Redescription of Rhopalurus abudi (Scorpiones, Buthidae), with first description of the male and first record from mainland Hispaniola.
Abstract: Rhopalurus abudi Armas & Marcano Fondeur 1987 was originally described on the basis of a single female specimen from Isla Saona, La Romana Province, off the southeast coast of the Dominican Republic. The species is redescribed here based on a series of new specimens including 19 adult males and 14 adult females collected at two nearby localities on the eastern side of Parque Nacional del Este, La Altagracia Province, southeastern Dominican Republic. These specimens represent the first records of R. abudi on mainland Hispaniola and the first male specimens of the species to be collected. Keywords: Alacran, Caribbean, Dominican Republic, Parque Nacional del Este, taxonomy, biogeography
Article Type: Report
Subject: Zoology (Identification and classification)
Zoology (Research)
Authors: Prendini, Lorenzo
Esposito, Lauren A.
Huff, Jeremy C.
Volschenk, Erich S.
Pub Date: 05/01/2009
Publication: Name: Journal of Arachnology Publisher: American Arachnological Society Audience: Academic Format: Magazine/Journal Subject: Biological sciences; Zoology and wildlife conservation Copyright: COPYRIGHT 2009 American Arachnological Society ISSN: 0161-8202
Issue: Date: May, 2009 Source Volume: 37 Source Issue: 2
Topic: Event Code: 310 Science & research
Geographic: Geographic Scope: Dominican Republic Geographic Code: 5DOMN Dominican Republic
Accession Number: 238834226
Full Text: The buthid scorpion genus Rhopalurus Thorell 1876 comprises 18 species and three subspecies (one nominotypical) of relatively large, lapidicolous (Prendini 2001a) scorpions with a discontinuous distribution in the Greater Antilles (Cuba and Hispaniola) and northern South America (Brazil, Colombia, Guyana, and Venezuela) (Appendix 1). These scorpions are unique in possessing the ability to stridulate audibly by scraping nodules and/or ridges on the dorsal surfaces of their pectines against granules on the ventral surfaces of mesosomal sternite III, a remarkable behavior that presumably functions to deter would-be predators (Pocock 1904; Lourenco & Cloudsley-Thompson 1995; Armas 2001; Lourenco 2007). Lourenco (1986) considered the stridulation organ to be synapomorphic for Rhopalurus, a hypothesis that has yet to be tested cladistically.

The taxonomic distinction between Rhopalurus and another New World buthid scorpion genus, Centruroides Marx 1890, distributed from the southwestern USA throughout Mexico, Central America, the Greater and Lesser Antilles, to northern South America (Colombia, Ecuador, and Venezuela), remains unclear. The two genera are separated primarily according to the presence, in Rhopalurus, of the stridulation organ on opposing surfaces of sternite III and pectines, which is absent in Centruroides (Lourenco 1979; Sissom 1990). The stridulation organ is variably developed within the genus, however, and the species of Rhopalurus form a rather heterogeneous assemblage in other respects. Evidence from ovariuterine morphology (Volschenk et al. 2008) and DNA sequences (L.A. Esposito, E.S. Volschenk & L. Prendini, in prep.) suggests that Rhopalurus may be paraphyletic with respect to Centruroides.

Rhopalurus was last revised by Lourenco (1982). Numerous changes to its composition have been made since then (Lourenco 1984, 1986, 2002, 2007; Armas & Marcano Fondeur 1987; Lourenco & Pinto-da-Rocha 1997; Armas 1999; Lourenco et al. 2004; Lenarducci et al. 2005; Teruel 2006; Teruel & Armas 2006; Teruel & Roncallo 2008; Teruel & Tietz 2008; Lourenco 2008). These include the description of 10 new species, one of which was subsequently synonymized, and two new subspecies; the resurrection of a species previously placed in synonymy; the elevation of a subspecies to species rank; the resurrection of a monotypic genus, Physoctonus Mello-Leitao 1934, to accommodate a species once placed in Rhopalurus; and the creation of another monotypic genus, Troglorhopalurus Lourenco et al. 2004, to accommodate a new troglomorphic species. The validity of Physoctonus and Troglorhopalurus is presently unclear. The systematics of Rhopalurus and related genera warrants reinvestigation, including detailed morphological revision and rigorous cladistic analysis based on morphological and molecular data.

Three species of Rhopalurus are endemic to Hispaniola (Armas 1999, 2001; Fet & Lowe 2000; Teruel 2005, 2006; Fig. 1). Rhopalurus abudi Armas & Marcano Fondeur 1987 (Figs. 2, 5A, B, 6A, 7A, 8, 11) and Rhopalurus bonettii Armas 1999 (Figs. 3, 5C, D, 6B, 7B, 9) are endemic to the Dominican Republic (DR), whereas Rhopalurus princeps (Karsch 1879) (Figs. 4, 5E, F, 6C, 7C, 10) also occurs in Haiti. Rhopalurus abudi, described on the basis of a single female specimen from Isla Saona, La Romana Province, off the southeast coast of the DR, is the least known of the three species and among the least known species in the genus. No new records of this species have been reported in the literature since the original description (Armas & Marcano Fondeur 1987; Armas et al. 1999; Teruel 2005, 2006). Lourenco & Pinto-da-Rocha (1997:181) suggested that it may be a junior synonym of R. princeps (see also Fet & Lowe 2000:217).

In July 2004, an expedition to collect arachnids in the DR was conducted by EV and JH. During the course of that expedition, a series of new specimens of R. abudi, including 19 adult males and 14 adult females, was collected in humid coastal forest at two nearby localities on the eastern side of Parque Nacional del Este, La Altagracia Province, southeastern DR. These specimens represent the first records of R. abudi on mainland Hispaniola, the first male specimens of the species to be collected, and the first records of a Rhopalurus species from a humid coastal forest habitat. On the basis of this new material, we provide a detailed redescription of R. abudi, including a comparison with the other two species of Rhopalurus endemic to Hispaniola.

METHODS

Specimens were collected using ultraviolet (UV) light detection at night or by rolling limestone boulders during the day. Geographical coordinates and elevation were recorded with a portable Garmin GPS V Personal Navigator device, using the WGS84 datum. Most specimens were preserved in the field in 75% ethanol. One specimen from each locality was preserved in 95% ethanol for future DNA isolation.

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Specimens were examined using a Nikon SMZ1500 dissection stereomicroscope. Hemispermatophores were dissected following the method described by Prendini et al. (2006) and the soft paraxial tissues dissected away from the capsule area using minuten entomology pins prior to examination in 75% ethanol. Specimens for which tissue could not be completely removed from the capsule area were dehydrated in ethanol (80% for 10 min, 95% for 10 min), followed by isopropanol (100% for 10 min), and then cleared in clove oil for ~ 20 min. Specimens were measured using Mitutoyo[R] digital calipers and an ocular micrometer. Ultraviolet fluorescence and conventional light photomicrographs were prepared, following a modified version of the method outlined by Volschenk (2005), using a Microptics[TM] ML-1000 digital imaging system, and the digital images subsequently edited and prepared into plates with the aid of Adobe Photoshop and Corel Draw.

Specimens of R. abudi, other species of Rhopalurus and related taxa studied for comparison (Appendix 2) are deposited in the following collections: American Museum of Natural History (AMNH), New York, USA, incorporating the Alexis Harington (AH) Collection; Natur-Museum Senckenberg, Frankfurt (SMF), Germany; Zoologisches Museum der Humboldt-Universitat, Berlin (ZMB), Germany; Zoologisches Museum der Universitat Hamburg (ZMH), Germany. Reference numbers (ESV and LP), provided on labels with the specimens, correspond to entries in the specimen databases of the author with the corresponding initials.

General anatomy follows Hjelle (1990) and Sissom (1990), trichobothria follows Vachon (1974), carination follows Prendini (2001b), and hemispermatophore follows reinterpretation of the character system in Buthidae, to be described fully elsewhere. Ovariuterine anatomy follows Volschenk et al. (2008). Measurements follow Stahnke (1970), Lamoral (1979), and Prendini (2001b).

TAXONOMY Family Buthidae C.L. Koch 1837 Genus Rhopalurus Thorell 1876 Rhopalurus abudi Armas & Marcano Fondeur 1987 (Figs. 2, 5A, B, 6A, 7A, 8, 11) Rhopalurus abudi Armas & Marcano Fondeur 1987:19-20, fig. 4, pl. II, tab. 10; Rudloff 1994:9; Lourenco & Pinto-daRocha 1997:181; Kovarik 1998:118; Armas 1999:127; Armas et al. 1999:30-32; Armas 2001:246, tab. 1; Fet & Lowe 2000:217; Fet et al. 2003:3, tab. 1; Teruel 2005:165; Armas 2006:6; Teruel 2006:50, 51, fig. 12 e; Teruel et al. 2006:220, 221, 223, fig. 1; Volschenk et al. 2008:653, 658, 659, 663, 664, 674, fig. 1D, tab. 1, tab. 2.

Material examined.--DOMINICAN REPUBLIC: La Altagracia Province: Parque Nacional del Este: Cabo Flaso (entrance zone), 18[degrees]22'25"N, 68[degrees]37'01"W, 14 July 2004, E.S. Volschenk & J. Huff, 67.7 m, 1 [male] (AMNH [ESV6091]); Track between ranger station (at Boca de Yuma) and Punta Faustino, 18[degrees]21'17.2"N, 68[degrees]36'52.3"W, 14 July 2004, E.S. Volschenk & J. Huff, 3.3 m, dense canopy humid forest, hand collected at night with blacklights, from limestone outcrops, especially along an old rock wall along the start of the track, 1 [female], 48 first instars (AMNH [ESV6010]), 1 [female], 22 first instars (AMNH [ESV6019]), 1 [female] 32 first instars (AMNH [ESV6039]), 11 [male],4 [female], 1 subad. [female], 1 subad. [female], 2juv. (AMNH [ESV6072]), 1 [male],1 [female] (AMNH [ESV7110]), 1 [male],1 [female] (AMNH [ESV7117]), 1 [male], (AMNH [ESV7120]), 1 [male],1 [female] (AMNH [ESV7242]), 1 [male],1 [female] (AMNH [ESV7303]), 1 [female] (AMNH [ESV7306]), 1 [male],1 [female] (AMNH [ESV7705]), 1 [male],1 [female] (AMNH [ESV7937]), 3 juv. (AMNH), 1 juv. (AMNH [LP 3268]).

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Relationships.--Based on morphological similarity, R. abudi appears to be most closely related to R. bonettii, and was compared directly with the latter by Armas (1999: 127). When compared to R. princeps, the third Rhopalurus species occurring on Hispaniola, R. abudi and R. bonettii are similar in carinal development and shape of the carapace (Figs. 5A, C, E; Tables 1-3); development of the pectines (Figs. 5B, D, F); carinal development and length of the pedipalp chela manus (Figs. 6, 7; Tables 1-3); and length of the metasomal segments (Figs. 8-11; Tables 1-3). Lourenco & Pinto-daRocha (1997:181) suggested that R. abudi may be a junior synonym of R. princeps but the two species differ in many respects (Figs. 2, 4, 5A, B, E, F, 6A, C, 7A, C). Rhopalurus princeps appears to be more closely related to Rhopalurus species on Cuba than to R. abudi and R. bonettii.

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Diagnosis.--Rhopalurus abudi differs conspicuously from R. bonettii on the basis of the sexually dimorphic pedipalp chelae of the adult male. This dimorphism is well developed in R. abudi: the chela manus of the adult male is incrassate and the fingers strongly curved proximally (fixed finger curved dorsally, movable finger curved ventrally), such that only the distal portion of the fingers connect and a distinctive gap is present between them proximally, when closed (Fig. 6A). The chela manus of female R. abudi is not incrassate and the fingers are not curved proximally, such that the fingers connect along most of their length and little to no gap is present between them proximally, when closed (Fig. 7A). This dimorphism is considerably less developed in R. bonettii,in which the male and female chelae are similar, the manus of the male being only slightly incrassate, relative to the female, and the fingers not curved proximally, such that the fingers connect along most of their length and little to no gap is present between them proximally, when closed (Figs. 6B, 7B).

The two species differ further in development of the pectines. The pectines ofR. bonettii are very broad proximally, with a more pronounced basal plate (Armas 1999), and the first 6-7 pectinal teeth are noticeably larger than the rest (Fig. 5D), compared to R. abudi, in which the pectines are narrower proximally, with a less pronounced basal plate, and the pectinal teeth similar in size (Fig. 5B). These morphological differences appear to be associated with differences in behavior. In the field, R. bonettii was observed to stridulate more loudly than R. abudi (and R. princeps, in which the pectines are even less developed).

Other differences between the two species are as follows. The coloration of R. abudi is darker, due to extensive infuscation, than R. bonettii, which is pale (Armas 1999; Figs. 2, 3). The carapace, pedipalp chelae, legs and tergites are noticeably infuscated in R. abudi but pale in R. bonettii. The metasoma and telson of R. abudi are strongly infuscated laterally and ventrally, especially on segments II-IV, becoming more so distally, with each segment darker than the preceding one and segment V darkest. The metasoma and telson of R. bonettii are weakly infuscated on segments III-V or IV and V only. The carapace and tergites are more coarsely and densely granular in R. abudi than in R. bonettii. The submedian sulci of sternite III are convergent in R. abudi and subparallel in R. bonettii (Armas & Marcano Fondeur 1987; Armas 1999; Figs. 15, 17). The pale, raised posteromedial surface of sternite V in the male is more prominent in R. bonettii than in R. abudi. The metasomal segments of R. abudi are shorter and broader (i.e., the width/length ratio is smaller) than those of R. bonettii, which are longer and narrower (i.e., the width/length ratio is greater) (Armas 1999; Figs. 8, 9, Tables 1, 2). The granulation, ventromedian, and ventrolateral carinae of metasomal segment V are less developed compared with those of the preceding segments such that the segment has a shinier, rounded appearance in R. abudi (Fig. 8). The granulation, ventromedian, and ventrolateral carinae of metasomal segment V are more developed in R. bonettii such that the segment has a matt, angular appearance (Fig. 9).

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Description.--The following description is based on the specimens illustrated in Figs. 2, 5A, B, 6A, 7A, 8, 11 and listed in Table 1.

Coloration: Chelicerae brownish-yellow with finely reticulate infuscation on manus, becoming more intense distally; fingers brownish-yellow, not infuscated, teeth darker due to sclerotization. Carapace brownish-yellow with darker tanbrown patches of infuscation around median ocelli (Figs. 2A, C); lateral surfaces, carinae and lateral ocular tubercles with blackish-brown infuscation. Pedipalp femur and patella brownish-yellow, carinae noticeably darker; femur lightly and uniformly infuscated; patella not infuscated; chela manus reddish-brown, darker than femur and patella, entirely infuscated, becoming gradually darker towards base of fingers; chela fingers infuscated, becoming gradually paler distally. Legs pale brownish-yellow; external surfaces of femur and patella lightly and uniformly infuscated. Mesosoma brownish-yellow with broad, transverse band across each tergite; pretergites infuscated, tan-brown; post-tergites with reticulate infuscation concentrated near carinae, becoming paler posteriorly; posterior margins pale brownish-yellow, not infuscated. Sternites tannish-brown, without infuscation; VII with darker carinae. Metasomal segments infuscated, becoming gradually darker ventrally and posteriorly, carinae darker than intercarinal surfaces; each segment darker than preceding segment, I and II, tan-brown, III, reddish-brown, IV and V, dark to very dark tan. Telson dark reddish-brown, not infuscated; aculeus black distally.

Chelicerae: Movable finger, ventral surface with two subdistal teeth; distal external and distal internal teeth equal, opposable. Fixed finger, ventral surface with single denticle; ventral surface with dense brush of long, fine macrosetae.

Carapace: Carapace coarsely and sparsely granular, mainly on interocular and lateral surfaces. Anterior and posterior margins of carapace procurved; anterior margin with shallow median notch (emargination), without median projection (epistome) (Fig. 5A). Lateral ocular tubercles each with three macro-ocelli and one (anterior) micro-ocellus, situated dorsal to posterior macro-ocellus; posterior micro-ocellus absent. Median ocelli considerably larger than lateral ocelli, situated anteromedially. Median ocular macrosetae fine, acuminate; lateral ocular macrosetae absent. Ocular tubercle with pair of costate-granular superciliary carinae, protruding slightly above median ocelli. Five other pairs of costate-granular carinae present, disconnected. Anteromedian sulcus moderately deep, ovate; posteromedian sulcus narrow, shallow anteriorly, deep posteriorly; posterolateral sulci shallow, wide, curved; posteromarginal sulcus deep, narrow.

Sternum: Subtriangular (Fig. 5B). Median longitudinal sulcus Y-shaped, shallow anteriorly, deep and narrow posteriorly.

Genital operculum: Completely divided longitudinally; macrosetae evenly distributed. Genital papillae present ([male]), absent ([female]).

Pectines: Pectines broad (Fig. 5B), dorsal surfaces with stridulatory nodules. First proximal median lamella unmodified in [female]. Fulcra prominent. Proximal pectinal teeth not noticeably larger than others in [male] and [female]. Pectinal tooth counts, 17-24 ([male]), 19-22 ([female]).

Pedipalps: Femur with five distinct carinae; dorsoexternal, dorsointernal, ventrointernal and externomedian carinae continuous, costate-granular; internomedian carina discontinuous, comprising row of isolated spiniform granules; externomedian and dorsoexternal carinae each with an acuminate macroseta distally; intercarinal surfaces finely and uniformly granular.

Patella with seven distinct carinae; dorsomedian, dorsointernal, ventrointernal, ventroexternal, externomedian, dorsoexternal carina continuous, costate-granular; internomedian carina discontinuous, comprising several large, well-spaced spiniform granules proximally, becoming smaller distally; proximal tubercle moderately developed; dorsointernal carina not fused with ventrointernal carina; intercarinal surfaces smooth, except for ventral surface which is finely granular.

Chela manus ([male]) incrassate, length along ventroexternal carina 33-51% greater than manus width, 32-37% greater than manus height (Table 1), fingers strongly curved proximally (fixed finger curved dorsally, movable finger curved ventrally), such that only connect distally and distinctive gap present between them proximally, when closed (Fig. 6A); manus (9) not incrassate, length along ventroexternal carina 72-85% greater than manus width, 69-71 % greater than manus height (Table 1), fingers not curved proximally, such that connect along most of length and little to no gap present between them proximally when closed (Fig. 7A). Chela with five distinct carinae; dorsomedian, dorsal secondary and ventroexternal carinae continuous, costate-granular (Figs. 6A, 7A); digital carina continuous, costate-granular, becoming obsolete proximally; dorsointernal carina discontinuous, comprising row of small granules distally, becoming obsolete proximally; other carinae absent; intercarinal surfaces smooth, except for internal surface where several low granules present. Movable finger with small lobe (eminence) proximally; movable finger length 72-89% (L) or 99-105% (9) greater than length along ventroexternal carina (Table 1); dentate margins of fixed and movable fingers each with eight oblique denticle rows, in addition to short apical row of four denticles; each row terminating in large denticle at proximal and distal ends; rows slightly imbricated, terminal denticle of each row displaced distally from the main row by space of one or more denticles; internal and external supernumerary denticles present in addition to internal and external accessory denticles; fingers each with an enlarged terminal denticle.

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Trichobothria: Orthobothriotaxic, Type A, [alpha] configuration (femoral trichobothria [d.sub.1] and [d.sub.4] situated closer to dorsoex ternal carina than [d.sub.3]), with the following segment totals: femur II (5 dorsal, 4 internal, 2 external), patella 13 (5 dorsal, 1 internal, 7 external), and chela 15 (8 manus, 7 fixed finger). Total number of trichobothria per pedipalp, 39. Femoral trichobothrium [d.sub.2] similar in size to [d.sub.1,] situated internal to dorsointernal carina; [d.sub.4] smaller than [d.sub.1;] [d.sub.5] situated distinctly proximal to [e.sub.1;] [e.sub.1] considerably smaller than [e.sub.2.] Patellar trichobothrium [d.sub.2] considerably smaller than [d.sub.1;] [d.sub.3] situated external to dorsomedian carina. Chela trichobothrium E[b.sub.1] smaller than E[b.sub.2] and E[b.sub.3;] E[b.sub.1-E[b.sub.3]] situated proximally on manus; [V.sub.2] larger than, and situated close to [V.sub.1;] Est smaller than Em and Et, which are similar in size; esb smaller than eb; esb and eb situated near base of fixed finger; db situated between est and et; dt situated distal to et.

Legs: I and II, tibiae and basitarsi each with paired rows of fine, acuminate macrosetae on pro- and retrolateral surfaces. III and IV, tibiae without spurs; basitarsi prolateral pedal spur with one acuminate seta, basal lobe pointed and stout; retrolateral pedal spur asetose. I-IV, telotarsi each with paired ventrosubmedian rows of fine, acuminate macrosetae; laterodistal lobes truncated; median dorsal lobes extending to ungues; ungues short, distinctly curved, equal in length.

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Mesosoma: Tergites entirely granular, finely on pretergites, coarsely on post-tergites, becoming more so distally; I-VII each with a strongly developed, granular dorsomedian carina; VII additionally with distinct pairs of costate-granular dorsosubmedian and dorsolateral carinae. Sternites III-VI smooth, acarinate, each with pair of narrow, slit-like respiratory spiracles (Figs. 2B, D); III with smooth, raised ridge medially, with stridulatory granules submedially; V with prominent pale, raised surface posteromedially in adult L, and 6-10 evenly spaced short, acuminate macrosetae along posterior margin; VII finely granular laterally and medially, with pair of costate-granular ventrosubmedian and ventrolateral carinae.

Metasoma: Segments I-V progressively increasing in length (Fig. 8; Table 1), segment V 51-57% ([male]) or 52-59% ([female]) longer than segment I; segments stout, width/length segment I, 65-69% (L) or 69-71% ([female]), II, 57-58% ([male]) or 55-57% ([female]), III, 51-56% (L) or 51-57% ([female]), IV, 51-54% ([male]) or 49-52% ([female]), and V, 43-44% ([male]) or 34-41 % ([female]). Intercarinal surfaces uniformly finely granular. Segments I-IV, paired dorsosubmedian and dorsolateral carinae continuous, costate-granular, granules gradually becoming larger posteriorly, without associated macrosetae; paired ventrolateral and ventrosubmedian carinae continuous, costate-granular, granules subequal; median lateral carinae continuous, costate-granular, fully developed on segment I, obsolete, granular, restricted to the posterior two-thirds of segment II, absent on segments III-V. Segment V, dorsosubmedian carinae absent; dorsolateral and ventrolateral carinae continuous, costate-granular, granules subequal; ventrosubmedian carinae obsolete, granular, reduced to anterior half of segment; ventromedian carina continuous, costate-granular, granules subequal, without posterior bifurcation.

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Telson: Vesicle globose, height/length 52-57%, with flat dorsal surface and rounded ventral surface, slightly compressed anteroventrally (Table 1); slightly narrower than metasomal segment V, width 59-60% ([male]) or 63-77% ([female])of segment V. Subaculear tubercle absent (Fig. 8B). Ventrolateral and ventrosubmedian carinae absent; ventromedian carina continuous, granular. Vesicle surfaces with scattered granules, sparse microsetae, and fewer than 16 macrosetae. Aculeus long, 73-80% ([male]) to 81-85% ([female]) of vesicle length (Table 1), strongly curved.

Male hemispermatophore: Flagelliform, flagellum gradually tapering along its length, folded against shaft (Fig. 11); basal process lobate longitudinally; distal process terminating adjacent to base of flagellum (in dorsal aspect), rib-like and extending longitudinally; distal lobe represented by shelf at base of flagellum; median lobe not developed; internobasal inflection absent; external lobe present, separated from distal process; with small, longitudinally-oriented costate process.

Female reproductive system: Ovariuterine network comprising three longitudinal and ten transverse tubules, forming eight "cells."

Geographic variation: The single male specimen from Cabo Flaso is similar to those from the track between Boca de Yuma and Punta Faustino.

Ontogenetic variation: As in other species of Rhopalurus, male closely resembles female until the final instar; however, juveniles and subadults may be sexed by examination of the pectines and genital aperture.

Sexual dimorphism: In addition to aforementioned characters, adult males are proportionally longer than adult females. The increased length of the male is attributed mainly to the longer metasomal segments, which sum to 72-78% of the total length of males, but to 69-72% of the total length of females. Adult males are slightly more slender than adult females: sternite VII length is 37-51 % greater than its width in males and 49-53% greater in females (Table 1). The coloration of adult females is similar to but darker than that of adult males.

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Distribution.--Rhopalurus abudi was described from Catuano, Isla Saona, off the southeast coast of the DR (Armas & Marcano Fondeur 1987). No new records of this species have been reported in the literature since the original description (Armas & Marcano Fondeur 1987; Armas et al. 1999; Teruel 2005, 2006). The records reported here, therefore, represent the first for this species on mainland Hispaniola. Based on published records and those obtained during our expedition, R. abudi appears to be restricted to humid coastal forest in the southeast of mainland DR and Isla Saona (Fig. 1). Rhopalurus princeps inhabits dry scrub in the central part of Hispaniola, including the valley of the Yaque del Norte River, the Neiba Valley, the Sierra de Baoruco, Sierra de Martin Garcia, and Sierra de Ocoa (Teruel 2006). Rhopalurus bonettii is restricted to dry spiny forests south of the Sierra de Baoruco in the western part of mainland DR and Isla Beata, the type locality. The plotted locality data agree with Teruel's (2006: 51, fig. 12) map illustrating the approximate distributions of the three species.

Ecology.--Rhopalurus abudi is probably restricted to humid forests, a habitat not previously reported for any Rhopalurus species. Although collections were made on the western and eastern sides of Parque Nacional del Este during the course of our expedition, no specimens were found on the western side, which is drier and dominated by dense, spiny forest. Whereas the South American species of Rhopalurus appear to be restricted to savannas (Lourenco 1996, 2008), those of the Caribbean are also found in other vegetation zones, including forest (Armas 2001). During our expedition, R. abudi was collected in lowland coastal humid forest on limestone, R. bonettii in dry spiny forests on limestone, and R. princeps in dry scrub on mixed substrata. All specimens of R. abudi were collected at night using UV light detection. None were found during the day, unlike R. bonettii, which was commonly found sheltering between slabs of rock (though never under bark or wood), and R. princeps, which was found under bark, wood and stones, as well as in dead and dry agave plants. The holotype of R. abudi was collected from under a stone (Armas & Marcano Fondeur 1987).

Appendix 1. Currently recognized species and subspecies of Rhopalurus Thorell 1876 and related genera, with countries, departments (Colombia, Haiti), provinces (Cuba, Dominican Republic), regions (French Guiana) and states (Brazil, Venezuela) of known distribution (data from Gonzailez-Sponga 1996; Fet & Lowe 2000; Floirez 2001; Teruel 2006; Teruel & Roncallo 2008; Teruel & Tietz 2008; Lourenco 2008; this study). New records reported in this study are marked with an asterisk.

Physoctonus debilis (C.L. Koch 1840): Brazil (Bahia, Ceara, Pernambuco*, Piauf). This species was originally placed in the nonbuthid genus Vaejovis C.L. Koch 1836. It was transferred to Rhopalurus by Borelli (1910) and remained there until Lourenco (2002) resurrected the genus Physoctonus Mello-Leitao 1934, earlier synonymized with Rhopalurus by Francke (1977).

Rhopalurus abudi Armas & Marcano Fondeur 1987: Dominican Republic (La Altagracia, La Romana).

Rhopalurus acromelas Lutz & Mello 1922: Brazil (Bahia, Ceara, Tocantins, Maranhao*, Pernambuco, Piaui).

Rhopalurus agamemnon (C.L. Koch 1839): Brazil (Bahia, Ceara, Tocantins, Mato Grosso, Pernambuco, Piaui).

Rhopalurus amazonicus Lourenco 1986: Brazil (Para).

Rhopalurus bonettii Armas 1999: Dominican Republic (Pedernales).

Rhopalurus caribensis Teruel & Roncallo 2008: Colombia (Atlantico, La Guajira, Magdalena), Venezuela (Zulia). Lourenco (2008) suggested that this species might be more appropriately recognized as a subspecies of Rhopalurus laticauda Thorell 1876.

Rhopalurus crassicauda Caporiacco 1947: Brazil (Amazonas*, Roraima), Guyana. This species was synonymized with Rhopalurus pintoi Mello-Leitao 1933 by Lourenco (1982) and reinstated by Lourenco (2002). Teruel & Tietz (2008) demonstrated that R. pintoi is a distinct species but questioned whether R. crassicauda can be regarded as distinct from R. laticauda. In our opinion, R. crassicauda is probably a junior synonym of R. laticauda. Lourenco (2008) rejected the suggestion that R. crassicauda may be synonymous with R. laticauda, suggesting instead that it might be a subspecies of the latter. Lourenco (2008) also created two new subspecies of R. crassicauda. The distinction between R laticauda, R crassicauda and its two subspecies warrants further investigation.

Rhopalurus crassicauda kourouensis Lourenco 2008: French Guiana (Kourou).

Rhopalurus crassicauda paruensis Lourenco 2008: Brazil (Para).

Rhopalurus garridoi Armas 1974: Cuba (Guantanamo)

Rhopalurus gibarae Teruel 2006: Cuba (Holguin).

Rhopalurus granulimanus Teruel 2006: Cuba (Holguin).

Rhopalurus guanambiensis Lenarducci, et al. 2005: Brazil (Bahia).

Rhopalurus junceus (Herbst 1800): Cuba (Camaguey, Cienfuegos, Ciego de Avila, Granma, Guantanamo, Havana, Holguin, Isla de la Juventud, Las Tunas, Matanzas, Pinar del Rio, Santiago de Cuba, Sancti Spiritus, Villa Clara). Records of this species from Haiti and Venezuela (see, e.g. Fet & Lowe 2000: 220) are probably erroneous (Armas 2001:248).

Rhopalurus lacrau Lourenco & Pinto-da-Rocha 1997: Brazil (Bahia).

Rhopalurus laticauda Thorell 1876: Colombia (Arauca, Boyacai, Casanare, Cesar, Meta, La Guajira, Magdalena, Norte de Santander, Vichada), Venezuela (Amazonas, Anzoaitegui, Apure, Aragua, Barinas, Bolivar, Carabobo, Cojedes, D.F., Falcon, Guarico, Lara, Meirida, Miranda, Monagas, Nueva Esparta, Portuguesa, Sucre, Taichira, Vargas, Yaracuy, Zulia).

Rhopalurus melloleitaoi Teruel & Armas 2006: Cuba (Granma).

Rhopalurus pintoi Mello-Leitao 1933: Brazil (Roraima), Guyana, ?Venezuela (Bolivar). This species was relegated to a subspecies of R. laticauda by Lourenco (1982) until reinstated by Lourenco (2002). Teruel (2006) suggested that it might be a senior synonym of Rhopalurus piceus Lourenco & Pinto-da-Rocha 1997 and this was confirmed by Teruel & Tietz (2008). Lourenco (2008) agreed with the recognition of R. pintoi as a distinct species, but suggested that R. piceus may yet prove to be valid. We agree with the decision of Teruel & Tietz (2008).

Rhopalurus princeps (Karsch 1879): Dominican Republic (Azua, Barahona, Baoruco, Independencia, Montecristi, Pedernales, Peravia), Haiti (Deipartement du l'Ouest). Records of this species from Cuba (listed by Fet & Lowe 2000:221) are erroneous.

Rhopalurus rochae Borelli 1910: Brazil (Bahia, Ceara, Paraiba, Pernambuco, Piaui, Rio Grande de Norte, Sergipe*). Borelli (1910) named the species after Francisco Diaz da Rocha, but his original spelling was rochae. Fet & Lowe (2000) noted that the correct spelling is rochai and changed it accordingly. Although the corrected spelling has been adopted by others (e.g., Teruel 2006:52), we use Borelli's (1910) original spelling.

Troglorhopalurus translucidus Lourenco, et al. 2004: Brazil (Bahia). In our opinion, this monotypic genus is a junior synonym of Rhopalurus. As twice noted by Lourenco et al. (2004:1153, 1156), when comparing Troglorhopalurus with Rhopalurus: "It may be that all modifications presented by the new troglobitic scorpion are the result of adaptation to a cave dwelling life."

Appendix 2. Material examined for comparison with Rhopalurus abudi Armas & Marcano Fondeur, 1987. Specimens are deposited in the following collections: American Museum of Natural History (AMNH), New York, USA, incorporating the Alexis Harington (AH) Collection; Natur-Museum Senckenberg, Frankfurt (SMF), Germany; Zoologisches Museum der Humboldt-Universitat, Berlin (ZMB), Germany; Zoologisches Museum der Universitat Hamburg (ZMH), Germany. Reference numbers (ESV and LP), provided on labels with the specimens, correspond to entries in the specimen databases of the author with the corresponding initials.

Physoctonus debilis (C.L. Koch, 1840): BRAZIL: Pernambuco: Exu, 5 km N, 4 October 1977, L.J. Vitt, 1 [female] (AMNH), 18 January 1978, L.J. Vitt & K.E. Streilein, 1 [female] (AMNH); Exu, 18 km N, 5 March 1977, L.J. Vitt, under leaf of granite on boulder, caatinga habitat, 1 [female] (AMNH); Fazenda Batente, 13 km E Exu, 10 November 1977, L.J. Vitt & K.E. Streilein, 1 9 (AMNH); Fazenda Caterino, 10 km NE Exu, 9 July 1977, L.J. Vitt, 1 [female] (AMNH), 25 September 1977, L.J. Vitt, 1 [female] (AMNH).

Rhopalurus acromelas Lutz & Mello, 1922: BRAZIL: Maranhao: Municipio de Loreto: Santa Barbara, on shore of Rio Parnoiba, June 1962, G. Eiten, 1 [male] (AMNH). Pernambuco: Exu, 10 km N, 13 March 1977, L.J. Vitt, rocky habitat within thorn scrub forest, 1 [female], 1 subad. [female], 4 juv. (AMNH), 14 March 1977, L.J. Vitt, rocky habitat in thorn scrub, 1 [male],1 [female] (AMNH [ESV7532]); Exu, 10 km NE, 28 April 1977, L.J. Vitt, 1 [male],1 [female], 2 subad. [female], 2 subad., 1 juv. (AMNH), 25 September 1977, L.J. Vitt, 1 [male],1 [female] (AMNH [ESV7244]); Exu, 15 km NE, 14 May 1977, L.J. Vitt, high caatinga, under bark of tree, 1 subad. [female] (AMNH); Exu, 20 km E, 30 March 1977, L.J. Vitt, 1 juv. L (AMNH); Fazenda Caterino, 10 km NE Exu, 9 July 1977, L.J. Vitt, 1 subad. [male] (AMNH), 1 August 1977, L.J. Vitt, 1 juv. [male] (AMNH).

Rhopalurus agamemnon (Herbst, 1800): BRAZIL: Bahia: Salvador, February 1972, Weinkselbaum, 1 [female] (AMNH [ESV7405]).

Rhopalurus bonettii Armas, 1999: DOMINICAN REPUBLIC: Pedernales Province: Parque National Jaragua: Cabo Rojo, 17[degrees]53'45.2"N, 71[degrees]39'35.8'W, 9 July 2004, E.S. Volschenk & J. Huff, 15 m, dry cactus and spiny forest on limestone karst, hand collected at night with blacklights, 3 [male],10 [female], 4 subad., 2 juv. (AMNH [ESV6005]), 1 [male] (AMNH [ESV7126]), 1 [male],1 [female] (AMNH [ESV7127]), 1 subad. [male] (AMNH), 1 juv. [male] (AMNH [LP 3267]); Road to Fondo Paradi, 1.8 km from Highway 44, 17[degrees]48.692'N, 71[degrees]26.600'W, 12 January 2004, J. Huff, 302 ft, found between rocks, 1 [female] (AMNH [LP 2471]), 1 [female] (AMNH [LP 3265]); Track into park, between Manuell Goa and Oviedo, 17[degrees]48'41.5"N, 71[degrees]26'35.9"W, 9 July 2004, E.S. Volschenk & J. Huff, 83.3 m, deciduous forest and thorny scrub, hand collected from between stones during the day and with blacklights at night, 13 [male],7 [female], 1 subad., 1 juv. (AMNH [ESV6011]), I [male],1 [female] (AMNH [ESV7112]), 1 [male] (AMNH [ESV7129]), 1 juv. (AMNH [LP 3266]).

Rhopalurus caribensis Teruel & Roncallo, 2008: COLOMBIA: Magdalena Department: Bahia de Guairaca, Tayrona Park, 31 October 1985, H.-G. Muller, 1 9 (SMF 37027); Pozo Colorado, II km W Santa Marta, 18-30 April 1968, B. Malkin, 1 9, 1 subad., 19 first instars (AMNH); Puente de Los Clavos, 15 km E Pueblo Bello, Sierra Nevada de Santa Marta, 13 June 1968, B. Malkin, 1500 m, 1 subad. [female] (AMNH); Santa Marta, 29 June-31 July 1966, 2 [female] (SMF 39120).

Rhopalurus crassicauda Caporiacco, 1947: BRAZIL: Amazonas: Rio Branco, Amazonasgebiet, 1912, E. Ule, 1 juv. [female] (ZMB 14867). Roraima: Mt. Roraima, 2 [male],1 [female], 1 subad. (AMNH 29180).

Rhopalurus junceus (Herbst, 1800): CUBA: July 2007, C. Hamilton, 1 juv. (AMNH [LP 7009]); 'Antillen?,' 1 [male],2 [female] (ZMB 7370); 'Portorico', Stahl, 2 [female], 1 juv. (ZMB 7280 [ESV7001]); Gundlach, 2 [female] (ZMB 2637), 1 [male],1 [female] (ZMB 7380 [ESV7224]), 1 juv. (ZMB, 7343); Arroyo Bermejo, near Fibacoa, 31 May 1967, Kleiderschrank, 1 [male] (ZMB 31020), 15 June 1967, im zelt. wiese auf sandboden, 1 [female] (ZMB 31021), June 1967, 1 juv. (ZMB 31022); 1 L (ZMH), Santiago de las Caballeros, P. Thumb, 1936. Havana Province: Havana, 1 [female] (AMNH), April 1941, Dr E. Weiss, 1 [female], 1 subad. (AMNH). Holguin Province: August 2000, Heist, captive bred, 1 juv. (AMNH [LP 1928]); near Bafios [Banes], May 1918, 2 [male] (AMNH); Guardalavaca, 29 March 1993, W. Altmann, captive bred, 1 [male] (AMNH [LP 1565]); Mountains near Guisa, October 1936, P. Thumb, 1 [female], 28 juv. (ZMH); Moa, September 1937, P. Thumb, 1 [male] (ZMH), 1938, P. Thumb, 4 [female] (ZMH). Isla de la Juventud Province: Isle of Pines, 1 [male] (AMNH). Pinar del Rio Province: Guanahacabiles, Akad.-stat. El Beral, December 1967, G. Peters, 1 subad. (ZMB 31023); Sierra de Anafe, 23 February 1947, M. Barro, 2 subad. (AMNH); Vinales Valley, 1940, Osorio, 1 [female] (AMNH). Santiago de Cuba Province: La Socapa, 10 km SW of Santiago de Cuba, 9 April 1999, R. Teruel, 1 [male] (AMNH), 1 [female] (AMNH [LP 1509]), 1 juv. [female] (AMNH [LP 1517]), 1 [female] (AMNH [LP 1518]); Santiago de Cuba, 1 [male], 2 juv. (AMNH). Sancti Spiritus Province: Trinidad, August 1978, B. Acosta, 1 [male] (AMNH AH 4514 [ESV7041]).

Rhopalurus lacrau Lourenco & Pinto-da-Rocha, 1997: BRAZIL: Bahia: Municipio Itaete: Trail between Caves "Lapa do Bode" and "Lapa Escondida," 12[degrees]56'9.1"S, 41[degrees]3'56.2"W, 21 January 2007, C.I. Mattoni, R. Pinto-da-Rocha & H. Yamaguti, under rocks, 2 [female] (AMNH), 1 subad. [female], 4 juv. (AMNH [LP 7637]).

Rhopalurus laticauda Thorell, 1876: 2 [female] (ZMB 14865); "Mexico,": Dr v. Hubl, 1 L (ZMB 14866). VENEZUELA: F. Kummerow, 1 [male],1 [female] (ZMB 8226). San Jose de Guaviare, December 1955, Meden, 1 [female] (SMF 39252). Aragua: Maracay, 1 subad. [male] (SMF 29208), Fahren holz, 1 [male],1 [female], 1 subad. (SMF 8876/218). Bolivar: Ciudad Bolivar, 20 February 1903, 2 [female] (ZMH); La Paragua, M.A. de Verde, 1 [male] (AMNH); Upata, February 1973, A. Bordes, 1 [female] (AMNH). Carabobo: Valencia, F. Kummerow, 29 December 1904, 1 [female] (ZMB 31024), September 1958, H. Ardelt, 2 [female] (ZMH). Distrito Federal: Caracas, March 1999, C. Siederman, 2 [female], 20 first instars (AMNH [ESV7444]), 2001, C. Siederman, 1 [male] (AMNH [LP 2462]). Guarico: Calabozo and San Fernando de Apure (about halfway between), 30 November 1967, M.A. de Verde, 1 [male] (AMNH); 'Hato Masaguarat,' 45 km S Calabozo, 7 April 1978, Y. Lubin, 1 [male] (AMNH [ESV7816]). Merida:Merida, 2 L,3 [female] (SMF 5712/27). Miranda: Guatire, 29 April 2004, R.C. West, under rocks, dry forest, 1 [male] (AMNH [LP 2845]), 1 [female] (AMNH); Hda. Santa Rosa, 3 km N Guatire, 10 January 1973, M.A. GonzailezSponga, 450 m, 1 [male],1 [female], 2 juv. (AMNH). Nueva Esparta: Isla Margarita, N of Peninsula de Macanao, 11[degrees]02.618'N, 64[degrees]21.542'E, 4 September 2005, S. Huber, 1 [female] (AMNH [LP 4221]). Trujillo: Valera region, N, October 2005, S.E. Bazo Abreu, 1 [female] (AMNH [LP 5504]), 1 [female] (AMNH [LP 5505]).

Rhopalurus pintoi Mello-Leitao, 1933: GUYANA: Roraima Province: Rununui region, SW Guyana, near Venezuelan border, ex A. Tietz, March 2008, 1 juv. [male] (AMNH [LP 8278]).

Rhopalurus princeps (Karsch, 1879): DOMINICAN REPUBLIC: Independencia Province: Isla Cabritos, 18[degrees]30.019'N, 71[degrees]43.228'W, 7 January 2004, J. Huff, 110 ft, under rock, coral, 1 [male],1 [female], 16 juv. (AMNH), 5 [male],3 [female], 3 subad., 1 juv. (AMNH), 3 juv. (AMNH [LP 2470]), 1 subad., 2 juv. (AMNH [LP 3260]); Ranger station for Parque Nacional Isla Cabritos, 18[degrees]33'45"N, 71[degrees]41'50"W, 8 July 2004, E.S. Volschenk & J. Huff,--19 m, dry forest, hand collected from under stones and logs, and with blacklights, 3 [male],7 [female], 5 subad., 2 juv. (AMNH [ESV6006]), 1 subad. [male] (AMNH), 1 subad. (AMNH [LP 3264]); Parque Nacional Isla Cabritos, behind Ranger Station, 18.56287[degrees]N, 71.69762[degrees]W, 8 August 2005, L. Esposito,--23 m, mixed dry forest with succulents, UV detection, 35[degrees]C, 2 [male],8 [female], 1 subad. [female],32 first instars (AMNH), 2 [male], 1 subad. [female] (AMNH), 1 [male] (AMNH [LP 5102]); Parque Nacional Sierra de Baoruco, road between Rabo de Gato and Duverge, 18[degrees]19'38"N, 17[degrees]33'55"W, 7 July 2004, E.S. Volschenk & J. Huff, 447 m, arid thorny scrub, hand collected from under stones and in dead and dry agaves, 3 [male],3 [female], 3 juv. (AMNH [ESV6033]), 1 juv. (AMNH), 1 [female] (AMNH [LP 3263]); Puerto Escondido, Sierra de Baoruco, 18[degrees]19.762'N, 71[degrees]33.502'W, 6 January 2004, J. Huff, 1592 ft, under dead agave, 1 [male],3 [female], 1 juv. (AMNH), 1 juv. (AMNH [LP 3261]); Road to Puerto Escondido, 18[degrees]20.376'N, 71[degrees]33.345'W, 6 January 2004, J. Huff, 1388 ft, under rocks in gravel quarry, 1 [female] (AMNH), 1 juv. (AMNH [LP 3262]). Pedernales Province: Manuel Goja, 3.9. km N, 9 May 1998, D. Huber, 1 [female] (AMNH [LP 1566]); Oviedo to Pedernales, 11.5 km N, 8 May 1998, D. Huber, 1 [male] (AMNH [LP 1516]). HAITI: Departement de l'Ouest: Port-au-Prince, Ehrenberg, holotype [male] (ZMB 116).

Rhopalurus rochae Borelli, 1910: BRAZIL: Bahia: Municipio Ceraima: Guanambi, 7 km S, 14[degrees]17'5.6"S, 42[degrees]47'2.2"W, 24 January 2007, C. Mattoni, R. Pinto-da-Rocha & H. Yamaguti, 533 m, UV sampling, modified savanna, cloudy and raining, 1 juv. (AMNH [LP 7638]); Fazenda du Fabiano, 8 km NE Guanambi,14[degrees]10'17.6"S, 42[degrees]43'56.4"W, 24 January 2007, C. Mattoni, R. Pinto-da-Rocha & H. Yamaguti, 539 m, under rocks, rocky hill and surrounds, open savanna modified, 1 [male], 2 juv. (AMNH [LP 7639]), 1 [female] (AMNH); Guanambi, 16 km SE, 14[degrees]17'19"S, 42[degrees]41'31.1"W, 25 January 2007, C. Mattoni, R. Pinto-da-Rocha, H. Yamaguti, 559 m, UV sampling and under leaf litter, banana plantation and surrounds, 1 juv. (AMNH [LP 7655]). Paraiba: Soledade, 07[degrees]02.118'S, 36[degrees]27.311'W, 16 March 1999, A. Kury & A. Giupponi, 575 m, 1 [male] (AMNH [LP 1581]), 1 [female] (AMNH [LP 1582]), 1 [male] (AMNH [LP 1775]). Pernambuco: Escola Aquicola, Exu, 30 March 1977, L.J. Vitt, caatinga, 1 [male] (AMNH [ESV7248]), 27 June 1977, L.J. Vitt, 1 [male] (AMNH); Exu, 3 kmNW, 10 March 1977, L.J. Vitt, 2 [male],1 [female], 3 juv. (AMNH); Exu, 3 km W, 30 May 1977, L.J. Vitt, 2 [male],4 [female], 4 juv. (AMNH), 1 June 1977, L.J. Vitt, 1 [female] (AMNH); Exu, 5 km N, 6 April 1977, L.J. Vitt, caatinga, 1 [male], 1 juv. (AMNH), 18 January 1978, L.J. Vitt & K.E. Streilein, 1 juv. (AMNH); Exu, 5 km E, 8 May 1977, L.J. Vitt, 1 juv. (AMNH); Exu, 6 km N, 15 March 1977, L.J. Vitt, open fields (cotton), under fallen logs, 1 [female], 1 juv. L (AMNH); Exu, 6 km NE, 16 March 1977, L.J. Vitt, under rock on larger rock, caatinga habitat, 1 [female], 49 first instars (AMNH); Exu, 18 km E, 5 March 1977, L.J. Vitt, under leaf of granite on boulder, caatinga habitat, 1 [female], 29 first instars (AMNH), 1 [female], 39 first instars (AMNH [ESV7210]); Exu, 20 km E, 30 March 1977, J.L.Vitt, 1 [male],1 [female] (AMNH [ESV7625]); Fazenda Batente, 5 km NE Exu, 29 March 1977, L.J. Vitt, 1 juv. (AMNH); Fazenda Caterino, 10 km NE Exu, 1 August 1977, L.J. Vitt, 2 [male], 3 juv. (AMNH), 5 [male],3 [female] (AMNH); Fazenda Chelonia, 8 km S Exu, 28 July 1977, L.J. Vitt, 2 juv. (AMNH); Fazenda Guarani, 3 km N Exu, 14 July 1977, L.J. Vitt, 1 [male],3 [female], 1 subad. 3 juv. (AMNH); Fazenda Guarani, 5 km N Exu, 29 July 1977, L.J. Vitt, 1 [female], 3 juv. (AMNH), 19 February 1978, L.J. Vitt, 1 [female] (AMNH). Sergipe: Municipio Lagarto: near Genipapo, July 1982, O.F. Francke, 1 [male],2 [female] (AMNH).

ACKNOWLEDGMENTS

We are grateful to the Department de Investigaciones de la Subsecretaria de Areas Protegidas y Biodiversidad, Government of the Dominican Republic, for Permit Number 01496 to collect and export scorpions from the country. Kelvin Guerrero kindly assisted with the permit application and provided valuable advice on collecting in the DR (he was the first to observe R. abudi in the Parque Nacional del Este). We thank the following for assistance with the study of material at their institutions: Peter Jager and Julia Altmann (SMF), Jason Dunlop and Shahin Nawai (ZMB), Hieronymus Dastych (ZMH); the following for donating specimens to L. Prendini that were examined during the course of this study: Santos Bazo Abreu, Dietmar Huber, Siegfried Huber, Adriano Kury, Charles Siederman, Rolando Teruel Ochoa, Alex Tietz, Rick C. West; and the following for the participating in fieldwork during which specimens, examined during the course of this study, were collected: Camilo I. Mattoni, Ricardo Pinto-da Rocha, Humberto Yamaguti. The 2004 field expedition to the DR, during which the series of R. abudi and comparative material of R. bonettii and R. princeps was collected, was funded by a Genomics Postdoctoral Research Fellowship from the AMNH to E.S. Volschenk and National Science Foundation grant EAR 0228699 to L. Prendini. Fieldwork by C.I. Mattoni in Brazil and by L.A. Esposito in the DR, during which other material examined for this study was collected, was funded by grants from the National Science Foundation (EAR 0228699) and the Richard Lounsbery Foundation to L. Prendini. We thank Steve Thurston (AMNH) for assistance with preparing the plates for this contribution, and Mark Harvey and an anonymous reviewer for comments on a previous draft of the manuscript. While at the AMNH, E.S. Volschenk was supported by a Genomics Postdoctoral Research Fellowship, supplemented by a grant from the Richard Lounsbery Foundation to L. Prendini; L.A. Esposito was supported by a National Science Foundation GK-12 Fellowship, a City University of New York MAGNET Fellowship, and a City University of New York/NSF AGEP Fellowship.

Manuscript received 8 July 2008, revised 5 November 2008.

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Teruel, R., V. Fet & M.R. Graham. 2006. The first mito chondrial DNA phylogeny of Cuban Buthidae (Scorpiones: Buthoidea). Boletin de la Sociedad Entomoloigica Aragonesa 39:219-226.

Teruel, R. & C.A. Roncallo. 2008. Rare or poorly known scorpions from Colombia. III. On the taxonomy and distribution of Rhopalurus laticauda Thorell, 1876 (Scorpiones: Buthidae), with description of a new species of the genus. Euscorpius 68:1-12.

Teruel, R. & A.K. Tietz. 2008. The true identity of Rhopalurus pintoi Mello-Leitao, 1932, with notes on the status and distribution of Rhopalurus crassicauda Caporiacco, 1947 (Scorpiones: Buthidae). Euscorpius 70:1-14.

Vachon, M. 1973 [1974]. Etude des caracteres utilises pour classer les familles et les genres de scorpions (Arachnides). 1. La trichobothriotaxie en arachnologie. Sigles trichobothriaux et types de trichobothriotaxie chez les scorpions. Bulletin du Museium National d'Histoire Naturelle, Paris (3) 140:857-958.

Volschenk, E.S. 2005. A new technique for examining surface morphosculpture of scorpions. Journal of Arachnology 33: 820-825.

Volschenk, E.S., C.I. Mattoni & L. Prendini. 2008. Comparative anatomy of the mesosomal organs of scorpions (Chelicerata, Scorpiones), with implications for the phylogeny of the order. Zoological Journal of the Linnean Society 154:651-675.

Lorenzo Prendini, Lauren A. Esposito, Jeremy C. Huff and Erich S. Volschenk: Scorpion Systematics Research Group, Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, USA. E-mail: lorenzo@amnh.org
Table 1.--Meristic data for three male and three female specimens of
Rhopalurus abudi Armas & Marcano Fondeur 1987 deposited in the
collection of the American Museum of Natural History (AMNH), New York.
(1) Sum of carapace, tergites I-VII, metasomal segments I-V, and
telson; (2) sum of tergites I-VII; (3) sum of metasomal segments I-V
and telson; (4) measured along an axis parallel to the dorsal surface;
(5) measured from base of condyle to tip of fixed finger.

               Sex                     [male]    [male]    [male]

               Repository                AMNH      AMNH      AMNH

               Reference number        ESV7937   ESV7303   ESV7117
Total
length (1)                              67.45     69.71     68.75
Carapace       length                    7.50      8.50      8.01
               anterior width            4.67      6.90      6.03
               posterior width           7.65      8.82      8.08
               eye diameter              0.63      0.74      0.63
               interocular distance      0.63      0.71      0.53
Mesosoma       total length  (2)        18.96     21.77     17.85
Sternite VII   length                    5.08      5.93      4.89
               width                     6.98      8.16      7.36
Metasoma       total length (3)         48.42     54.51     53.08
Metasoma I     length                    6.36      7.32      7.22
               width                     4.37      5.03      4.69
               height                    3.69      4.12      3.19
Metasoma II    length                    7.48      8.30      8.13
               width                     4.23      4.84      4.70
               height                    3.52      4.01      3.73
Metasoma III   length                    8.13      8.95      8.71
               width                     4.18      5.03      4.81
               height                    3.84      4.09      4.00
Metasoma IV    length                    8.58      9.39      9.25
               width                     4.39      5.03      4.89
               height                    3.59      4.10      3.92
Metasoma V     length                    9.99     11.32     10.88
               width                     4.28      4.90      4.74
               height                    3.54      4.12      3.79
Telson         total length (4)          7.88      9.23      8.89
               vesicle length            4.44      5.26      5.15
               vesicle width             2.55      2.90      2.84
               vesicle height            2.55      2.91      2.67
               aculeus length            3.54      3.86      3.76
Pedipalp       total length (5)         29.96     35.72     33.57
               trochanter length         3.74      4.24      3.49
Femur          length                    6.92      7.49      7.32
               width                     2.25      2.28      2.37
               height                    1.43      1.83      1.71
Patella        length                    7.45      8.92      8.66
               width                     2.95      3.25      3.06
               height                    2.06      2.35      2.24
Chela          length                   11.85     15.07     14.10
               width                     3.09      3.96      4.06
               height                    3.39      4.03      4.00
               fixed finger length       6.73      8.26      7.56
               ventroexternal carina     4.66      5.32      5.38
                 length
               movable finger length     8.67     10.06      9.23
Pectines       total length              7.57      7.79      7.12
               length along dentate      6.86      7.24      6.43
                 margin
               tooth count              23/23     23/23     22/23
                 (left/right)

               Sex                     [female]   [female]   [female]

               Repository                AMNH       AMNH       AMNH

               Reference number        ESV7937    ESV7303    ESV7117
Total
length (1)                              77.48      91.67      87.83
Carapace       length                    8.75      10.23      10.28
               anterior width            6.80       8.18       7.98
               posterior width           9.32      10.84      11.29
               eye diameter              0.72       0.79       0.73
               interocular distance      0.58       0.66       0.74
Mesosoma       total length  (2)        23.32      28.27      28.59
Sternite VII   length                    5.86       7.08       6.92
               width                     8.71      10.86      10.30
Metasoma       total length (3)         54.16      63.36      63.36
Metasoma I     length                    7.08       8.59       8.47
               width                     5.00       5.73       5.82
               height                    4.00       3.91       4.09
Metasoma II    length                    8.29       9.70       9.91
               width                     4.73       5.40       5.45
               height                    4.06       4.69       4.63
Metasoma III   length                    9.02       9.52      10.25
               width                     4.59       5.41       5.48
               height                    4.11       4.56       4.77
Metasoma IV    length                    9.07      10.76      10.63
               width                     4.72       5.29       5.46
               height                    3.99       4.50       4.57
Metasoma V     length                   11.27      13.33      12.87
               width                     4.63       4.47       5.33
               height                    4.01       4.53       4.79
Telson         total length (4)          9.43      11.46      11.23
               vesicle length            5.22       6.43       6.38
               vesicle width             2.94       3.46       3.59
               vesicle height            2.90       3.44       3.49
               aculeus length            4.22       5.33       5.41
Pedipalp       total length (5)         35.69      42.36      41.93
               trochanter length         3.92       4.47       4.72
Femur          length                    8.02       9.05       8.74
               width                     2.32       2.84       2.89
               height                    1.81       2.54       2.05
Patella        length                    8.99      10.98      10.57
               width                     3.35       3.81       3.76
               height                    2.55       2.85       2.84
Chela          length                   14.76      17.86      17.90
               width                     2.87       3.49       3.22
               height                    2.97       3.53       3.49
               fixed finger length       8.99      10.75      10.74
               ventroexternal carina     5.02       6.00       5.96
                 length
               movable finger length    10.01      12.16      12.22
Pectines       total length              7.36       8.74       8.43
               length along dentate      6.66       7.91       7.54
                 margin
               tooth count              20/20      22/21      20/20
                 (left/right)

Table 2.--Meristic data for three male and three female specimens of
Rhopalurus bonettii Armas 1999 deposited in the collection of the
American Museum of Natural History (AMNH), New York. (1) Sum of
carapace, tergites I-VII, metasomal segments I-V, and telson; (2) sum
of tergites I-VII; (3)sum of metasomal segments I-V and telson; (4)
measured along an axis parallel to the dorsal surface; (5) measured
from base of condyle to tip of fixed finger.

               Sex                     [male]    [male]    [male]

               Repository                AMNH      AMNH      AMNH

               Reference number        ESV7112   ESV7127   ESV7126
Total
length (1)                              64.64     63.42     63.48
Carapace       length                    7.33      7.22      7.35
               anterior width            5.55      5.38      5.33
               posterior width           7.16      6.99      7.51
               eye diameter              0.55      0.57      0.55
               interocular distance      0.52      0.56      0.50
Mesosoma       total length (2)         18.10     18.35     18.93
Sternite VII   length                    4.50      4.41       4.8
               width                     6.53      6.09      6.53
Metasoma       total length (3)         47.02     46.52     47.48
Metasoma I     length                    6.31      6.28      6.14
               width                     3.87      3.65      4.04
               height                    3.35      3.28      3.59
Metasoma II    length                    7.31      7.35      7.32
               width                     3.89      3.60      3.80
               height                    3.30      3.50      3.38
Metasoma III   length                    7.84      7.90      8.05
               width                     3.78      3.70      3.91
               height                    3.36      3.38      3.42
Metasoma IV    length                    8.18      7.79      8.29
               width                     3.83      3.68      4.06
               height                    3.36      3.29      3.44
Metasoma V     length                    9.56      9.64      9.85
               width                     3.77      3.63      4.03
               height                    3.40      3.17      3.34
Telson         total length (4)          7.82      7.56      7.83
               vesicle length            4.52      4.50      4.57
               vesicle width             2.59      2.46      2.73
               vesicle height            2.69      2.50      2.70
               aculeus length            3.22      3.77      3.16
Pedipalp       total length (5)         32.64     31.82     33.49
               trochanter length         3.36      3.53      3.64
Femur          length                    7.13      6.77      6.97
               width                     2.10      1.98      2.08
               height                    1.58      1.45      1.46
Patella        length                    8.49      8.16      9.00
               width                     2.79      2.63      3.05
               height                    2.06      2.02      2.08
Chela          length                   13.66     13.36     13.88
               width                     2.84      2.85      3.20
               height                    2.98      2.94      3.28
               fixed finger length       7.55      6.87      7.54
               ventroexternal carina     4.44      4.65      4.76
                 length
               movable finger length     9.02      9.04      9.32
Pectines       total length              7.05      6.35      6.53
               length along dentate      6.43      6.00      6.22
                 margin
               tooth count              24/23     22/23     22/21
                 (left/right)

               Sex                     [female]   [female]   [female]

               Repository                AMNH       AMNH       AMNH

               Reference number        ESV7127    ESV7112    ESV6005
Total
length (1)                              70.31      76.96      71.72
Carapace       length                    7.63       9.30       7.89
               anterior width            5.91       7.09       6.10
               posterior width           7.67       9.51       7.58
               eye diameter              0.62       0.58       0.64
               interocular distance      0.61       0.53       0.55
Mesosoma       total length (2)         21.24      23.48      21.44
Sternite VII   length                    5.22       6.16       5.43
               width                     7.48       8.76       7.56
Metasoma       total length (3)         48.74      56.53      48.84
Metasoma I     length                    6.54       7.66       6.84
               width                     4.05       4.60       4.02
               height                    3.52       4.02       3.46
Metasoma II    length                    7.66       8.76       7.76
               width                     4.02       4.43       3.67
               height                    3.35       3.87       3.34
Metasoma III   length                    7.85       9.18       8.04
               width                     3.82       4.32       3.90
               height                    3.50       3.96       3.32
Metasoma IV    length                    8.52       9.86       7.79
               width                     4.01       4.35       3.74
               height                    3.69       3.77       3.41
Metasoma V     length                    9.86      11.26       9.92
               width                     3.62       4.25       3.68
               height                    3.59       3.90       3.43
Telson         total length (4)          8.31       9.81       8.49
               vesicle length            4.56       5.67       4.87
               vesicle width             2.75       3.27       2.88
               vesicle height            2.59       3.22       2.79
               aculeus length            3.98       4.99       3.74
Pedipalp       total length (5)         34.96      40.85      35.35
               trochanter length         3.95       4.49       3.97
Femur          length                    7.34       8.71       7.46
               width                     2.20       2.71       2.26
               height                    1.61       1.73       1.83
Patella        length                    9.16      10.69       9.11
               width                     2.91       3.28       3.01
               height                    2.24       2.53       2.13
Chela          length                   14.51      16.96      14.81
               width                     2.76       3.07       2.77
               height                    2.76       3.21       3.10
               fixed finger length       8.63      10.21       8.33
               ventroexternal carina     4.88       5.71       4.72
                 length
               movable finger length     9.93      11.59       9.96
Pectines       total length              6.89       7.50       6.47
               length along dentate      5.93       6.82       5.93
                 margin
               tooth count              21/20      20/20      19/19
                 (left/right)

Table 3.--Meristic data for three male and three female specimens of
Rhopalurus princeps (Karsch 1879) deposited in the collection of the
American Museum of Natural History (AMNH), New York. (1) Sum of
carapace, tergites I-VII, metasomal segments I-V, and telson; (2) sum
of tergites I-VII; (3) sum of metasomal segments I-V and telson; (4)
measured along an axis parallel to the dorsal surface; (5) measured
from base of condyle to tip of fixed finger.

               Sex                        [male]    [male]    [male]

               Repository                   AMNH      AMNH      AMNH

               Locality or number    Is. Cabritos   ESV6033   LP 3260

Total
length (1)                                 59.90     47.27     51.19
Carapace       length                       6.62      5.63      6.38
               anterior width               5.33      4.65      5.12
               posterior width              6.97      5.84      6.95
               eye diameter                 0.58      0.46      0.45
               interocular                  0.40      0.43      0.47
                 distance
Mesosoma       total length (2)            17.67     14.00     17.06
Sternite VII   length                       4.65      3.59      4.36
               width                        6.43      5.51      6.60
Metasoma       total length (3)            42.90     35.94     41.52
Metasoma I     length                       5.50      4.76      5.08
               width                        4.19      3.24      4.11
               height                       3.39      2.98      3.54
Metasoma II    length                       6.55      5.47      6.17
               width                        4.11      3.26      4.13
               height                       3.39      2.86      3.43
Metasoma III   length                       7.16      5.86      6.73
               width                        4.16      3.53      4.24
               height                       3.52      2.96      3.49
Metasoma IV    length                       7.45      6.05      7.28
               width                        4.65      3.67      4.51
               height                       3.58      3.02      3.61
Metasoma V     length                       9.03      7.46      8.91
               width                        4.67      3.54      4.50
               height                       3.40      2.89      3.42
Telson         total length (4)             7.21      6.34      7.35
               vesicle length               4.70      3.93      4.69
               vesicle width                2.51      2.41      2.62
               vesicle height               2.51      2.29      2.52
               aculeus length               3.40      2.48      3.46
Pedipalp       total length (5)            28.26     23.46     26.93
               trochanter length            3.21      2.71      2.87
Femur          length                       6.14      4.73      5.54
               width                        1.97      1.68      1.80
               height                       1.72      1.40      1.61
Patella        length                       6.98      6.17      6.94
               width                        2.75      2.24      1.98
               height                       2.10      1.84      2.44
Chela          length                      11.93      9.85     11.58
               width                        3.51      2.83      3.52
               height                       3.54      2.81      3.39
               fixed finger length          5.45      4.34      5.06
               ventroexternal               4.95      4.15      4.78
                 carina length
               movable finger               7.07      6.05      6.94
                 length
Pectines       total length                 6.34      5.14      6.67
               length along                 6.10      5.03      6.18
                 dentate margin
               tooth count                 25/24     24/25     25/26
                 (left/right)

               Sex                     [female]     [female]   [female]

               Repository                  AMNH       AMNH       AMNH

               Locality or number   Is. Cabritos    ESV6033    LP 3260

Total
length (1)                                69.19      66.32      55.21
Carapace       length                      7.90       7.18       7.03
               anterior width              6.59       5.81       5.59
               posterior width             8.44       7.45       7.24
               eye diameter                0.62       0.52       0.48
               interocular                 0.46       0.63       0.48
                 distance
Mesosoma       total length (2)           21.87      20.58      19.23
Sternite VII   length                      5.53       5.02       4.31
               width                       8.36       7.81       7.68
Metasoma       total length (3)           49.00      45.32      45.63
Metasoma I     length                      6.30       5.91       5.26
               width                       4.90       4.30       4.50
               height                      3.85       3.69       3.61
Metasoma II    length                      7.24       6.72       6.85
               width                       4.59       4.14       4.39
               height                      3.89       3.68       3.62
Metasoma III   length                      7.56       7.24       7.55
               width                       4.74       4.22       4.30
               height                      3.97       3.70       3.72
Metasoma IV    length                      7.87       7.39       8.08
               width                       4.93       4.52       4.55
               height                      3.96       3.69       3.69
Metasoma V     length                     11.37       9.51       9.62
               width                       4.86       4.30       4.35
               height                      3.81       3.63       3.60
Telson         total length (4)            8.66       8.55       8.27
               vesicle length              5.41       5.45       4.85
               vesicle width               3.27       3.06       2.99
               vesicle height              3.02       2.94       2.87
               aculeus length              3.77       4.05       4.61
Pedipalp       total length (5)           32.75      29.12      28.23
               trochanter length           4.20       3.40       3.44
Femur          length                      7.12       6.28       5.86
               width                       2.36       2.05       1.74
               height                      1.96       1.82       1.74
Patella        length                      7.97       7.45       7.18
               width                       3.23       2.91       2.91
               height                      2.56       2.22       2.25
Chela          length                     13.46      11.99      11.75
               width                       3.33       2.95       2.98
               height                      3.40       3.02       2.89
               fixed finger length         6.58       6.00       6.08
               ventroexternal              4.85       4.76       4.40
                 carina length
               movable finger              8.25       7.98       7.28
                 length
Pectines       total length                6.99       6.25       5.75
               length along                6.50       5.37       5.22
                 dentate margin
               tooth count                22/23      21/22      21/20
                 (left/right)
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