeoscorpiones S93
       |    |--Archaeoctonoidea S93
       |    |    |--Archaeoctonus glaber (Peach 1883) S93
       |    |    |--Pseudoarchaeoctonus denticulatus (Kjellesvig-Waering 1986) S93
       |    |    `--Loboarchaeoctonus squamosus (Kjellesvig-Waering 1986) S93
       |    `--Proscorpioidea S93
       |         |--Proscorpius osborni Whitefield 1885 LSE13, F04
       |         |--Archaeophonus eurypteroides Kjellesvig-Waering 1966 S93
       |         |--Stoermeroscorpio delicatus Kjellesvig-Waering 1986 S93
       |         `--Labriscorpio alliedensis Leary 1980 S93
       `--+--Neoscorpionina S93
          |    |--Orthosterni LSE13
          |    `--Paleosterni S93
          |         |--Allobuthiscorpius major (Wills 1960) S93
          |         `--Buthiscorpius pescei Vachon & Heyler 1985 S93
          `--+--Dolichophonus loudonensis (Laurie 1899) GE02, S93
             `--Palaeophonidae GE02
                  |--Allopalaeophonus GE02
                  `--Palaeophonus Thorell & Lindström 1884 LSE13, F04
                       |--P. hunteri Pocock 1902 F04
                       |--P. londonensis Laurie 1889 F04
                       `--P. nuncius Thorell & Lindström 1885 F04

Scorpiones incertae sedis:
  Mesoscorpionina S93
    |--Phoxiscorpio peachi Kjellesvig-Waering 1986 S93
    `--Mesophonus maculatus Brauer et al. 1889 S93
  Dacurus galbineus (Koch 1838) [=Centrurus galbineus, C. galbinus (l. c.), Caucon galbineus] K92
  Broteochactas V-MDR07
    |--B. mapuera Lourenço 1988 L-M-HG08
    `--B. trezzi V-MDR07
  Pseudouroctonus apacheanus (Gertsch & Soleglad 1972) AS-N08
  Alacran tartarus Francke 1982 L95
  Liassoscorpionides G88
  Waeringoscorpio GD14
    |--W. hefteri GD14
    `--W. westerwaldensis GD14
  Brontoscorpius anglicus Kjellesvig-Waering 1972 F14
  Hubeiscorpio gracilitarsis Walossek et al. 1990 S93
  Prearcturus gigas Woodward 1871 S93
  Butheolus thalassinus S89

*Type species of generic name indicated

REFERENCES

[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.

[F14] Francke, O. F. 2014. Biodiversidad de Arthropoda (Chelicerata: Arachnida ex Acari) en México. Revista Mexicana de Biodiversidad 85 (Suppl.): 408–418.

[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.

[GE02] Giribet, G., G. D. Edgecombe, W. C. Wheeler & C. Babbitt. 2002. Phylogeny and systematic position of Opiliones: a combined analysis of chelicerate relationships using morphological and molecular data. Cladistics 18: 5–70.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[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.

[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.

[L-M-HG08] Lo-Man-Hung, N. F., T. A. Gardner, M. A. Ribeiro-Júnior, J. Barlow & A. B. Bonaldo. 2008. The value of primary, secondary, and plantation forests for Neotropical epigeic arachnids. Journal of Arachnology 36 (2): 394–401.

[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.

[S89] Simon, E. 1889. Butheolus thalassinus. Annales de la Société Entomologique de France, 6^e série 9: xcviii–xcix.

[V-MDR07] Villarreal-Manzanilla, O., C. DoNascimiento & C. J. Rodríguez. 2007. Two new species of the enigmatic genus Stenophareus (Opiliones: Laniatores: Stygnidae) from the Venezuelan Guiana Shield. Zootaxa 1471: 43–51.

Last updated: 13 March 2022.

Orthosterni

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 mapesii SF01
    |         Palaeoburmesebuthus Lourenço 2002 S-BF04
    |           `--P. grimaldii Lourenço 2002 S-BF04
    |--+--Pulmonoscorpius kirktonensis LSE13, GD14
    |  `--Anthracoscorpii F04
    |       |--Cyclophthalmus senior Corda 1835 F04
    |       |--Microlabis sternbergii Corda 1839 F04
    |       |--Anthracoscorpio juvenis Kušta 1888 [=Anthracomartus juvenis] F04
    |       |--Mazonia woodiana F04
    |       |--Eobuthus Fritsch 1904 F04
    |       |    `--*E. rakovnicensis Fritsch 1904 F04
    |       |--Feistmantelia Fritsch 1904 F04
    |       |    `--*F. ornata Fritsch 1904 F04
    |       |--Glyptoscorpius [incl. Centromachus] F04
    |       |    `--G. caledonicus Peach 1881–1882 F04
    |       |--Isobuthus Fritsch 1904 F04
    |       |    |--I. kralupensis [=Cyclophthalmus kralupensis] F04
    |       |    `--I. nyranensis Fritsch 1904 F04
    |       `--Eoscorpius Meek & Worth 1868 LSE13, F04
    |            |--E. anglicus Woodward 1876 F04
    |            `--E. carbonarius LSE13
    `--+--Palaeopisthacanthidae SF01
       |    |--Cryptoscorpius americanus SF01
       |    |--Compsoscorpius LSE13
       |    |    |--C. buthiformis GD14
       |    |    `--C. elegans Petrunkevitch 1949 S93
       |    `--Palaeopisthacanthus SF01
       |         |--P. schucherti SF01
       |         `--P. vogelandurdeni SF01
       `--+--Archaeobuthus Lourenço 2001 SF01, S-BF04 [Archaeobuthidae]
          |    `--*A. estephani Lourenço 2001 S-BF04
          `--+--Iurida SF15
             `--Buthida [Chaerilida, Pseudochactida] SF15
                  |--+--Buthoidea SF15
                  |  `--Troglokhammouanus steineri BSL19
                  `--+--Chaerilus Sim. 1877 T94 [Chaerilidae SF15, Chaerilini, Chaeriloidea]
                     |    |--C. cavernicola [incl. Timogenes sumatranus Simon 1880] K77
                     |    |--C. celebensis Pocock 1893 T94
                     |    `--C. chapmani (Vachon & Lourenço 1985) L95
                     `--Pseudochactidae [Pseudochactoidea] SF15
                          |--Vietbocap lao SF15
                          |--Toglokhammouanus steineri SF15
                          `--Pseudochactas Gromov 1998 SF01
                               `--P. ovchinnikovi Gromov 1998 SF01

*Type species of generic name indicated

REFERENCES

[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.

Last updated: 17 June 2020.

Isometrus

Isometrus melanodactylus, photographed by azn4114.
Belongs within: Buthoidea.

Isometrus is a genus of scorpions found primarily in the Old World tropics, except for the pantropical I. maculatus.

See also: The world's scorpion.

Characters (from Kovařík 2003): Trichobothrial patterns orthobothriotaxic. Dorsal trichobothria of femur arranged in beta-configuration. Tibial spurs absent on all legs. Sternum subtriangular. Movable fingers of pedipalps with six rows of granules and external and internal granules. Tibiae and tarsomeres of legs I–III with setae not arranged into bristlecombs. Carapace granulated, without carinae. Mesosoma dorsally with one median carina. Telson with subaculear tooth pointed or rounded. Total length under 70 mm.

Isometrus Hemprich & Ehrenberg 1829 K92
    |--I. antillanus T89
    |--I. atomarius T89
    |--I. feae Thorell 1889 T89
    |--I. fuscus T89
    |--I. maculatus (De Geer 1778) K92 (see below for synonymy)
    |--I. melanodactylus (Koch 1867) [incl. I. papuensis Werner 1916] J98
    |--I. melanophysa (Koch 1867) [=Lychas melanophysa; incl. I. gracilis Thorell 1876] KK84
    |--I. perfidus Koch & Keyserling 1884-1885 KK84
    |--I. phipsoni T89
    |--I. scutilus [=Lychas scutilus] T89
    |--I. shoplandii Oates 1888 T89
    |--I. thorellii Koch & Keyserling 1884-1885 KK84
    `--I. variatus Thorell 1876 KK84

Isometrus maculatus (De Geer 1778) K92 [=Scorpio maculatus K92, Lychas maculatus KK84; incl. Lychas americanus Koch 1845 KK84, Centrurus (Isometrus) americanus KK84, Scorpio dentatus Herbst 1800 KK84, Buthus (Isometrus) filum Hemprich & Ehrenberg 1829 KK84, S. (Atreus) filum KK84, S. (Lichas) gabonensis Lucas 1858 KK84, S. guineensis Lucas 1858 KK84]

*Type species of generic name indicated

REFERENCES

[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.

[KK84] Koch, L., & E. Keyserling. 1884-1890. Die Arachniden Australiens nach der Natur beschrieben und abgebildet vol. 2. Bauer & Raspe: Nürnberg.

Kovařík, F. 2003. A review of the genus Isometrus Ehrenberg, 1828 (Scorpiones: Buthidae) with descriptions of four new species from Asia and Australia. Euscorpius 10: 1-19.

[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.

Scorpionoidea

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

REFERENCES

[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.

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Last updated: 16 June 2020.