Buthidae C.L. Koch, 1837

Prendini, Lorenzo, Ehrenthal, Valentin L. & Loria, Stephanie F., 2021, Systematics Of The Relictual Asian Scorpion Family Pseudochactidae Gromov, 1998, With A Review Of Cavernicolous, Troglobitic, And Troglomorphic Scorpions, Bulletin of the American Museum of Natural History 2021 (453), pp. 1-153 : 120-122

Family Buthidae C.L. Koch, 1837 .

Alayotityus delacruzi Armas, 1973 : CUBA. Habitat: inside cave. Troglomorphies: median and lateral ocelli reduced; pigmentation and sclerotization slightly reduced. Previous assessments: une espèce néotroglobionte en train d’inféodation dans le milieu cavernicole (Vachon, 1977); mais dans tous les cas, elles ont été considérées uniquement comme des trogloxènes réguliers ou, à la rigueur, comme des troglophiles (Lourenço and Francke, 1985); troglobite (Armas, 2000; Gallão and Bichuette, 2016); nontroglomorph troglobite (Teruel and Kovařík, 2012); guanobitic-troglobitic (Deharveng and Bedos, 2018). Current assessment: hypogean: troglophile (eutroglophile). Citations: Armas (1973); Vachon (1977); Lourenço and Francke (1985); Armas (2000); Teruel and Kovařík (2012); Gallão and Bichuette (2016); Deharveng and Bedos (2018).

Alayotityus juraguaensis Armas, 1973 : CUBA. Habitat: under stones near seashore, arid area. Previous assessments: neotroglobiont (Vachon, 1977); troglobio-guanobio (Decou, 1981); troglófila del primer orden (Armas and Alayón Garcia, 1984); trogloxene/troglophile (Lourenço and Duhem, 2010a). Current assessment: epigean, hypogean: accidental. Citations: Armas (1973); Armas and Alayón Garcia (1984); Lourenço and Duhem (2010a); Teruel and Kovařík (2012).

Babycurus johnstonii Pocock, 1896 : CAM- EROON. Habitat: surface, forest. Previous assessments: trogloxene/troglophile (Lourenço and Duhem, 2010a). Current assessment: epigean: accidental. Citations: Lourenço and Duhem (2010a).

Centruroides gracilis (Latreille, 1804) : CUBA. Habitat: inside cave; surface. Previous assessments: troglófila del primer orden (Armas and Alayón Garcia, 1984); accidental (Sissom and Reddell, 2009). Current assessment: epigean: accidental. Citations: Armas and Alayón Garcia (1984); Sissom and Reddell (2009).

Heteroctenus junceus (Herbst, 1800) : CUBA. Habitat: inside cave; surface. Previous assessments: troglófila del primer orden (Armas and Alayón Garcia, 1984). Current assessment: epigean: accidental. Citations: Armas and Alayón Garcia (1984); Armas (2000).

Lychas hosei Pocock, 1891 : MALAYSIA. Habitat: inside cave; surface (forest). Previous assessments: probablement troglophile mais, il se pourrait qu’elle soit uniquement trogloxène (Vachon and Lourenço, 1985); trogloxene/troglophile (Lourenço and Duhem, 2010a); troglophile (Beron, 2015). Current assessment: epigean, hypogean: trogloxene (subtroglophile). Citations: Vachon and Lourenço (1985); Lourenço (2007c); Lourenço and Duhem (2010a); Beron (2015); Kovařík (2019). A monotypic genus of dubious validity, Spelaeolychas Kovařík, 2019 , was erected to accommodate this species; it is retained in the genus Lychas C.L. Koch, 1845 , in the present contribution. The subspecies, Lychas hosei cavernicola Lourenço, 2007 appears to be a junior synonym of the nominotypical form.

Reddyanus deharvengi (Lourenço and Duhem, 2010) : VIETNAM. Habitat: inside cave; in litter. Previous assessments: troglophile (Lourenço and Duhem, 2010a; Beron, 2015). Current assessment: epigean, hypogean: accidental. Citations: Lourenço and Duhem (2010a); Beron (2015). Remarks: The holotype male closely resembles the males of other epigean Reddyanus Vachon, 1972 , including Reddyanus petrzelkai Kovařík, 2003 , with which it may be conspecific. Reddyanus are typically attenuate, consistent with the lithophilous ecomorphotype (Prendini, 2001b).

Tityopsis inexpectata (Moreno, 1940) : CUBA. Habitat: inside cave; surface. Previous assessments: troglófila del primer orden (Armas and Alayón Garcia, 1984). Current assessment: epigean: accidental. Citations: Armas and Alayón Garcia (1984); Armas (2000); Teruel and Rodríguez-Cabrera (2020).

Tityopsis sheylae Teruel and Rodríguez- Cabrera, 2020: CUBA. Habitat: inside cave. Troglomorphies: pigmentation and sclerotization slightly reduced; pedipalps, legs, and metasoma attenuate. Previous assessments: troglobite (Teruel and Rodríguez-Cabrera, 2020). Current assessment: hypogean: troglophile (eutroglophile). Citations: Teruel and Rodríguez-Cabrera (2020).

Tityus demangei Lourenço, 1981 : ECUADOR. Habitat: inside cave; surface, forest. Previous assessments: trogloxene (Lourenço, 1981); not a troglobite (Volschenk and Prendini, 2008); trogloxene/troglophile (Lourenço and Duhem, 2010a). Current assessment: epigean, hypogean: accidental. Citations: Lourenço (1981); Lourenço and Francke (1985); Volschenk and Pren- dini (2008); Lourenço and Duhem (2010a); Botero-Trujillo and Flórez (2014). Remarks: According to Lourenço (1981) and Lourenço and Duhem (2010a), this species is a trogloxene, but only the holotype was recorded in a cave, with other specimens collected far from caves (Botero- Trujillo and Flórez, 2014). The yellowish coloration of this species resembles that of many epigean species of the genus and meets the criterion of an accidental.

Tityus jussarae Lourenço, 1988 : ECUADOR. Habitat: inside cave. Current assessment: hypogean: trogloxene (subtroglophile). Citations: Lourenço (1988).

Tityus grottoedensis Botero-Trujillo and Flórez, 2014 : COLOMBIA. Habitat: inside cave, under stone; outside cave. Previous assessments: eutroglophile/subtroglophile (Botero-Trujillo and Flórez, 2014). Current assessment: epigean, hypogean: trogloxene (subtroglophile). Citations: Botero-Trujillo and Flórez (2014).

Tityus magnimanus Pocock, 1897 : BRAZIL, COLOMBIA, VENEZUELA. Habitat: surface, forest; inside cave and surrounding areas. Previous assessments: mais dans tous les cas, elles ont été considérées uniquement comme des trogloxènes réguliers ou, à la rigueur, comme des troglophiles (Lourenço and Francke, 1985); troglophile (Galan and Herrera, 2006); trogloxene/ troglophile (Lourenço and Duhem, 2010a). Current assessment: epigean, hypogean: accidental. Citations: González-Sponga (1974, 1996); Lourenço and Francke (1985); Linares and Bordon (1987); Rojas-Runjaic (2004); Galan and Herrera (2006); Lourenço and Duhem (2010a). Remarks: Tityus falconensis González-Sponga, 1974 is a junior synonym of T. magnimanus , which is why the species was listed as hypogean by Lourenço and Duhem (2010a). Cave records and citations from Lourenço and Francke (1985), Linares and Bordon (1987), Rojas-Runjaic (2004), and Galan and Herrera (2006) refer to T. falconensis .

Tityus monaguensis González-Sponga, 1974 : VENEZUELA. Habitat: forested areas and caves. Previous assessments: mais dans tous les cas, elles ont été considérées uniquement comme des trogloxènes réguliers ou, à la rigueur, comme des troglophiles (Lourenço and Francke, 1985); troglophile (Galan and Herrera, 2006); trogloxene/ troglophile (Lourenço and Duhem, 2010a). Current assessment: epigean, hypogean: accidental. Citations: González-Sponga (1974, 1996); Lourenço and Francke (1985); Linares and Bordon (1987); Rojas-Runjaic (2004); Galan and Herrera (2006); Lourenço and Duhem (2010a).

Troglorhopalurus lacrau (Lourenço and Pinto-da-Rocha, 1997): BRAZIL. Habitat: inside caves. Troglomorphies: pigmentation and sclerotization slightly reduced. Previous assessments: trogloxene; non-cavernicolous (Lourenço and Pinto-da-Rocha, 1997); obligatory cave-dwelling life habitat, troglobite (Gallão and Bichuette, 2016); troglophile (Esposito et al., 2017). Current assessment: hypogean: troglophile (eutroglophile). Citations: Lourenço and Pinto-da-Rocha (1997); Gallão and Bichuette (2016); Esposito et al. (2017).

Troglorhopalurus translucidus Lourenço et al., 2004 : BRAZIL. Habitat: inside cave. Troglomorphies: median and lateral ocelli reduced; pigmentation and sclerotization reduced; pedipalps, legs and metasoma attenuate; prolateral and retrolateral pedal spurs reduced. Previous assessments: troglobite (Lourenço et al., 2004; Volschenk and Prendini, 2008; Trajano and Bichuette, 2010; Botero-Trujillo and Flórez, 2014; Gallão and Bichuette, 2016; Esposito et al., 2017); unequivocal troglobitic characteristics (Lourenço and Duhem, 2010a; Lourenço and Pham, 2013). Current assessment: hypogean: troglobite. Citations: Lourenço et al. (2004); Volschenk and Prendini (2008); Lourenço and Duhem (2010a); Ochoa et al. (2010); Trajano and Bichuette (2010); Lourenço and Pham (2013); Botero-Trujillo and Flórez (2014); Gallão and Bichuette (2016); Esposito et al. (2017).

Troglotityobuthus gracilis (Fage, 1946) : MADAGASCAR. Habitat: inside cave. Troglomorphies: pigmentation and sclerotization unknown (old, faded specimen; pale coloration likely due to preservation); pedipalps, legs, and metasoma attenuate; telson vesicle slightly enlarged. Previous assessments: troglobie (Fage, 1946); troglobite (Lourenço, 2000b; Lourenço and Goodman, 2008; Lourenço and Duhem, 2010a; Gallão and Bichuette, 2016); perhaps a troglobite (Volschenk and Prendini, 2008); unequivocal troglobitic characteristics (Lourenço and Pham, 2013). Current assessment: hypogean: troglophile (eutroglophile). Citations: Fage (1946); Lourenço (2000b); Lourenço and Goodman (2008); Volschenk and Prendini (2008); Lourenço and Duhem (2010a); Lourenço and Pham (2013); Gallão and Bichuette (2016). Remarks: Although this species may be troglomorphic, it seems likely the pale coloration is an artefact of age and preservation. If so, only the attenuation and presence in a cave would be consistent with its classification in the Schiner-Racovitza system. However, many lithophilous epigean buthids are highly attenuated (Volschenk and Prendini, 2008).