Urophonius tumbensis Cekalovic, 1981

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 : 28-39

Urophonius tumbensis Cekalovic, 1981

Figures 1 , 2E, 3E, 6C, D, 10A, B, 11C, F, 12C, F, 20, 21C, 22–24, 25A, B; tables 1, 2

Urophonius tumbensis Cekalovic, 1981: 197–199 ; Lowe and Fet, 2000: 46 (complete reference list until 1998); Ojanguren-Affilastro et al., 2010: 2; Pizarro-Araya et al., 2011: 166–169.

TYPE MATERIAL: Holotype Ƌ ( MZUC 259 ), CHILE: Región VII (Bío-Bío): Concepción Province: Tumbes Peninsula, Caleta Leandro [36°38′S 73°05′W], 15.VI.1965, G. Sanhueza . Paratypes: same data except “ 15.V.1966,” 2 Ƌ ( MZUC 614 , 663 ). According to the MZUC curator (J.N. Artigas, personal commun.), these specimens cannot be found although they were not formally loaned, and must therefore be lost .

NEW RECORDS: Same data as paratypes, 1 Ƌ ( MZUC 345 ). CHILE: Región VII (Bío-Bío): Concepción Province : Tumbes Peninsula [36°38′S 73°05′W], 4.xii.1972 , T. Cekalovic, 1 Ƌ ( AMNH); Hualpén Botanical Reserve , 36°47′51.2″S 73°09′29.3″W, 150 m, 24.vii.2009, A.A. Ojanguren- Affilastro, J. Pizarro-Araya, F.M. Alfaro, O. Vergara, D. Valdivia, and C. Grismado, 2 ♀ ( MACN), 1 ♀ ( AMNH), 15.vii.2010, A.A. Ojanguren-Affilastro, J. Pizarro-Araya, L. Piacentini, E. Soto, and D. Valdivia, 2 Ƌ, 3 ♀ ( MACN), 2 Ƌ, 3 ♀ ( MHNS), 2 subad., 1 juv. ( AMNH [ LP 10649 ]) .

DIAGNOSIS: Urophonius tumbensis is most similar morphologically to U. transandinus : the hemispermatophore is similar; the VM and VSM carinae of metasomal segment V are well developed, occupying the entire length of the segment; and the chela manus is robust, with distinct carinae in the male. The two species can be separated as follows. One macroseta (M1), associated with the d and e trichobothria of the pedipalp femur, is present in U. tumbensis , compared with two macrosetae (M1, M2) in U. transandinus . The VSM carinae of metasomal segments I and II form two transverse carinae, one at the anterior margin of the segment, the other in its posterior third, in U. tumbensis , whereas the VSM carinae are diverging at the anterior margin of the segment and subparallel posteriorly, in U. transandinus .

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

Total length: 26–31 mm (n = 6; mean = 29.17 mm) in Ƌ; 29–36.5 mm (n = 9; mean = 32.11 mm) in ♀.

Color: Base color yellowish, with dark brown spots of pigmentation (fig. 20). Cheliceral manus with dense reticulate pigmentation; fingers densely pigmented in distal half and near point of articulation. Carapace, anterior two-thirds densely pigmented; anterior margin densely pigmented medially and around lateral ocelli; anteromedian longitudinal sulcus, ocular tubercle, and anterior margin of postocular sulcus densely pigmented; two broad, dark stripes extending from lateral ocelli to anterior margin of posteromedian longitudinal sulcus; paired dark spots laterally and posterolaterally, posterolateral spots connected medially by dark narrow stripe at posterior margin. Tergites I–VI, each with paired dark spots laterally in posterior two-thirds of segment, not reaching lateral margins, and paired dark spots submedially in posterior half, contiguous at anterior margin, lateral and submedian spots contiguous at posterior margin in most specimens; VII with dense reticulate pigmentation in posterolateral two-thirds of segment, contiguous medially at anterior margin. Sternum, genital opercula and pectines unpigmented. Sternite III unpigmented; IV–VII each with small spot at anterolateral margins, VII additionally with faint reticulate pigmentation in posterior third of segment. Metasomal segment I, dorsal surface with paired triangular spots submedially and at posterior margin, not contiguous, leaving unpigmented stripe medially; lateral margins densely pigmented in posterior half of segment ventral to LSM carinae, with reticulate pigment elsewhere; VL stripes restricted to anterior third of segment and posterior margin; VL stripes absent or reduced to faint pigmentation medially (fig. 3E). Metasomal segments II and III, dorsal and lateral surfaces as for segment I; VL stripes restricted to anterior margin and posterior half of segment, contiguous with lateral spots in posterior quarter; VSM stripes restricted to posterior half of segment, contiguous medially, and connecting to VL stripes in posterior third of segment. Metasomal segment IV, dorsal margin with paired dark spots posterolaterally, leaving unpigmented area medially, and reticulate pigmentation elsewhere; lateral margins with dark spot in posterior quarter of segment, contiguous with dorsal and ventral pigmentation at posterior margin, and reticulate pigmentation elsewhere; ventral surface as for segment III. Metasomal segment V, dorsolateral margins densely pigmented, especially in posterior third, leaving unpigmented area medially; lateral margins densely pigmented in posterior third, with reticulate pigmentation elsewhere; paired VL and VSM stripes extending entire length of segment, VL stripes connected to VSM stripes and to lateral pigmentation by dense reticulate pigmentation, VSM stripes separated, additional VM stripe evident between them in densely pigmented specimens. Telson vesicle faintly pigmented, except for narrow VM and paired narrow VSM stripes; aculeus unpigmented basally, apex dark brown. Pedipalps, trochanter with dark spot dorsally; femur pigmented dorsally and externally, especially near articulation with patella, unpigmented ventrally; patella densely pigmented dorsally, dorsoexternally and dorsointernally, unpigmented ventrally; chela manus, external surface with reticulate pigmentation and five dark longitudinal stripes, internal surface unpigmented, except at base of fixed finger and near articulation of movable finger, which are densely pigmented. Legs, coxa unpigmented, trochanter faintly pigmented, femur external surface with two dark spots medially and near articulation with patella;

patella dorsoexternal surface with two dark spots medially and near articulation with tibia; tibia with dark spot medially; basitarsi with dark spot at articulation with tibia; telotarsi unpigmented.

Carapace: Surfaces smooth or finely granular, usually more so in Ƌ. Anterior margin with shallow median notch. Anteromedian longitudinal, interocular, posteromedian longitudinal, and posterolateral sulci well developed. Median ocular tubercle shallow,

ocelli larger in Ƌ than in ♀, approximately two diameters apart, each with two longitudinally aligned microsetae anteriorly and one macroseta posteriorly. FIGURE 22. Urophonius tumbensis Three pairs of small lateral ocelli on each side of cara- Cekalovic, 1981, Ƌ (MACN), dextral pedipalp patella. A. Dorsal aspect. B. External pace, anterior and median ocelli situated very close

aspect. C. Ventral aspect. Scale bar = 1 mm. 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. Segments I–VI each with anteromedian pair of microsetae, and one to three pairs of macrosetae, some or all of which may be reduced to microsetae, at posterior margin.

Sternites: Surfaces III–VI smooth, with small, elliptical spiracles; VII, anterior half smooth,

posterior half granular, with well developed VSM and VL carinae, posterior margin with welldeveloped carina.

Metasoma: Metasomal segment I, dorsal surface sparsely granular; DL carinae granular,

extending entire length of segment, with larger granules near posterior margin; LSM carinae weakly granular, extending entire length of segment; surface between DL and LSM carinae densely granular; LIM carinae restricted to posterior two-thirds of segment; one pair of LIM macrosetae; lateral margins sparsely granular; paired VL and VSM carinae, VL carinae subparallel, VSM carinae forming two transverse carinae, one well developed near anterior margin, other less developed in median part of segment (fig. 10A, B), some granules forming two subparallel VSM carinae between them; four pairs of VSM macrosetae, anterior pair situated on external margin of anterior transverse carina, not in same axis as other pairs;

three pairs of VL macrosetae. Segment II as for I, except with carinae slightly less developed;

one pair of LSM macrosetae and three pairs of VSM macrosetae, anterior pair absent. Segment III as for II, except carinae much less developed; DL and LSM carinae extending entire length of segment but comprising small scattered granules; one pair of DL and LSM macro-

pedipalp chela. A–D. Ƌ (MACN). D. ♀ (MACN). A. Dorsal aspect. B.

External aspect. C. Ventral aspect. D, E. Internal aspect. Scale bar = 1 mm.

setae; LIM carinae restricted to posterior third of segment, with one pair of LIM macrosetae; VL carinae granular, extending entire length of segment, with three pairs of VL macrosetae; VSM carinae comprising few scattered granules, anterior transverse carina well developed, posterior transverse carina reduced to few more pronounced granules (♀) or absent (Ƌ), with three pairs of VSM macrosetae. Segment IV, DL, LSM, and VL carinae obsolete, reduced to slight curvature of surface along entire length of segment, LSM carinae barely visible medially; LIM carinae absent; VSM carinae reduced to few scattered low granules, barely visible (♀) or absent (Ƌ); one pair of DL macrosetae; one or two pairs of LSM macrosetae; three pairs of VL and VSM macrosetae. Segment V elongated (fig. 11C, F); length/width ratio 2.22–2.57 (n = 6; mean = 2.47) in Ƌ, 1.85–2.08 (n = 9; mean = 1.95) in ♀ ; length/height ratio 2.8–3.23 (n = 9; mean = 3.01) in Ƌ, 2.07–2.47 (n = 9; mean = 2.26) in ♀; DL carinae reduced to small bulge at anterior margin of segment (♀) or absent (Ƌ); two pairs of DL macrosetae; LSM carinae represented only by three or four pairs of macrosetae; LIM carinae absent; VL carinae granular, extending entire

FIGURE 25. Urophonius Pocock, 1893 , telotarsi. A, length of segment (♀) or restricted to posterior

B. U. tumbensis Cekalovic, 1981 , ♀ (MACN). A. half (Ƌ); five pairs of VL macrosetae, posterior

Sinistral telotarsus III, retrolateral aspect, with pair situated posterior to posterior transverse

detail of ungues. B. Sinistral telotarsus IV, retrolat-

carina; VSM and VL carinae obsolete; VM carina

eral aspect. C. U. brachycentrus (Thorell, 1876) , Ƌ

(MACN), dextral telotarsus IV, prolateral aspect. extending entire length of segment, bifurcating

Scale bars = 0.2 mm (A) and 0.5 mm (B, C). in posterior third; four pairs of VM macrosetae, posterior pair situated posterior to posterior transverse carina.

Telson: Vesicle shallow, ventral surface broader anteriorly in Ƌ than ♀ (fig. 12C, F), length/

height ratio 3.23–3.63 (n = 6; mean = 3.39) in Ƌ, 3.27–3.58 mm (n = 9; mean = 3.50) in ♀;

dorsal surface 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 comprising discontinuous row of small granules along entire length of segment (Ƌ) or reduced to few scattered granules (♀) (fig.

2E); one macroseta (M1) associated with d and e trichobothria; trichobothrium e situated proximal to macroseta M 1 in some specimens and distal to M 1 in others. Patella with DI, DE, and VI carinae obsolete, reduced to slight curvature of surface, along entire length of segment (fig. 22). Chela manus slender, more robust in Ƌ (fig. 23), length/width ratio 2.80– 3.52 (n = 6; mean = 3.14) in Ƌ; 3.84–4.5 (n = 9; mean = 4.03) in ♀; length/height ratio 2.82–3.18 (n = 6; mean = 3.03) in Ƌ, 3.48–4 (n = 9; mean = 3.67) in ♀; acarinate; 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 (Ƌ) (fig. 23D); fingers elongated, median denticle row uneven medially, forming double row in parts, with five pairs of internal and external accessory denticles.

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, with following counts on telotarsus I: 1/1, II: 2/2, III: 4–5/5, IV: 5/5–6; only pair of spiniform macrosetae on I and first pair on II–IV subspiniform, others stout spiniform (figs. 24, 25A, B). Ungues strongly curved, equal in length, with acute tip (fig. 25A). Pseudonychium well developed, apex curved and very acute. Median dorsal lobe protruding ca. 20%–30% length of unguis.

Pectines: Tooth count: 13–15 (n = 6; mode = 14) in Ƌ; 12–14 (n = 9; mode = 12) in ♀. Cekalovic (1981) mentions 14–15 for a male of the type series.

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 not connected to distal lamina (fig. 6D), external denticle ca. 50% larger than internal denticle. Lobe region well developed (fig. 6C), basal lobe very well developed, protruding; internal surface forming deep, concave excavation. Genital plug very well developed, covering most of posterior part of basal lobe; area near concave excavation with grooves.

DISTRIBUTION: All known records of Urophonius tumbensis occur on the Tumbes Peninsula in the Concepción Province of Región VIII (Bío-Bío), southern Chile (fig. 1). We examined a female Urophonius specimen from “Cueva de Pincheiras” in Ñuble Province, also in Región VIII, which appears to be closely related. This specimen exhibits minor differences from the Tumbes specimens, however, and we prefer not to speculate on its identity until additional specimens, including adult males, become available.

ECOLOGY: All records of this species occur in dense, temperate to cold deciduous humid forests, near the Pacific coast, an area that belongs to the “Bosque Caducifolio de Concepción” of the “Bosque Caducifolio” botanical region ( Gajardo, 1993).

We collected this species in the Hualpén Botanical Reserve, one of few areas in southern Chile where scorpions have been previously recorded ( Cekalovic, 1976). Personally collected specimens were discovered with UV light detection at night, during the winter months of July and August. All other known specimens were collected during May and September. These records suggest a winter activity period for U. tumbensis . This species was collected in sympatry with two other bothriurid species in primary forest: a species of Phoniocercus Pocock, 1893 , probably Phoniocercus sanmartini Cekalovic, 1968 , and a species of Centromachetes Lönnberg, 1897 , probably Centromachetes pocockii ( Kraepelin, 1894) . Only the Centromachetes species was present in adjacent patches of grassland within the reserve, and no specimens of U. tumbensis were found in nearby coastal areas, or plantations of Pinus sp. , close to the reserve boundaries.

Urophonius tumbensis is the only arboreal (corticolous) species of the family Bothriuridae , and the only arboreal scorpion species recorded from the temperate forests of southern Chile ( Pizarro-Araya et al., 2011). Most specimens were collected from 2 to 6 m above ground in “Peumo” trees, Cryptocara alba (Molina) Looser (fig. 21), where they were observed walking on the main trunk and, in some cases, feeding on spiders. Few specimens were collected on the ground surface.

The telotarsal ungues and pseudonychium of U. tumbensis are more strongly curved, with a more acute tip, than those of other ground-dwelling species of Urophonius (figs. 19, 24, 25). This character, observed in arboreal (corticolous) scorpion species of several other families ( Prendini, 2001), may be associated with the arboreal habitat of U. tumbensis . Other bothriurid species, e.g., U. tregualemuensis , have been observed foraging on shrubs, bamboo, and other plants at night ( Ojanguren-Affilastro et al., 2010), and may occasionally use the base of trees as a daytime shelter ( Maury, 1973). However, no other bothriurid species was previously observed to be active on trees at such heights.

Figures

FIGURE 1. Map of central Chile, plotting known locality records for three species of Urophonius Pocock, 1893: U. mondacai, n. sp. (circles); U. transandinus Acosta, 1998 (triangles); U. tumbensis Cekalovic, 1981 (squares).