Heteroctenus junceus (Herbst, 1800)

Figures 2 B, 3 A, 11 A, 14D, 17A, 18C, D, 21B, 22B, 23H–K, 27A, C, 28D, 29D, 30D, 34

Scorpio junceus Herbst, 1800: 65–67, pl. III, fig. 2; Latreille, 1804: 126 –127.

Scorpio (Atreus) hemprichii Gervais, 1844a: 218, fig. 18 (synonymized by Pocock, 1893: 392); Gervais, 1843: 130 (nomen nudum); 1844b: 39, 54; Lucas, 1851: 70, pl. V, fig. 5–5c; Gervais, 1859: 41, pl. I, fig. 2, 2a–b.

Rhopalurus hemprichii: Karsch, 1879b: 119 .

Centrurus hemprichii: Kraepelin, 1891: 123, 135– 137, figs. 30, 33; Thorell, 1893: 372, 373.

Centrurus junceus: Kraepelin, 1891: 135, figs. 30, 33; Thorell, 1893: 372; Kraepelin, 1895: 95; 1899: 89, 94; 1901: 270; 1908: 187, 190, 193, 194.

Heteroctenus junceus: Pocock, 1893: 392 .

Rhopalurus junceus: Pocock, 1902a: 37, 38, pl. VIII, fig. 5, 5a, pl. IX, fig. 1, 1a; Banks, 1909: 172; Herrera, 1917: 271; Lampe, 1917: 197; Franganillo, 1930a: 95; 1930 b: 119; Mello- Leitão, 1932: 14; Meise, 1934: 29; Franganillo, 1935: 21; 1936: 164, figs. 86, 87; Prado, 1940: 26 –28; Roewer, 1943: 219; Esquivel de Verde, 1968: 67; Esquivel de Verde and Machado-Allison, 1969: 33; Bücherl, 1971: 327; Armas, 1974b: 2, figs. 1A, 2A, 3A; 1977: 3; Stahnke and Calos, 1977: 119; Lourenço, 1979: 215, fig. 8; Armas, 1982a: 4; 1982 b: 5, table 2; Lourenço, 1982a: 110, 114, 136, figs. 10, 11, 14–23, 78, table 1; Armas, 1983: 3, fig. 1; Lourenço, 1986a: 133, fig. 17; 1986b: 165, fig. 7; Armas, 1988: 68 –70, 93, figs. 27, 36; Lourenço, 1992: 55; Rudloff, 1994: 9; Kovařík, 1997: 181; Lourenço, 1997a: 590; Kovařík, 1998: 118; Fet and Lowe, 2000: 219, 220; Teruel, 2003: 149, 150, figs. 1, 2; 2006: 51–53; Teruel and Armas, 2006: 179; 2012 a: 153–167, figs. 1–16, tables 1 –7; 2012b: 215, fig. 7; Kamenz and Prendini, 2008: 9, table 2, pl. 41; Prendini et al., 2009: 222, 223; Teruel and Kovařík, 2012: 123 –140, figs. 29, 40, 41, 48, 265, 267–302, 534–545; Rodríguez- Cabrera and Teruel, 2014: 121; 2015: 85, 86, fig. 1.

Rhopalurus junceus ravidus Franganillo, 1930a: 95, figs. 19–21 (synonymized by Lourenço, 1982a: 114); 1935: 21; 1936: 164; Moreno, 1939a: 65 –66; Jaume, 1954: 1091.

Centruroides (Rhopalurus) junceus: Werner, 1934: 274, fig. 33b.

Rhopalurus junceus junceus: Moreno, 1939a: 65; 1939 b: 124–128, pl. XIX–XXI; Jaume, 1954: 1090; Armas, 1973: 7.

Rhopalurus junceus cadenasi Moreno, 1939a: 66 –67, pl. 6, fig. 3 (synonymized by Lourenço, 1982a: 114); 1940b: 129–130, pl. XXII; Jaume, 1954: 1090.

Rhopalurus melloleitaoi Teruel and Armas, 2006: 175–179, figs. 1–4, tables 1, 2; Armas, 1974b: 6; 1981 a: 52 (part); 1982a: 4; Teruel and Montano, 2005: 221 –223, 225–227, figs. 10, 14, tables 2, 4; Prendini et al., 2009: 222; Teruel and Armas, 2012b: 215, figs. 6, 7.; Teruel and Kovařík, 2012: 141, figs. 303– 305, 552–555; syn. nov.

Rhopalurus aridicola Teruel and Armas, 2012b: 210, 212, 214, 215, figs. 1–6, tables 1–3; Armas, 1974b: 6; 1982 a: 4; Prendini et al., 2009: 222; Teruel and Kovařík, 2012: 116, figs. 546–551; syn. nov.

TYPE MATERIAL: CUBA: Scorpio (Atreus) hemprichii, holotype [sex?] (MNHN). Scorpio junceus: Holotype [lost], “Brazil” [dubious] ; ♂ neotype, Villa Clara Prov.: Municipio Placetas: Loma del Vigía. Rhopalurus j. ravidus: Holotype [sex?] (IZACC) [lost], Pinar del Río Prov.: Sierra del Cuzco, Pinar del Río. Rhopalurus j. cadenasi: subad . ♀ holotype (IZACC), Havana Prov.: Loma de la Universidad, La Habana. Rhopalurus melloleitaoi: Granma Prov.: Municipio Niquero: Holotype ♂, paratype ♀ (RTO), El Guafe (19°57′18″N 6°45′27″W), 2 km N of Cabo Cruz, 50 m, 19. iv.1996; N. Navarro, A. Fernández, and L.M. Díaz . Paratypes: same data except 8–12.vii.2000, R. Teruel, L. Montano, Y. Cala, and R. Escalona, 1 ♀, 1 juv. (IES), 1 juv. (BIOECO), 1 juv. (RTO), 27. vi.2001, L. Montano and R. Escalona, 1 juv. (RTO), 4.iv.2002, L. Montano and R. Escalona, 1 juv. (RTO), 23.i.2003, L. and M. Montaño, 1 juv. (RTO); Bosque Castillo (19°54′14″N 77°31′15″W), Alegría de Pío, 270 m, iii.2002, Jacinto , 1 juv. (RTO); Los Muertos (19°54′0 5″N 77°30′33″W), Alegría de Pío, 280 m, 20.vi.2002, L. Montano, R. Escalona and Y. Cala , 2 juv. (RTO); Sendero Morlotte-Fustete (19°52′48″N 77°33′0 8″W), 5 km SW of Alegría de Pío, 150 m, 21.vi.2002, Y. Cala and Jacinto, 2 juv. (RTO); Ensenada del Real (19°51′14″N 77°35′40″W), 5 m, 19.iv.1996, N. Navarro, A. Fernández, and L. Díaz , 1 juv. (RTO), iii.2002, Jacinto, 1 ♂, 1 ♀, 1 juv. (RTO). Rhopalurus aridicola: Guantánamo Prov.: Holotype ♂ (RTO), Municipio Maisí: Punta de Maisí (20°14′42″N 74°08′38″W), 18–19.iv. 1998, 5 m, R. Teruel, A. Sánchez, N. Navarro, and A. Fong. Paratypes: Municipio Baracoa : Dolina de la Cueva de La Majana (20°20′14″N 74°27′22″W), 50 m, 10.xi.2007, R. Teruel , 1 juv. (BIOECO). Municipio Maisí: Sabana, Santa Rosa (20°17′35″N 74°15′45″W), 14.iv.1998, R. Teruel, 1 ♂ (RTO); Punta de Maisí (20°14′42″N 74°08′38″W), 20.iv.1969, O. Tapia , 1 ♀ (RTO), 6.x.1973, L.F. de Armas and L.R. Hernández, 2 ♂, 2 ♀, 1 juv. (IES), viii.1979, L.F. de Armas, 1 ♂ (IES), 18–19.iv.1998, R. Teruel, A. Sánchez, N. Navarro, and A. Fong, 3 ♂, 8 ♀, 1 ♀ with 15 1 st instars, 4 juv. (RTO), 6–8. viii.1999, A. Fong, A. Sánchez, and D. Maceira , 1 ♀ with 41 1 st instars (BIOECO Sc-104) , 6 juv. (BIOECO Sc-105), 8.ii.2004, A. Sánchez, 2 juv. (BIOECO); Punta de Maisí, Cueva El Molino, 31.i.2006, D. de la Nuez , 1 ♀ (IES); Bahía de Ovando, Punta Negra (20°05′39″N 74°14′46″W), 8.viii.1998, A. Sánchez, D. Maceira, and A. Fong , 1 juv. (BIOECO Sc-103) .

DIAGNOSIS: Heteroctenus junceus differs from the other two species of Heteroctenus occurring on Cuba, H. garridoi and H. gibarae, primarily in its larger size and typically darker coloration. Heteroctenus junceus varies from 54–107 mm in total length, whereas H. garridoi and H. gibarae vary from 56–69 mm. Heteroctenus junceus varies from yellow to dark reddish brown, typically the interocular surface is infuscate, and the metasomal segments IV and V are noticeably darker than segments I–III, whereas H. garridoi and H. gibarae are more uniformly yellow to yellowish brown in color, with less contrast between the metasomal segments. Additionally, in H. junceus, there is a ventral infuscate stripe on metasomal segments I–III which is not the case in H. garridoi and H. gibarae . As in H. garridoi, the pedipalp chela manus of H. junceus is similar in color or only slightly darker than the pedipalp femur and patella, unlike H. gibarae, in which the manus is darker than the femur and patella. Additionally, the chela of H. junceus is more markedly sexually dimorphic, because the curvature of the fixed and movable fingers of the adult male is more pronounced, resulting in a larger gap between them proximally, when closed (fig. 27A), and the manus of the adult female is relatively more slender, than in H. garridoi and H. gibarae . The basal expansion of the pectines is also more pronounced in H. junceus than in H. garridoi and H. gibarae . Finally, H. junceus differs from H. garridoi and H. gibarae in the more coarsely granular intercarinal surfaces of the carapace and pedipalp chela manus.

DISTRIBUTION: Heteroctenus junceus is endemic to Cuba, and has been recorded across the island and on several offshore islets in the following provinces: Camagüey, Cienfuegos, Ciego de Avila, Granma, Guantánamo, Havana, Holguín, Isla de la Juventud, Las Tunas, Matanzas, Pinar del Río, Santiago de Cuba, Sancti Spíritus, Villa Clara. The known localities range in elevation from 55 to 716 m (fig. 3A). Historical reports from Haiti, Puerto Rico and Venezuela (see, e.g., Fet and Lowe, 2000: 220) are erroneous (Armas, 2001; Prendini et al., 2009).

ECOLOGY: Heteroctenus junceus inhabits a wide range of habitats across the island, including human-altered environments, savanna, secondary vegetation, wetlands, broadleaf forest, and deciduous dry forest (fig. 2B; Teruel and Armas, 2012a). It has been found in elevations ranging from sea level to 1175 m (Rodríguez-Cabrera and Teruel, 2014). This species is typically found in rock crevices, under stones or logs, and under the bark of trees (Teruel and Armas, 2012a). Its habitat and habitus are consistent with the lapidicolous ecomorphotype (Prendini, 2001b). On account of its broad distribution, many Cuban scorpion species have been recorded in sympatry with H. junceus .

REMARKS: The evidence supporting R. aridicola and R. melloleitaoi, as distinct from H. junceus, is underwhelming. Rhopalurus aridicola, described from four localities, overlapping known records of H. junceus, appears to be morphologically indistinguishable from the latter except for its dark reddish color and more granular carapace and tergites, characters known to be variable in the widespread H. junceus . Teruel and Armas (2012b) offer the additional evidence that the pectinal tooth counts of R. aridicola are greater than those of H. junceus, yet the counts provided for the two species (19–25 and 15–23, respectively), are statistically insignificant. Little genetic divergence was evident between topotypes of R. aridicola and other samples conspecific with H. junceus (Esposito et al., in review). Rhopalurus melloleitaoi, described from a single locality in close proximity to many known locality records of H. junceus, appears to differ from the latter solely on the basis of coloration, the pedipalp chelae being darker than the patella and femur, which is to be expected because the habitat in which it was collected has a dark, red substrate (L.A.E., personal obs.). Based on the lack of convincing morphological differences between R. aridicola, R. melloleitaoi, and H. junceus, their overlapping distributions and, in the case of R. aridicola, the low genetic divergence between topotypes thereof and other samples conspecific with H. junceus (see Esposito et al., in review), we propose the following synonyms: Rhopalurus melloleitaoi Teruel and Armas, 2006 = Heteroctenus junceus (Herbst, 1800), syn. nov.; Rhopalurus aridicola Teruel and Armas, 2012 = Heteroctenus junceus (Herbst, 1800), syn. nov.

MATERIAL EXAMINED: “Antillen?,” 1 ♂, 2 ♀ (ZMB 7370). “Portorico” [erroneous], Stahl, 2 ♀, 1 juv. (ZMB 7280). CUBA: Gundlach, 1 ♂, 1 ♀ (ZMB 738), 2 ♀ (ZMB 2637), 1 juv. (ZMB 7343); vii.2007, C. Hamilton, 1 juv. (AMCC [LP 7009]). Oriente Prov., 1938, P. Thumb , 4 ♀ (ZMH). Artemisa Prov.: near Baños [probably Santiago del los Baños], v.1918 , 2 ♂ (AMNH). Camagüey Prov.: Sierra de Cubitas, Limones-Tuabaquey Ecological Reserve, area around field station, 21°32.887′N 77°46.705′W, 55 m, 13–14.iv.2012, CarBio team , 2 ♂ (AMCC [LP 12619, 12620]), 2 subad. ♀ (AMCC [LP 12621, 12622]), 1 juv. ♂ (AMCC [LP 12623]). Guantánamo Prov.: Alejandro Humboldt National Park: near El Yunque de Baracoa, 20°19.907′N 74°34.151′W, 74 m, 4. iv.2012, CarBio team , 1 ♀, 26 juv. (AMCC [LP 12432]), 20°20.701′N 74°33.985′W, 370 m, 5. iv.2012, CarBio team , 1 ♂ (AMCC [LP 12613]), 20°19.64′N 74°35.59′W, 530 m, 6–7.iv.2012, Car- Bio team , 1 ♂, 2 ♀ (AMNH), 1 ♂ (AMCC [LP 12614]), 4 ♀ (AMCC [LP 12615–12618]). Havana Prov.: Havana, 1 ♀ (AMNH), iv.1941, E. Weiss, 1 ♀, 1 subad. (AMNH). Holguín Prov.: viii.2000, Heist, captive bred, 1 juv. (AMCC [LP 1928]); Guardalavaca, 29.iii.1993, W. Altmann, captive bred , 1 ♂ (AMCC [LP 1565]); Mayari, Parque Nacional “Mensura-Piloto,” 1 km después de la carretera al Hotel “Mayari,” 716 m, 10.v.2013, F. Cala-Riquelme and A. Deler-Hernández, secondary riverine forest with abundance of pines, collected in the evening and at night on vegetation and trunks , 1 juv. ♂ (AMCC [LP 12896]). Isla de la Juventud Prov: Isle of Pines, 1 ♂ (AMNH). Mayabeque Prov.: Arroyo Bermejo, near Fibacoa [Jibacoa], 15.vi.1967 , 1 ♀ (ZMB 31021), vi.1967, 1 juv. (ZMB 31022), 31.v.1967, Kleiderschrank, 1 ♂ (ZMB 31020); Guisa, mountains near, x.1936, P. Thumb, 1 ♀, 28 juv. (ZMH), Moa, ix.1937, P. Thumb, 1 ♂ (ZMH); Santiago de las Caballeros, 1936, P. Thumb, 1 ♂ (ZMH). Pinar del Río Prov.: Guanahacabiles, Akad.-stat. El Beral, xii.1967, G. Peters, 1 subad. (ZMB 31023); Sierra de Anafe, 23.ii.194, M. Barro , 2 subad. (AMNH); Sierra del Rosario, near Aspuru, xi.1937, H.H. Voelckers , 1 ♂, 1 ♀, 1 juv. ♀ (ZMH); Viñales National Park, near Dos Hermanas, 22°37.265′N 83°44.3′W, 130 m, 18.iv.2012, CarBio team , 2 juv. ♂ (AMCC [LP 12624, 12625]), 1 juv. ♀ (AMCC [LP 12626]), 22°39.424′N 83°42.097′W, 280 m, 20–21.iv.2012, CarBio team, 6 juv . ♂ (AMCC [LP 12627–12632]), 1 juv. ♀ (AMCC [LP 12633]); Viñales Valley, 1940, Osorio , 1 ♀ (AMNH). Sancti Spíritus Prov.: Trinidad, viii.1978, B. Acosta, 1 ♂ (AMNH [AH 4514]). Santiago de Cuba Prov.: La Socapa, 10 km SW of Santiago de Cuba, 9.iv.1999, R. Teruel, 1 ♂ (AMNH), 3 ♀ (AMCC [LP 1509, 1517, 1518]); Santiago de Cuba, 1 ♂, 2 juv. (AMNH) .