Centruroides Marx, 1890

Buthus exilicauda Wood, 1863 (= Centruroides exilicauda (Wood, 1863)), type species by monotypy.

Centrurus (nec Ehrenberg, 1829): Thorell, 1876a: 9; Thorell, 1876b: 83; Karsch, 1879a: 18; Pocock, 1890: 120, 121, 127; Kraepelin, 1891: 119 –124 (part); Pocock, 1893: 375, 385, 386; Laurie, 1896: 131; Lönnberg, 1897: 196, 197, 208; Kraepelin, 1899: 87 (part); Banks, 1900: 425; Borelli, 1909: 222; Comstock, 1912: 25, 27, fig. 31; Birula, 1917a: 164; Birula, 1917b: 54, 107; Ochoterena, 1920: 223; Mello-Campos, 1924a: 246; 1924 b: 312; Comstock, 1940: 27, fig. 31 (lapsus calami); Millot and Vachon, 1949: 427; Díaz Nájera, 1966: 110, 111, pl. 1; 1970: 113.

Centruro: Karsch, 1879b: 120 (lapsus calami).

Centruroides Marx, 1890: 211; Pocock, 1902a: 19, 20; 1902b: 365; Kraepelin, 1912: 69–71; 1914: 22; Hoffmann, 1932: 244, 245; Mello-Leitão, 1932: 27; 1934 a: 4, 5; Franganillo, 1936: 158; Hoffmann, 1937: 201 –203; Moreno, 1939a: 63; Comstock, 1940: 25, 27; Moreno, 1940a: 164; Mello-Leitão, 1942: 126; 1945: 240, 250– 252; Scorza, 1954a: 190; Bücherl, 1964: 59; 1967: 113; Muma, 1967: 2, 4; Bücherl, 1969: 767; Aguilar and Meneses, 1970: 3; Bücherl, 1971: 327; Stahnke, 1971: 282; 1972: 125, fig. 9; Armas, 1974a: 25; Vachon, 1974: 906, 908; 1975: 1598; Francke, 1977b: 75; Stahnke and Calos, 1977: 111; Vachon, 1977: 294; Lourenço, 1979: 214; Williams, 1980: 2, 4; Armas, 1984: 2; González-Sponga, 1984: 64, 65; Francke and Stockwell, 1987: 7; Armas, 1988: 44, 91, 95; Stockwell, 1988: 3; Sissom, 1990: 94, 101; Nenilin and Fet, 1992: 9, 12–14; Stockwell, 1992: 412, 419; Francke, 1985: 7, 15; González-Sponga, 1996: 118, 119, 124, 125, figs. 285–287, 289, 292; Armas, 1998: 50; Kovařík, 1998: 106; Fet and Lowe, 2000: 98 –122; Towler et al., 2001: 161 –163; Prendini and Wheeler, 2005: 481, table 10; Kamenz and Prendini, 2008: 6, 8, 22, 40, tables 1, 2, pl. 13–17; Volschenk et al., 2008: 654, 656, 658, 659, 663, 664, 674, fig. 1C, tables 1, 2; Armas et al., 2012: 106, 112; Loria and Prendini, 2014: 3, 9, 10, 24, 25, fig. 2D, table 5; Ponce Saavedra and Francke, 2014: 54, figs. 11, 14;

Centruroides (Centruroides): Werner, 1934: 273.

Centruoroides: Díaz Nájera, 1970: 117 (lapsus calami).

Centruriodes: Lourenço and Eickstedt, 1988: 7 (lapsus calami).

DIAGNOSIS: Centruroides differs from Heteroctenus, Ischnotelson gen. nov., Jaguajir, gen. nov., and Rhopalurus by the linear, parallel-sided metasoma that does not increase markedly in width posteriorly; from Heteroctenus, Jaguajir, gen. nov., and Rhopalurus by the absence of a pecten-sternite stridulatory organ; and from Physoctonus and Rhopalurus by the separate (unfused) central lateral and posterior central submedian carinae of the carapace. Centruroides differs further from Heteroctonus by the presence or absence of two lateral depressions on the pectinal plate and the absence of macrosetae on the dorsobasal surface of the pectinal teeth; from Ischnotelson by the separate (unfused) lateral ocular and central lateral carinae of the carapace, and the telson vesicle not laterally compressed; from Jaguajir by the separate (unfused) lateral ocular and anterior central submedian carinae of the carapace; from Physoctonus by the bifurcate prolateral pedal spur of leg I, and the oblique subrows of primary denticles on the pedipalp chela fingers flanked closely by pro- and retrolateral accessory (supernumerary) denticles; and from Troglorhopalurus by the distinct retromedian carina on the pedipalp chela manus, and the adjacent prodorsal and proventral carinae of the pedipalp patella.

DESCRIPTION: A revision of Centruroides will be presented elsewhere. The following general description outlines characters common to the species of this diverse genus.

Total length: Varying from small, gracile to very large, robust scorpions (total length, 35–110 mm).

Color: Varying from uniformly pale yellow to uniformly black, often maculate or variegated; legs and pedipalps often paler in color than carapace, tergites, metasoma, and telson; coxosternal region, pectines, and sternites usually paler than carapace, tergites, metasoma, and telson; telson may be darker or paler than metasomal segments.

Chelicerae: Base, dorsal surface with medial transverse row of well-developed tubercles.

Carapace: Median ocular tubercle raised; two median ocelli; three pairs of lateral macroocelli; one pair of lateral microocelli. Anteromedian, median ocular and posteromedian sulci well developed, forming single, almost continuous, longitudinal sulcus. Lateral ocular, central lateral, anterior central submedian and posterior central submedian carinae distinct, finely to coarsely granular or costate-granular and separate (unfused).

Pedipalps: Pedipalp femur retrolateral accessory carinae usually absent. Pedipalp chela manus of adult male slender to slightly incrassate, fixed and movable fingers may be slightly curved proximally (fixed finger curved dorsally, movable finger curved ventrally), such that proximal dentate margin emarginate, small gap present between fingers proximally, when closed, manus of female not incrassate, fixed and movable fingers not curved proximally, such that proximal dentate margin sublinear, little or no gap present between them proximally, when closed; fixed and movable fingers, median denticle rows each comprising 7–9 oblique subrows of primary denticles flanked closely by pro- and retrolateral accessory (supernumerary) denticles; movable finger with proximal lobe. Pedipalps orthobothriotaxic Type A, α configuration; femur with five dorsal trichobothria, trichobothrium d 2 situated on prolateral surface; patella trichobothrium d 3 situated retrolateral to dorsomedian carina; chela fixed finger trichobothrium db aligned with or distal to trichobothrium et.

Legs: Legs III and IV, tibial spurs absent; I–IV, basitarsi each with bifurcate prolateral pedal spur; telotarsi each with irregular tufts of fine, acuminate macrosetae.

Pectines: Pectinal plate with or without two lateral depressions (male), anterior margin with sulcus. Pectines not proximally expanded; proximal dorsal fulcra setose or asetose; pectinal teeth almost straight, slightly curved laterally, proximal teeth not enlarged, dorsal surfaces without nodules or striations, dorsobasal surfaces without macrosetae; pectinal sensillae peg shaped.

Mesosoma: Tergites IV–VI same width or wider than I–III and VII; I–VI tricarinate, dorsomedian and dorsosubmedian carinae granular to costate-granular, restricted to posterior two thirds of segment. Tergite VII pentacarinate, dorsomedian carina restricted to anterior two thirds. Sternites smooth, carinae obsolete, more developed on VI and VII; sternite III, lateral margins not forming smooth, raised carina, ventromedian carina not elevated anteriorly, ventrosubmedian surfaces not forming paired depressions, finely and irregularly granular; respiratory spiracles (stigmata) width more than 5× length.

Metasoma: Metasoma slender, usually not increasing in width posteriorly, segments I and V similar width in both sexes. Segment I with 10 distinct, granular to costate-granular carinae, II with eight or 10 distinct, granular to costategranular carinae, III and IV each with eight distinct, granular to costate-granular carinae, V with seven distinct but less pronounced, granular carinae; dorsosubmedian carinae absent or obsolete, reduced to rows of granules on dorsal surfaces of segments I–IV; dorsolateral carinae complete on segments I–IV, often terminating in prominent, spiniform granules posteriorly on III and IV, absent on V; lateral supramedian carinae complete on segments I–V; lateral inframedian carinae complete on segment I, partial or absent on II, absent on III–V; ventrosubmedian carinae complete on segments I–IV, restricted to anterior third or absent on V; ventromedian carina absent on segments I–IV, complete on V. Intercarinal surfaces finely granular.

Telson: Vesicle spherical to elongate, not laterally compressed, usually similar in width or slightly narrower than metasoma V; anterodorsal lateral lobes reduced or absent; lateral and ventral surfaces smooth or granular; subaculear tubercle usually present.

Hemispermatophore: Flagelliform.

DISTRIBUTION: Centruroides is widely distributed from the United States, throughout Mexico, Central America, and the West Indies, to northern South America. One species is endemic to the Galápagos Islands. The list of countries and territories from which Centruroides has been recorded is as follows: Anguilla; Antigua and Aruba; Barbuda; Bahamas; Belize; British Virgin Islands; Colombia; Costa Rica; Cuba; Curaçao; Dominican Republic; Dominica; Ecuador (including the Galápagos Islands); El Salvador; Guadeloupe; Guatemala; Haiti; Honduras; Jamaica; Mexico (Aguascalientes, Baja California, Baja California Sur, Campeche, Chiapas, Chihuahua, Coahuila, Colima, Distrito Federal, Durángo, Estado de México, Guanajuato, Guerrero, Hidalgo, Jalisco, Michoacán, Morelos, Nayarit, Nuevo León, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Sinaloa, Sonora, Tabasco, Tamaulipas, Tlaxcala, Yucatán, Veracruz, Zacatecas); Martinique; Netherlands Antilles (Bonaire, Saba, St. Eustatius); Nicaragua; Panama; Puerto Rico; St. Barthélemy; St. Kitts and Nevis; St. Martin/Sint Maarten; Turks and Caicos; Venezuela; and the United States of America (Alabama, Arizona, Arkansas, California, Colorado, Florida, Georgia, Illinois, Kansas, Kentucky, Louisiana, Mississippi, Missouri, Nebraska, New Mexico, Nevada, South Carolina, Tennessee, Texas, Utah, U.S. Virgin Islands, with introduced populations in New Jersey, North Carolina, and Virginia). Records from Argentina and Peru (Fet and Lowe, 2000) are dubious.

ECOLOGY: Centruroides occur in diverse habitats ranging from semidesert to tropical rainforest, from sea level to 1700 m elevation. All species of the genus are eurytopic, most being lapidicolous or corticolous (Prendini, 2001b), sheltering under stones, peeling tree bark, cracks and crevices in rock faces and earthen walls, and holes in tree trunks.

INCLUDED SPECIES: The genus Centruroides currently includes 90 described species and three subspecies: Centruroides alayoni Armas, 1999; Centruroides altagraciae Teruel, Armas and Kovařík, 2015; Centruroides anchorellus Armas, 1976; Centruroides arctimanus Armas, 1976; Centruroides baergi Hoffmann, 1932; Centruroides balsasensis Ponce-Saavedra and Francke, 2004; Centruroides bani Armas and Marcano Fondeur, 1987; Centruroides baracoae Armas, 1976; Centruroides barbudensis Pocock, 1898; Centruroides bertholdii (Thorell, 1876); Centruroides bicolor (Pocock, 1898); Centruroides bonito Quijano-Ravell, Teruel and Ponce- Saavedra, 2016; Centruroides caral Armas and Trujillo, 2013; Centruroides chamela Ponce-Saavedra and Francke, 2011; Centruroides chamulaensis Hoffmann, 1932; Centruroides chiapanensis Hoffmann, 1932; Centruroides edwardsii (Gervais, 1843); Centruroides elegans (Thorell, 1876); Centruroides elegans insularis Pocock, 1902; Centruroides exilicauda (Wood, 1863); Centruroides exilimanus Teruel and Stockwell, 2002; Centruroides exsul (Meise, 1933); Centruroides fallassisimus Armas and Trujillo, 2010; Centruroides farri Armas, 1976; Centruroides flavopictus (Pocock, 1898); Centruroides flavopictus meridionalis Hoffmann, 1932; Centruroides franckei Santibañez-López and Contreras-Félix, 2013; Centruroides fulvipes (Pocock, 1898); Centruroides galano Teruel, 2001; Centruroides gracilis (Latreille, 1804); Centruroides granosus (Thorell, 1876); Centruroides griseus (C.L. Koch, 1845); Centruroides guanensis Franganillo, 1930; Centruroides hentzi (Banks, 1900); Centruroides hirsuticauda Teruel, 2011; Centruroides hirsutipalpus Ponce-Saavedra and Francke, 2009; Centruroides hoffmanni Armas, 1996; Centruroides huichol Teruel, Ponce-Saavedra and Quijano-Ravell, 2015; Centruroides infamatus (C.L. Koch, 1844); Centruroides insulanus (Thorell, 1876); Centruroides ixil Trujillo and Armas, 2016; Centruroides jaragua Armas, 1999; Centruroides jorgeorum Santiago-Blay, 2009; Centruroides koesteri Kraepelin, 1912; Centruroides limbatus (Pocock, 1898); Centruroides limpidus (Karsch, 1879); Centruroides luceorum Armas, 1999; Centruroides lucidus Teruel, Armas and Kovařík, 2015; Centruroides marcanoi Armas, 1981; Centruroides margaritatus (Gervais, 1841); Centruroides mariaorum Santiago-Blay, 2009; Centruroides mascota Ponce-Saavedra and Francke, 2011; Centruroides meisei Hoffmann, 1938; Centruroides melanodactylus Teruel, 2001; Centruroides navarroi Teruel, 2001; Centruroides nigrescens (Pocock, 1898); Centruroides nigrimanus (Pocock, 1898); Centruroides nigropunctatus Teruel, 2006; Centruroides nigrovariatus (Pocock, 1898); Centruroides nitidus (Thorell, 1876); Centruroides nitidus taino Armas and Marcano Fondeur, 1987; Centruroides noxius Hoffmann, 1932; Centruroides ochraceus (Pocock, 1898); Centruroides orizaba Armas and Martín-Frías, 2003; Centruroides ornatus Pocock, 1902; Centruroides pallidiceps Pocock, 1902; Centruroides panamensis Quintero and Esposito, 2014; Centruroides platnicki Armas, 1981; Centruroides pococki Sissom and Francke, 1983; Centruroides polito Teruel, 2007; Centruroides poncei Teruel et al., 2015; Centruroides rileyi Sissom, 1995; Centruroides robertoi Armas, 1976; Centruroides rodolfoi Santibañez- López and Contreras-Félix, 2013; Centruroides ruana Quijano-Ravell and Ponce-Saavedra, 2016; Centruroides sanandres Armas, Sarmiento and Flórez, 2012; Centruroides sasae Santiago- Blay, 2009; Centruroides schmidti Sissom, 1995; Centruroides sculpturatus Ewing, 1928; Centruroides serrano Santibañez-López and Ponce- Saavedra, 2009; Centruroides simplex (Thorell, 1876); Centruroides sissomi Armas, 1996; Centruroides spectatus Teruel, 2006; Centruroides stockwelli Teruel, 2001; Centruroides suffusus (Pocock, 1902); Centruroides tapachulaensis Hoffmann, 1932; Centruroides tecomanus Hoffmann, 1932; Centruroides testaceus DeGeer, 1778; Centruroides thorellii (Kraepelin, 1891); Centruroides tuxtla Armas, 1999; Centruroides underwoodi Armas, 1976; Centruroides villegasi Baldazo-Monsivaiz, Ponce-Saavedra and Flores- Moreno, 2013; Centruroides vittatus (Say, 1821).

REMARKS: The name Centruroides was first introduced by Marx (1890: 211) for two species: Centruroides exilicauda (Wood, 1863) and Centruroides luctifer Marx, 1890. Because C. luctifer is a nomen nudum, C. exilicauda is the type species by monotypy (Fet and Lowe, 2000). As Marx (1890) also used the name Centrurus, Centruroides was not introduced as a replacement name for Centrurus.

The name “ Centrurus Ehrenberg, 1829 ” was incorrectly used for many years to denote species of Centruroides Marx, 1890. However, these two names are not synonymous (Braunwalder and Fet, 1998; Fet and Lowe, 2000). The type species of Centrurus was not originally designated by Ehrenberg (1829), and therefore Centrurus Ehrenberg, 1829, is a nomen nudum (Francke, 1985). Thorell (1876a: 9) designated Androctonus biaculeatus Lucas, 1835 (= Centruroides gracilis (Latreille, 1804)), as the type species. However, the priority in type designation belongs to C.L. Koch (1838) who first used the name in combination with the description of a species, Centrurus galbineus C.L. Koch, 1838, a junior synonym of Heterometrus longimanus (Herbst, 1800). Therefore, the correct designation and synonymy is Centrurus C.L. Koch, 1838 = Heterometrus Ehrenberg, 1828 (Scorpionidae).