Urophonius transandinus Acosta, 1998

Ojanguren-Affilastro, Andrés A., Pizarro-Araya, Jaime & Prendini, Lorenzo, 2011, New data on Chilean Urophonius Pocock, 1893 (Scorpiones, Bothriuridae), with description of a new species, American Museum Novitates 2011 (3725), pp. 1-44 : 16-28

Urophonius transandinus Acosta, 1998

11B, E, 12B, E, 16–19; tables 1, 2

Urophonius transandinus Acosta, 1998: 157–164 ; Ojanguren-Affilastro et al., 2010: 2, 3.

Urophonius brachycentrus: Kraepelin, 1894 (part): 221 [ZMH specimen from Valparaíso; misidentfied]; Cekalovic, 1983 (part): 51 [reference to Kraepelin, 1894].

Urophonius granulatus: Kraepelin, 1899 (part): 194 [presumably same ZMH specimen from Valparaíso, with corrected determination label; misidentified]; Cekalovic, 1983a (part): 52 [reference to Kraepelin, 1899].

Urophonius corderoi Mello-Leitão, 1931 : Cekalovic, 1981 (part): 200; 1983b: 189; 1983a: 52 [misidentified].

Urophonius sp. [ brachycentrus group]: Acosta, 1988: 25 ( Fig. 4 ), 28.

TYPE MATERIAL: Holotype Ƌ ( ZMH), CHILE: Región V (Valparaíso) : Valparaíso Province: Valparaíso [33°03′S 71°38′W], 15.v.1893, J. Michelsen . Paratypes: CHILE: Región V (Valparaíso) : Valparaíso Province: Road from Casablanca to Algarrobo , 15.vii.1966, J. Solervicens, 1 Ƌ ( MACN 9602 ), allotype ♀ ( MACN 9601 ), 1 ♀ ( CDA); El Arrayan [33°37′26.5″S 71°36′40.1″W, 19 m], 4.ix.1963, Ecología Animal, 1 Ƌ ( MZUC 392 ); El Canelo [33°08′11.2″S 71°39′54.2″W, 212 m], 3.x.1967, G. López, 1 ♀ ( MZUC 608 ) .

NEW RECORDS: CHILE: Región V (Valparaíso): Marga-Marga Province: “La Campana” National Park, Granizo sector, 32°58′53.1″S 71°07′38.2″W, 446 m, 16.vii.2009, A.A. Ojanguren- Affilastro, J. Pizarro-Araya, M. Izquierdo and F.M. Alfaro, 1 Ƌ, 3 ♀, 5 juv. ( AMNH) , 1 Ƌ, 3 ♀, 5 juv. ( MHNS) , 1 Ƌ, 3 ♀ ( MACN) , 22–23.vii.2010, A.A. Ojanguren-Affilastro, J. Pizarro-Araya, D. Valdivia, L. Piacentini and E. Soto, 5 Ƌ, 12 ♀ ( MACN) , 4 Ƌ, 14 ♀ ( MNHNS) , 1 ♀ ( AMNH [ LP 10614 ]) . Región Metropolitana de Santiago: Santiago Province: Cerro Manquehue [33°20′S 70°35′W], viii.1979, Jacobsohn, 1 ♀, ( MACN) ; Farellones [33°20′38.98″S 70°22′38.44″W], 12.xi.1973, Valencia, 2 ♀, 1 juv. ( MZUC) ; Río Clarillo [33°43′50.47″S 70°27′59″W], vii.1983, Lewin, 1 ♀ ( MACN) . Región VI (Libertador Bernardo de O’Higgins): Colchagua Province: La Palmilla de Reto [34°30′37.71″S 71°34′07.92″W], 1.vi.2002, Cornejo, 1 ♀, ( MHNS) ; Rancagua- Machali [34°11′11.64″S 70°41′46.9″W], viii.2007, Castro-Martínez, 1 Ƌ, 4 ♀ ( MHNS) ; “Río los Cipreses” National Reserve , “Las Arpas” sector, 34°16′52.1″S 70°27′4.2″W, 1066 m, 20.vii.2010, A.A. Ojanguren-Affilastro, J. Mondaca, J. Pizarro-Araya, D. Valdivia, L. Piacentini, and E. Soto, 5 Ƌ, 12 ♀ ( MACN) , 5 Ƌ, 12 ♀ ( MNHNS) , 1 Ƌ, 1 ♀, 2 subad., 1 juv. ( AMNH [ LP 10669 ]) . Región VII (Maule): Curico Province: Los Niches , Curico , 35°03′47.4″S 71°07′22.3″W, 299 m, 19.vii.2009. A.A. Ojanguren-Affilastro, J.E. Barriga-Tuñón, J. Pizarro-Araya, M. Izquierdo and F.M. Alfaro, 1 Ƌ, 10 ♀ ( AMNH) , 1 Ƌ, 10 ♀ ( MACN) , 12 ♀ ( MNHNS) , 18.vii.2010, A.A. Ojanguren-Affilastro, J.E. Barriga-Tuñón, J. Pizarro-Araya, D. Valdivia, L. Piacentini and E. Soto, 5 Ƌ, 12 ♀ ( MACN) , 5 Ƌ, 12 ♀ ( MNHNS) , 2 ♀ ( AMNH [ LP 10653 ]) ; Yacal , 35°10′23.9″S 71°07′25.1″W, 421 m, 19.vii.2010 A.A. Ojanguren-Affilastro, J. Pizarro-Araya, D. Valdivia, L. Piacentini, and E. Soto, 4 Ƌ, 11 ♀ ( MACN) , 4 Ƌ, 13 ♀ ( MNHNS) .

DIAGNOSIS: Urophonius transandinus is most similar morphologically to U. tumbensis and occurs in sympatry with U. mondacai , n. sp., U. pizarroi , and U. tregualemuensis . It may be separated from these species by the presence of two macrosetae (M1, M2) associated with the d and e trichobothria of the pedipalp femur, only one (M1) of which is present in the other species. Additionally, the pedipalp chela manus of U. transandinus is more robust, with length/ width ratio of 2.53–3.08 (n = 20; mean = 2.78) in Ƌ and 3.27–3.77 (n = 20; mean = 3.51) in ♀, than that of U. mondacai , n. sp., with length/width ratio of 4.36 Ƌ, and 5.07 in ♀.

Urophonius transandinus can be further separated from U. mondacai , n. sp., and U. tregualemuensis by the shape of the hemispermatophore. In U. transandinus , the proximal part of the basal lobe is well developed, situated proximal to the capsular concavity, and the basal lobe terminates in a conspicuous elongated structure, whereas in U. mondacai , n. sp.,

mondacai , n. sp., holotype Ƌ (AMNH). B. U. transandinus Acosta, 1998 , Ƌ

(MACN). C. U. tumbensis Cekalovic, 1981 , Ƌ (MACN). D. U. mondacai , n. sp.,

paratype ♀ ( MACN). E. U. transandinus , ♀ ( MACN). F. U. tumbensis , ♀

(MACN). Scale bars = 1 mm.

Group A

Urophonius eugenicus (Mello-Leitão, 1931) ARGENTINA: Santa Cruz

Urophonius exochus (Penther, 1913) ARGENTINA: Mendoza, Neuquén , Río Negro Urophonius mahuidensis Maury, 1973 ARGENTINA: Buenos Aires

Urophonius martinezi Ojanguren-Affilastro and Cheli, 2009 ARGENTINA: Chubut

Group B

Urophonius achalensis Abalos and Hominal, 1974 ARGENTINA: Córdoba

Urophonius brachycentrus (Thorell, 1876) ARGENTINA: Buenos Aires, Córdoba, La Pampa ,

La Rioja, Río Negro, San Juan, San Luís, Santiago del

Estero, Tucumán

Urophonius granulatus Pocock, 1898 ARGENTINA: Chubut, Santa Cruz; CHILE: Magallanes

Urophonius iheringii Pocock, 1893 ARGENTINA: Buenos Aires ; BRAZIL: Río Grande do Sul ; URUGUAY: Lavalleja, Maldonado, Montevideo, Tacuarembó

Urophonius mondacai , n. sp. CHILE: Metropolitana de Santiago, Valparaíso

Urophonius pizarroi Ojanguren-Affilastro et al., 2010 CHILE: Metropolitana de Santiago

Urophonius somuncura Acosta, 2003 ARGENTINA: Río Negro

Urophonius transandinus Acosta, 1999 CHILE: Metropolitana de Santiago, Valparaíso, Libertador Bernardo de O’Higgins, Maule

Urophonius tregualemuensis Cekalovic, 1981 CHILE: Araucanía, Bío-Bío, Maule, Libertador Bernardo de O’Higgins

Urophonius tumbensis Cekalovic, 1981 CHILE: Bío-Bío and U. tregualemuensis , the proximal part of the basal lobe is less developed, situated in line with or distal to the capsular concavity, and the proximal part of the basal lobe terminates in a short laminar structure.

Urophonius transandinus can be further separated from U. pizarroi , U. tregualemuensis , and U. tumbensis by the orientation of the VSM carinae on metasomal segments I and II or I–III. The VSM carinae are subparallel in the posterior two-thirds of the segment, diverging in the anterior third, in U. transandinus but subparallel along its entire length in U. pizarroi and U. tregualemuensis , and they form a transverse carina in U. tumbensis .

Urophonius transandinus can be further separated from U. pizarroi and U. tregualemuensis by the pigmentation pattern on the ventral surface of metasomal segments I–IV. Urophonius transandinus exhibits paired VL and VSM stripes, and occasionally a single VM stripe, whereas U. pizarroi and U. tregualemuensis exhibit a single VM and paired VL stripes.

DESCRIPTION: Based on Ƌ and ♀ specimens (AMNH, MACN, MZUC).

Total length: 33–43 mm (n = 20; mean = 37.77 mm) in Ƌ; 36.5–47 mm (n = 34; mean = 41.21 mm) in ♀.

♀ ( MACN). A, C. Dorsal aspect. B, D. Ventral aspect. Scale bars = 10 mm .

Color: Base color yellowish, with dark brown spots of pigmentation (fig. 16). Cheliceral fingers densely pigmented; manus densely pigmented at base of fixed finger and articulation of movable finger, with reticulate pigmentation elsewhere. Carapace, anterior two-thirds densely pigmented; anterior margin pigmented medially; lateral ocelli, anteromedian longitudinal sulcus, median ocular tubercle, and anterior margin of postocular sulcus densely pigmented; posterior two-thirds of postocular sulcus unpigmented; paired broad, dark stripes extending laterally from anterior margin to posteromedian longitudinal sulcus, complete or nearly so in densely pigmented specimens but disappearing near anterior margin in faintly pigmented specimens; paired dark spots laterally and posterolaterally. Tergites I–VII, each with paired dark spots laterally, leaving unpigmented stripe medially, but not reaching lateral margins, spots restricted to posterior two-thirds or posterior half of segment in faintly pigmented specimens, but occupying most of lateral surface, reaching anterior part of segment in densely pigmented specimens; median unpigmented stripe complete on I–III, lateral spots usually contiguous at anterior margins of IV–VII. Sternum, genital opercula, and pectines unpigmented. Sternite III with faint sparse spots; IV–VI each with small spot at anterolateral margin and additional faint sparse spots; VII, lateral margins densely pigmented in anterior half, with three dark stripes ventrally, usually connected by reticulate pigmentation in median part of segment, two broad stripes laterally, usually restricted to posterior half of segment, and narrow stripe medially, usually extending entire length of segment. Metasomal segment I, dorsal surface with paired dark spots at articulation points, paired triangular spots (very faint in less pigmented specimens) submedially, not contiguous medially, and paired spots at posterior margin, not contiguous medially, leaving unpigmented stripe medially; lateral margins densely pigmented, each with dark spot in posterior two-thirds of segment, ventral to LSM carinae, reaching VL carinae and dorsal spots posteriorly; VL stripes restricted to posterior half of segment, contiguous with ventral part of lateral spot; VSM stripes absent or reduced to faint pigmentation in median part of segment; VM stripe very narrow, faint, and restricted to median part of segment in specimens from northern populations (fig. 3C), well developed, extending entire length of segment in specimens from southern populations (fig. 3D). Metasomal segments II and III as for I, except dorsal surface with median spots contiguous, forming single triangular spot that may be faint in less pigmented specimens; lateral surfaces each with dark lateral spot restricted to posterior third of segment, remainder of lateral margin reticulate; VL stripes extending entire length of segment, posterior margins contiguous with dark lateral spot; VSM stripes well developed, occupying most of segment in specimens from northern populations. Metasomal segment IV as for II and III, except dorsal margin with median spot reduced to narrow stripe; lateral margins with lateral spot restricted to posterior third of segment; VL stripes extending entire length of segment, VSM stripes well developed, contiguous with VL stripes in posterior third of segment, and connected to VM stripe in anterior half by reticulate pigmentation. Metasomal segment V, dorsal surface with dorsolateral margins densely pigmented, especially in posterior half of segment, leaving unpigmented area medially; lateral margins densely pigmented at posterior margin, with reticulate pigmentation elsewhere; VL and VSM stripes extending entire length of segment, VL stripes becoming wider and more diffuse in posterior two-thirds, VSM stripes contiguous with VL stripes in posterior two-thirds; VM stripe extending entire length of segment, connected to VSM stripes in posterior half by reticulate pigmentation. Telson vesicle, dorsal surface unpigmented; ventral and lateral surfaces densely pigmented, except for VM stripe and paired VSM and lateral unpigmented stripes; aculeus reddish brown, apex dark brown. Pedipalps, trochanter with dark reticulate pigmentation dorsally; femur densely pigmented dorsally, especially near articulation points, external surface densely pigmented near articulation with patella, unpigmented elsewhere; patella densely pigmented dorsally, externally and dorsointernally, unpigmented ventrally; chela with reticulate pigmentation and seven dark longitudinal stripes, internal surface less pigmented and with two faint stripes,

area near articulation of fixed and movable fingers, and base of fingers densely pigmented. Legs, coxa unpigmented; trochanter with dark spot; femur, external surface with two dark spots medially and near articulation with patella; patella, dorsoexternal surface with two dark spots near articulation with femur and tibia; tibia with two dark spots, medially and near articulation with basitarsus; basitarsus with spot medially; telotarsus unpigmented. Carapace: Surfaces finely granular in ♀, more coarsely granular in Ƌ. Anterior margin with very shallow median notch. Anteromedian longitudinal, interocular, posteromedian longitudinal, and posterolateral sulci well developed. Median ocular tubercle well developed, superciliary carinae more pronounced FIGURE 14. Urophonius mondacai , n. sp., and median ocelli larger in Ƌ; ocelli approximately two holotype Ƌ (AMNH), dextral pedipalp patella. A. Dorsal aspect. B. External diameters apart, each with one microseta situated ante-

aspect. C. Ventral aspect. Scale bar = 1 riorly and one macroseta situated posteriorly, anterior mm. microsetae replaced by macrosetae in some specimens. Three pairs of small lateral ocelli on each side of carapace, anterior and median ocelli situated very close together, in same horizontal axis, posterior ocellus situated slightly dorsal to others, one diameter apart. Tergites: Surfaces I – VI smooth; VII with paired submedian and lateral carinae, comprising medium-sized granules, lateral carinae restricted to posterior two-thirds of segment, submedian carinae to posterior half. Tergite I with four posterior macrosetae, II – VI each with six posterior macrosetae; III –VII each with paired submedian macrosetae in anterior half of segment. Sternites: Surfaces III – VI smooth, each with small elliptical spiracles; VII, anterior half smooth, posterior half granular, with four carinae, paired VSM and VL, well developed in ♀, obsolete in Ƌ; posterior margin with well developed carina. Metasoma: Metasomal segment I, dorsal surface smooth to sparsely granular; DL carinae granular, more coarsely so near posterior margin, extending entire length of segment, with one pair of DL macrosetae; LSM carinae granular, extending entire length of segment, with one pair of LSM macrosetae; surface between DL and LSM carinae sparsely granular; LIM carinae restricted to posterior two-thirds of segment, with one pair of LIM macrosetae; VL carinae subparallel, VSM carinae diverging obliquely at anterior margin, subparallel at posterior margin, anterior part almost always oblique, VSM carinae creating V-shape in northern populations (fig. 8A, B), but forming transverse anterior carina in southern populations (fig. 9A, B); three pairs of VL macrosetae; four pairs of VSM macrosetae, anterior pair situated on anterolateral margin of VSM carina, not in same axis as other pairs. Segment II as for I, except with slightly less granular carinae; LIM carinae restricted to posterior third of smooth, with (Ƌ) or without (♀) elliptical median depression, corresponding to telson gland; ventral surface slightly granular. Aculeus short, shallowly curved .

Pedipalps: Femur with DI, DE, and VI carinae granular (Ƌ) or obsolete, reduced to slight curvature of surface (♀), extending entire length of segment; two dorsal macrosetae (M1, M2) associated with d and e trichobothria (fig. 2C); trichobothrium e situated proximal to macroseta M 1 in some specimens and distal to M 1 in others. Patella with DI and VI carinae granular, extending entire length of segment; DE and VE carinae obsolete, reduced to slight elevation of surface along entire length of segment (fig. 17). Chela manus robust (Ƌ) or slender (♀), length/ width ratio 2.53–3.08 (n = 20; mean = 2.78) in Ƌ, 3.27–3.77 (n = 20; mean = 3.51) in ♀; length/ height ratio 2.33–2.79 (n = 20; mean = 2.56) in Ƌ, 2.88–3.31 (n = 20; mean = 3.12) in ♀; E, VM, DS, DM, DI, D, and IM carinae obsolete, more developed in Ƌ than in ♀ (fig. 18); internal surface with slight bulge near articulation of movable finger (♀) or with pronounced, subtriangular projection, shallow depression, and group of 4 or 5 granules near base of fixed finger, with one or two additional granules between this group of granules and median denticle row of fixed finger (Ƌ); fingers elongated, median denticle row uneven medially, forming double row, with five or six pairs of internal and external accessory denticles; fixed finger, median denticle row with basal denticles usually fused.

Legs: Surfaces smooth. Basitarsi each with two well-developed, equal-length pedal spurs. Telotarsi elongated, shallow, each with well-developed ventromedian row of hyaline spinules and paired pro- and retroventral rows of spiniform macrosetae; spinules similar in length to spiniform macrosetae near base of telotarsus, twice their length at distal margin; spiniform macrosetae with following counts in pro- and retroventral rows on telotarsus I: 1/1, II: 2/2, III: 5–6/6–7, IV: 6–7/7–8; only pair of spiniform macrosetae on I and first pair on II–IV subspiniform, others stout spiniform (fig. 19). Ungues curved, equal in length (fig. 19C, E, G). Pseudonychium well developed, apex curved. Median dorsal lobe protruding approximately 20%–30% length of unguis.

Pectines: Tooth count: 14–17 (n = 29; mode = 16) in Ƌ; 12–16 (n = 74; mode = 13) in ♀.

Hemispermatophore: Basal portion very well developed. Distal lamina well developed, elongated, similar in length to basal portion; distal crest almost straight, oriented in same direction as frontal margin on distal lamina; frontal crest (distal posterior flexure) barely distinguishable from frontal margin; internal lobe with two well-developed denticles forming bicusp, not connected to distal lamina (fig. 5H), external denticle slightly larger than internal denticle. Lobe region well developed (fig. 5G), basal lobe very well developed, protruding, internal surface forming deep, concave excavation. Genital plug well developed, with internal longitudinal flexure and median longitudinal excavation.

DISTRIBUTION: According to Acosta (1998), U. transandinus is restricted to a narrow area extending from the Región Metropolitana de Santiago to Región V (Valparaíso), central Chile. Based on additional material examined in Chilean collections and collected during several winter expeditions to central and southern Chile, this species is more widely distributed, however. In the present contribution, U. transandinus is recorded from four regions of Chile (table 2). The specimens described by Acosta (1998) appear to represent the northernmost populations of this species.

ECOLOGY: The habitat in central Chile, where the type material was collected, is a mixture of forests and shrub steppes ( Gajardo, 1993). In this area, we collected U. transandinus in dry woods at “La Campana” National Park. We also collected this species further south, in shrub steppe at approximately 1100 m in the foothills of the Andes, at “Río los Cipreses” National Reserve. The ground surface on this occasion was covered by a layer of snow, approximately 5 cm deep, accumulated several hours earlier. Based on observations with UV light detection, we found that scorpions were active on the surface, in spite of the weather conditions, but avoided snow-covered areas by climbing over small rocks instead. We also collected U. transandinus near Curicó, in Region VII (Maule), the southernmost record of the species, more than 200 km south of the type locality and other records listed by Acosta (1998). These specimens were collected on plains and small hills, in a mixed grass/shrub steppe. VARIATION: Based on the additional material available, several morphological differences are evident among northern and southern populations of U. transandinus . Specimens from northern populations exhibit two dark VSM stripes and, in some specimens, traces of a VM stripe on the ventral surface of metasomal segments I–III (fig. 3C) whereas those from southern populations exhibit a welldefined VM stripe and traces of VSM stripes (fig. 3D). The VSM carinae on metasomal segments I and II are oblique and diverging in the anterior FIGURE 17. Urophonius transandinus Acosta,

1998, Ƌ (MACN), dextral pedipalp patella. A. third of the segment in northern populations (fig.

Dorsal aspect. B. External aspect. C. Ventral 8A), but form a transverse carina in the anterior aspect. Scale bar = 1 mm. third in most specimens from southern populations (fig. 9A). Finally, the denticles of the internal lobe of the hemispermatophore are slightly less developed in some males from southern populations. None of these differences appear to be fixed within the respective populations, however. We therefore consider these populations to be conspecific.

Figures

FIGURE 4. Species of Urophonius Pocock, 1893, carapace, dorsal aspect, showing pigmentation pattern. A. U. pizarroi Ojanguren-Affilastro et al., 2010, paratype ♀ (MZUC). B. U. tregualemuensis Cekalovic, 1981, ♀ (AMNH). Scale bars = 1 mm.