Diagnosis

The genus Orobothriurus comprises small, slender scorpions, varying from 18.3 mm (male O. wawita) to 51.5 mm (female O. quewerukana) in total length. Orobothriurus may be distinguished from other bothriurid genera by the distinctive morphology of the hemispermatophore (fig. 10), which is slender with a reduced lobe region. The frontal crest of the hemispermatophore is divided in two parts: the proximal part is oblique and the distal part straight with a lateral projection on both the ental and ectal surfaces. The basal lobe forms a tortuous stem with a typical subdistal spatulate dilation.

The genus may be further distinguished on the following diagnostic characters. The male pedipalp chela manus possesses an acuminate conical apophysis on the internal surface. Trichobothrium ib is situated at the base of this apophysis. The VSM carinae of metasomal segment V are arranged in parallel and situated close to the VL carinae. The female genital opercula are more elongated posteriorly, with more rounded anterior edges, than those of other bothriurid genera.

Remarks

The characters shared by most Orobothriurus species are as follows. The coloration is usually yellowish. The pigmentation pattern of the mesosoma typically comprises two sublateral spots on each tergite, leaving an unpigmented median stripe along the length of the mesosoma (fig. 13C, E–G), except in O. ampay, in which tergites I–VII are completely pigmented (fig. 12D), and O. grismadoi, in which tergite VII is almost completely pigmented (fig. 13G).

The metasomal segments usually exhibit a single VM and paired VL stripes (figs. 11B, C, 13A). The VM stripe is contiguous with the paired VL stripes in the posterior third of the segment (fig. 11A) in O. ampay, and absent in O. wawita (fig. 13D). The carapace anterior margin is almost straight, either with a weak median projection (epistome) or a shallow median notch. The anteromedian longitudinal sulcus is complete but weakly developed, the posteromedian longitudinal sulcus well developed, the median ocular tubercle low, and situated anteromedially on the carapace (figs. 14–Fig. 15.16). The metasomal segments are elongated, with variable carinal development (figs. 17–Fig. 18.Fig. 19.Fig. 20.Fig. 21.22). The DL carinae occupy the entire length of segments I–IV, and usually comprise granules of medium size. The ML, LIM, and VL carinae are variable, usually present on segments I and II. The LSM carinae are absent or obsolete. The VSM carinae are present on segments I and II in most species (figs. 17–Fig. 18.19), extending to IV in O. ampay (fig. 17B). The VM carina of metasomal segment V is granular (figs. 20, 21A, B, D–F, 22), except in the male of O. wawita, in which it is absent (fig. 21C). The VSM carinae are arranged in parallel and situated close to the VL carinae, except in O. atiquipa and O. wawita, where they are absent (figs. 21A, C, 22F). Metasomal segment I possesses 2 or 3 pairs of VL and VSM macrosetae, II and III each possess 3 pairs, IV, 3–5 pairs, and V, 3 or 4 pairs (fig. 6). The male telson vesicle is narrower than that of the female, with a weakly developed dorsal gland that is not always apparent (fig. 23).

The sternum is transverse with 2 or 4 macrosetae. The genital operculum is divided longitudinally, comprising two subtriangular sclerites, joined by flexible tissue in the anterior half, each sclerite tapering posteriorly, the sclerites of females being more elongated posteriorly, with the anterior edges more rounded (fig. 9) than those of males.

Each pecten comprises more than 10 teeth, fulcra, and a single row of median lamellae, the first being dilated (fig. 9). The cheliceral manus possesses 2 or 3 macrosetae near the movable finger articulations. The movable finger possesses two subdistal teeth and is more strongly curved in the male than the female. The ventrointernal margin of the fixed finger and ventral surface of the movable finger are setose basally.

The basitarsi possess two well-developed, symmetric pedal spurs. The telotarsi are elongated, with a ventromedian row of elongated spinules, and well-developed pro- and retroventral rows of spiniform macrosetae with the following formula on legs I–IV: 1/1, 2/2, 3/3, 3/3, except in O. atiquipa, which has 3/4 spiniform macrosetae in the pro- and retroventral on legs III and IV. The telotarsal ungues are curved and symmetric.

The pedipalps are usually elongated, especially in adult males, with a slender, prismatic chela in most species. The male chela manus of all species, except O. ampay and O. wawita, possesses an acuminate conical apophysis on the internal surface (fig. 8B). The pedipalp chelal carinae are usually absent or obsolete, evident only by differences in the angles between surfaces, but at least the D, DS, DMA, DI, and VM carinae may be identified, depending on the species, with vestigial VI and IM carinae also evident in several species (Ochoa, 2004a). The median denticle row of the chela fingers comprises 4–6 pairs of internal and external accessory denticles.

The pedipalp trichobothriotaxy is neobothriotaxic major Type C (Vachon, 1974; figs. 7, 8). The femur possesses three trichobothria (d, i, e). The patella possesses 19 trichobothria, one of which is petite (esb₂): one internal (i), two dorsal (d_{1, 2}), three ventral (v_1–3), and 13 external (et_1–3, est, em_{1, 2}, esb_{1, 2}, eb_1–5). The chela possesses 27 trichobothria, including one accessory trichobothrium in the V series, and two petite trichobothria (Et₄, Esb): 18 trichobothria are situated on the chela manus (Et_1–5, Est, Esb, Eb_1–3, V_1–5, Dt, Db, db) and nine on the fixed finger (et, est, esb, eb, dt, dst, dsb, it, ib). Macroseta M₁ is situated near trichobothrium e on the pedipalp femur (fig. 7A). On the pedipalp chela, trichobothrium db is situated medially; Et₃ is situated either in the same same axis as, or proximal to Est; Esb is situated either between Eb₂ and Eb₃ or dorsal to Eb₂; V₁, V₂, and V₃ are situated in the same axis; dst is situated proximal to est; Db is equidistant between Dt and Eb₃ (except in O. huascaran); Et₅ forms an obtuse angle with eb and Et₄ (fig. 8A); and ib is situated at the base of the apophysis on the internal surface of the chela manus of adult males, where it is not visible in ventral view (fig. 8C).

The hemispermatophore is lamelliform and slender (fig. 10). The elongated distal lamina is separated into two parts, a basal portion delimited by the frontal crest, and a distal portion termed the apex (Ochoa, 2004a). The apex comprises two parts, a narrow base termed the pedicel, and a subtriangular distal part that includes the distal crest. The frontal crest is further divided into two parts: the proximal part is oblique and possesses two lateral folds, the distal part is straight and usually parallel to the ventral margin of the lamina, with a lateral projection on both the ental and ectal surfaces. The distal edge exhibits a small incision and, in most species, there is a constriction along the border at the point of inflexion. The shape and length of the apex and frontal crest vary among the species (figs. 27, 28, 32, 36, 40, and 47). The lobe region is greatly reduced, the capsular concavity is shorter than the fold of the internal lobe, and the basal lobe forms a tortuous stem with a subdistal spatulate dilation ending in a terminal process (fig. 10).

Included taxa

Fifteen species (table 1). Orobothriurus lourencoi 86Ojanguren Affilastro, 2003, will be placed in another genus based on phylogenetic analysis (fig. 5, appendix 2; Mattoni et al., in litt.).

Fig. 5.

Most parsimonious tree (length  =  167; CI: 0.479; RI: 0.813), obtained by cladistic analysis of 65 morphological characters (appendix 2) scored for 15 species of Orobothriurus Maury, 1976, and 15 outgroup taxa with the implied weighting regime that maximized average support (Mattoni et al., in litt.). Synapomorphies optimized with accelerated transformation indicated with bars. Black bars indicate uniquely derived apomorphic states, white bars indicate parallel derivations of apomorphic states. Numbers above bars indicate characters, numbers below indicate states.

Fig. 6.

Orobothriurus Maury, 1976, metasomal segments I–V showing distribution of macrosetae. A. Lateral aspect. B. Ventral aspect. Annotations: dorsolateral (DL); median lateral (ML); ventrolateral (VL); ventrosubmedian (VSM).

Fig. 7.

Orobothriurus Maury, 1976, dextral pedipalp femur (A) and patella (C–D), showing trichobothrial pattern. A. Femur, dorsal aspect. B. Patella, dorsal aspect. C. Patella, external aspect. D. Patella, ventral aspect. Annotations: dorsal (d); external (e); external basal (eb); external medial (em); external suprabasal (esb); external subterminal (est); external terminal (et); internal (i); ventral (v). Scale bar  =  1 mm.

Distribution

Orobothriurus is endemic to Argentina, Bolivia, Chile and Peru (table 1). Five of the described species are endemic to Argentina, two to Chile and seven to Peru.

Fig. 8.

Orobothriurus Maury, 1976, dextral pedipalp chela, showing trichobothrial pattern. A. External aspect, with dorsal aspect of fixed finger. B. Ventral aspect. C. Ventrointernal aspect. Annotations: dorsal basal (db, Db); dorsal suprabasal (dsb); dorsal subterminal (dst); dorsal terminal (dt, Dt); external basal (eb, Eb); external suprabasal (esb, Esb); external subterminal (est, Est); external terminal (et, Et); internal basal (ib); internal terminal (it); ventral (V). Scale bar  =  1 mm.

Orobothriurus alticola (Pocock, 1899)

Figures 13F, 15B, 18D, 20A, 21A, 36I, J, 54

Cercophonius brachycentrus var. β bivittatus Thorell, 1877: 183 [synonymized by Ojanguren Affilastro et al., 2009: 35].

Bothriurus alticola Pocock 1899: 357–358, fig. 1; Kraepelin, 1911: 91, 97; Mello-Leitão, 1931: 82, 92; 1932: 34; 1934: 63, 65; 1935: 93; 1937: 103; 1945: 138, 144, 145; Bücherl, 1959a: 23, 24, fig. 4; Abalos, 1959: 591; 1963: 113; Cekalovic, 1966: 3; Bücherl, 1969: 769; Maury, 1973: 110; Roig Alsina, 1973: 198; Masnú de Moreno, 1991: 184, 189, map 1.

Bothriurus (Andibothriurus) alticola: Bücherl et al., 1963: 217, 224, fig. 8.

Urophonius brachycentrus bivittatus: Mello-Leitão 1931: 99–100; 1932: 35; 1934: 48, 51; 1938: 94–95; 1945: 213, 215; Acosta and Maury, 1998: 559; Lowe and Fet, 2000: 44.

Urophonius brachicentrus bivittatus: Mello-Leitão, 1939: 612; Abalos, 1959: 592; 1963: 117; Roig Alsina, 1973: 200 [incorrect subsequent spelling].

Orobothriurus bivitattus: Acosta, 2005: 3–12, figs. 1–8; Ojanguren Affilastro, 2005: 181–183, 220, 241; Acosta, 2006: 20, 21; ICZN, 2008: 69–70.

Orobothriurus alticola: Maury, 1976: 17–18, figs. 1–10, table 1 (part); 1979: 717, map 10 (part); 1981: 98 (part); Acosta and Maury, 1998: 559 (part); Kovařík, 1998: 101; Acosta and Ochoa, 2000: 135, 136, 143; Lowe and Fet, 2000: 35; Acosta and Ochoa, 2001: 203–205, 208, 209; Acosta, 2002: 176, 177; Ochoa, 2004a: 44, 49, 50, 52, 55, figs. 1, 2, 21, table 1; Acosta, 2005: 1, 2, 8, 9, 12; Ojanguren Affilastro, 2005: 176, 178, 179, 180, 220, 241; Kamenz and Prendini, 2008: 39, pl. 111, table II; Ojanguren Affilastro et al., 2009: 35–40, figs. 2, 18, 19, 25–35, table 2.

Type Material

ARGENTINA: Mendoza Province: Las Heras Department: 1 ♂, 1 ♀ syntypes (BMNH), Puente del Inca [32°49′09.15″S 69°55′01.82″W, 2721 m]. Cercophonius brachycentrus bivittatus: 1 juv. holotype (NRS), San Juan Province.

New Records

ARGENTINA: Mendoza Province: Las Heras Department: Puente del Inca, 32°49.568′S 69°54.606′W, 2759 m, 2.xi.2003, J.A. Ochoa, L. Prendini and C.I. Mattoni, alpine vegetation (low spiny bushes) on hard, rocky ground, under stone, 1 juv. (AMNH [LP 2386]). San Juan Province: Calingastra Department: Cerro el Tontal, 31°31′24.7″S 69°12′23.3″W, 3600 m, 25.i.2006, A. Ojanguren Affilastro, L. Compagnucci and L. Piacentini, 2 juv. (AMNH [LP 5848]); between Paso de Agua Negra and Aduana, vega and surrounds, 30°17′33.1″S 69°46′45.6″W, 4005 m, 27.i.2005, C.I. Mattoni and A. Ojanguren Affilastro, UV sampling, 1 subad. ♂, 1 subad. ♀, 1 juv. ♂ (AMNH [LP 4309]).

Diagnosis

Orobothriurus alticola is similar to O. grismadoi in pigmentation pattern, external morphology, and hemispermatophore. The two species can be separated by the shape of the hemispermatophore: the apex is narrower, and the angle formed by the apex with the rest of the distal lamina smaller in O. grismadoi (fig. 36D, E) than in O. alticola (fig. 36I, J). Males can also be separated by the shape of the telson, which is more slender in O. grismadoi. Additionally, the dorsal surface of the telson vesicle is concave in males of O. grismadoi, whereas it is flat in males of O. alticola (fig. 23A, E). Tergite VII is almost entirely pigmented, without an unpigmented median stripe, in O. grismadoi (fig. 13G), but exhibits paired lateral spots of pigmentation, delimiting an unpigmented median stripe, in O. alticola (fig. 13F). On the other hand, O. alticola can be separated from O. compagnuccii by the shape of the lamina of the hemispermatophore: the apex comprises 52.17%–54.54% (n  =  3; mean  =  52.98%) of the lamina in O. compagnuccii and 40.87%–46.08% (n  =  20; mean  =  43.88%) in O. alticola. Orobothriurus alticola and O. ramirezi can be separated by the distal crest of the hemispermatophore, which is less developed, occupying only half the apex of the lamina, in O. alticola, than in O. ramirezi, in which it occupies almost the entire apex (figs. 36I, J, 47A, B). The telson vesicle of O. alticola is narrower, especially in females (telson length/width ratio: ♂, 2.87–3.21, mean  =  3.07; ♀, 2.44–2.55, mean  =  2.49; fig. 23A) than that of O. ramirezi (telson length/width ratio: ♂, 2.65–2.94, mean  =  2.78; ♀, 2.13–2.37, mean  =  2.27; figs. 23G, 24E). The dorsal surface of the femur and ventral surface of the metasoma are less pigmented, with narrower stripes, in O. alticola than O. ramirezi. The metasomal carinae, especially the VL and VSM carinae of segments I and V, and the DL carinae of segment V, are less developed in males of O. alticola than O. ramirezi (fig. 20A, C).

Remarks

Acosta (2002) transferred Cercophonius brachycentrus bivittatus to Orobothriurus, and subsequently redescribed it as a valid species based on a single male specimen (Acosta, 2005). Based on an examination of additional material, however, Ojanguren Affilastro et al. (2009) were unable to identify any characters by means of which this species could be consistently separated from O. alticola, and it was therefore synonymized.

Distribution

Orobothriurus alticola is endemic to central-western Argentina, from the central part of Mendoza Province (Laguna Diamante) to the central part of San Juan Province (Paso del Agua Negra; fig. 54). It occurs at 2700–4000 m in the Andes (fig. 3B) and nearby El Tontal mountain range of the Precordillera (fig. 3C, Ojanguren Affilastro et al., 2009).

Ecology

The area inhabited by O. alticola belongs to the Puna and the Altoandina phytogeographic provinces (Cabrera and Willink, 1980). This species was collected in sympatry with another bothriurid, Brachistosternus montanus Roig Alsina, 1977.

Orobothriurus ampay Ochoa and Acosta, 2003

Figures 4A, 11A, 12D, 17B, 22A, 25A, 27A, 52

Orobothriurus ampay Ochoa and Acosta, 2003: 2–6, figs. 1–14; Ochoa, 2004a: 43, 52, 54, 73, figs. 1, 2, 21, table 1; 2005: 55, figs. 7, 9, table 2.

Type Material

PERU: Apurímac Department: Abancay Province: Holotype ♂ (MUSM), Tamburco, Sector Arapato, Santuario Nacional Ampay, 13°35′S 72°52′W, 3580 m, 25.viii.1998, R. Aimituma. Paratypes: Santuario Nacional Ampay, Bosque Nativo del Ampay [13°35′41″S 72°52′48″W], 3100–3300 m, 26.viii.1998, J.C. Chaparro and J.A. Ochoa, 2 ♀ (MHNC), 1 ♀ (CDA 088), 1 ♀ (MACN-Ar 10039); Santuario Nacional Ampay, Ccorhuani [ca. 13°35′S 72°52′W], 3280 m, 18.ix.1998, R. Aimituma, 1 ♀ (MUSM).

New Records

PERU: Apurímac Department: Abancay Province: Santuario Nacional Ampay, Bosque Nativo del Ampay, 13°35′S 72°52′W, 3200 m, 17.vii.2000, J.A. Ochoa, in Podocarpus glomeratus forest, 1 ♀ (AMNH [LP 1918]).

Diagnosis

Orobothriurus ampay differs from all other species in the genus by its well-developed pigmentation pattern. The tergites of this species are entirely, densely pigmented (fig. 12D) and the paired VL stripes contiguous with the VM stripe posteriorly on metasomal segments I–V (fig. 11A) whereas, in other species, tergites I–VI each possess two lateral spots delimiting an unpigmented median stripe along the mesosoma (fig. 13E–G) and the paired VL stripes of the metasomal segments are not contiguous with the VM stripe posteriorly, at least on segments IV and V (figs. 11B–D, 13A, D). Orobothriurus ampay may be further distinguished on the basis of metasomal carination, trichobothriotaxy and pedipalp chela ornamentation. The VSM carinae, present on metasomal segment IV in O. ampay (fig. 17B), are absent in other species of Orobothriurus (fig. 17A). The angle formed between pedipalp chela trichobothria eb–Et₅–Et₄ is less than 90° in O. ampay (fig. 25A), but greater than 90° in other species (figs. 25B, D, 26B, C). The acuminate conical apophysis on the internal surface of the male pedipalp chela manus, observed in all other Orobothriurus species (except O. wawita, in which it is absent altogether), is reduced to a few granules in O. ampay. The hemispermatophore of O. ampay resembles that of O. parvus and O. wawita in possessing an elongated apex and a short frontal crest, which is less than half the length of the lamina. However, the ventral margin of the apex is inclined to the dorsal surface and curved distally in O. ampay (fig. 27A) and O. parvus (fig. 27B), but straight in O wawita (fig. 27C). Finally, O. ampay possesses the lowest pectinal tooth count in the genus (12–15).

Distribution

Orobothriurus ampay is known only from the Santuario Nacional Ampay, near Abancay in the Apurímac Department of southeastern Peru, at 3100–3580 m (figs. 3E, 52).

Ecology

Orobothriurus ampay appears to be endemic to an inter-Andean forest of Podocarpus glomeratus Don (Podocarpaceae) (fig. 3E). All known records of this species were collected inside the forest under stones on moist soil. No other scorpion species are known from this forest. The closest records are two other bothriurids, Brachistosternus peruvianus Piza, 1974, and Pachakutej iskay (Acosta and Ochoa, 2001), and a buthid, Tityus footei Chamberlin, 1916, which inhabit an adjacent dry valley in Tamburco.

Orobothriurus atiquipa Ochoa and Acosta, 2002

Figures 11D, 18A, 21A, 25D, 28A, 53

Orobothriurus atiquipa Ochoa and Acosta, 2002a: 99–102, figs. 1–9, table 1; Ojanguren Affilastro, 2003a: 121, fig. 14; Ochoa, 2004a: 43, 52, 55, 73, figs. 1, 2, 21, table 1; 2005: 55, figs. 7, 9, table 2; Rein, 2007: 5.

Orobothriurus aff. paessleri: Lourenço and Dastych, 2001: 54 [probable misidentification: specimens from Chala].

Orobothriurus paessleri: Lourenço and Dastych, 2001: 54 [probable misidentification: specimens from Atico and Road between Chala and Cháparra].

Type Material

PERU: Arequipa Department: Caraveli Province: Holotype ♂ (MACN-Ar 10010), paratype ♂ (MHNC), Lomas Atiquipa, Cerro Lloque, 15°45′S 74°22′W, 950 m, 13.ix.1999, H. Zeballos and R. Gutiérrez.

New Records

PERU: Arequipa Department: Caraveli Province: Lomas Atiquipa, Sector Conchara, 15°46′16″S 74°22′42″W, 750 m, 6.iii.2004, J.L. Velasquez and J.A. Ochoa, 3 ♂, 3 juv. (MHNC), 2 juv. (AMNH [LP 3056]).

Diagnosis

Orobothriurus atiquipa may be distinguished from all other species of the genus by means of the telotarsal setation of the legs and the granulation on the ventral surface of metasomal segment V. In O. atiquipa, the telotarsi of legs III and IV each possess 3/4 spiniform macrosetae in the pro- and retroventral rows, and the VSM and VM carinae are not discernible from the dense granulation on the ventral surface of segment V (fig. 21A) whereas, in other species of Orobothriurus, the telotarsi of legs III and IV each possess 3/3 spiniform macrosetae in the pro- and retroventral rows, and the VM and VL carinae are well defined on segment V (figs. 20A, D, E, 21D). Additionally, the hemispermatophore of O. atiquipa is proportionally longer than that of all other species. This species possesses the most elongated frontal crest in the genus (fig. 28A).

Distribution

Orobothriurus atiquipa is known only from the type locality in northern Arequipa Department, southern Peru, at 750–950 m (figs. 2D, 53).

Ecology

Lomas biotopes are isolated patches of green vegetation on hillsides below 1000 m, surrounded by hyperarid desert, along the western coast of southern Peru and northern Chile (fig. 2D). Climatic patterns produce fog zones that allow plant communities to exist in areas where the rainfall is very low. The flora and fauna of Lomas have closer affinities with that of the high Andean peaks than the surrounding desert biota (Herrera, 1930; Péfaur, 1981; Ochoa, 2005). The vegetation in the habitat of O. atiquipa is characterized by shrubs, herbaceous plants, and trees like Myrcianthes ferreyrae (McVaugh) McVaugh (Myrtaceae), Caesalpinia spinosa (Molina) Kuntze (Fabaceae), Acacia macracantha Humboldt and Bonpland ex Willdenow (Fabaceae), and Carica candicans A. Gray (Caricaceae). An iurid in the genus Hadruroides was recorded in sympatry with O. atiquipa (Ochoa and Prendini, 2010).

Remarks

Specimens from three localities (Atico, Chala and Road between Chala and Cháparra) in the Arequipa Department, Peru, were listed under O. paessleri and “O. aff. paessleri” by Lourenço and Dastych (2001: 54), who noted that the specimens from Chala are “possibly a new species.” Although we have not examined these specimens, they are probably conspecific with O. atiquipa, not O. paessleri, based on the known distribution of this species.

Orobothriurus calchaqui, n. sp.

Figures 14A, 18B, 21D, 23B, 24A, 29–Fig. 30.31, 32A, B, 36K–O, 54; table 3

Table 3

Selected measurements (mm) of type specimens of six new species of Orobothriurus Maury, 1976: Orobothriurus calchaqui, n. sp., Orobothriurus compagnuccii, n. sp., Orobothriurus huascaran, n. sp., Orobothriurus quewerukana, n. sp., Orobothriurus ramirezi, n. sp., and Orobothriurus tamarugal, n. sp. Abbreviations: AMNH  =  American Museum of Natural History, New York; MACN  =  Museo Argentino de Ciencias Naturales “Bernardino Rivadavia,” Argentina; MHNC  =  Museo de Historia Natural, Universidad Nacional de San Antonio Abad del Cusco, Peru; LBRE  =  Laboratorio de Biología Reproductiva y Evolución, Universidad Nacional de Córdoba, Argentina; MZUC  =  Museo de Zoología de la Universidad de Concepción, Chile.

Orobothriurus alticola: Maury, 1976: 18 (part) [misidentification].

Type Material

ARGENTINA: Tucumán Province: Tafí del Valle Department: Holotype ♂ (MACN-Ar), El Infiernillo (Tafí del Valle), 26°44′13.3″S 65°45′51.2″W, 2700 m, 30.x.2004, A. Ojanguren Affilastro and L. Compagnucci. Paratypes: same data, 20 ♂, 2 ♀ (MACN-Ar), 2 ♂ (AMNH), 2 ♂ (AMNH [LP 5195]), 2 ♂ (LBRE), 2 ♂ (MHNC); Cumbres Calchaquies, Huaca Huasi, 4250 m, E. Domínguez, 1 ♀ (MACN-Ar); Vaca Huasi, 4300 m, 16.xi.1974, S. Hallow, 1 ♀ (MACN-Ar).

Etymology

The specific name is a noun in apposition referring to the Cumbres Calchaquies mountain range in the Andes, where this species occurs.

Diagnosis

Orobothriurus calchaqui is most closely related to O. famatina (fig. 5). The two species can be separated by a combination of characters, several of which overlap in some specimens. Orobothriurus calchaqui possesses two or three ML macrosetae on metasomal segment IV, and 70% of specimens possess one DL macroseta on segment III (absent in 30%), whereas O. famatina possesses one ML macroseta on segment IV and only 10% of specimens possess one DL macroseta on segment III (absent in 90%). There are also differences in the trichobothrial pattern. Pedipalpal chela trichobothria db and Et₃ are always located proximal to Est in O. calchaqui (fig. 31C) whereas, in O. famatina, db is located in the same axis as Et₃ (in 70% of the specimens) or slightly distal to it (30%). Pedipalp patella trichobothrium em₂ is located proximal to em₁ in 80% of the specimens of O. calchaqui (fig. 30C), but in the same axis as, or distal to em₁ in 70% of the specimens of O. famatina. There are also slight differences in the shape of the hemispermatophore in most specimens. The pedicel of the apex is usually narrower and the angle formed by the apex with the rest of the distal lamina usually greater in O. calchaqui than O. famatina (fig. 36F–H, K–M). However, the hemispermatophore of some specimens of O. calchaqui is indistinguishable from that of O. famatina (fig. 36N, O).

Description

Based on holotype ♂ and paratypes. Measurements of holotype ♂ and paratype ♀ recorded in table 3.

Total length: ♂, 23–30 mm (n  =  10, mean  =  25.9 mm); ♀, 33.5–36 mm (n  =  4, mean  =  34.45 mm).

Color: General color yellowish with dark brown spots (fig. 29). Carapace, anterior margin faintly pigmented; lateral margins with two large spots medially and posteriorly, remaining area with reticulate pigmentation; median ocular tubercle and lateral ocelli dark brown to black; posterior half of anteromedian longitudinal sulcus, median ocular tubercle, and postocular sulcus pigmented. Chelicerae, distal surface of manus, and movable finger with reticulate pigmentation. Pedipalp coxa and trochanter faintly pigmented; femur densely pigmented near articulation with patella; patella densely pigmented near articulations; chela manus with six complete longitudinal stripes along carinae, contiguous at base of movable finger; fingers and articulation of fingers pigmented. Legs, femur, patella, tibia, and basitarsus faintly pigmented, especially near articulations and along dorsal margin. Tergites I–VI each with two dark spots sublaterally along almost entire margin, broader near anterior margin, delimiting broad, unpigmented median stripe (fig. 29A, B); VII with four dark spots, two larger spots posterolaterally, occupying area between dorsolateral carinae and external margins of sclerite, and two small dark triangular spots submedially, extending along submedian carinae, delimiting unpigmented median stripe. Sternum, genital opercula and pectines unpigmented. Sternites III–VI usually unpigmented, but lateral margins of segments V and VI faintly pigmented in some specimens; VII with two narrow dark stripes sublaterally (fig. 29B, D). Metasomal segment I, dorsal surface unpigmented or with two small faint spots medially; lateral surfaces densely pigmented between ML and LIM carinae (fig. 29); ventral surface with two narrow VL stripes, becoming slightly broader in posterior half. Segment II, dorsal surface with two subtriangular spots medially; lateral surface as for segment I; ventral surface with one narrow VM and two narrow VL stripes, becoming slightly broader in posterior half. Segment III, dorsal surface as for segment II; lateral surfaces densely pigmented below ML carinae; ventral surface as for segment II. Segment IV, dorsal surface with reticulate pigmentation medially; lateral and ventral surfaces as for segment III. Segment V, dorsal and lateral surfaces as for segment IV; ventral surface with three dark narrow stripes (one VM and two VL), becoming slightly broader in posterior half but not contiguous with lateral pigmentation at posterior margin. Telson vesicle with narrow VM and two broad, dark VL stripes, separated by two narrow unpigmented stripes; aculeus sclerotized, dark reddish brown.

Chelicerae: Movable finger with two subdistal teeth.

Carapace: Surfaces smooth medially, finely granular near lateral margins. Anterior margin with shallow median notch (fig. 14A). Anteromedian longitudinal sulcus weakly developed; median ocular, posteromedian longitudinal and posterolateral sulci well developed. Median ocular tubercle raised, situated anteromedially; median ocelli two ocular diameters apart.

Pedipalps: Femur, DI and VI carinae complete, finely granular; DE carina complete, finely granular proximally (fig. 30A); internal surface finely and sparsely granular medially; ventral surface finely granular proximally; other surfaces smooth. Patella, DI and VI carinae complete, finely granular (fig. 30B–D); DPP carina finely granular; VPP carina vestigial, comprising two or three small granules proximally; internal surface with prominent granule adjacent to trichobothrium i near DI carina. Chela manus slender, fingers relatively elongated (fig. 31); length/width ratio: ♂, 3.97–4.51 (n  =  10; mean  =  4.17), ♀, 3.47–3.73 (n  =  4, mean  =  3.63); length/height ratio: ♂, 3.40–4.03 (n  =  10; mean  =  3.71), ♀, 3.07–3.31 (n  =  4, mean  =  3.18); D, DS, DMA, DI, and VM carinae obsolete; intercarinal surfaces smooth; internal surface with acuminate apophysis (♂) or low bulge (♀) near articulation of movable finger (fig. 31A, D); fingers, dentate margins each with median denticle row and 4–5 pairs of internal and external accessory denticles.

Trichobothria: Femur with 3 trichobothria, patella with 19, chela with 27 (fig. 30, 31). Patella trichobothrium em₂ situated proximal to em₁ in 80% of specimens (fig. 30C). Chela trichobothrium Et₃ situated proximal to Est (fig. 31C).

Tergites: Tergites I–VI, surfaces finely granular, becoming smooth posterolaterally. Tergite VII tetracarinate, paired DL carinae restricted to posterior two-thirds of segment, paired DSM carinae to posterior half; intercarinal surfaces coarsely granular, other surfaces finely granular.

Legs: Femur and patella, prolateral surfaces finely granular, retrolateral surfaces smooth. Femur, ventral carinae weakly developed; other carinae absent. Patella acarinate. Telotarsi, pro- and retroventral rows of spiniform macrosetae with following counts on leg I, 1/1; II, 2/2; III and IV, 3/3.

Pectines: Pectinal tooth count: ♂, 18–21 (n  =  21, mode  =  18); ♀, 14 (n  =  4, mode  =  14).

Sternites: Sternites III–VI, surfaces smooth; spiracles small, narrow. Sternite VII, surface smooth; VSM and VL carinae well developed (♀, fig. 17B) or obsolete (♂).

Metasoma: Segment I, DL carinae complete, moderately granular; ML carinae complete, moderately granular in posterior two-thirds; one pair of ML macrosetae; LIM carinae complete, moderately granular in posterior half; VL and VSM carinae complete, well developed (especially in ♀), granular (fig. 17B); three pairs of VL and two pairs of VSM macrosetae. Segment II, DL carinae complete, granular; ML carinae complete, granular in posterior two-thirds; one pair of ML macrosetae; LIM carinae restricted to posterior third; VL and VSM carinae complete, well developed (♀) or obsolete (♂); three pairs of VL and VSM macrosetae. Segment III, DL carinae complete, granular; ML carinae complete, granular in posterior third; one pair of ML macrosetae; LIM carinae obsolete, restricted to posterior quarter; VL and VSM carinae complete, well developed (♀) or absent (♂); three pairs of VL and VSM macrosetae. Segment IV, DL carinae complete, granular; one pair of DL macrosetae; ML carinae obsolete, restricted to posterior margin; one pair of ML macrosetae; LIM carinae absent; VL and VSM carinae absent; three pairs of VL and VSM macrosetae. Segment V, length/width ratio, ♂, 2.33–2.73 (n  =  10; mean  =  2.52), ♀, 1.93–1.98 (n  =  4; mean  =  1.96); DL carinae granular, restricted to anterior half; ML carinae absent; four or five pairs of ML macrosetae; lateral margin smooth; VL carinae complete (♂) or restricted to posterior three-quarters (♀); VL and VSM carinae situated close together, fused in anterior and posterior thirds; four pairs of VL and VSM macrosetae; two pairs of macrosetae along posterior margin; VM carina complete, obscured by surface granulation in posterior half (fig. 21D).

Telson: Length/height ratio: ♂, 3.52–3.89 (n  =  10, mean  =  3.74); ♀, 2.93–3.13 (n  =  4, mean  =  3.01). Vesicle elongated (♂, fig. 23B) or globose (♀, fig. 24A); dorsal surface slightly concave, gland not apparent (♂); ventral surface smooth (♂) or granular (♀). Aculeus short and curved, more so in ♀.

Hemispermatophore: Apex very well developed; distal crest curved like ventral margin. Frontal crest weakly developed, less than half length of lamina; basal part oblique; distal part short, parallel to ventral margin of lamina; lateral projections slightly undulated. Basal lobe, terminal process extending almost to distal end of frontal crest (fig. 32A, B).

Remarks

We examined some Orobothriurus specimens from other Andean localities in Salta and Tucumán provinces of northern Argentina. However, specimens from these localities are so scarce and the morphological differences between them and O. calchaqui so subtle that at this time it is unclear whether or not they are conspecific. More specimens from these Orobothriurus populations are required to establish their identity.

Distribution

This species is known only from the type locality, Infiernillo Mountain (Tucumán Province, Argentina), belonging to the Cumbres Calchaquies, part of the Andes (fig. 54). Altitude records from Huaca Huasi (4250 m) and Vaca Huasi (4300 m) should be confirmed, because, at this latitude, the environment above 4000 m is usually too extreme for scorpions to survive.

Ecology

All personally collected specimens were found at night in a rocky area near a small mountain river, at 2700 m. The area is a typical Puna habitat (Cabrera and Willink, 1980). Most specimens collected were on the ground, but some were taken from near-vertical rock walls besides the river, almost 2 m from its base. No specimens could be found in the surrounding area, which has a sandy soil without rocks. Most of the specimens collected were adult males, moving very rapidly, probably searching for females, suggesting that late November, the start of spring in the Southern Hemisphere, is the start of the active period for this species. No other scorpion species were collected in sympatry with O. calchaqui. However, another bothriurid, Brachistosternus intermedius Lönnberg, 1902, has been recorded in the vicinity (Ojanguren Affilastro, 2005).

Orobothriurus compagnuccii, n. sp.

Figures 14B, 18E, F, 20B, 21C, 24B, 32C, D, 33–Fig. 34.35, 36A–C, 54; table 3

Type Material

ARGENTINA: La Rioja Province: General Lamadrid Department: Holotype ♂ (MACN-Ar), Laguna Brava Provincial Park, road to Laguna Brava, El Peñón refuge, 28°32′18.6″S 68°45′11.9″W, 3200 m, i.2003, A. Ojanguren Affilastro and P. Korob (MACN-Ar). Paratypes: Laguna Brava Provincial Park, road to Laguna Brava, 28°27′17.2″S 68°50′39.9″W, 3835 m, 4.iii.2002, A. Ojanguren Affilastro and P. Korob, 2 ♀ (MACN-Ar), 28°30′26.5″S 68′47′57.0″W, 3600 m, 23.i.2007, A. Ojanguren Affilastro, L. Compagnucci, and J.J. Martínez, 2 ♂ (MACN-Ar), 28°25′54.1″S 68°50′34.7″W, 3900 m, 27.i.2006, A. Ojanguren Affilastro, L. Compagnucci, and L. Piacentini, 2 ♀ (MACN-Ar), 28°25′50.3″S 68°50′31.3″W, 3900 m, 27.i.2006, A. Ojanguren Affilastro, L. Compagnucci, and L. Piacentini, 1 juv. ♂ (AMNH [LP 5847]).

Etymology

This species is named after Argentine entomologist Luis Compagnucci (MACN), who participated in several expeditions and collected part of the type material of this species.

Diagnosis

Orobothriurus compagnuccii is similar to O. calchaqui and O. famatina. The three species can be separated by the shape of the lamina of the hemispermatophore, which has a more developed apex in O. calchaqui and O. famatina: the apex comprises 52.17%–54.54% (n  =  3; mean  =  52.98%) of the lamina in O. compagnuccii (fig. 36A–C), 66.6%–68.48% (n  =  3, mean  =  67.02%) in O. famatina (fig. 36F–H), and 69.0%–71.56% (n  =  6; mean  =  70.43%) in O. calchaqui (fig. 36K–O). Orobothriurus compagnuccii can be separated from O. alticola and O. grismadoi by means of the same characters: the apex comprises 40.87%%–46.08% (n  =  20; mean  =  43.88%) of the lamina in O. alticola and 42.70–45.69% (n  =  7; mean  =  44.17%) in O. grismadoi. Orobothriurus compagnuccii can be distinguished from other Argentine species of the genus, in which the VL and VSM carinae of sternite VII and metasomal segment I are well developed (fig. 18B, D), by the weak development or absence of these carinae, which are less developed in males (fig. 18E, F).

Description

Based on holotype ♂ and paratypes. Measurements of holotype ♂ and paratype ♀ recorded in table 3.

Total length: ♂, 23.91–25.6 mm (n  =  3, mean  =  24.86 mm); ♀, 24.5–30 mm (n  =  4, mean  =  27.75 mm).

Color: General color yellowish with dark brown spots. Carapace, anterior margin faintly pigmented (fig. 33A, C); lateral margins with two large spots medially and posteriorly, intermediate area with reticulate pigmentation; lateral margins with two large spots laterally and posterolaterally, remaining area with reticulate pigmentation; median ocular tubercle and lateral ocelli dark brown to black; posterior half of anteromedian longitudinal sulcus, median ocular tubercle, and anterior half of postocular furrow pigmented. Chelicerae, distal margin of movable finger with reticulate pigmentation. Pedipalp femur and patella, dorsal margin densely pigmented; chela manus with seven complete longitudinal stripes along carinae, contiguous at base of movable finger; fingers and articulation of fingers pigmented. Legs, especially femur, with reticulate pigmentation near articulations and along dorsal margins. Tergites I–VI each with two dark spots sublaterally along almost entire margin and pretergites, broader near anterior margin, delimiting broad, unpigmented median stripe (fig. 33A, C); VII with four dark spots, two spots posterolaterally, occupying area between dorsolateral carinae and external margins of sclerite, and two small dark triangular spots submedially, extending along submedian carinae, delimiting unpigmented median stripe. Sternum, genital opercula, and pectines unpigmented. Sternites III–VI usually unpigmented, but lateral margins of segments V and VI faintly pigmented in some specimens; VII with two dark narrow stripes sublaterally (fig. 33B, D). Metasomal segment I, dorsal surface with two small, faint spots medially, separated by narrow unpigmented median stripe; lateral surfaces densely pigmented between ML and LIM carinae (fig. 33B, D); ventral surface with three narrow stripes, two VL and one VM, becoming slightly broader, but not contiguous, in posterior half. Segment II similar to segment I, but with dorsal spots joining in some specimens. Segments III and IV similar to segment I but with dorsal spots contiguous, and DL carinae faintly pigmented. Segment V, dorsal surface with two dark stripes, contiguous in posterior third, faint or absent in some specimens; lateral surfaces each with faint narrow stripe in anterior half; ventral surface as for segments I–IV. Telson vesicle with three broad dark stripes (one VM and two VL), separated by two narrow unpigmented stripes; dorsal surface slightly pigmented on lateral margins; aculeus sclerotized, dark reddish brown.

Chelicerae: Movable finger with two subdistal teeth.

Carapace: Surfaces finely granular (♂) or smooth (♀). Anterior margin sublinear, with very shallow median notch (fig. 14B), more developed in ♀. Anteromedian longitudinal sulcus incomplete, absent at anterior margin and weakly developed toward median ocular tubercle; median ocular sulcus well developed; posteromedian longitudinal and posterolateral sulci obsolete (fig. 14B). Median ocular tubercle shallow, situated anteromedially; median ocelli two ocular diameters apart.

Pedipalps: Femur, DI and VI carinae complete, granular; DE carina granular, restricted to proximal three-quarters of segment (fig. 34A); internal surface finely and sparsely granular. Patella, DI and VI carinae complete, granular (♂) or obsolete (♀) (fig. 34B–D); DPP comprising small granules proximally; VPP vestigial, reduced to one or two small granules medially (♂) or absent (♀); internal surface with prominent granule adjacent to trichobothrium i near DI carina; other intercarinal surfaces smooth. Chela manus slender, fingers relatively elongated (fig. 35); length/width ratio: ♂, 3.81–4.47 (n  =  3, mean  =  4.17), ♀, 3.75–4.19 (n  =  4, mean  =  3.97); length/height ratio: ♂, 3.31–3.47 (n  =  3, mean  =  3.39), ♀, 3.17–3.40 (n  =  4, mean  =  3.26); DMA, DI, and VM carinae obsolete; intercarinal surfaces smooth; internal surface with acuminate apophysis (♂) or low bulge (♀) near articulation of movable finger (fig. 35A, D); fingers, dentate margins each with median denticle row and 4–5 pairs of internal and external accessory denticles.

Trichobothria: Femur with 3 trichobothria, patella with 19, chela with 27 (fig. 34, 35). Chela trichobothrium Et₃ situated in same axis as, or slightly proximal to Est (fig. 35C).

Tergites: Tergites I–VI, surfaces smooth (♀) or finely granular (♂), more coarsely so near lateral margins. Tergite VII tetracarinate, paired DL carinae restricted to posterior two-thirds of segment, paired DSM carinae to posterior third; intercarinal surfaces coarsely granular, other surfaces finely granular.

Legs: Femur and patella, prolateral surfaces finely granular (♂), retrolateral surfaces smooth. Femur, ventral carinae weakly developed; other carinae absent. Patella acarinate. Telotarsi, pro- and retroventral rows of spiniform macrosetae with following counts on leg I, 1/1; II, 2/2; III and IV, 3/3.

Pectines: Pectinal tooth count: ♂, 21–23 (n  =  3); ♀, 16–18 (n  =  4, mode  =  17).

Sternites: Sternites III–VI, surfaces smooth (♀) or densely granular medially (♂); spiracles small, narrow. Sternite VII, surface with six ventral macrosetae, smooth in anterior half, densely granular in posterior half (♂); VL and VSM carinae obsolete (indistinct from granulation) or absent (fig. 18E, F).

Metasoma: Segment I, DL and ML carinae complete, moderately granular; one pair of ML macrosetae; LIM carinae restricted to posterior two-thirds, moderately granular; VL and VSM carinae complete but indistinct, sparsely and finely (♂) or coarsely (♀) granular (fig. 18E, F); two pairs of VL and VSM macrosetae. Segment II, DL and ML carinae complete, granular; one pair of DL and ML macrosetae (DL macrosetae absent in some specimens); LIM carinae restricted to posterior quarter; VL and VSM carinae absent (♂, some ♀) or obsolete (some ♀); two pairs of VL and three pairs of VSM macrosetae (fig. 18E, F). Segment III as for segment II, but ventral intercarinal surfaces smooth and carinae less developed. Segment IV, DL carinae complete, granular; one pair of DL macrosetae; ML carinae obsolete, restricted to posterior margin; one pair of ML macrosetae; LIM carinae absent; VSM and VL carinae absent; three pairs of VSM and VL macrosetae. Segment V, length/width ratio: ♂, 2.92–3.17 (n  =  3, mean  =  3.10), ♀, 2.03–2.14, (n  =  4, mean  =  2.09); DL carinae granular, complete (♀) or restricted to anterior half (♂); DL macrosetae absent; ML carinae absent; four pairs of ML macrosetae; lateral margin smooth; VL carinae complete (♂) or restricted to posterior three-quarters (♀); VL and VSM carinae situated close together, fused in anterior and posterior thirds; four pairs of VL and VSM macrosetae; two pairs of macrosetae along posterior margin; VM carina complete, obscured by surface granulation in posterior half (fig. 20B).

Telson: Length/height ratio: ♂, 3.57–4.12 (n  =  3, mean  =  3.76); ♀, 2.81–2.97 (n  =  4, mean  =  2.90). Vesicle elongated (♂, fig. 23C) or globose (♀, fig. 24B); dorsal surface flat, gland not apparent (♂); ventral surface smooth (♂) or granular (♀). Aculeus short and curved, more so in ♀.

Hemispermatophore: Apex well developed, slightly longer than frontal crest, comprising 52.17–54.54% (n  =  3; mean  =  52.98%) of lamina; distal crest curved like ventral margin. Frontal crest divided into two parts, basal part oblique, distal part parallel to ventral margin of lamina; lateral projections slightly undulated and larger than basal part. Basal lobe, terminal process extending to median part of frontal crest (fig. 32C, D).

Distribution

Orobothriurus compagnuccii is known only from the type locality, Laguna Brava Provincial Park, in the western Andres of La Rioja Province, northwestern Argentina (fig. 54). All specimens were collected on the ascent to Pircas Negras international pass, on the border between Argentina and Chile, at 3200–3900 m.

Ecology

The habitat at the type locality is intermediate between Puna and high Andean habitats (Cabrera and Willink, 1980), with very sparse vegetation. All specimens were collected in areas with very pronounced slopes, on high rocks or spiny shrubs. We conducted several expeditions to the type locality of this species, in different years and at different times during the summer, when Orobothriurus are presumed to be most active at this latitude. However, despite considerable effort, only a few specimens were collected each time. Most were very difficult to find, hiding in inaccessible places, e.g., in vertical rocks more than two or three meters above ground level. In the same area, we collected many Brachistosternus montanus, a bothriurid species more than twice the size of O. compagnuccii. The scarcity of O. compagnuccii during summer expeditions, and its habitat preference for places inaccessible to larger scorpions adapted to sandy soils like Brachistosternus, may be explained by the abundance of B. montanus. It is possible that the active period of O. compagnuccii starts in early spring, before that of B. montanus, to avoid competition and predation by the larger species.

Orobothriurus curvidigitus (Kraepelin, 1911)

Figures 11B, 12C, 13E, 14C, 19A, 22C, 26C, 28B, D, 53

Bothriurus curvidigitus Kraepelin, 1911: 91, 97–99, fig. 7a, 7b; Mello-Leitão, 1931: 82, 92; 1932: 34; 1934: 65; 1935: 93; 1937: 103; 1945: 161; Aguilar and Meneses, 1970: 2; Maury, 1973: 110; 1981: 104.

Bothriurus lampei Werner, 1916: 92 [synonymized by Maury, 1973: 110]; Lampe, 1917: 203; Mello-Leitão, 1931: 88; 1932: 34; 1945: 175; Aguilar and Meneses, 1970: 2; Maury, 1973: 110; 1981: 104.

Bothriurus (Andibothriurus) curvidigitus: Bücherl et al., 1963: 216.

Bothriurus (Andibothriurus) lampei: Francke, 1974: 219.

Orobothriurus curvidigitus: Maury, 1976: 18, 19, figs. 11–24, table I; 1980: 338; 1981: 101; Francke, 1977: 75; Kovařik, 1998: 101; Acosta and Ochoa, 2000: 136, 143; Lowe and Fet, 2000: 35, 36; Acosta and Ochoa, 2001: 205; Ochoa and Acosta, 2002a: 102, fig. 9; Ochoa, 2004a: 52, 55, 73, figs. 1, 2, 13, 19, 21, table 1; 2005: 55, figs. 7, 10, table 2.

Type Material

PERU: Arequipa Department: Arequipa Province: 1 ♂, 1 subad. ♀ syntypes (ZMH), Yura [16°12′53″S 71°42′33″W, 2642 m], 20.vi.1909.

New Records

PERU: Arequipa Department: Arequipa Province: Agua del Milagro, Characato [16°28′S 71°28′W, 2460 m], 12.x.1983, E. Maury, 4 ♀, 8 juv. (MACN-Ar); Arequipa, 1937, P. Crawford, 1 ♀ (MACN-Ar); Canchimayo, Chiguata [16°24′13″S 71°23′34″W, 2950 m], 18.x.1998, J.A. Ochoa, 4 juv. (MHNC); Charcani, Arequipa, 2800 m, xi.1947, W. Weyrauch, 3 ♀, 2 juv. (MACN-Ar); entrance to Charcani Quinto, near Arequipa city, Chili (Radio Azul) [16°19′33″S 71°31′47″W, 2550 m], 28.ii.1998, E. Ponce and J.A. Ochoa, 1 juv. (MHNC); Espíritu Santo, Chiguata [16°24′28″S 71°23′07″W, 2900 m], 7.iii.1998, E. Ponce and J.A. Ochoa, 2 ♀, 1 juv. (MHNC); Miraflores, near Chiguata [16°23′S 71°21′W, 3295 m], 7.iii.1998, E. Ponce and J.A. Ochoa, 12 ♀, 3 juv. (MHNC), 3 ♀ (CDA 166); near Mollebaya river [16°28′56″S 71°28′01″W, 2513 m], 3.iii.1998, E. Ponce, D. Muñiz and J.A. Ochoa, 2 ♂, 6 ♀, 2 juv. (MHNC), 1 ♂, 1 ♀ (CDA 167); Ribera del Rio Mollebaya [ca. 16°28′56″S 71°28′01″W, 2513 m], 8.ii.2004, R. Cardeñas and J.A. Ochoa, 2 juv. (AMNH [LP 3064]); Socabaya [16°28′01″S 71°31′18″W, 2300 m], 1948, W. Weyrauch, 1 ♂ (MUSM), 2329 m, 5.ix.1998, U. Zanabria, 1 ♀ (MHNC); Yura [16°12′53″S 71°42′33″W, 2642 m], 7.viii.1939, K.P. Schmidt, 2 ♂, 1 juv. (FMNH), 8.viii.1939, K.P. Schmidt, 1 ♂, 1 ♀ (FMNH), 10.viii.1939, K.P. Schmidt, 1 ♂, 1 juv. (AMNH), 9.v.1998, U. Zanabria, 1 ♀, 2 juv. (MHNC); Yura, Río Yura, 20 m del Cementerio, 16°13′59″S 71°42′27″W, 2426 m, 29.xii.2007, R. Gutiérrez, A. Quiroz and J.A. Ochoa, Serrania Esteparia, collected at night with UV light, 3 juv. (AMNH [LP 8357]). Cailloma Province: Calera Chivay [15°38′34″S 71°36′01″W, 3640 m], 27.ix.1998, S. Rivera, 1 ♀ (MHNC); Curiña, Yanque [15°39′34″S 71°42′15″W, 3390 m], 20.xii.1996, J.A. Ochoa, 10 ♂, 3 juv. (MHNC); Curiña [3600 m], 21.xii.1996, J.A. Ochoa, 2 ♀, 4 juv. (MHNC).

Diagnosis

Orobothriurus curvidigitus is most closely related to O. paessleri, O. quewerukana, and O. tamarugal (fig. 5), all of which have the following characters: anterior margin of carapace with weak median projection (epistome; figs. 14C, 15A, C, D); pedipalp chela movable finger of male curved (figs. 26B, C, 43C, 51A); pedipalp femur and patella of male more elongated than that of female. Orobothriurus curvidigitus may be separated from O. paessleri and O. tamarugal by the shape of the lamina of the hemispermatophore (figs. 28B, D), the frontal crest of which is more developed and elongated in O. curvidigitus, but shorter in O. paessleri and O. tamarugal (fig. 28C, E, 47D). The hemispermatophore of O. curvidigitus differs from that of O. quewerukana in the slightly shorter frontal crest and absence of folds in the distal crest (figs. 28B, 40E). Orobothriurus curvidigitus also differs from O. quewerukana in the shape and granulation of the male telson vesicle: the length/height ratio is 3.72–4.23 (mean  =  3.96) and only the anterior third of the ventral surface is granular in O. curvidigitus, whereas the length/height ratio is 3.59–4.00 (mean  =  3.76) and the ventral surface is entirely granular in O. quewerukana (fig. 23F). The male pedipalp chela carinae are finely and sparsely granular to smooth in O. curvidigitus, smooth in O. paessleri, and finely and densely granular in O. quererukana and O. tamarugal (figs. 43D, E, 51B, C). The pigmentation pattern is the most conspicuous difference between O. curvidigitus and the other three species. Orobothriurus curvidigitus exhibits well-developed pigmentation on the carapace, tergites, metasoma, and pedipalps (figs. 11B, 12C, 13E), compared to O. quewerukana, in which the pigmentation, especially that of the carapace and tergites, is faint or barely discernible (fig. 41) and O. tamarugal, which is unpigmented (fig. 48). Orobothriurus curvidigitus and O. paessleri may be further distinguished by the dimensions of metasomal segment V and the setation of metasomal segment I: segment V is relatively elongated, its length greater than twice its width, in O. curvidigitus (fig. 22C), but shorter, its length less than twice its width, in O. paessleri (fig. 22B); two pairs of VSM macrosetae are present on segment I in O. curvidigitus (fig. 19A), whereas three pairs are present in O. paessleri (fig. 19B).

Distribution

Orobothriurus curvidigitus inhabits the western slopes of the Andes at 2300–3600 m (figs. 2A, 53), in the Arequipa Department, southern Peru.

Ecology

The vegetation in the areas inhabited by this species falls within the Serrania Estaparia Ecoregion, characterized by dry zones with herbaceous vegetation, cacti, shrubs, and few trees (Brack, 1986; Ochoa, 2005; fig. 2A). Three other bothriurids, Brachistosternus ehrenbergii Gervais, 1841, and two undescribed species of Brachistosternus, also occur in this area.

Remarks

We examined populations of an undetermined Orobothriurus species, apparently related to O. curvidigitus, from Cotahuasi and Chuquibamba in the Arequipa Department, Peru, and Lucanas in the Ayacucho Department. Lourenço and Dastych (2001: 54) listed the record from Chuquibamba under “O. aff. curvidigitus” and stated that it is “probably a new species.” We cannot establish the identity of these specimens until more material becomes available, however.

Orobothriurus famatina Acosta, 2001

Figures 36F–H, 54

Orobothriurus alticola: Maury 1976: 17 (part); Cei, 1982: 660, 669 [misidentification].

Orobothriurus famatina Acosta in Acosta and Ochoa, 2001: 205–211, figs. 4–14; Ochoa, 2004a: 43, 52, 55, 73, table 1; Ojanguren Affilastro, 2005: 177–181, figs. 458–466, 477, 488, 663, table 15.

Type Material

ARGENTINA: La Rioja Province: Famatina Department: Holotype ♂ (MACN-Ar 9932), Sierra de Famatina, Rio Oro canyon, path to Mina El Oro, 29°06′S 67°42′W, 2450 m, 6.xii.1998, L. Acosta, M. Acosta and G. Repossi. Paratypes: same data, 1 ♂, 1 ♀ (LEA 240), 1 ♂ (CDA 051), 1 ♂ (MHNC); Famatina, iv.1951, R. González Amorío, 1 ♀ (MACN-Ar 6844); Mina El Oro, Chilecito, ii.1956, M.E. Galiano, 1 ♀ (MACN-Ar 6843); Sierra de Famatina, path to Mina La Mejicana, 3060 m, 5.xii.1998, L. Acosta, M. Acosta, and G. Repossi, 1 ♀ (CDA 050), 1 ♀ (MACN-Ar 9933), 1 ♀ (MHNC).

New Records

ARGENTINA: La Rioja Province: Famatina Department: Sierra de Famatina, Cuevas de Noronha (path to La Mejicana mine), 28°55′31″S 67°40′23″W, 2846 m, 29.i.2006, L. Piacentini, L. Compagnucci, and A. Ojanguren Affilastro, 2 ♂, 6 ♀, 12 juv. (MACN-Ar), 2 ♀ (AMNH [LP 5846]).

Diagnosis

Orobothriurus famatina is most closely related to O. calchaqui (fig. 5), from which it differs in the following respects. Orobothriurus famatina possesses one ML macroseta on metasomal segment IV and only 10% of the specimens possess one DL macroseta on segment III (absent in 90%), whereas O. calchaqui possesses two or three macrosetae on segment IV and 70% of the specimens possess one DL macroseta on segment III (absent in 30%). There are also differences in the trichobothrial pattern. Pedipalpal chela trichobothria db and Et₃ are located in the same axis as, or slightly distal to Est in 70% and 30% of the specimens of O. famatina, respectively, but always located proximal to Est in O. calchaqui. Pedipalp patella trichobothrium em₂ is located in the same axis as, or distal to em₁ in 70% of the specimens of O. famatina, but proximal to em₁ in 80% of the specimens of O. calchaqui. Orobothriurus famatina may be differentiated from O. compagnucci by the more elongated apex of the hemispermatophore lamina: the apex comprises 66.6–68.48% (n  =  3, mean  =  67.02%) of the lamina in O. famatina (fig. 36F–H) and 52.17%–54.54% (n  =  3; mean  =  52.98%) in O. compagnuccii (fig. 36A–C). The two species can also be separated by the VL and VSM carinae of metasomal segment I and sternite VII in the male, which are well developed in O. famatina, and weakly developed to absent in O. compagnuccii (fig. 18F).

Distribution

Orobothriurus famatina is known only from high altitudes (2500–3200 m) of the Sierra Famatina, a mountain range close to the Andes, in La Rioja Province of central-western Argentina (figs. 3G, 54).

Ecology

This species inhabits an area of grassland and shrub steppe that belongs to the Prepuna and Altoandina phytogeographic provinces (Cabrera and Willink, 1980; fig. 3G). It occurs in sympatry with another bothriurid, Bothriurus olaen Acosta, 1997 (Ojanguren Affilastro, 2005).

Orobothriurus grismadoi Ojanguren Affilastro et al., 2009

Figures 4B, 13G, 21E, 23E, 36D, E, 54

Orobothriurus grismadoi Ojanguren Affilastro et al., 2009: 29–35, figs. 1, 3–13, 15–17, 20–24, 35, tables 1, 2.

Type Material

ARGENTINA: Mendoza Province: Malargüe Department: Holotype ♂ (MACN-Ar 17986), Cerro Nevado, 35°35′45.06″S 68°30′24.12″W, 3130 m, 25.ii.2006, F. Fernández Campón and S. Lagos Silnik. Paratypes: same data, 6 ♂, 2 ♀ (MACN-Ar 17987), 2 ♂ (CDA), 1 juv. (IADIZA); Cerro Nevado, 35°36′04.08″S 68°30′44.28″W, 2900 m, 16.xi.2004, G. Flores, 1 ♀ (CDA), 2949 m, 7.i.2006, F. Fernández Campón and S. Lagos Silnik, 1 juv. (IADIZA); Cerro Nevado, 35°36′02.46″S 68°30′40.92″W, 2953 m, ii.2006, F. Fernández Campón and S. Lagos Silnik, 1 ♂ (AMNH), same data except 25.ii.2006, 5 ♂ (IADIZA), 1 ♂ (AMNH).

New Records

ARGENTINA: Mendoza Province: Malargüe Department: Cerro Nevado, road to antennas, SW face of mountain, 35°35′58.6″S 68°30′43.9″W, 2984 m, 23.iv.2011, C.I. Mattoni and M. Vivanco, UV sampling on rocky slopes, with small bushes and shrubs, no moon, 3°C, winds up to 45 km/h, wind chill factor −10°C, specimens walking or sitting on rocks, 5 ♂, 6 ♀, 4 juv. (LBRE), 4 juv. (AMNH), 1 juv. (AMNH [LP 10968]).

Diagnosis

Orobothriurus grismadoi is similar to O. alticola in pigmentation pattern, external morphology, and hemispermatophore. The two species may be separated by the shape of the hemispermatophore: in O. grismadoi, the apex is more slender and the angle it forms with the rest of the distal lamina more acute (fig. 36D, E) than in O. alticola (fig. 36I, J). Additionally, the apex comprises 42.70%–45.69% (n  =  7; mean  =  44.17%) of the lamina in O. grismadoi and 40.87–46.08% (n  =  20; mean  =  43.88%) in O. alticola. Males of O. grismadoi may also be recognized by the slender telson vesicle, with a concave dorsal surface; the vesicle is broader, with a flat dorsal surface, in males of O. alticola (fig. 23A, E). Orobothriurus grismadoi is also more densely pigmented, with tergite VII entirely pigmented (fig. 13G), compared to O. alticola, in which tergite VII exhibits paired lateral spots of pigmentation delimiting an unpigmented median stripe (fig. 13F).

Distribution

Orobothriurus grismadoi is the southernmost species of the genus. It is known only from the Cerro El Nevado, an isolated mountain range, 200 km east of the Andes, in central-western Mendoza Province, Argentina (fig. 54). The El Nevado range, with a maximum altitude of 3833 m, extends north to south between 34°S and 36°S, parallel to the Andes, and is separated from it by a plateau of 1800 m (Ojanguren Affilastro et al., 2009). El Nevado is an extinct, eroded, stratovolcano, unrelated to and younger than the Andes. The age of these volcanoes is estimated at between 2.5 and 0.01 MA (Caminos, 1999).

Ecology

Orobothriurus grismadoi was collected in a high-altitude shrub steppe at 2900–3130 m (fig. 3D), in an area belonging to the Altoandina and Puna phytogeographic provinces (Cabrera and Willink, 1980). Arthropod surveys conducted using pitfall traps in the summers of 2004 and 2006 did not collect other scorpion species. A recent expedition to the area confirmed that this species is the only scorpion present and active during the summer at this altitude on the El Nevado volcano. All specimens were collected at night in rocky areas (mostly basalt and pumice), and efforts to find the species on sandy slopes (with ash and pumice fragments) surrounding these rocky habitats were unsuccessful. Despite the harsh conditions, with very low temperatures and strong winds during the night, the species is apparently active on the surface from mid-November to the end of April.

Orobothriurus huascaran, n. sp.

Figures 4D, 11C, 12A, 14D, 19D, 21F, 22D, 23D, 24C, 37–Fig. 38.39, 40A–C, 52; table 3

Orobothriurus crassimanus: Polis, 1990: 252; Lourenço, 1997: 588 (part); Coddington and Colwell, 2001: 207; Lourenço, 2002: 400; 2003: 227; Lourenço and Qi, 2006: 290. [misidentification].

Orobothriurus paessleri: Lourenço and Dastych, 2001: 54 (part) [misidentification: 1 ♀, from Quebrada Yanganuco].

Orobothriurus sp. 2: Ochoa, 2004a: 50, 73, figs. 1, 2, 14, 21, table 1.

Type Material

PERU: Ancash Department: Huaraz Province: Holotype ♂ (MHNC), Cordillera Blanca, Huascarán National Park, Ishinca, 09°22′49″S 77°27′39″W, 4100 m, 19.vi.1998, J.A. Ochoa. Paratypes: Huaraz Province: Cordillera Blanca, Huascarán National Park, Ishinca, from 09°22′49″S 77°27′39″W, 4050 m, to 09°22′28″S 77°24′36″W, 4910 m, 18–19.vi.1998, J.A. Ochoa, 3 ♂, 3 ♀, 2 juv. ♂ (MHNC); Quebrada Querococha, Catac [09°32′S 77°24′W], 3900 m, vi.1981, W.R. Lourenço, 1 ♂, 2 ♀ (MHNG); Querococha, 3850 m, 16.iii.1994, M. Etonti, 1 ♂ (MRSN); Pastoruri [09°51′S 77°12′W], 4900 m, 18.iii.1994, M. Etonti, 1 ♂ (MRSN). Huaylas Province: Laguna Parón [08°59′43″S 77°40′41″W, 4160 m], 16.v.2010, R. Pinto da Rocha and D. Silva, 2 ♂ (MZSP), 1 ♂, 1 ♀ (MUSM), 2 juv. (AMNH [LP 10709, 10713]). Yungay Province: Llanganuco, 4000 m, 2.i.1976, O. F. Francke, 1 ♂, 5 ♀ (MACN-Ar 17889); same locality, 3600–3850 m, 15.iii.1994, M. Etonti, 2 ♀ (MRSN); same locality, 3810 m, vi.1997, F. Pribik, 1 ♀, 1 juv. (FKPC); Llanganuco, Huascarán National Park, 09°04′42″S 77°38′57″W, 3800–4000 m, 28–29.ix.1998, J.A. Ochoa, Polylepis forest, 1 ♂, 3 ♀ paratypes (MHNC), 1 ♂, 1 ♀ (AMNH), 1 ♂, 1 ♀ (LBRE); Chinancocha, Llanganuco, 09°04′18″S 77°38′38″W, 3844 m, 15.v.2010, R. Pinto da Rocha and D. Silva, 1 ♀ (MZSP).

Additional Material

PERU: Ancash Department: Carhuaz Province: Huascarán National Park, Aquilpo, Porganillo, 09°19′02″S 77°27′32″W, 3730 m, 15.vi.1998, J.A. Ochoa, 1 ♂, 3 ♀, 2 juv. ♂, 1 juv. ♀ (MHNC). Huaraz Province: Cordillera Blanca, Huascarán National Park, Ishinca ravine, 09°22′49″S 77°27′39″W, 4100 m, 19.vi.1998, J.A. Ochoa, 1 ♂, 3 juv. (MHNC). Huaylas Province: Laguna Parón [08°59′58″S 77°41′09″W, 4180 m], ii.1981, W.R. Lourenço, 1 juv. ♀ (MHNG). Recuay Province: Huama, 4200 m, 22.viii.1972, P. Brignoli, 6 ♀ [mislabeled “2 ♂, 4 ♀”], 14 first instar juv. (MHNG). Yungay Province: Llanganuco, 4300 m, ii.1981, W.R. Lourenço, 3 ♀ (MHNG), 4400–5500 m [see discussion on the altitude record in Scorpiones], ii.1981, W.R. Lourenço, 2 ♀ (MHNG); Llanganuco, Huascarán National Park, 09°04′42″S 77°38′57″W, 3800–4000 m, 28–29.ix.1998, J.A. Ochoa, Polylepis forest, 1 ♂, 2 ♀ (MHNC).

Etymology

The specific name is a noun in apposition derived from the Huascarán National Park, Ancash Department, central Peru, where the type locality of this species is situated. This area belongs to the Cordillera Blanca, the world's highest tropical mountain range.

Diagnosis

Orobothriurus huascaran may be distinguished from other species of the genus by the following combination of characters. The distal border of the apex of the hemispermatophore is slightly rounded, the internal lateral projection of the frontal crest vestigial, and the external lateral projection well developed in O. huascaran (fig. 40A–C) compared to other species in which the distal border of the apex is subtriangular, and both lateral projections of the frontal crest are well developed. Pedipalp chela trichobothrium Db is situated slightly closer to Dt than to Eb₃ in O. huascaran (fig., 39C), but equidistant between Dt and Eb₃ in other species. The VM and VL carinae are restricted to the posterior two-thirds of metasomal segment V in O. huascaran (fig. 21F), absent in O. wawita and complete in all other species. Orobothriurus huascaran is similar to four other Peruvian species, O. atiquipa, O. curvidigitus, O. paessleri, and O. quewerukana, in several respects. Pedipalp chela trichobothrium Et₃ is situated proximal to Est, and Esb situated between Eb₂ and Eb₃ in all these species, except O atiquita, in which Esb is situated dorsal to Eb₂. The hemispermatophore of these species exhibits an elongated frontal crest, with a strongly curved, S-shaped apex and a basal lobe with a well-developed terminal process. Orobothriurus huascaran may be distinguished by the presence of VM and VSM carinae on sternite VII and metasomal segment I (fig. 19D), which are absent in O. atiquipa, O. curvidigitus, O. paessleri, and O. quewerukana (figs. 18A, 19A, C). Additionally, the pedipalp chela movable finger of the male is straight in O. huascaran (fig. 39C) and curved in O. curvidigitus, O. paessleri, and O. quewerukana (fig. 26B, C, 43C).

Description

Based on holotype ♂ and paratypes. Measurements of holotype ♂ and paratype ♀ recorded in table 3.

Total length: ♂, 26.43–30.79 mm (n  =  9, mean  =  27.97 mm); ♀, 30.05–33.21 mm (n  =  10, mean  =  31.65 mm).

Color: General color yellowish with dark brown spots (fig. 4D). Carapace densely pigmented, especially laterally and posteriorly; anterior margin with narrow stripe (fig. 37A, C); median lateral surfaces densely pigmented; posterolateral surfaces with dense reticulate pigmentation; median ocular tubercle and lateral ocelli dark brown to black; posterior half of anteromedian longitudinal sulcus, median ocular tubercle and anterior half of posteromedian longitudinal sulcus densely pigmented; posterolateral sulci unpigmented. Chelicerae, dorsal surfaces with fine reticulate pigmentation, contiguous distally near base of movable finger; movable finger external surface pigmented. Pedipalp coxa and trochanter faintly pigmented; femur densely pigmented along margins with small unpigmented areas in proximal half; patella densely and irregularly pigmented; chela manus with faint longitudinal stripes along carinae, contiguous at base of movable finger. Legs, femur, and patella densely pigmented prolaterally; tibia and basitarsus faintly pigmented. Tergites I–VI each with two dark spots sublaterally along entire margin and pretergites, delimiting broad, unpigmented median stripe; VII with similar pattern and additional reticulate pigmentation (fig. 12A). Sternum, genital opercula and pectines unpigmented. Sternites III–VI unpigmented, VII with two dark stripes sublaterally (fig. 11C). Metasomal segments I–III, dorsal surfaces each with two subtriangular spots medially, separated by narrow unpigmented line on I, contiguous or separated on II, contiguous on III, becoming broader posteriorly and often connected to median pigmentation by fine reticulate pigmentation, with additional pigmentation along DL carinae; lateral surfaces densely pigmented, especially in anterior half, posterior half with reticulate pigmentation (figs. 11C, 12A); ventral surface with narrow VM stripe, not contiguous with lateral pigmentation, and with two dark VL stripes, becoming broader in posterior half and extending to LIM carinae. Segment IV as for segment III, but with dorsal surface more densely reticulate. Segment V, dorsal surface faintly pigmented along DL carinae; lateral surfaces with dense reticulate pigmentation; ventral surface with three dark stripes, two VL and one VM, becoming slightly broader in posterior half but not contiguous with lateral pigmentation (fig. 11C). Telson vesicle with narrow VM and two broad VL stripes, separated by two narrow unpigmented stripes; aculeus sclerotized, dark reddish-brown.

Chelicerae: Movable finger with two subdistal teeth.

Carapace: Surfaces smooth in anterior third, finely granular elsewhere. Anterior margin linear, without median notch (fig. 14D). Anteromedian longitudinal sulcus complete, well developed, more so in ♂; median ocular and posteromedian longitudinal sulci well developed; posterolateral sulci obsolete. Median ocular tubercle raised, situated anteromedially; median ocelli two ocular diameters apart.

Pedipalps: Femur, DI and VI carinae complete, finely granular; DE carina obsolete, smooth (fig. 38A); internal surface finely and sparsely granular medially; other intercarinal surfaces smooth. Patella, DI and VI carinae obsolete, finely granular (♂) or smooth (♀) (fig. 38B–D); internal surface with two prominent granules distally, the larger adjacent to trichobothrium i; other intercarinal surfaces smooth. Chela manus slender, fingers relatively elongated (fig. 39); length/width ratio: ♂, 3.46–4.14 (n  =  11, mean  =  3.76), ♀, 3.37–4.14 (n  =  20, mean  =  3.74); D, DS, DMA, DI, and VM carinae obsolete; intercarinal surfaces smooth; internal surface with acuminate apophysis (♂) or low bulge (♀) near articulation of movable finger (fig. 39A, D); fingers, dentate margins each with median denticle row and 4–5 pairs of internal and external accessory denticles.

Trichobothria: Femur with 3 trichobothria, patella with 19, chela with 27 (figs. 38, 39). Chela trichobothrium Et₃ situated proximal to Est; Esb situated between Eb₂ and Eb₃ (fig. 39).

Tergites: Tergites I–VI, surfaces finely granular. Tergite VII tetracarinate, paired DL carinae restricted to posterior two-thirds of segment, paired DSM carinae to posterior half; surfaces more coarsely and densely granular in posterior half.

Legs: Femur and patella, prolateral surfaces finely granular, retrolateral surfaces smooth. Femur, ventral carinae weakly developed; other carinae absent. Patella acarinate. Telotarsi, pro- and retroventral rows of spiniform macrosetae with following counts on leg I, 1/1; II, 2/2; III and IV, 3/3.

Pectines: Pectinal tooth count: ♂, 17–20 (n  =  30, mode  =  18); ♀, 14–17 (n  =  56, mode  =  15).

Sternites: Sternites III–VI, surfaces smooth; spiracles small, narrow. Sternite VII, surface smooth in anterior half, granular in posterior half; VL and VSM carinae well developed (♀, fig. 19D) or obsolete (indistinct from granulation) to absent (♂).

Metasoma: Segment I, DL carinae complete, more developed in ♀; usually one pair of DL macrosetae; ML carinae complete; two pairs of ML setae; LIM carinae obsolete, entirely smooth or with few granules in posterior third; VL and VSM carinae complete, obsolete, smooth (♂) or granular (♀); three pairs of VL and two or three pairs of VSM macrosetae (fig. 19D). Segments II and III, DL carinae complete; one pair of DL macrosetae; ML carinae restricted to posterior half (♂) or complete but less developed than on segment I (♀); two pairs of ML macrosetae; LIM carinae reduced to one or two granules posteriorly (♀) or absent (♂) on segment II, absent on III; VL carinae as for segment I, but less developed (♀) or absent (♂) on II, obsolete (♀) or absent (♂) on III; VSM carinae as for segment I, but less developed on II, obsolete (♀) or absent (♂) on III; three pairs of VL and VSM macrosetae. Segment IV, DL carinae complete; one or (usually) two pairs of DL macrosetae; ML carinae reduced to few small granules posteriorly; LIM carinae absent; VL and VSM carinae absent; three pairs of VL and four pairs of VSM macrosetae. Segment V, length/width ratio: ♂, 2.07–2.40 (n  =  5, mean  =  2.17), ♀, 1.71–2.02 (n  =  7, mean  =  1.87); DL carinae reduced to three or four prominent granules anteriorly; lateral intercarinal surfaces smooth (♂) or finely and sparsely granular medially near VL carinae (♀); VL carinae usually restricted to posterior two-thirds (but complete, weakly granular in anterior third in some ♀), with terminal granules more developed; VSM and VM carinae usually obscured by surface granulation in posterior two-thirds (fig. 21F), but VM carinae faintly evident in some ♀ (fig. 21D); four pairs of VL and VSM macrosetae; two pairs of macrosetae along posterior margin; ventral intercarinal surfaces densely granular in posterior two-thirds, more so in ♀, granules more acute in ♂.

Telson: Length/height ratio: ♂, 2.86–3.52 (n  =  11, mean  =  3.24); ♀, 2.61–3.39 (n  =  19, mean  =  3.04). Vesicle slightly elongated (♂, fig. 23D) or globose (♀, fig. 24C); dorsal surface smooth, flat, gland not apparent (♂); ventral surface smooth (♂) or granular in anterior third (♀). Aculeus short and curved.

Hemispermatophore: Apex shorter than frontal crest, distal border slightly rounded; distal crest curved like ventral margin. Frontal crest elongated; basal part oblique; distal part parallel to ventral margin of lamina, lateral projections slightly undulated and larger than basal part, ental lateral projection vestigial, ectal lateral projection complete. Basal lobe, terminal process extending to constriction of frontal crest (fig. 40A–C).

Distribution

Orobothriurus huascaran is endemic to the Ancash Department of central Peru (fig. 52). All known localities of this species occur in the Cordillera Blanca mountain range of the western Andes, the world's highest tropical mountain range (fig. 3F). This species has been collected from 3730 m (Porganillo, trekking route to Aquilpo mountain) to 4910 m (Ishinca ravine). It is present above 4050 m in the Ishinca area, but more abundant at 4100 m; only four specimens were found above 4500 m. The record of this species from Laguna Parón is currently the northernmost record for the genus Orobothriurus. The record from the Ishinca ravine is the world's highest record for a scorpion.

Ecology

The area inhabited by O. huascaran represents typical Puna vegetation with shrub steppe, grasses and small patches of Polylepis spp. forest (“queñua” in Quechua), situated at 3800–4200 m (fig. 3F). All specimens were found under stones during the day. No other scorpion species have been recorded in the area.

Remarks

The female specimen from Quebrada Yanganuco, listed under O. paessleri by Lourenço and Dastych (2001: 54), is probably conspecific with O. huascaran, n. sp., based on the known distribution of this species.

Orobothriurus paessleri (Kraepelin, 1911)

Figures 13A, B, 15A, 19B, 21B, 22B, 26B, 28C, E, 53

Bothriurus paessleri Kraepelin, 1911: 91, 98, 99, figs. 8, 9a, 9b; Mello-Leitão, 1931: 92; 1932: 34; 1934: 63; 1945: 181; Bücherl, 1953: 117; Aguilar and Meneses, 1970: 2; Maury, 1973: 110.

Bothriurus (Andibothriurus) paessleri: Bücherl et al., 1963: 216.

Orobothriurus paessleri: Maury, 1976: 19, 20, figs. 25–33, table I; 1981: 105; Francke, 1977: 75; Kovařík, 1998: 101; Acosta and Ochoa, 2000: 136, 143; Lowe and Fet, 2000: 36; Acosta and Ochoa, 2001: 205; Ochoa and Acosta, 2002a: 102, fig. 9; Ochoa, 2004a: 52, 55, 73, figs. 1, 2, 21, table 1; Ochoa, 2005: 55, figs. 7, 10, table 2.

Orobothriurus curvidigitus: Lourenço and Dastych, 2001: 54 [misidentification: 3 ♀ from Mejía].

Type Material

PERU: Arequipa Department: Islay Province: Syntypes: 1 ♂, 3 ♀ (ZMH), “Kataringo” [Catarindo], near Mollendo [17°00′58″S 72°02′03″W, 34 m?], 25.ii.1907, R. Paessler.

New Records

PERU: Arequipa Department: Camaná Province: S of Camaná, 7.viii.1977, L. Peña, 1 ♀ (MACN-Ar). Islay Province: Lomas de Mejía [17°02′S 71°51′W, 530 m], 1–3.ii.2000, J.A. Ochoa, 1 ♂, 2 ♀, 3 juv. (MHNC), 1 ♂, 1 ♀ (CDA 168); Lomas de Mejía, Challascapa [17°02′S 71°51′W, 550 m], 3.iii.2004, R. Gutierrez and J.A. Ochoa, 2 juv. (AMNH [LP 3057]); near Mejía, 17°07′S 71°55′W, 350–500 m, 15.i.1956, Koepcke, Park loma, Kp 1350, 1 ♂, 1 subad. ♂, 2 juv. ♂, 2 juv. ♀ (ZMH A18/101) [misidentified by W.R. Lourenço as “O. curvidigitus”]; Lomas de Yuta, Matarani, 16°57′16″S 72°04′10″W, 570 m, 12.iii.1998, E. Ponce and J.A. Ochoa, 1 ♀ (MHNC).

Diagnosis

Orobothriurus paessleri is most closely related to O. curvidigitus, O. quewerukana and O. tamarugal (fig. 5), from which it may be separated as follows. Metasomal segment V is markedly shorter, its length less than twice its width, in the female of O. paessleri (fig. 22B), than in the females of other species, in which its length is greater than or equal to twice its width. Orobothriurus paeslleri possesses three pairs of VSM macrosetae on metasomal segment I (fig. 19B), whereas the other species possess two (fig. 19A). The pedipalp chela movable finger of the male is slightly curved in O. paessleri (fig. 26B), but strongly curved in the other species (figs. 26C, 43C, 51A). Orobothriurus paessleri may be further separated from O. curvidigitus and O. quewerukana by the shape of the lamina of the hemispermatophore (figs. 28B, C, 40E), the frontal crest of which is shorter and less developed in O. paessleri (fig. 28E) than in O. curvidigitus and O. quewerukana (fig. 28D). Additionally, the ventral surfaces of metasomal segments I–III are finely granular in O. paessleri (fig. 19B), whereas only the ventral surfaces of segment I in males are finely granular in O. curvidigitus and O. quewerukana.

Distribution

Orobothriurus paessleri appears to be endemic to the area near Mejía and Yuta (Matarani) in the Arequipa Department of southern Peru (fig. 53). A single record from Toquepala (17°14′S 70°36′W, 3000 m) in the Tacna Department (Maury, 1976), probably based on a misidentification, requires confirmation.

Ecology

As with O. atiquipa, this species has only been collected in Lomas biotopes, isolated patches of green vegetation on hillsides below 1000 m, sustained by coastal fog, and surrounded by hyperarid desert. The Lomas vegetation near Mejía and Yuta is dominated by herbaceous shrubs, e.g., Nolana L.f., Palaua Cav., Portulaca L., Weberbauerella Ulbr., and the tree Caesalpinia spinosa (Molina) Kuntze (Péfaur, 1981; Dávila Flores, 1982).

Remarks

Specimens of O. paessleri from Mejía deposited in the ZMH collection were examined, and found to be misidentified as O. curvidigitus by Lourenço and Dastych (2001: 54). Currently two Orobothriurus species have been recorded from Lomas formations: O. atiquipa and O. paessleri. In addition, we examined a single female deposited at MHNC from Lomas de Sama, southern Peru. This specimen is poorly preserved and difficult to identify with certainty.

Orobothriurus parvus Maury, 1976

Figures 4C, 9, 12B, 17A, 22E, 26A, 27B, 52

Fig. 9.

Orobothriurus parvus Maury, 1976, sternum and pectines. A. ♂ (MHNC). B. ♀ (MHNC). Scale bars  =  1 mm.

Fig. 10.

Orobothriurus Maury, 1976. Sinistral hemispermatophore showing morphology and measurements. A. Ental aspect. B. Dorsal aspect. C. Ectal aspect. Annotations: basal lobe (bl); capsular concavity (cc); constriction (cns); distal crest (Dc); external lobe (el); external lateral projection (elp); frontal crest (Fc); internal lobe (il); internal lateral projection (ilp); incision (in); Lamina (Lam); pedicel (Ped); terminal process (tp); trunk (Tr); tortuous stem (ts); apex length (x); length of basal portion of lamina (y).

Orobothriurus parvus Maury, 1976: 17, 21–23, figs. 45–54, table I; Galiano and Maury, 1979: 327; Maury, 1980: 338, fig. 12; Kovařík, 1998: 101; Acosta and Ochoa, 2000: 136, 143; Lowe and Fet, 2000: 36; Acosta and Ochoa, 2001: 205; Lourenço and Dastych, 2001: 54; Ochoa 2004a: 52, 55, 73, table 1, figs. 1, 2, 21.

Bothriurus borellianus Mello-Leitão, 1934: Bücherl, 1959b: 273 [misidentification].

Bothriurus chilensis (Molina, 1782): Aguilar and Meneses, 1970: 3 [misidentification].

Bothriurus (Andibothriurus) peruvianus Mello-Leitão, 1948: Francke, 1974: 217, 218, figs. 1–5, table 1 [misidentification].

Type Material

PERU: Junín Department: Yauli Province: Holotype ♂ (MACN-Ar 6837), Abra de Anticona, Oroya [11°36′31″S 76°11′48″W], 4750 m, 28.xi.1974, A. Martínez. Paratypes: same data, 1 ♂, 1 juv. (MACN-Ar 6838); Acaya, Río Mantaro [11°49′S 75°36′W], 3480 m, 1.iii.1957, W. Weyrauch, 1 ♀ (IML 388).

New Records

PERU: Junín Department: Jauja Province: Acolla, near Jauja, [11°44′17″S 75°32′35″W, 3400 m], 8.viii.1953, F. Blancas, 4 ♀, 12 juv. (MUSM), ii.1965, F. Blancas, 2 ♀ (MUSM). Junín Province: Ondores [11°04′06″S 76°09′21″W, 4300 m], 6.v.2000, E. Ponce, 2 ♂, 7 ♀ (MHNC); Quebrada Antacocha, between Junín and Carhuamayo [11°03′S 76°00′W, 4090 m], viii.1952, F. Blancas, 1 juv. (MUSM). Tarma Province: Tarma, 2 km E [11°24′58″S 75°39′59″W, 3400 m], 4.i.1973, N.F. Hadley and O.F. Francke, 1 ♀ (AMNH). Yauli Province: Casaragra?, xi.1947, H. de Madedo, 1 ♀ (MUSM); Cerro Quinan, Tupac Amaru, Yauli, La Oroya [11°32′S 75°54′W, 3968 m], 26.viii.2005, W. Paredes, 1 ♀ (MHNC), 1 juv. (AMNH [LP 6257B]).

Diagnosis

Orobothriurus parvus is most closely related to O. wawita (fig. 5). Both species share similar hemispermatophore morphology, including an elongated lamina apex and a short frontal crest (fig. 27B, C). The reticulate pigmentation along the metasomal DL carinae (figs. 12B, 13C) and the macrosetal counts on the ventral surface of the metasoma are also similar. The following characters separate the two species. The pedipalp chela manus of the male O. parvus possesses an apophysis on the internal surface, and the fixed finger is slightly curved, creating a small gap with the movable finger when the fingers are closed (fig. 26A), whereas the apophysis is absent and the fixed finger straight, such that no gap is evident when the fingers are closed, in the male of O. wawita (fig. 25B, C). The VL and VM carinae of metasomal segment V are complete in O. parvus (fig. 22E), but absent (♂) or restricted to the distal third of the segment (♀) in O. wawita (figs. 21C, 22F). Metasomal segments II–V exhibit a distinct VM pigmentation stripe in O. parvus that is absent in O. wawita (fig. 13D). The two species also differ in the shape of the ventral margin of the hemispermatophore apex, which is inclined to the dorsal surface and curved distally in O. parvus (fig. 27B), but straight in O. wawita (fig. 27C).

Distribution

Orobothriurus parvus is endemic to the central Andes, recorded from 3400–4750 m, in the Junín and Lima departments, Peru (figs. 2E, 52).

Ecology

The area inhabited by O. parvus corresponds to the Puna ecoregion or Puna biogeographical province (Brack, 1986; Ceballos Bendezu, 1976). The typical Puna vegetation of this area comprises shrub, steppe, and grasses (fig. 2E). No other scorpion species have been recorded in sympatry with O. parvus.

Remarks

Orobothriurus parvus was misidentified in several papers as Bothriurus borellianus, Bothriurus chilensis or Bothriurus (Andibothriurus) peruvianus (Bücherl, 1959b; Aguilar and Meneses, 1970; Francke, 1974). A record from Colcabamba (Lourenço and Dastych, 2001: 54), based on a single female, requires confirmation.

Orobothriurus quewerukana, n. sp.

Figures 15C, 19C, 20D, 23F, 24D, 40D, E, 41–Fig. 42.43, 53; table 3

Orobothriurus dumayi: Maury, 1976: 21, figs. 40–44; Acosta and Ochoa, 2000: 136, 143. [misidentification].

Type Material

CHILE: Región I (Tarapacá): Parinacota Province: Holotype ♂ (AMNH), Putre, 6 km W, 18°13′26.8″S 69°32′47.1″W, 3732 m, 16.i.2005, A. Ojanguren Affilastro, C.I. Mattoni and J.A. Ochoa, UV and under stones. Paratypes: CHILE: Región I (Tarapacá): Iquique Province: Termas de Mamiña, 8.viii.1967, J.C. Ortiz, 4 juv. (MACN-Ar 6846); Mamiña, 20°04′33.6″S 69°12′05.4″W, 2822 m, 19.i.2005, A. Ojanguren Affilastro, C.I. Mattoni and J.A. Ochoa, UV and under stones, small wet valley with old terrace cultivation, 2 ♂, 2 ♀, 14 juv. (LBRE), 2 ♂, 2 ♀, 8 juv. (MACN-Ar), 2 ♂, 1 ♀, 1 subad. ♀ (MHNC), 1 ♂, 1 ♀, 6 juv. (MZUC). Parinacota Province: same data as holotype, 2 ♂, 2 ♀, 6 juv. (MACN-Ar), 2 ♂, 1 ♀, 3 juv. (LBRE), 1 ♂, 1 ♀, 3 juv. (MHNC), 1 subad. ♂ (AMNH). PERU: Tacna Department: Tacna Province: Palca, 17°46′45″S 69°57′04″W, 3230 m, 26.ii.2000, J.A. Ochoa, 1 ♀ (AMNH), 1 ♂, 2 ♀ (MHNC).

Additional Material

CHILE: Región I (Tarapacá): Iquique Province: Mamiña, 20°04′33.6″S 69°12′05.4″W, 2822 m, 19.i.2005, A. Ojanguren Affilastro, C.I. Mattoni, and J.A. Ochoa, UV and under stones, small wet valley with old terrace cultivation, 1 subad. ♂, 1 subad. ♀, 2 juv. ♂, 2 juv. ♀ (AMNH [LP 4307]), 5 juv. (AMNH), 5 juv. (MHNC). Parinacota Province: Putre, 6 km W, 18°13′26.8″S 69°32′47.1″W, 3732 m, 16.i.2005, A. Ojanguren Affilastro, C.I. Mattoni and J.A. Ochoa, UV and under stones, 1 subad. ♂, 4 juv. ♂, 1 juv. ♀ (AMNH [LP 4306]), 4 juv. (LBRE), 3 juv. (MHNC).

Etymology

The specific name is a noun in apposition composed of two Quechua words, quewe, meaning “curved,” and rukana, meaning “finger.” It refers the strong curvature of the movable finger of the pedipalp chela in the adult male.

Diagnosis

Orobothriurus quewerukana is closely related to O. curvidigitus and O. tamarugal (fig. 5), with which it shares the following characters: anterior margin of carapace with weak median projection (epistome; figs. 14C, 15C, D); pedipalp chela shape similar, with movable finger strongly curved in male (figs. 26C, 43C, 51A); pedipalp femur and patella more elongated in male than female; sternite VII with VSM carinae absent or obsolete (fig. 19A, C); metasomal segments I–IV with DL and ML carinae complete, granular, and VL carinae obsolete, smooth. Orobothriurus quewerukana may be separated from O. tamarugal by the shape of the lamina of the hemispermatophore (figs. 40E, 41D). The pedicel of the apex is narrower, and the frontal crest more developed and elongated in O. quewerukana; the ventral border of the distal portion of the apex is slightly curved in O. quewerukana, but straight in O. tamarugal; and one or two small folds, observed in the distal crest of 80% of the specimens of O. quewerukana, are absent in the distal crest of O. tamarugal. The hemispermatophore of O. quewerukana is similar to that of O. curvidigitus, except for the slightly shorter frontal crest and the absence of folds in the distal crest (figs. 28B, 40E). The two species may be distinguished by the shape and granulation of the male telson vesicle, which is slightly deeper, with a length/height ratio of 3.59–4.00 (mean  =  3.76), and granulation on the entire ventral surface (fig. 23F), in O. quewerukana, compared with O. curvidigitus, in which the length/height ratio is 3.72–4.23 (mean  =  3.96) and granulation is present only on the anterior third of the ventral surface. The male pedipalp chela carinae are finely and densely granular in O. quewerukana and O. tamarugal (figs. 43D, E, 51B, C), but finely and sparsely granular to smooth in O. curvidigitus. These species may also be separated by the pigmentation pattern. Orobothriurus curvidigitus exhibits well-developed pigmentation on the carapace, tergites, metasoma, and pedipalps (figs. 11B, 12C, 13E), compared to O. quewerukana in which the pigmentation, especially that of the carapace and tergites, is faint or barely discernible (fig. 41) and O. tamarugal, in which it is absent (fig. 48).

Fig. 11.

Orobothriurus Maury, 1976, sternite VII and metasomal segments I–V, ventral aspect, showing pigmentation pattern. A. Orobothriurus ampay Ochoa and Acosta, 2003, allotype ♀ (MUSM). B. Orobothriurus curvidigitus (Kraepelin, 1911), ♀ (MHNC). C. Orobothriurus huascaran, n. sp., paratype ♀ (MHNC). D. Orobothriurus atiquipa Ochoa and Acosta, 2002, ♂ paratype (MHNC). Scale bars  =  1 mm.

Fig. 12.

Orobothriurus Maury, 1976, tergite VII (A, C, D only) and metasomal segments I–IV, dorsal aspect, showing pigmentation pattern. A. Orobothriurus huascaran, n. sp., paratype ♀ (MHNC). B. Orobothriurus parvus Maury, 1976, ♀ (MHNC). C. Orobothriurus curvidigitus (Kraepelin, 1911), ♀ (MHNC). D. Orobothriurus ampay Ochoa and Acosta, 2003, allotype ♀ (MUSM). Scale bars  =  1 mm.

Fig. 13.

Orobothriurus Maury, 1976, tergites, sternites and metasomal segments I–V, showing pigmentation pattern. A, B. Orobothriurus paessleri (Kraepelin, 1911), ♀ (MHNC), metasomal segments I–V, ventral aspect (A), tergite VII and metasomal segments I–IV, dorsal aspect (B). C, D. Orobothriurus wawita Acosta and Ochoa, 2000, ♀ (MHNC), tergite VII and metasomal segments I and II, dorsal aspect (C), sternite VII and metasomal segments I–V, ventral aspect (D). E. Orobothriurus curvidigitus (Kraepelin, 1911), ♀ (MHNC), tergite V, dorsal aspect. F. Orobothriurus alticola (Pocock, 1899), ♀ (MACN-Ar), tergites VI and VII, dorsal aspect. G. Orobothriurus grismadoi Ojanguren-Affilastro et al., 2009, tergites VI and VII, dorsal aspect. Scale bars  =  1 mm.

Description

Based on holotype ♂ and paratypes. Measurements of holotype ♂ and paratype ♀ recorded in table 3.

Total length: ♂, 38.09–47.00 mm (n  =  8, mean  =  41.59 mm); ♀, 37.20–51.50 mm (n  =  8, mean  =  43.35).

Color: General color yellowish with light brown spots on carapace, tergites, and pedipalps; metasomal segments, ventral pigmentation slightly darker, especially in specimens from Palca (fig. 41). Carapace faintly spotted, especially laterally and posteriorly; anterior margin with two spots submedially (fig. 41A, C); posterolateral surfaces faintly pigmented; median ocular tubercle and lateral ocelli dark brown to black; anterior third of anteromedian longitudinal sulcus pigmented along borders; posteromedian longitudinal sulcus faintly pigmented or unpigmented; posterolateral sulci unpigmented. Chelicerae unpigmented. Pedipalp coxa, trochanter and femur usually unpigmented or faintly pigmented along DI and DE carinae (some juveniles); patella external and, usually, internal surfaces faintly spotted; chela unpigmented. Legs, only ventral margin of femur, and dorsal and ventral margins of patella pigmented (unpigmented in some adults). Tergites I–VI each with two faint spots sublaterally, extending along anterior two-thirds, becoming broader near anterior margins, delimiting broad, unpigmented median stripe (fig. 41A, C); VII unpigmented (adults) or with two faint spots sublaterally (juveniles). Sternum, genital opercula, pectines and sternites III–VII unpigmented (fig. 41B, D). Metasomal segments I–III, dorsal surfaces each with two faint subtriangular spots medially, separated by narrow unpigmented line (juveniles) or unpigmented (adults, fig. 41); lateral surfaces unpigmented, except near VL carinae; ventral surface with narrow VM stripe on III (absent on I and II), not contiguous with lateral pigmentation, and two VL stripes on I–III (occasionally faint or absent on I) becoming broader in posterior third; specimens from Mamiña usually less pigmented: VL stripe absent on segments I and II; VM stripe often absent on III. Segment IV, dorsal surface unpigmented; lateral and ventral surfaces as for segment III. Segment V, dorsal surface unpigmented or faintly pigmented posteriorly; lateral surfaces pigmented in distal third; ventral surface with three stripes, two VL and one VM, becoming slightly broader in posterior half but not joining with lateral pigmentation. Telson vesicle unpigmented or with lateral surfaces faintly pigmented; aculeus sclerotized, dark reddish brown.

Chelicerae: Movable finger with two subdistal teeth.

Carapace: Surfaces finely granular, more coarsely so along anterior margin, anteromedian longitudinal and median ocular sulci (♂) or smooth in anterior third, finely granular elsewhere (♀). Anterior margin with weak median projection (epistome; fig. 14C). Anteromedian longitudinal sulcus complete, moderately developed; median ocular and posteromedian longitudinal sulci well developed; posterolateral sulci obsolete. Median ocular tubercle raised, situated anteromedially; median ocelli two ocular diameters apart.

Fig. 14.

Orobothriurus Maury, 1976, carapace, dorsal aspect. A. Orobothriurus calchaqui, n. sp., paratype ♂ (MACN-Ar). B. Orobothriurus compagnuccii, n. sp., paratype ♂ (MACN-Ar). C. Orobothriurus curvidigitus, ♂ (MHNC). D. Orobothriurus huascaran, n. sp., paratype ♂ (MACN-Ar). Scale bars  =  1 mm.

Pedipalps: Femur, DI, DE, and VI carinae complete (fig. 42A), DI carina more strongly developed than DE and VI carinae; VM carina vestigial, finely granular proximally; EM carina complete, granular (♂) or obsolete, finely granular (♀, juveniles); dorsal surface finely granular; internal surface sparsely granular medially. Patella, DI, VI, and VE carinae complete, granular (fig. 42B–D); DE carina complete, less developed than DI carina (♂) or obsolete (♀); EM carina vestigial, reduced to few small granules medially (♂) or absent (♀); DPP carina comprising prominent granule and additional small granules proximally; VPP carina vestigial, comprising two or three small granules proximally; internal surface with prominent granule adjacent to trichobothrium i near DI carina. Chela manus slightly rounded, fingers relatively elongated (fig. 43); length/width ratio: ♂, 3.42–4.19, (n  =  9, mean  =  3.71), ♀, 3.54–4.22 (n  =  6, mean  =  3.87); length/height ratio: ♂, 2.95–3.32, (n  =  9, mean  =  3.07), ♀, 2.87–3.34 (n  =  6, mean  =  3.16); most carinae obsolete, finely granular (less so in ♀) or absent; VM carina restricted to proximal two-thirds of manus, VI carina to proximal third; DS, DMA, and DI carinae complete, finely and densely granular, especially near base of fixed finger (♂) or smooth (♀); D carina smooth; internal surface with acuminate apophysis (♂) or low bulge (♀) near articulation of movable finger (fig. 43A, D); movable finger (♂) strongly curved, creating small gap with fixed finger when fingers closed (fig. 43C); fingers, dentate margins each with median denticle row and 4–5 pairs of internal and external accessory denticles.

Trichobothria: Femur with 3 trichobothria, patella with 19, chela with 27 (figs. 42, 43). Chela trichobothrium Et₃ situated proximal to Est (in same axis as Est in one specimen); Esb situated dorsal to Eb₂.

Tergites: Tergites I–VI, surfaces finely granular, more coarsely and densely so in ♂. Tergite VII tetracarinate, paired DL carinae restricted to posterior two-thirds of segment, paired DSM carinae to posterior half; surfaces more coarsely and densely granular in posterior half.

Legs: Femur and patella, prolateral surfaces finely granular, retrolateral surfaces smooth. Femur, VI and EM carinae complete in ♂, less developed in ♀; DE and DI carinae vestigial. Patella acarinate. Telotarsi, pro- and retroventral rows of spiniform macrosetae with following counts on leg I, 1/1; II, 2/2; III and IV, 3/3.

Pectines: Pectinal tooth count: ♂, 20–24 (n  =  34, mode  =  22); ♀, 19–22 (n  =  24, mode  =  20).

Sternites: Sternites III–VI, surfaces smooth (♀) or matt in posterior three-quarters (♂); spiracles small, narrow. Sternite VII, surface smooth (♀) or finely granular (♂) (fig. 19C), acarinate.

Metasoma: Segment I, DL carinae complete; ML carinae complete, posterior granules more prominent than adjacent granulation; one pair of ML macrosetae; LIM carinae reduced to few granules in posterior third; VL carinae obsolete; VSM carinae absent (fig. 19C); two pairs of VL and VSM macrosetae. Segments I and II, ventral intercarinal surfaces finely granular (♂). Segments II–IV, surfaces between DL and ML carinae finely and sparsely granular, granulation decreasing posteriorly. Segments II and III, DL carinae complete; one pair of DL macrosetae in 14% and 50% of specimens, respectively; ML carinae as for segment I; LIM carinae reduced to few granules posteriorly on segment II, restricted to posterior third (♂) or absent (♀) on segment III; VL carinae as for segment I but less developed; VSM carinae absent; three pairs of VL and VSM macrosetae. Segment IV, DL carinae complete, granular; one pair of DL macrosetae; ML carinae complete, granular; LIM carinae absent; VL carinae as for segment III (absent in some subadult and juvenile specimens); VSM carinae absent; usually three pairs of VL and VSM macrosetae. Segment V, length/width ratio: ♂, 2.18–2.44 (n  =  6, mean  =  2.31), ♀, 2.01–2.20 (n  =  4, mean  =  2.10); DL carinae complete, granular; lateral surfaces sparsely granular near DL carinae (fig. 20D); VL carinae complete; VSM carinae reduced to few granules medially; VM carina complete, obscured by surface granulation in posterior half; usually three pairs of VL and VSM macrosetae; two pairs of macrosetae along posterior margin; ventral intercarinal surfaces, granulation decreasing anteriorly.

Telson: Length/height ratio: ♂, 3.59–4.00 (n  =  9, mean  =  3.76); ♀, 2.90–3.17 (n  =  6, mean  =  3.03). Vesicle slightly elongated (♂, fig. 23F) or globose (♀, fig. 24D); dorsal surface smooth, flat or slightly concave, gland not apparent (♂); ventral surface sparsely granular. Aculeus short and curved.

Hemispermatophore: Apex approximately half the length of lamina, distal border subtriangular and slightly curved to ventral margin; distal crest curved like ventral margin, with one or two small folds in 80% of specimens. Frontal crest elongated; basal part oblique; distal part parallel to ventral margin of lamina, lateral projections complete, slightly undulated, and larger than basal part. Basal lobe, terminal process extending to constriction of frontal crest (fig. 40D, E).

Distribution

Orobothriurus quewerukana is endemic to the western slopes of the Andes, at 2822–3732 m, in the Tacna Department of southern Peru and the Tarapacá Region of northern Chile (fig. 53).

Ecology

The area inhabited by O. quewerukana belongs to the Serrania Esteparia ecoregion (Brack, 1986) or Estepa Altoandina botanical region (Gajardo, 1993), the vegetation of which is characterised by abundant cacti and some shrubs, like Baccharis L., Lupinus L., and Senecio L. Specimens were collected under stones during the day and with UV light detection at night. Orobothriurus quewerukana is sympatric with another bothriurid, Brachistosternus quiscapata Ochoa and Acosta, 2002, at Palca and Putre.

Orobothriurus ramirezi, n. sp.

Figures 16A, B, 18C, 20C, F, 21G, 24E, 44–Fig. 45.46, 47A, B, 54; table 3

Type Material

CHILE: Region IV (Coquimbo): Elqui Province: Holotype ♂ (MACN-Ar), Paso del Agua Negra, between Juntas and international border, 30°12′11.5″S 69°54′57.5″W, 3734 m, 27.i.2005, C.I. Mattoni and A. Ojanguren Affilastro, UV detection, specimens walking actively near a “vega” (small stream), extremely wet. Paratypes: same data, 4 ♂, 1 ♀ (AMNH), 2 ♂, 1 ♀ (LBRE), 1 ♂, 1 ♀ (MACN-Ar), 1 ♂ (MHNC); same data, except 30°16′14.5″S 69°58′27.9″W, 3295 m, 1 ♂, 1 ♀, 5 juv. (AMNH), 1 ♀, 2 subad. ♂ (AMNH [LP 4305]), 1 ♂, 1 ♀, 3 juv. (LBRE), 1 ♂, 1 ♀, 2 juv. (MACN-Ar), 1 ♀, 2 juv. (MHNC), 1 ♂, 2 juv. (MZUC).

Additional Material

CHILE: Region IV (Coquimbo): Elqui Province: Mine El Indio, Cancha Sky base camp [29°51′38″S 70°02′12″W, 3225 m], v.1996, H. Vásquez C., pitfall trap, 1 ♀, 1 juv. (ULS); Mine El Indio, Mangueras, ca. 4000 m, 1 juv. (AMNH), 1 juv. (LBRE); Mine El Indio, Pastos Largos [29°49′S 70°03′W, 3600 m], i.1995, H. Vásquez C., pitfall trap, 1 ♀, 1 juv. (ULS), 6.xi.2003, L. Prendini, C.I. Mattoni and J.A. Ochoa, UV detection, very cold and windy, full moon, alpine vegetation on scree slope near small stream, specimen sitting in open ground, 1 subad. (AMNH [LP 2402]); Mine El Indio, Quebrada El Negro [29°47′06″S 70°00′00″W, 3808 m], 6.xi.2003, C.I. Mattoni, L. Prendini, and J.A. Ochoa, UV detection and diurnal rock rolling, puna vegetation, low bushes and grass tufts, hard, rocky ground with bare patches, stones in places, at night, very cold and windy, full moon, specimens on rocky slope (juv. under stone), 1 ♀, 1 juv. (AMNH [LP 2401]); Mine El Indio, Tambo [29°48′S 69°58′W, 4030 m], ii.1997, H. Vásquez C., pitfall trap, 1 ♂, 1 ♀, 1 juv. (ULS); Paso del Agua Negra, between Juntas and international border, 30°12′11.5″S 69°54′57.5″W, 3350 m, 6.iii.2006, A. Ojanguren Affilastro, L. Compagnucci, and C. Cuezzo, UV detection, 1 ♂, 1 ♀, 1 subad. ♀, 1 juv. (MACN-Ar).

Etymology

This species is named in honor of the Argentine arachnologist Martín J. Ramírez (MACN) for his contributions on the biodiversity and systematics of South American arachnids.

Diagnosis

Orobothriurus ramirezi is closely related to O. alticola (fig. 5), based on similar pigmentation pattern, metasomal carination, and hemispermatophore morphology. The two species may be separated by the distal crest of the hemispermatophore, which is comparatively more developed, occupying almost the entire apex of the lamina, in O. ramirezi, than in O. alticola, in which it occupies only half the apex (figs. 36I, J, 47A, B). The telson vesicle of O. ramirezi is more rounded, especially in females (telson length/width ratio: ♂, 2.65–2.94, mean  =  2.78; ♀, 2.13–2.37, mean  =  2.27; figs. 23G, 24E) than that of O. alticola (telson length/width ratio: ♂, 2.87–3.21, mean  =  3.07; ♀, 2.44–2.55, mean  =  2.49; fig. 23A). The dorsal surface of the femur and ventral surface of the metasoma are more pigmented, with broader stripes, in O. ramirezi than O. alticola. The metasomal carinae, especially the VL and VSM carinae of segments I and V, and the DL carinae of segment V, are more developed in males of O. ramirezi than O. alticola (fig. 20A, C).

Description

Based on holotype ♂ and paratypes. Measurements of holotype ♂ and paratype ♀ recorded in table 3.

Total length: ♂, 27.3–36.0 mm (n  =  13, mean  =  31.6 mm); ♀, up to 37.8 mm.

Color: General color yellowish with dark brown spots. Carapace, anterior margin unpigmented (fig. 44A, C); lateral margins with two spots laterally and two smaller spots posterolaterally, intermediate area with faint reticulate pigmentation; median ocular tubercle and lateral ocelli dark brown or black; posterior half of anteromedian longitudinal sulcus, median ocular tubercle, and posteromedian longitudinal sulcus pigmented. Chelicerae unpigmented. Pedipalp coxa unpigmented; trochanter, dorsal and lateral surfaces faintly pigmented; femur, ventral surface unpigmented, dorsal surface pigmented, except for oval unpigmented area, occupying up to two-thirds of segment distally; internal and external surfaces unpigmented, except for faint stripes along VL and VI carinae; chela manus with six complete longitudinal stripes along carinae, joining at base of movable finger; fingers unpigmented except basally. Legs, coxa and trochanter unpigmented; femur, dorsal and prolateral surfaces sparsely spotted, spots becoming larger near articulation with patella; patella, prolateral, dorsal and ventral surfaces sparsely pigmented; basitarsi and telotarsi, dorsal and prolateral surfaces faintly spotted. Tergites I–VI each with two faint spots sublaterally, reaching pretergites on III–VI and delimiting unpigmented median stripe; VII with two small, faint, spots of pigmentation submedially (fig. 44A, C). Sternum, genital opercula, pectines, and sternites III–VI unpigmented; sternite VII with two, often faint, spots sublaterally. Metasomal segment I, dorsal surfaces each with faint subtriangular spot medially; lateral surfaces pigmented between ML and LIM carinae; ventral surface with two narrow VL stripes becoming slightly broader in posterior half (fig. 44). Segments II–IV, dorsal surfaces each with small subtriangular spot medially; lateral surfaces densely pigmented below ML carinae; ventral surfaces each with three separate, narrow stripes (one VM and two VL), becoming broader in posterior half. Segment V, dorsal surface unpigmented; lateral surfaces with weak reticulate pigmentation; ventral surface with three separate stripes (one VM and two VL), becoming broader posteriorly. Telson vesicle, dorsal surface unpigmented (♀) or with faint longitudinal stripe (♂); ventral surface with three broad dark stripes (two VL and one VM), separated by two narrow unpigmented stripes; aculeus sclerotized, dark reddish brown.

Chelicerae: Movable finger with two subdistal teeth.

Carapace: Median surfaces smooth; lateral margins finely granular, less so in ♀. Anterior margin linear, without median notch (fig. 16A, B). Anteromedian longitudinal sulcus obsolete; median ocular, posteromedian longitudinal and posterolateral sulci well developed. Median ocular tubercle raised, situated anteromedially; median ocelli two ocular diameters apart.

Pedipalps: Femur, DI and VI carinae complete, finely granular; DE carina obsolete, smooth (fig. 38A); internal surface finely and sparsely granular medially; other intercarinal surfaces smooth. Patella, DI and VI carinae obsolete, finely granular (♂) or smooth (♀) (fig. 38B–D); internal surface with two prominent granules distally, the larger adjacent to trichobothrium i; other intercarinal surfaces smooth.

Pedipalps: Femur, DI and VI carinae complete, granular (fig. 45A); DE carina complete, granular in distal third to distal half of segment, other carinae smooth; ventral surface sparsely granular, more densely granular in proximal half. Patella, DI and VI carinae complete, granular, less so in ♀ (fig. 45B–D); DPP carina comprising small granules proximally; VPP carina absent; internal surface coarsely granular with prominent granule adjacent to trichobothrium i near DI carinae; other surfaces smooth. Chela manus slender, fingers relatively elongated (fig. 46); length/width ratio: ♂, 3.58–4.14 (n  =  12; mean  =  3.84), ♀, 3.31–3.9 (n  =  6, mean  =  3.62); length/height ratio: ♂, 2.94–3.75 (n  =  12; mean  =  3.3), ♀, 3.06–3.4 (n  =  6, mean  =  3.29); DMA, DI, and VM carinae obsolete, other carinae absent; intercarinal surfaces smooth; internal surface with acuminate apophysis (♂) or low bulge (♀) near articulation of movable finger (fig. 46A, D); fingers, dentate margins each with median denticle row and 4–5 pairs of internal and external accessory denticles.

Trichobothria: Femur with 3 trichobothria, patella with 19, chela with 27. Chela trichobothrium Et₃ situated proximal to Est (fig. 46).

Tergites: Tergites I–VI, surfaces finely granular (♂) or smooth (♀). Tergite VII tetracarinate, paired DL carinae restricted to posterior two-thirds of segment, paired DSM carinae to posterior half; intercarinal surfaces coarsely granular, other surfaces finely granular.

Legs: Femur and patella, prolateral surfaces finely granular, retroventral surfaces smooth. Femur, ventral carinae weakly developed; other carinae absent. Patella acarinate. Telotarsi, pro- and retroventral rows of spiniform macrosetae with following counts on leg I, 1/1; II, 2/2; III and IV, 3/3.

Pectines: Pectinal tooth count: ♂, 21–26 (n  =  33, median  =  23); ♀, 15–20 (n  =  34, median  =  18).

Sternites: Sternites III–VI, surfaces smooth (♀) or finely corrugated (♂), especially in posterior half; spiracles small, narrow. Sternite VII, surface smooth; VL and VSM carinae obsolete (fig. 18C).

Metasoma: Segment I, DL carinae complete, moderately granular; ML carinae complete, moderately granular in posterior two-thirds; one pair of ML macrosetae; LIM carinae complete, moderately granular in posterior half; VL and VSM carinae well developed (especially in ♀, fig. 19C), complete, granular; two pairs of VSM and VL macrosetae. Segment II, DL carinae complete, granular; ML carinae complete, granular; one pair of ML macrosetae; LIM carinae restricted to the posterior third; VSM and VL carinae well developed, complete (♀, fig. 18C) or obsolete (♂); usually two pairs of VL and three pairs of VSM macrosetae. Segment III, DL and ML carinae complete, moderately granular; one pair of ML macrosetae; LIM carinae reduced to two or three granules in posterior quarter; VL and VSM carinae obsolete; two or three pairs of VL and three pairs of VSM macrosetae. Segment IV, DL carinae complete, granular; one pair of DL macrosetae; ML carina reduced to few distal granules (♀) or obsolete (♂); one or two pairs of ML macrosetae; LIM, VL and VSM carinae absent; three pairs of VL and three or four pairs of VSM macrosetae. Segment V, length/width ratio, ♂, 2.29–2.69 (n  =  12; mean  =  2.44), ♀, 1.9–2.2 (n  =  6; mean  =  2.02); DL carinae complete, more evident in anterior half; lateral margin smooth; ML carinae absent; four pairs of ML macrosetae; VL carinae complete (♂) or restricted to posterior three-quarters (♀); VL and VSM carinae situated close together, fused in posterior third; VM carina complete, obscured by surface granulation in posterior half (fig. 20C); four pairs of VL and VSM macrosetae; two pairs of macrosetae along posterior margin.

Telson: Length/width ratio: ♂, 2.65–2.94 (n  =  13, mean  =  2.78); ♀, 2.13–2.37 (n  =  10, mean  =  2.27). Length/height ratio: ♂, 3.33–3.89 (n  =  13, mean  =  3.53); ♀, 2.63–3 (n  =  10, mean  =  2.86). Vesicle elongated (♂, fig. 23G, 24G) or globose (♀, fig. 24E); dorsal surface smooth, slightly concave, gland not apparent (♂); ventral surface smooth (♂) or granular (♀). Aculeus short and curved, more so in ♀.

Hemispermatophore: Apex very well developed; distal crest very well developed, occupying almost three quarters of apex, curved like ventral margin. Frontal crest weakly developed, less than half length of lamina; basal part oblique; distal part short, parallel to ventral margin of lamina, slightly undulated. Basal lobe, terminal process extending almost to constriction of frontal crest (fig. 47A, B).

Distribution

Orobothriurus ramirezi is known only from localities above 3200 m in the Cordillera de Doña Ana of the Andes, in the Elqui Province of Region IV (Coquimbo), Chile (fig. 54).

Ecology

Most specimens were collected near wetland ecosystems termed “vega” or “humedal” (Squeo et al., 1994; Cepeda-Pizarro, 2004) in the high Andes. These habitats are characterized by an accumulation of water resulting in permanently flooded soil. The temperature at 3750 m varies from −17 °C in winter, to more than 24 °C in summer, but temperatures below zero are possible throughout the year (Cepeda-Pizarro, 2004). Precipitation in the form of snow, which may accumulate up to 8 m in some areas, occurs mainly during winter. The vegetation corresponds to a steppe with diverse altitudinal levels: shrub steppe (or sub-Andean floor) at 2700–3500 m, subshrub steppe at 3500–4250 m, and high-Andean steppe at 4250–4450 m, above which there is no vegetation (Squeo et al., 1994; Cepeda-Pizarro, 2004). Orobothriurus ramirezi has not been found in the upper altitudinal level, where the “vega” ecosystem is absent. It is sympatric with another bothriurid, Brachistosternus perettii Ojanguren Affilastro and Mattoni, 2006, that apparently prefers rocky areas (Ojanguren Affilastro and Mattoni, 2006). The most closely related species, O. alticola, inhabits similar habitats on the western side of the Andes. Despite the proximity of the nearest locality of O. alticola (only 16 km as the crow flies), the two species are allopatric, separated by the highest peaks of the Andes, the lowest pass being 4700 m, where life is almost absent. Most adult males of O. ramirezi were collected at night with UV light detection, sitting or walking near streams, probably in search of females, suggesting that the period when they where collected (late January, midsummer in the Southern Hemisphere) is the active season for this species.

Orobothriurus tamarugal, n. sp.

Figures 15D, 19F, 20E, 21H, 24F, 47C, D, 48–Fig. 49.Fig. 50.51, 53; table 3

Fig. 15.

Orobothriurus Maury, 1976, carapace, dorsal aspect. A. Orobothriurus paessleri (Kraepelin, 1911), ♂ (CDA 186). B. Orobothriurus alticola (Pocock, 1899), ♂ (AMNH). C. Orobothriurus quewerukana, n. sp., paratype ♂ (AMNH). D. Orobothriurus tamarugal, n. sp., holotype ♂ (MACN-Ar). Scale bars  =  1 mm.

Type Material

CHILE: Region I (Tarapacá): Iquique Province: Holotype ♂ (MACN-Ar), La Tirana, 2 km W, 20°19′59.8″S 69°40′07.6″W, 999 m, 18.i.2005, A. Ojanguren Affilastro, C. Mattoni and J.A. Ochoa, UV sampling in Prosopis tamarugo forest. Paratypes: Pampa del Tamarugal National Park, Salar de Pintados (salt lake), near rangers' office and campsite, 20°26′16.1″S 69°45′55.2″W, 1014 m, 18.i.2005, A. Ojanguren Affilastro, C. Mattoni, and J.A. Ochoa, UV sampling in Prosopis tamarugo forest and under salt plates, 1 ♂ (AMNH), 1 ♀ (MZUC), 1 juv. (LBRE).

Additional Material

CHILE: Region I (Tarapacá): Iquique Province: Pampa del Tamarugal National Park, Salar de Pintados (salt lake), near rangers' office and campsite, 20°26′16.1″S 69°45′55.2″W, 1014 m, 18.i.2005, A. Ojanguren Affilastro, C. Mattoni, and J.A. Ochoa, UV sampling on Prosopis tamarugo forest and under salt plates, 1 ♀, 4 juv. (AMNH [LP 4308]).

Etymology

The specific name is a noun in apposition referring to the Pampa del Tamarugal, where this species was collected.

Diagnosis

Orobothriurus tamarugal is most closely related to O. curvidigitus, O. paessleri, and O. quewerukana (fig. 5). The carapace possesses a median projection (epistome) at the anterior margin, and a marked anteromedian longitudinal sulcus (figs. 14C, 15C, D), and the males exhibit a curved movable finger on the pedipalp chela (fig. 51A) and a more elongated femur and patella than the females, in all four species. These species may be separated by the shape of the lamina of the hemispermatophore. The frontal crest is more developed and elongated, and the pedicel of the apex narrower in O. curvidigitus and O. quewerukana (figs. 28B, 40E) than in O. paessleri and O. tamarugal (figs. 28C, D, 47D). Most specimens of O. quewerukana possess one or two small folds in the distal crest (fig. 40E), which are absent in O. curvidigitus, O. paessleri, and O. tamarugal (figs. 28A, B, 47D). Additionally, the ventral border of the distal portion of the apex is straight in O. tamarugal, but slightly curved in O. quewerukana. The metasomal carinae of O. curvidigitus, O. quewerukana, and O. tamarugal are similarly developed: the DL and ML carinae are complete and granular on segments I–IV, the VL carinae obsolete and smooth on I–IV, and the VSM carinae almost completely fused with the VL carinae on V. Orobothriurus tamarugal can be separated from O. curvidigitus, O. paessleri, and O. quewerukana by the near complete absence of pigmentation, which is present on the carapace, tergites, and ventral surfaces of the metasoma in the other species. The carinae of the pedipalp femur are less developed in O. tamarugal: the DE carinae, in particular, are incomplete in O. tamarugal (fig. 49A), but complete and comprising larger granules in O. curvidigitus and O. quewerukana (fig. 42A). The IM carina of the male pedipalp chela is present and finely granular in O. tamarugal, but absent in O. curvidigitus and O. quewerukana. The ventral surfaces of sternite VII and metasomal segment I are densely granular in O. tamarugal (fig. 19F), finely granular to smooth in O. quewerukana (fig. 19C), and entirely smooth in O. curvidigitus (fig. 19A).

Description

Based on holotype ♂ and paratypes. Measurements of holotype ♂ and paratype ♀ recorded in table 3.

Total length: ♂, 23.5–35.3 mm (n  =  2); ♀, 30.1 mm (n  =  1).

Color: General color yellowish, with faint dark brown spots. Carapace almost unpigmented (adults), except for median ocular tubercle and lateral ocelli (fig. 48) or faintly pigmented (juveniles), one spot near anterior margin, two spots median laterally and two faint spots posterolaterally, with median ocular tubercle, lateral ocelli, anteriomedian and posteriomedian longitudinal sulci more densely pigmented. Tergites almost unpigmented, except for two large faint spots laterally, delimiting broad, unpigmented median band (fig. 48). Chelicerae, pedipalps, legs, sternum, genital operculum, pectines, sternites, metasoma, and telson unpigmented; aculeus sclerotized, dark reddish brown.

Chelicerae: Movable finger with two subdistal teeth, very small in ♂.

Carapace: Anterior margin granular (♂) or smooth (♀); anteromedian surfaces smooth; lateral margins finely granular, more so in ♂. Anterior margin linear (♀, juv.) or with small median projection (epistome) (♂, fig. 15D). Anteromedian longitudinal, median ocular, and posteromedian sulci well developed; posterolateral sulci obsolete. Median ocular tubercle raised, situated anteromedially; median ocelli two ocular diameters apart.

Pedipalps: Femur, more elongated in ♂, length/width ratio: ♂, 4.05–4.27 (n  =  2), ♀, 3.13; DI and VI carinae complete, granular; DE carina almost complete, coarsely granular in proximal three-quarters, smooth elsewhere; internal surface sparsely and coarsely granular (fig. 49A). Patella, more elongated in ♂, length/width ratio: ♂, 3.45–4.10 (n  =  2), ♀, 2.95; DI and VI carinae almost complete, granular (fig. 49B–D); DE and VE carinae vestigial, finely and sparsely granular in ♂; internal surface sparsely granular, with prominent granule adjacent to trichobothrium i near DI carina. Chela manus slightly rounded, fingers relatively elongated (figs. 49, 50); length/width ratio: ♂, 3.18–4.07 (n  =  2), ♀, 3.83; length/height ratio: ♂, 2.90–3.62 (n  =  2), ♀, 3.83; carinae obsolete, finely granular (less so in ♀) or absent; VM carina restricted to proximal three-quarters of manus; DS, DMA, DI carinae complete, finely and densely granular (♂) or smooth (♀); D carina finely and sparsely granular; IM carina finely and sparsely granular in proximal half (fig. 51B); intercarinal surfaces finely granular (♂, fig. 51) or smooth (♀, fig. 50); internal surface with acuminate apophysis (♂) or low bulge (♀) near articulation of movable finger (♀) (figs. 50A, 51B); movable finger (♂), strongly curved, creating small gap with fixed finger when fingers closed (fig. 51A); fingers, dentate margins each with median denticle row and 5–6 pairs of internal and external accessory denticles.

Trichobothria: Femur with 3 trichobothria, patella with 19, chela with 27 (figs. 49–Fig. 50.51). Chela trichobothrium Et₃ situated in same axis as, or slightly basal to Est (figs. 50, 51).

Tergites: Tergites I–VI, surfaces finely granular, more coarsely and densely so near distal margins of III–VI (♂) or smooth (♀). Tergite VII tetracarinate, paired DL carinae restricted to posterior two-thirds of segment, paired DSM carinae to posterior third; intercarinal surfaces granular.

Legs: Femur and patella, prolateral surfaces finely granular, retrolateral surfaces smooth. Femur, ventral carinae weakly developed; other carinae absent. Patella acarinate. Telotarsi, pro- and retroventral rows of spiniform macrosetae with following counts on leg I, 1/1; II, 2/2; III and IV, 3/3.

Pectines: Pectinal tooth count: ♂, 19–21 (n  =  6; mode  =  20); ♀, 17–18 (n  =  2).

Sternites: Sternites III–VI, surfaces smooth; spiracles small, elliptical, and narrow. Sternite VII, surface smooth in anterior half, sparsely granular in posterior half; VL carinae obsolete, slightly more developed in holotype ♂ (fig. 19F); VSM carinae absent.

Metasoma: Segment I, DL carinae complete, moderately to coarsely granular; ML carinae well developed, becoming more coarsely granular in posterior two-thirds; LIM carinae moderately to coarsely granular, restricted to posterior two-thirds, oriented obliquely to antero-dorsal surface, almost joining ML carinae; one pair of macrosetae situated medially between ML and LIM carinae; VL carinae complete, obsolete, finely granular; VSM carinae weakly developed, sparsely granular; two pairs of VSM and VL macrosetae (fig. 19F). Segments II–IV, DL carinae complete, coarsely granular; one pair of DL macrosetae; ML carinae complete, coarsely granular; one pair of ML macrosetae; surfaces between ML and LIM carinae granular on segments II and III; LIM carinae reduced to few granules in posterior third of segment II, less developed on III, absent on IV; VL carinae vestigial, becoming absent toward segment IV; VSM absent or reduced to few scattered granules on segment II (♀); three pairs of VL and VSM macrosetae. Segment V, length/width ratio, ♂, 1.95–2.24, ♀, 1.86; DL carinae reduced to few granules in anterior half; one pair of DL macrosetae; ML carinae obsolete, granular; two pairs of ML macrosetae; lateral intercarinal surfaces finely and sparsely granular; VL and VM carinae complete, granular; VL and VSM carinae fused, only separated medially (fig. 20E); ventral intercarinal surfaces granular; three pairs of VL and VSM macrosetae; two pairs of macrosetae along posterior margin.

Telson: Length/height ratio: ♂, 2.53–3.06 (n  =  2); ♀, 2.3 (n  =  1). Vesicle elongated (♂, fig. 22H) or oval (♀, fig. 24F); dorsal surface smooth, slightly concave medially, gland not apparent (♂); ventral surface coarsely granular. Aculeus short and curved, more so in ♀.

Hemispermatophore: Apex very well developed; distal crest almost straight, well developed, restricted to distal half of apex. Frontal crest weakly developed, less than half length of lamina; basal part oblique; distal part short, parallel to ventral margin of lamina, lateral projections slightly undulated with minute granulation. Basal lobe, terminal process extending almost to median part of frontal crest (fig. 47C, D).

Distribution

All known records of this species are located in the Pampa del Tamarugal of Iquique Province, Region I (Tarapacá), Chile (figs. 3A, 53).

Ecology

The Pampa del Tamarugal is a desert area characterized by the near absence of rainfall (0.2–1 mm per year), high-salinity soils (mostly of alluvial origin from the Andes), depressions with salt lakes, and a forest dominated by Prosopis tamarugo and Prosopis chilensis (Mol.) Stuntz trees (fig. 3A). Prosopis tamarugo is a deciduous open-crowned tree up to 18 m tall, with a trunk up to 80 cm in diameter, a dense mat of lateral roots and a deep taproot (up to 6 m deep on trees 15 m tall; Habit et al., 1981; Serra, 1997). Temperatures in the Pampa del Tamarugal vary from −12 °C during winter nights to 36 °C on summer days, with a daily range of more than 35 °C in summer. The salt lakes are dry, surface water is absent year-round, and the groundwater may be more than 60 m below the surface. The only humidity is provided by sporadic fog. Prosopis tamarugo forests occur only where groundwater is between 2–40 m below the surface (Serra, 1997). Specimens of O. tamarugal were collected at night, with UV light detection, inside the forest. The two males collected were active on the surface, sitting below large trees. The females and juveniles were inactive, resting under large plates of hard soil and salt (one of the forests is located near a dry salt lake). Brachistosternus donosoi Cekalovic, 1974, a larger and more active bothriurid species, was collected in sympatry.

Orobothriurus wawita Acosta and Ochoa, 2000

Figures 13C, D, 16C, D, 19E, 21C, 22F, 25B, C, 27C, 52

Fig. 16.

Orobothriurus Maury, 1976, carapace, dorsal aspect. A, B. Orobothriurus ramirezi, n. sp. A. Holotype ♂ (MACN-Ar). B. Paratype ♀ (MACN-Ar). C, D. Orobothriurus wawita Acosta and Ochoa, 2000. C. Paratype ♂ (CDA 019). D. Paratype ♀ (CDA 019). Scale bars  =  1 mm.

Fig. 17.

Orobothriurus Maury, 1976, sternite VII and metasomal segments I–IV, ventral aspect. A. Orobothriurus parvus Maury, 1976, ♀ (MHNC). B. Orobothriurus ampay Ochoa and Acosta, 2003, paratype ♀ (MHNC). Scale bars  =  1 mm.

Fig. 18.

Orobothriurus Maury, 1976, sternite VII and metasomal segments I and II, ventral aspect. A. Orobothriurus atiquipa Ochoa and Acosta, 2003, paratype ♂ (MHNC). B. Orobothriurus calchaqui, n. sp., paratype ♀ (MACN-Ar). C. Orobothriurus ramirezi, n. sp., paratype ♀ (MACN-Ar). D. Orobothriurus alticola (Pocock, 1899), ♀ (MACN-Ar). E, F. Orobothriurus compagnuccii, n. sp. E. Paratype ♀ (MACN-Ar). F. Paratype ♂ (MACN-Ar). Scale bars  =  1 mm.

Fig. 19.

Orobothriurus Maury, 1976, sternite VII and metasomal segments I and II, ventral aspect. A. Orobothriurus curvidigitus (Kraepelin, 1911), ♀ (MHNC). B. Orobothriurus paessleri (Kraepelin, 1911), ♀ (MHNC). C. Orobothriurus quewerukana, n. sp., paratype ♂ (MHNC). D. Orobothriurus huascaran, n. sp., paratype ♀ (MACN-Ar). E. Orobothriurus wawita Acosta and Ochoa, 2000, paratype ♀ (MHNC). F. Orobothriurus tamarugal, n. sp., holotype ♂ (MACN-Ar). Scale bars  =  1 mm.

Fig. 20.

Orobothriurus Maury, 1976, metasomal segment V, ventral aspect. A. Orobothriurus alticola (Pocock, 1899), ♀ (MACN-Ar). B. Orobothriurus compagnuccii, n. sp., paratype ♂ (MACN-Ar). C. Orobothriurus ramirezi, n. sp., paratype ♂ (MACN-Ar). D. Orobothriurus quewerukana , n. sp., paratype ♂ (MHNC). E. Orobothriurus tamarugal, n. sp., holotype ♂ (MACN-Ar). F. O. ramirezi, n. sp., paratype ♀ (MACN-Ar). Scale bars  =  1 mm.

Fig. 21.

Orobothriurus Maury, 1976, metasomal segment V, ventral aspect. A. Orobothriurus atiquipa Ochoa and Acosta, 2002, paratype ♂ (MHNC). B. Orobothriurus paessleri (Kraepelin, 1911), ♂ (MACN-Ar). C. Orobothriurus wawita Acosta and Ochoa, 2000, paratype ♂ (MHNC). D. Orobothriurus calchaqui, n. sp., paratype ♂ (MACN-Ar). E. Orobothriurus grismadoi Ojanguren Afillastro et al., 2009, paratype ♂ (MACN-Ar). F. Orobothriurus huascaran, n. sp., paratype ♂ (MACN-Ar). Scale bars  =  1 mm.

Fig. 22.

Orobothriurus Maury, 1976, metasomal segment V, ventral aspect. A. Orobothriurus ampay Ochoa and Acosta, 2003, paratype ♀ (MHNC). B. Orobothriurus paessleri (Kraepelin, 1911), ♀ (MHNC). C. Orobothriurus curvidigitus (Kraepelin, 1911), ♀ (MHNC). D. Orobothriurus huascaran, n. sp., paratype ♀ (MACN-Ar). E. Orobothriurus parvus Maury, 1976, ♀ (MHNC). F. Orobothriurus wawita Acosta and Ochoa, 2000, paratype ♀ (MHNC). Scale bars  =  1 mm.

Orobothriurus wawita Acosta and Ochoa, 2000: 137–143, figs. 1–13; 2001: 205; 2002: 18; Ochoa, 2004a: 43, 52, 55, 73, figs. 1, 2, 21, table 1; 2005: 55, 56, figs. 7, 9, table 2; Rein, 2007: 5.

Type Material

PERU: Cusco Department: Urubamba Province: Holotype ♂ (MACN-Ar 9652), Pacchac, Pumahuanca, 13°13′S 72°06′W, 3800 m, 17.vii.1998, J.A. Ochoa. Paratypes: Cusco Department: Urubamba Province: same data as holotype, 1 ♂, 1 ♀ (CDA 019); same locality as holotype, 23.vii.1998, J. Flores and J.A. Ochoa, 2 ♂, 2 ♀ (MHNC), 1 ♀ (MACN-Ar 9653); Maras [13°19′53″S 72°09′22″W, 3360 m], 29.x.1997, O. Mujica and J.A. Ochoa, 1 ♂ (MHNC). Ayacucho Department: Cangallo Province: Común Pampa [13°37′S 74°08′W, 2700 m], 10.ii.1963, R. Garcia, 2 ♀ (MUSM).

New Records

PERU: Cusco Department: Quispicanchis Province: Lucre, Huacarpay [13°36′30″S 71°44′03″W], ca. 3000 m, 14.i.2004, J.A. Ochoa, 2 juv. (AMNH [LP 3059]). Urubamba Province: Ollantaytambo [13°15′18″S 72°15′46″W, 2861 m], 14.iii.1947, J.C. Palhsler, 1 ♀ (MACN-Ar), 13.ii.1983, S. and A. Roig, 1 ♀ (MACN-Ar); Ollantaytambo, Río Kusichaca (Inka trail) [13°14′45″S 72°25′46″W], 2750 m, 15.i.1983, S. and A. Roig, 1 ♀ (MACN-Ar).

Diagnosis

Orobothriurus wawita may be distinguished from other species of the genus by the following characters. This species lacks an apophysis on the internal surface of the pedipalp chela manus of the male (fig. 25C), which is present in all other species except O. ampay, in which it is reduced to vestigial granules. The pedipalp chela manus of O. wawita is slender (fig. 23B, C) with a greater length/width ratio, 5.6–6.5 (♂), 4.46–5.45 (♀), than all other species, 2.98–4.6 (♂), 3.2–4.22 (♀). The pigmentation on the ventral surfaces of the metasoma of O. wawita comprises irregular scattered spots, not forming a VM stripe (fig. 13D) that is present and well defined, at least on metasomal segments II–IV, in other species (figs. 11, 13A). Orobothriurus wawita is most closely related to O. parvus (fig. 5). Both species share similar reticulate pigmentation along the metasomal DL carinae (figs. 12B, 13C), macrosetal counts on the ventral surfaces of the metasoma, and hemispermatophore morphology, including an elongated lamina apex and a short frontal crest (figs. 27B, C). These two species may be distinguished by the following characters. The fixed finger of the pedipalp chela of the male is straight, such that no gap is evident when the fingers are closed, in O. wawita (fig. 25B, C), whereas it is slightly curved, creating a small gap when the fingers are closed, in O. parvus (fig. 26A). The VL and VM carinae of metasomal segment V are absent (♂) or restricted to the distal third of the segment (♀) in O. wawita (figs. 21C, 22F), but complete in O. parvus (fig. 22E). The ventral margin of the apex of the hemispermatophore is straight in O. wawita (fig. 27C), but curved distally to the dorsal surface in O. parvus (fig. 27B).

Fig. 23.

Orobothriurus Maury, 1976, telson, lateral aspect. A. Orobothriurus alticola (Pocock, 1899), ♂ (MACN-Ar). B. Orobothriurus calchaqui, n. sp., paratype ♂ (MACN-Ar). C. Orobothriurus compagnuccii, n. sp., paratype ♂ (MACN-Ar). D. Orobothriurus huascaran, n. sp., paratype ♂ (MACN-Ar). E. Orobothriurus grismadoi Ojanguren Afillastro et al., 2009, paratype ♂ (MACN-Ar). F. Orobothriurus quewerukana, n. sp., paratype ♂ (MHNC). G. Orobothriurus ramirezi, n. sp., paratype ♂ (MACN-Ar). H. Orobothriurus tamarugal, n. sp., holotype ♂ (MACN-Ar). Scale bars  =  1 mm.

Fig. 24.

Orobothriurus Maury, 1976, telson, lateral aspect. A. Orobothriurus calchaqui, n. sp., paratype ♀ (MACN-Ar). B. Orobothriurus compagnuccii, n. sp., paratype ♀ (MACN-Ar). C. Orobothriurus huascaran, n. sp., paratype ♀ (MACN-Ar). D. Orobothriurus quewerukana, n. sp., paratype ♀ (MHNC). E. Orobothriurus ramirezi, n. sp., paratype ♀ (MACN-Ar). F. Orobothriurus tamarugal, n. sp., paratype ♀ (MZUC). Scale bars  =  1 mm.

Fig. 25.

Orobothriurus Maury, 1976, dextral pedipalp chela. A. Orobothriurus ampay Ochoa and Acosta, 2003, paratype ♀ (MHNC), external aspect. B, C. Orobothriurus wawita Acosta and Ochoa, 2000, paratype ♂ (MHNC). B. External aspect. C. Ventral aspect. D. Orobothriurus atiquipa Ochoa and Acosta, 2002, paratype ♂ (MHNC), external aspect Scale bars  =  1 mm.

Fig. 26.

Orobothriurus Maury, 1976, dextral pedipalp chela, external aspect. A. Orobothriurus parvus Maury, 1976, ♂ (MHNC). B. Orobothriurus paessleri (Kraepelin, 1911), ♂ (MACN-Ar). C. Orobothriurus curvidigitus (Kraepelin, 1911), ♂ (MHNC), external aspect. Scale bars  =  1 mm.

Fig. 27.

Orobothriurus Maury, 1976, sinistral hemispermatophore, ectal aspect. A. Orobothriurus ampay Ochoa and Acosta, 2003, holotype ♂ (MUSM). B. Orobothriurus parvus Maury, 1976, ♂ (MHNC). C. Orobothriurus wawita Acosta and Ochoa, 2000, paratype ♂ (MHNC). Scale bar  =  0.5 mm.

Fig. 28.

Orobothriurus Maury, 1976, sinistral hemispermatophore. A. Orobothriurus atiquipa Ochoa and Acosta, 2002, holotype ♂ (MACN-Ar), ectal aspect. B, D. Orobothriurus curvidigitus (Kraepelin, 1911), ♂ (MHNC). C, E. Orobothriurus paessleri (Kraepelin, 1911), ♂ (MHNC). B, C. Ectal aspect. D, E. Frontal crest and lobe region, dorsal aspect. Scale bar  =  1 mm.

Fig. 29.

Orobothriurus calchaqui, n. sp., habitus. A, B. Paratype ♂ (MACN-Ar). C, D. Paratype ♀ (MACN-Ar). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bars  =  5 mm.

Fig. 30.

Orobothriurus calchaqui, n. sp., paratype ♂ (MACN-Ar), dextral pedipalp femur (A) and patella (B–D). A, B. Dorsal aspect. C. External aspect. D. Ventral aspect. Scale bar  =  1 mm.

Fig. 31.

Orobothriurus calchaqui, n. sp., dextral pedipalp chela. A, B. Paratype ♀ (MACN-Ar). A. Dorsal aspect. B. External aspect. C–E. Paratype ♂ (MACN-Ar). C. External aspect. D. Ventral aspect. E. Ventrointernal aspect. Scale bars  =  1 mm.

Fig. 32.

Orobothriurus Maury, 1976, sinistral hemispermatophores. A, B. Orobothriurus calchaqui, n. sp., paratype ♂ (MACN-Ar). C, D. Orobothriurus compagnuccii, n. sp., paratype ♂ (MACN-Ar). A, C. Ental aspect. B, D. Ectal aspect. Scale bar  =  1 mm.

Fig. 33.

Orobothriurus compagnuccii, n. sp., habitus. A, B. Paratype ♂ (MACN-Ar). C, D. Paratype ♀ (MACN-Ar). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bars  =  5 mm.

Fig. 34.

Orobothriurus compagnuccii, n. sp., paratype ♂ (MACN-Ar), dextral pedipalp femur (A) and patella (B–D). A, B. Dorsal aspect. C. External aspect. D. Ventral aspect. Scale bar  =  1 mm.

Fig. 35.

Orobothriurus compagnuccii, n. sp., dextral pedipalp chela. A, B. Paratype ♀ (MACN-Ar). C–E. Paratype ♂ (MACN-Ar). A. Dorsal aspect. B, C. External aspect. D. Ventral aspect. E. Ventrointernal aspect. Scale bars  =  1 mm.

Fig. 36.

Orobothriurus Maury, 1976, sinistral hemispermatophore, ectal aspect showing lamina. A–C. Orobothriurus compagnuccii, n. sp., ♂ holotype, ♂ paratypes (MACN-Ar). D, E. Orobothriurus grismadoi Ojanguren Affilastro et al., 2009, ♂ paratypes (MACN-Ar). F–H. Orobothriurus famatina Acosta, 2001, ♂ (MACN-Ar). I, J. Orobothriurus alticola (Pocock, 1899), ♂ (MACN-Ar). K–O. Orobothriurus calchaqui, n. sp., ♂ paratypes (MACN-Ar). Scale bar  =  1 mm.

Fig. 37.

Orobothriurus huascaran, n. sp., habitus. A, B. Paratype ♂ (MACN-Ar). C, D. Paratype ♀ (MACN-Ar). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bars  =  10 mm.

Fig. 38.

Orobothriurus huascaran, n. sp., paratype ♂ (MHNC), dextral pedipalp femur (A) and patella (B–D). A, B. Dorsal aspect. C. External aspect. D. Ventral aspect. Scale bar  =  1 mm.

Fig. 39.

Orobothriurus huascaran, n. sp., dextral pedipalp chela. A, B. Paratype ♀ (MACN-Ar). C–E. Paratype ♂ (MACN-Ar). A. Dorsal aspect. B, C. External aspect. D. Ventral aspect. E. Ventrointernal aspect. Scale bars  =  1 mm.

Fig. 40.

Orobothriurus Maury, 1976, sinistral hemispermatophores. A–C. Orobothriurus huascaran, n. sp., paratype ♂ (MACN-Ar). D, E. Orobothriurus quewerukana, n. sp., paratype ♂ (MACN-Ar). A, D. Ental aspect. B, E. Ectal aspect. C. Dorsal aspect. Scale bar  =  1 mm.

Fig. 41.

Orobothriurus quewerukana, n. sp., habitus. A, B. Holotype ♂ (AMNH). C, D. Paratype ♀ (MHNC). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bars  =  10 mm.

Fig. 42.

Orobothriurus quewerukana, n. sp., paratype ♂ (MHNC), dextral pedipalp femur (A) and patella (B–D). A, B. Dorsal aspect. C. External aspect. D. Ventral aspect. Scale bar  =  1 mm.

Fig. 43.

Orobothriurus quewerukana, n. sp., dextral pedipalp chela. A, B. Paratype ♀ (MHNC). C–E. Paratype ♂ (MHNC). A. Dorsal aspect. B, C. External aspect. D. Ventral aspect. E. Ventrointernal aspect. Scale bar  =  1 mm.

Fig. 44.

Orobothriurus ramirezi, n. sp., habitus. A, B. Paratype ♂ (MACN-Ar). C, D. Paratype ♀ (MACN-Ar). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bars  =  10 mm.

Fig. 45.

Orobothriurus ramirezi, n. sp., paratype ♂ (MACN-Ar), dextral pedipalp femur (A) and patella (B–D). A, B. Dorsal aspect. C. External aspect. D. Ventral aspect. Scale bar  =  1 mm.

Fig. 46.

Orobothriurus ramirezi, n. sp., dextral pedipalp chela. A, B. Paratype ♀ (MACN-Ar). C–E. Paratype ♂ (MACN-Ar). A. Dorsal aspect. B, C. External aspect. D. Ventral aspect. E. Ventrointernal aspect. Scale bar  =  1 mm.

Fig. 47.

Orobothriurus Maury, 1976, sinistral hemispermatophore. A, B. Orobothriurus ramirezi, n. sp., paratype ♂ (MACN-Ar). C, D. Orobothriurus tamarugal, n. sp., holotype ♂ (MACN-Ar). A, C. Ental aspect. B, D. Ectal aspect. Scale bar  =  1 mm.

Fig. 48.

Orobothriurus tamarugal, n. sp., habitus. A, B. Holotype ♂ (MACN-Ar). C, D. Paratype ♀ (LBRE). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bars  =  10 mm.

Fig. 49.

Orobothriurus tamarugal, n. sp., holotype ♂ (MACN-Ar), dextral pedipalp femur (A) and patella (B–D). A, B. Dorsal aspect. C. External aspect. D. Ventral aspect. Scale bar  =  1 mm.

Fig. 50.

Orobothriurus tamarugal, n. sp., paratype ♀ (MZUC), dextral pedipalp chela. A. Dorsal aspect. B. External aspect. C. Internal aspect. Scale bar  =  1 mm.

Fig. 51.

Orobothriurus tamarugal, n. sp., holotype ♂ (MACN-Ar), dextral pedipalp chela. A. External aspect. B. Ventral aspect. C. Ventrointernal aspect. Scale bar  =  1 mm.

Distribution

All except one record of this species are situated in inter-Andean valleys at 2700–3800 m in the Ayacucho and Cusco departments of southern Peru (figs. 2B, C, 52). A record from Potosi (19°34′S 65°28′W), 900 km from Cusco in Bolivia (fig. 1), is based on a single female, which, although morphologically similar to Peruvian material, possesses slight differences in pigmentation pattern and granulation of the metasomal segments (Acosta and Ochoa, 2002). This record is probably a mislabeling.

Fig. 52.

Orobothriurus Maury, 1976, locality records in central Peru. Orobothriurus ampay Ochoa and Acosta, 2003, cross; Orobothriurus huascaran, n. sp., circles; Orobothriurus parvus Maury, 1976, squares; Orobothriurus wawita Acosta and Ochoa, 2000, triangles.

Fig. 53.

Orobothriurus Maury, 1976, locality records in southern Peru and northern Chile. Orobothriurus atiquipa Ochoa and Acosta, 2002, square; Orobothriurus curvidigitus (Kraepelin, 1911), circles; Orobothriurus paessleri (Kraepelin, 1911), crosses; Orobothriurus quewerukana, n. sp., triangles; Orobothriurus tamarugal, n. sp., diamonds.

Fig. 54.

Orobothriurus Maury, 1976, locality records in Argentina and central-northern Chile. Orobothriurus alticola (Pocock, 1899), squares; Orobothriurus calchaqui, n. sp., circle; Orobothriurus compagnuccii, n. sp., star; Orobothriurus famatina Acosta, 2001, diamonds; Orobothriurus grismadoi Ojanguren Affilastro et al., 2009, triangle; Orobothriurus ramirezi, n. sp., crosses.

Ecology

This species is endemic to the Queswa biogeographical region (Marín Moreno, 1961; Ceballos Bendezú, 1976; Ochoa, 2005; fig. 2B, C) and syntopic with two other bothriurids, Brachistosternus andinus and Pachakutej oscari Ochoa, 2004.