Urophonius tregualemuensis Cekalovic, 1981

Ojanguren-Affilastro, Andrés A., Ochoa, José A., Mattoni, Camilo I. & Prendini, Lorenzo, 2010, Systematic Revision of the granulatus Group of Urophonius Pocock, 1893 (Scorpiones, Bothriuridae), with Description of a New Species from Central Chile, American Museum Novitates 2010 (3695), pp. 1-44 : 25-31

Urophonius tregualemuensis Cekalovic, 1981

Figures 1, 2C, 3C, 4C, 5C, 6C, 7C, G, 8E, F, 15–17; table 2

Urophonius tregualemuensis Cekalovic, 1981: 195 , 196, 198, 199; Acosta, 1999: 157, 158, 163; Lowe and Fet, 2000: 49 (complete reference list until 1998); 2003: 7, 8; Ojanguren-Affilastro, 2005: 133; Ojanguren-Affilastro and Cheli, 2009: 353, 355.

TYPE MATERIAL: CHILE: Holotype ♀ ( MZUC 603 ), Region VII (Maule): Maule Province: Tregualemu [36°03′ 51″S 72°45′ 58.4″W], 27.ii.1969, H. Moyano . Paratypes: Tregualemu [36°03′ 51″S 72°45′ 58.4″W], 27.ii.1969, H. Moyano, 1 ♀ ( MZUC 605 ), 14.iii.1967, T . Cekalovic, 3 ♀ ( MZUC 479 , 639 , 639 a). Region VI (Libertador Bernardo de O’Higgins) : Curicó Province: Llico [34°46′S 72°05′W], 16.iii.1967, T . Cekalovic, 1 ♀ ( MZUC 483 ) .

NEW RECORDS: CHILE: Region VI (Libertador Bernardo de O’Higgins): Curicó Province: Vegas del Flaco [34°57′S 70°26′W], 24.xi.1969 , L. Peña, 1 ♂ ( MZUC 715 ), 1 ♀ ( MZUC 713 ). Region VII (Maule): Cauquenes Province: Los Ruiles National Reserve , NW Cauquenes, 35°50′ 01.68″S 72°30′ 36.87″W, 146 m, 13.xii.2003 , C.I. Mattoni, J.A. Ochoa and L. Prendini, UV detection on cool, still, dark, humid night in Nothofagus forest on steep W-facing slope, small river at base, dense litter layer with rocks, specimens common leaf litter and also climbing bushes and tree trunks, syntopic with Centromachetes sp. , 51 ♂, 25 ♀, 1 juv. ( AMNH); 8 ♂, 13 ♀ ( LBRE); 2 ♂, 2 ♀ ( MHNC). Maule Province: Tregualemu [36°03′51″S 72°45′58.4″W], 27.ii.1969 , H. Moyano, 3 juv. ( MZUC 329 ), 8.xi.2003 , V. Cekalovic, 1 ♂ ( AMNH). Talca Province: Alto Vilches [35°36′ S 71°12′ W], 12.xii.1971 , M. Pino, 1 ♂ ( MZUC 633 ), 5.xii.1999 , J. Mondaca, 1 ♀ ( MHNS), 16–17.i.1984 , E. Maury, 1 ♀, 1 juv. ( MACN); Vilches Alto [35°36′S 71°12′ W], 2160 m, 4.xii.1999 , J. Mondaca, 1 ♀ ( MHNS), 18.xi.2000 , J. Mondaca, 1 ♀ ( MHNS), 29–30.xi.2003 , J. Mondaca, 1 ♂, 3 ♀ ( MACN); Vilches [35°36′ S 71°12′ W], 7–8.i.1989 , 1 ♀ ( MACN). Region VIII (Bio Bio): Ñuble Province : Piedras Comadres , 20 km W Chillán [36°41′ S 71°55′ W], 2.xii.2004 , J. Mondaca, 1 ♀ ( MACN); Recinto , 8 km E [36°44′ S 71°48′ W], 16.xi.1989 , 1 ♀ ( MACN). Region IX (Araucania): Malleco Province: Las Quilmas campsite and surrounds, El Manzano (between Vegas Blancas and Angol), 37°48′ 16.560″S 72°52′ 17.940″W, 599 m, under stones and UV in native Nothofagus forest, riverside of stream and Pinus forest, 2 ♀, 5 juv. ( AMNH), 6 ♀ ( LBRE), 16.i.2006 , C. Mattoni, M. and F. Vivanco.

DIAGNOSIS: Urophonius tregualemuensis is most similar morphologically to U. pizarroi , n. sp., from central Chile. Both species share the following combination of characters: the hemispermatophore distal lamina is elongated ( fig. 8E–H ); the carinae of metasomal segment V are weakly developed; pedipalp femoral trichobothrium e is situated proximal to dorsal macroseta M1 (fig. 19A).

Urophonius tregualemuensis may be distinguished from U. pizarroi , n. sp., by its paler pigmentation. The carapace anterior margin exhibits a wide, unpigmented triangle and the tergites a median unpigmented stripe, in U. tregualemuensis , whereas the carapace is almost completely pigmented, and there is no median unpigmented stripe on the tergites, in U. pizarroi , n. sp. The two species may also be distinguished by the shape of the lobe of the hemispermatophore, which is less developed, does not protrude, and has a slightly excavated, concave internal surface in U. tregualemuensis ( fig. 8E ), compared with U. pizarroi , n. sp., in which the lobe is well developed and protruding, without an internal laminar extension, and with a deeply excavated internal surface ( fig. 8G ).

Urophonius tregualemuensis is also morphologically similar to U. somuncura , from which it may be distinguished by the more elongated distal lamina of its hemispermatophore ( fig. 8C–F ) and the position of pedipalp femoral trichobothrium e, which is situated proximal to dorsal macroseta M1 (fig. 16A), rather than in the same axis (fig. 13A). Both species may also be distinguished by means of the pigmentation pattern of the carapace, the anterior margin of which exhibits a wide, unpigmented triangle in U. tregualemuensis ( fig. 2C ), but is densely pigmented in U. somuncura ( fig. 2B ). The two species may be further distinguished by means of the VL and VSM carinae of sternite VII and metasomal segments I and II, which are well developed in U. tregualemuensis ( figs. 3C , 4C ), compared with U. somuncura , in which the VL carinae are weakly developed, and the VSM carinae absent, or represented only by scattered granules ( figs. 3B , 4B ).

DESCRIPTION: Based on ♂ and ♀ specimens deposited in the LBRE.

Total length: 23–31.5 mm (n = 8; mean = 25.95 mm) in ♂; 26–33 mm (n = 10; mean = 28.76 mm) in ♀.

Color: Base color yellowish or reddish, with dark-brown spots of pigmentation ( fig. 15 ). Chelicerae with reticulate pigmentation on external surfaces of fingers and near articulation, in basal part of manus. Carapace, anterior margin with unpigmented triangle, and small black spot or narrow stripe anteromedially in most specimens ( fig. 2C ); two broad, dark stripes, extending from posterior margin of unpigmented triangle to anterior margin of posterior longitudinal sulcus; median ocular tubercle and area around lateral ocelli dark brown; two lateral stripes extending from lateral margins to posterior longitudinal sulcus; two dark spots posteriorly. Tergites I–VI, each with paired, dark spots laterally, not reaching lateral margins of segments, and leaving unpigmented stripe medially; in some specimens, anterior margin of spots on I–VI poorly developed, leaving unpigmented area medially, which forms unpigmented stripe, such that three unpigmented longitudinal stripes (one median on I–VII and two submedian on I–VI) are exhibited. Sternum, genital opercula, and pectines weakly pigmented, with some faint spots. Sternites III–VI, lateral margins weakly pigmented, with three broad, faint stripes, two submedially and one medially, absent in some specimens; VII densely pigmented on lateral margins, with three narrow, dark stripes, two submedially and one medially, extending entire length of segment. Metasomal segments I–III, dorsal surfaces each with two dark spots submedially, connected to two narrow stripes along DL carinae, becoming broader at posterior margins, and connecting to lateral stripes; lateral surfaces each with broad, dark stripe below LSM carinae, connected to lateral stripes by reticulate pigmentation; ventral surfaces each with three separate dark stripes (two broader VL and a narrow VSM stripe) along entire length of segment, not joined to posterior margin. Metasomal segment IV, similar to I– III but with DSM spots reduced to narrow lines connected to DL stripes. Metasomal segment V, dorsal surface with paired, narrow submedian stripes and broad, lateral stripes in anterior half, joining in posterior half; lateral surface with reticulate pigmentation joining with dorsal

♂ ( AMNH). E, F . ♀ ( AMNH). A. Dorsal aspect. B. External aspect. C, F. Ventral aspect .

D, E. Internal aspect. Scale bar = 1 mm.

Metasoma: Metasomal segment I, dorsal surface sparsely granular; DL and LSM carinae granular, extending entire length of segment; surface between DL and LSM carinae densely granular; LIM carinae restricted to posterior half of segment; one pair of LIM macrosetae; lateral margins sparsely granular; ventral surface with paired VL and VSM carinae, more developed in ♀, diverging slightly at anterior margins; surface between VL and VSM carinae sparsely granular; two pairs of VL and VSM macrosetae, sometimes with additional, intermediate row of 1 or 2 macrosetae. Segment II, similar to segment I, except with carinae less developed; one pair of LSM macrosetae and, in some specimens, one pair of DL macrosetae; LIM carinae restricted to posterior margin of segment; VL and VSM carinae well developed (♀) or weakly developed (♂); three pairs of VSM macrosetae. Segment III, similar to segment II, except with VL and VSM carinae weakly developed (♀) or obsolete (♂); one pair of DL macrosetae. Segment IV, DL carinae granular, extending entire length of segment, but weakly developed medially in some specimens, and connected to posterior margins of LSM carinae by scattered granules, forming accessory carina; LSM carinae vestigial, restricted to anterior and posterior margins of segment; LIM carinae absent; one pair of DL, LSM, and LIM macrosetae; ventral surface smooth; three pairs of VL and VSM macrosetae. Segment V elongated ( figs. 5C , 6C ); length/width ratio 2.24– 2.48 (n = 8; mean = 2.34) in ♂, 1.86–2.08 (n = 10; mean = 1.94) in ♀; length/height ratio 2.42–2.83 (n = 8; mean = 2.64) in ♂, 2.08–2.29 (n = 10; mean = 2.21) in ♀; dorsal and lateral surfaces finely and sparsely granular; DL carinae reduced to granules at anterior margin of segment; one or two pairs of DL macrosetae; LSM carinae represented by pair of macrosetae at posterior margin; LIM carinae represented by three pairs of macrosetae; ventral surface granular in posterior half (♂) or posterior three-quarters (♀) of segment; VL carinae reduced to posterior three-quarters (♀) or posterior half (♂) of segment, comprising larger granules near posterior margin; VL and VM carinae equally well developed; VSM carinae subparallel to VL carinae but diverging in posterior third; three or four pairs of VL macrosetae, three pairs of VSM macrosetae, and two pairs of macrosetae at posterior margin of segment.

Telson: Vesicle shallow, more globose in ♂ than ♀ ( fig. 7C, G ), length/height ratio 3.22– 3.44 (n = 8; mean = 3.34) in ♂, 3.22–3.68 mm (n = 10; mean = 3.49) in ♀; ventral surface granular (♀) or smooth (♂); dorsal surface smooth, with (♂) or without (♀) an elliptical median depression, corresponding to telson gland. Aculeus short, shallowly curved.

Pedipalps: Femur with DI, DE, and VI carinae comprising discontinuous row of small granules along entire length of segment (fig. 16A); trichobothrium e situated proximal to dorsal macroseta M1. Patella with DI, DE, and VI carinae obsolete, visible only as slight curvature of surface, along entire length of segment (fig. 16B–D). Chela manus slender (more robust in ♂), acarinate ( fig. 17 ), internal surface with pronounced, subtriangular projection and shallow depression, with group of 4 or 5 granules (and, in some specimens, 1 or more additional granules between this group of granules and median denticle row of fixed finger), near base of fixed finger (♂; fig. 17D ); fingers elongated, median denticle row medially uneven, forming double row in places, with five pairs of accessory granules.

Legs: Surfaces smooth. Basitarsi each with two well-developed, equal-length pedal spurs. Telotarsi elongated, shallow, each with well-developed ventromedian row of hyaline setae, and paired rows of ventrosubmedian spiniform setae, with following counts on each telotarsus: I: 1/1, II: 2/2, III: 5/5, IV: 5/6; the only pair of setae on I and first pair on II are setiform, whereas the rest are stout and spiniform. Ungues strongly curved and equal in length.

Pectines: Tooth count: 15–17 (n = 8; mode = 16) in ♂; 12–15 (n = 10; mode = 14) in ♀.

Hemispermatophore: Basal portion very well developed. Distal lamina well developed, elongated, similar in length to basal portion; distal crest straight, oriented in the same direction as principal axis of hemispermatophore; frontal crest (distal posterior flexure) present; internal lobe with two well-developed denticles ( fig. 8F ), external denticle ca. 50 % larger than internal denticle. Lobe region well developed ( fig. 8E ), basal lobe well developed, barely protruding, without internal laminar extension; internal surface forming broad, concave excavation. We examined the hemispermatophores of 10 specimens and observed no obvious variation. In recently collected specimens, we observed a thin lamina partially covering the external margin of the basal lobe ( fig. 8E ), which probably corresponds to half the genital plug ( Mattoni and Peretti, 2004). This structure is almost impossible to recover in poorly preserved specimens as it is extremely delicate and tends to break during dissection of the hemispermatophore.

DISTRIBUTION: Urophonius tregualemuensis is endemic to central and southern Chile (fig. 1), in regions VI (Libertador Bernardo de O’Higgins), VII (Maule) , VIII (Bio Bio), and IX (Araucania).

ECOLOGY: Records of this species are situated in humid forest habitats, from the coast to the base of the Andes at 2000 m, an area that belongs to the Bosque Caducifolio botanical region ( Gajardo, 1993). Specimens have been observed on the surface at night with UV light in spring and summer. At Los Ruiles National Reserve (13 December, 2003), we collected both sexes inside a mixed Nothofagus forest. Most specimens were located on small grasses or bamboos at about 30–70 cm from the ground, apparently hunting. They dropped down, trying to disappear into the dry foliage, at the slightest vibration. At Las Quilmas campsite (16 January, 2006), in a mixed but extremely disturbed Nothofagus forest, only females were captured, and all specimens were walking on the ground. Males appear to be active on the surface only during spring (late October–early December), as in the other species of the granulatus group. Urophonius tregualemuensis was collected in sympatry with a species of Centromachetes Lönnberg, 1897 , at Los Ruiles National Reserve.

Figures

FIGURE 2. Urophonius granulatus group, carapace, dorsal aspect, showing pigmentation pattern. A. Urophonius granulatus Pocock, 1898, ♂ (MACN). B. Urophonius somuncura Acosta, 2001, ♂ (MACN). C. Urophonius tregualemuensis Cekalovic, 1981, ♂ (MACN). D. Urophonius pizarroi, n. sp., paratype ♂ (MZUC). Scale bars = 1 mm.

FIGURE 3. Urophonius granulatus group, sternite VII and metasomal segments I and

FIGURE 4. Urophonius granulatus group, sternite VII and metasomal segments I and II, ventral aspect. A. Urophonius granulatus Pocock, 1898, ♀ (LBRE). B. Urophonius somuncura Acosta, 2001, ♀ (MACN). C. Urophonius tregualemuensis Cekalovic, 1981, ♀ (AMNH). D. Urophonius pizarroi, n. sp., paratype ♀ (MZUC). Scale bars = 1 mm.

FIGURE 5. Urophonius granulatus

FIGURE 6. Urophonius granulatus

FIGURE 7. Urophonius granulatus group, telson, lateral aspect. A. Urophonius granulatus Pocock, 1898, ♂ (MACN). B. Urophonius somuncura Acosta, 2001, ♂ (MACN). C. Urophonius tregualemuensis Cekalovic, 1981, ♂ (AMNH). D. Urophonius pizarroi, n. sp., paratype ♂ (MZUC). E. U. granulatus, ♀ (LBRE). F. U. somuncura, ♀ (MACN). G. U. tregualemuensis, ♀ (AMNH). H. U. pizarroi, n. sp., paratype ♀ (MZUC). Scale bars = 1 mm.

FIGURE 8. Urophonius granulatus group, sinistral hemispermatophore, ectal and ental aspects. A, B. Urophonius granulatus Pocock, 1898, ♂ (MACN). C, D. Urophonius somuncura Acosta, 2001, ♂ (MACN). E, F. Urophonius tregualemuensis Cekalovic, 1981, ♂ (MACN). G, H. Urophonius pizarroi, n. sp., paratype ♂ (MZUC). Scale bars = 1 mm.

FIGURE 15. Urophonius tregualemuensis Cekalovic, 1981, habitus. A, B. ♂ (AMNH). C, D. ♀ (AMNH). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bars = 10 mm.

FIGURE 17. Urophonius tregualemuensis Cekalovic, 1981, dextral pedipalp chela. A–D.