Centruroides edwardsii ( Gervais, 1843 ), 2011

de Armas, Luis F., Teruel, Rolando & Kovařík, František, 2011, On Centruroides margaritatus (Gervais, 1841) and closely related species (Scorpiones: Buthidae), Euscorpius 2011 (132), pp. 1-16 : 6-12

Centruroides edwardsii ( Gervais, 1843) ,

comb. nov.

( Figs. 3 , 4A–H , 5A–D , 6 A–E ; Table 1 )

Scorpio (Atreus) edwardsii Gervais, 1843: 130 ; Gervais, 1844: 208, 216–217, Pl. XI, figs. 13–15; Gervais, 1859: 41, Pl. I, fig. 1 (in part).

Scorpio (Atreus) degeerii Gervais, 1844: 217–218 , pl. 11, fig. 16–17, new synonymy. [Synonymized under C. margaritatus by Pocock (1893: 387)].

? Tityus ducalis C. L. Koch, 1844: 2–4 , pl. CCCLXI, fig. 851 [synonymized under C. margaritatus by Hoffmann (1932: 247)].

Centrurus gambiensis Karsch, 1879: 123–124 , new synonymy. [Synonymized under C. degeeri by Kraepelin (1891: 133)].

Centrurus margaritatus : Pocock, 1893: 387 (in part).

Centruroides margaritatus septentrionalis Hoffmann, 1932: 249 , 251–258, figs. 44–47, new synonymy. [Synonymized under C. margaritatus by Armas et al. (1995: 35)].

Rhopalurus danieli Prado et Rios-Patiño, 1940: 42–43 . New synonymy. [Synonymized under C. margaritatus by Sissom & Lourenço (1987: 16)].

Centruroides danieli : Mello-Leitão, 1945: 251, 253– 255, figs. 105–106; Flórez, 1990: 119.

Centruroides margaritatus : Mello-Leitão, 1945: 260– 265 (in part); Francke & Stockwell, 1987: 14–17, figs. 3, 39–47, 99 (misidentification); Armas & Hernández, 1989: 2–9 (misidentification); Armas, 1995: 2 (misidentification); Viquez, 1999: 46–47 (misidentification); Teruel, 2002: 87–88, figs. 1–2 (misidentification); Teruel & Stockwell, 2002: 119– 120, 126, fig. 10, 21, tables IV, VI (misidentification); Teruel & Roncallo, 2010: 1–2, figs. 1, 10 (misidentification); Teruel, 2011: 61–66, fig. 5 (misidentification).

Centruroides gracilis : Flórez, 1990: 117 (misidentification: records from Antioquia and Cundinamarca); Flórez, 2001: 28 (misidentification: Caribbean slope records); Gómez & Otero, 2007: 53, 54, 56, fig. 4 (misidentification).

Type data. Neotype adult ♂ (BIOECO, Ex– RTO: Sco-0395; herein designated, Fig. 4 A–H ): COLOMBIA, LA GUAJIRA, Riohacha City, Calle 6 esquina a Carrera 5, inside a box in house, 25 June 2008, leg. W. Ramos. Notes: This specimen was formerly determined as C. margaritatus by R. Teruel, 2008. It has 29/28 pectinal teeth, with one tooth missing for each pecten. All labels are written in Spanish.

Distribution. Mexico, El Salvador, Honduras, Nicaragua, Costa Rica, Colombia, and Cuba (introduced) ( Fig. 3 ). It has also been introduced in Senegal, Africa (see Comments below).

Diagnosis. A moderate to large species (60–110 mm). Body brown; pedipalp chelae, carapace, tergites I– VII, metasomal segment V and telson dark reddish brown (rarely, a yellowish longitudinal stripe is present over tergites I–VII, generally in immatures); pedipalp femur and patella, legs and venter yellowish brown; pectines light yellow; carinae on metasoma and pedipalps are dark brown ( Figs. 4 C, E, F ; 5 A–D , 6 AB ). Prosoma and tergites densely and coarsely granular, with large spiniform granules interspersed. Metasoma coriaceous, with all carinae strongly denticulate; telson with a spiniform subaculear tubercle directed toward the basal portion of the aculeus ( Figs. 4 H , 6 C–D ); segments II– IV with more than three (usually four) pairs of ventrolateral macrosetae. Pedipalps very densely hirsute; chela wider than patella and very robust in the adults, with dorsal marginal carina almost smooth, pilose ( Fig. 4 C–D , 6 E ); fingers with eight principal rows of granules, flanked internally and externally by numerous supernumerary granules, basal lobe/notch combination strong. Sexual dimorphism is evident: adult males have less inflated pedipalp chelae and a moderately slender metasoma, whereas adult females have more robust and shorter pedipalps, more inflated chelae, and nonelongate metasomal segments ( Figs. 5 A–D , 6 A–B ). Pectinal tooth counts 22–30 in ♀♀ (n = 110 combs), 26– 34 in ♂♂ (N = 90 combs). Measurements in Table 1 .

Comparisons. This species was largely confounded with C. margaritatus , from which it clearly differs by having pedipalps profusely pilose, hand with dorsal marginal carina almost smooth and very pilose ( Fig. 4 C–D ), telson with subaculear tubercle close to aculeus and directed towards its basal one-third ( Figs. 4 G , 6 C, D ), carinae on the pedipalp femur and patella with smaller and closer granules. Also, when same-sized adults of both species are compared side-by-side, it becomes evident that both sexes of C. edwardsii have pedipalps (especially chelae) and metasoma shorter and more robust than in C. margaritatus (see our Figs. 1–7 ).

With respect to its other close relatives, Centruroides edwardsii comb. nov. can be distinguished as follows: (1) Centruroides hirsuticauda Teruel, 2011 is remarkably smaller (♂♂ 69–74 mm, ♀♀ 53–57 mm), and has pedipalps and metasoma very densely hirsute (Teruel, 2011); (2) Centruroides exilimanus Teruel & Stockwell, 2001 is larger (♂♂ 89–150 mm, ♀ 98–116 mm), it has higher pectinal tooth counts (♂♂ 31–34, ♀♀ 30–32), and females have a deep discal pit in the basal plate of the pectines (Teruel & Stockwell, 2001; Víquez & Armas, 2005); (3) In C. chiapanensis and C. tapachulaensis , the pedipalps are attenuate, with chelae not globose, almost glabrous, having dorsal marginal carina granulose; (4) C. fallassisimus is a paler species, its pedipalp chelae are almost glabrous, with finely granular dorsal marginal carina; subaculear spine is moderately distant from the aculeus base and directed toward the aculeus tip.

Comments. Gervais (1843) briefly described Scorpio (Atreus) edwardsii from an unspecified number of syntypes from “ Colombia ”, reportedly deposited at MNHN, but Gervais (1844) provided a more precise description of this species, including an excellent colour figure, and mentioned “Carthagène de Colombie ” (p. 217) as the locality of the specimens examined by him.

Gervais (1844) based the original description of Scorpio (Atreus) degeerii upon another unspecified number of syntypes (measurements and figures of at least one adult female were actually given) from Chile and Colombia (Cartagena), also reportedly deposited at MNHN. The descriptions and figures of Gervais (1843, 1844), Hoffmann (1932) and Prado & Ríos-Patiño (1940) clearly show that Scorpio (Atreus) degeerii , Rhopalurus danieli and Centruroides margaritatus septentrionalis are all conspecific with Scorpio (Atreus) edwardsii , which is the senior synonym by page precedence ( CINZ, 2000). Tityus ducalis was described from Mexico, but it was synonymized under C. margaritatus by Hoffmann (1932); Fet & Lowe (2000: 112) included it with doubt as a junior synonymy of C. margaritatus , because the type specimens are lost, and the synonymy cannot be verified (Sissom & Lourenço, 1987: 16).

Sissom & Lourenço (1987: 16) stated that the original types of Gervais (1843, 1844) cannot be positively identified among the MNHN specimens labelled Centrurus edwardsii and Scorpio degeerii . Because of this reason, plus the fact that only with available namebearing types the taxonomic status of all members of this very complex group of closely related species can be clarified and stabilized, we have proceeded to designate a neotype for C. edwardsii .

Armas (1981a: 1, 8–9) recorded Centruroides margaritatus ssp. from Santo Domingo, Dominican Republic, on the basis of a lot of nymphs labelled “ Santo Domingo ”, without additional data and deposited at the Museum of Comparative Zoology, Harvard University, USA. As that species has never been found in Dominican Republic, we think that the correct locality of these immature specimens is Santo Domingo, Heredia Province, Costa Rica, but we are not sure that they belong to C. edwardsii .

We have seen excellent photos of the holotype of Centrurus gambiensis ( Fig. 7 A–D ) and concluded that it is actually a junior synonym of Centruroides edwardsii , as previously pointed out by Kraepelin (1891) as Centrurus degeerii . By the way, the type-locality of C. gambiensis has been erroneously stated to be the archipelago (and also independent country) of Cape Verde (i.e., Sissom & Fet, 2000: 113), but this is an incorrect transcription. Two of the original collecting labels accompanying the holotype clearly read “Cap Vert” (see our Fig. 7E ), but this toponym (either in German or French) actually corresponds to the Cape Vert peninsula in western central Senegal, mainland Africa (Microsoft Corporation, 2009).

Francke & Stockwell (1987, as C. margaritatus ) recorded pectinal tooth counts of 26–34 (mode 30) in ♂♂, and 24–33 (mode 28) in ♀♀ from Costa Rica. Nevertheless, Costa Rican populations are apparently composed of two species, which Víquez (1999: 47) referred to as “Central Valley variety” and “forest habitat variety”, without any further discussion on this matter. According to the specimens available to us, the most widespread and commonly collected is C. edwardsii , but the other taxon cannot be accurately identified yet, mostly because our sample is too small and not well-preserved.

As pointed out by Teruel & Roncallo (2010: 2), the Colombian population of C. edwardsii has the base colour of the body and appendages light yellowish, with a more contrasting dark pattern. But apart from this, we have not found any significant morphological differences with respect to the remaining specimens from Mexico, Central America, and Cuba.

Material examined. MEXICO: DURANGO: 2♀♀, 1♂ ( IES), Tayoltita , August 1993, leg. R. Herrera . NAYARIT: 1♂ ( FKCP), Rosatin (= Rosalía ?), 19 August 1975, leg. van den Berghe. HONDURAS: GRACIAS A DIOS: 1♀ (RTO: Sco-0168) , 2♀♀ ( USNM), Mocorón Army Base , 10 October 1990, leg. S. A. Stockwell ; 1♀ ( USNM), Puerto Lempira, 25 April 1984; 1♂♂ ( IES), León, 1991, leg. J.M. Maes. VALLE: Nacaome: 2♀♀, 3♂♂, 1 juvenile ( RTO, ex MM-S182), Jícaro Galán , 7–8 May 2000, leg. M. Montoya. NICARAGUA: REGIÓN AUTÓNOMA DEL ATLÁNTICO NORTE: Bosawas, 1♀♀, 2♂♂ ( FKCP), 18 km SW Ayapal, Bocay River watershed, Mt. Kum , 500 m, 28–31 May 1999, leg. van den Berghe. ESTELÍ: 2♀♀, 1 juvenile ♀, 1juvenile ♂ ( IES), El Rodeito, 16 km W Estelí , December 1985, leg. J. M. Maes. MATAGALPA: 1♂ ( IES), Las Tres Ceibas, road Telica to San Isidro Km 168, 21 August 1995, leg. L. F. Armas & J. M. Maes , eating Didymocentrus krausi under a stone, at 07:45 hr. LEÓN: 11♀♀, 6♂♂ ( IES), Base del volcán Momotombo , 6 August 1995, leg. Maes, Armas & Johnson , under bark in a wired fence post (one of the females carried 104 [one hundred and four – sic! – Eds.] offspring). 2♀♀, 1♂, 1 juvenile ( IES), Cerro Casitas , 15 August 1995, leg. Maes & Armas , under bark, 600 m; 1♂ ( IES) , León, March 1990, leg. B. Garcete ; 2♂♂ ( IES) , León, no additional data; 1♀ ( IES) , León, December 1990, leg. B. Garcete ; 1♀ ( IES) , León, June 1994, leg. Maes & Tellez ; 2♀♀ ( IES), El Sauce, December 1985, leg. L. R. Hernández . MANAGUA: Laguna Xiloá : 1♀, 1♂, 1 juvenile (RTO: Sco-0178), 1♀, 1♂, 3 subadults ( IES), 13 August 1995, leg. L. F. de Armas, J. M. Maes , T. Goodwin ; 1♂, 2 subadults ( IES), 2 0 April 1994, leg. E. van den Berghe , under stones; 1♂ ( FKCP), 8 December 1994, leg. van den Berghe ; 1♂ ( FKCP), 10 December 1994, leg. van den Berghe ; 1♂ ( FKCP), 29 December 1994, leg. van den Berghe ; 1♀ ( FKCP), 5 December 1995, leg. van den Berghe ; 1 ♀♀, 1 juvenile ( FKCP), March 1997, leg. van den Berghe ; 2♀♀, 1♂ ( FKCP), Chocoyero , October 2000, leg. van den Berghe ; 1♂ ( FKCP), El Crucero , 10 December 1994, leg. van den Berghe ; 2♀♀, 1♂ (RTO: Sco-0179) , Managua, 2 November 1989, leg. S. Jacobs ; 1♂ ( IES), Villa Nejapa, old road to León Km 10, October 1985, leg. L. R. Hernández . MASAYA: 2 ♀♀, 1♂, 1 immature ( FKCP), Laguna de Apoyo , 1 November 1994, leg. van den Berghe ; 1♀ ( IES), Mirador del Eco , Volcán Masaya, December 1985, leg. L. R. Hernández . GRANADA: 1♀ ( FKCP), Volcán Mombacho , 1100 m, 10 February 2001, leg. van den Berghe ; 1 ♂ ( FKCP), Reserva Domitila , 15 May 2001, leg. van den Berghe ; 3♀♀, 1♂ ( FKCP), Domitila Reserve , 27–28 December 2002, leg. van den Berghe. RIVAS: 1♀, 2♂♂ ( FKCP), Reserva La Flor , 25– 30 January 2003, leg. van den Berghe ; 2♀♀ ( FKCP), Reserva La Flor, 16–18 February 2001, van den Berghe; 3♀♀, 1♂ ( FKCP), La Flor vic., 4 September 2003, leg. van den Berghe ; 3♀♀, 2 immatures ( FKCP), Rancho Santana vic., 6 April 2003, leg. van den Berghe ; 1♀, 1♂, 2 immatures ( FKCP), Rancho Santana , December 2002, leg. van den Berghe. RÍO SAN JUAN: 1♀ ( IES), Solentiname , 30 July 1989, leg. F. Reinboldt. COSTA RICA: GUANACASTE: 1♂, 2 juveniles (RTO: Sco-0174), 1.5 km NE Puerto Carrillo, 11 April 1997, leg. C. Herrera ; 2♂♂ (RTO: Sco-0170), 29 September 1997, leg. C. Herrera ; Tilarán, 1♀, 1♂ ( RTO, ex MM-S009), Quebrada Grande , 3 February 1996, O. Araya, leg. I. Sánchez ; 1♂, 1juvenile ( IES, ex MM-S019), 20 February 1996, leg. M. González, I. Sánchez ; Tilarán, 1♀ ( IES, ex MMS022 ), Los Ayotes , 10 March 1996, leg. Y. Vargas, I. Sánchez ; 1♂ (RTO: Sco-0171), Estación Murciélago , 26 May 1996, leg. F. A. Quesada ; Santa Rosa National Park: fire cut to Playa Naranjo: 1♂ (RTO: Sco-0172), around campsite and Piñuelas trail, 6 February [year?]; 1♂ ( FKCP) , Guanacaste, Santa Rosa National Park , 310 m., June 2004, leg. S. Pokorný ; 2♀♀ ( FKCP), Boruca , March 2007, leg. V. Šejna & J. Hromadka . HEREDIA: 1♂ (RTO: Sco-0175), San Joaquín de Flores , 200 m N church, 30 March 1996 . ALAJUELA: Urbanización Punta del Este : 1 ♂ (RTO: Sco-0176), Desamparados, 15 February – July de 1997, leg. M. Castro. CUBA: MATANZAS: 1♂ ( RTO), Varadero , January 2000, leg. L. Velazco. COLOMBIA: MAGDALENA: 1 juvenile (RTO: Sco-0422), Santa Marta , January 2009, leg. J. A. Vargas. LA GUAJIRA: 1♀ (RTO: Sco-0360), Riohacha , 8 November 2006, leg. C. A. Roncallo ; 1♂ (RTO: Sco- 0375), 25 July 2007, leg. C. A. Roncallo ; 1 juvenile (RTO: Sco-0421), 5 October 2008, leg. C. A. Roncallo ; 1 ♂ (RTO: Sco-0420), August 2009, leg. C. A. Roncallo. Serranía de Macuira : 1♀ (RTO: Sco-0374), 3 km W Nazareth , 14 July 2007, leg. J. Echavarría.

Figures

Figure 1: Centruroides margaritatus. Female holotype. A, dorsal; B, ventral; C, detail of dorsal aspect of carapace; D, last segments of the metasoma and telson, lateral aspect; E, pectines; F, labels. Scale (A, B) = 10 mm. Photos courtesy of E.-A. Leguin.

Figure 2: Centruroides margaritatus from Guayaquil (A–B, D, F) and Loja (C, E), Ecuador. A–C, male: A, habitus, dorsal; B– C, telson, lateral aspect. D–F, female: D–E, pedipalp, dorsal aspect; F, metasomal segment V and telson, lateral aspect. Scale (in mm) is only for figure A. Photos C and E, courtesy of J. A. Ochoa Camara.

Figure 3: Geographical distribution of Centruroides edwardsii (black squares) and C. margaritatus (white squares).

Figure 4: Centruroides edwardsii. Male neotype. A, carapace and tergites; B, prosoma and mesosoma, ventral aspect; C, right chela, dorsal aspect; D, hand, dorsal aspect; E–H, metasoma: E, segments I–II, dorsal aspect; F, segments I–III, ventral aspect; G, segment V, lateral aspect; H, telson, lateral aspect.

Figure 5: Centruroides edwardsii. Males from Guanacaste, Costa Rica (A), La Guajira, Colombia (B), Varadero, Matanzas province, Cuba (C), and Durango, Mexico (D). Scale in millimeters only for D.

Figure 6: Centruroides edwardsii. Female. A–C from Sierra de Macuira, La Guajira, Colombia (RTO: Sco-0374); D–E from Tilarán, Guanacaste, Costa Rica (MM-S009). A, dorsal; B, ventral aspect; C–D, telson, lateral aspect. E, pedipalp chela, dorsal aspect.

Figure 7: Centrurus gambiensis. Male holotype. A-B, dorsal (A) and ventral (B) aspect, except metasomal segments II-V and telson); C, metasomal segments II-V and telson, dorsolateral aspect; D, metasomal segment V and telson, lateral aspect; E, labels. Photos courtesy of Anja Friederichs.