Butheolus gallagheri Vachon, 1980

Lowe, Graeme, 2018, The genera Butheolus Simon, 1882 and Xenobuthus gen. nov. (Scorpiones: Buthidae) in Oman, Euscorpius 261, pp. 1-73 : 5-15

Butheolus gallagheri Vachon, 1980

( Figs. 1–66 , 133–162 , 330, 332 , 338 , Tabs. 1 , 3–5 )

Butheolus (Butheolus) gallagheri Vachon, 1980: 253– 255 , fig. 24, pl. B, C.

Butheolus gallagheri: Vachon & Kinzelbach, 1987: 100 ; Fet & Lowe, 2000: 88–89; Kovařík, 2003: 137; Kovařík, 2004: 4 , 25; Santiago-Blay, Soleglad & Fet, 2004: 8; Fet, Soleglad & Lowe, 2005: 2; Lourenço, 2005: 27–28, fig. 32; Lourenço & Qi, 2006: 93–94, tab. 1; Kamenz & Prendini, 2008: 8, tab. 2, pl. 9; Kovařík & Lowe, 2012: 2, 18–23, figs. 76, 79, 82, 85, 88, 91, 93, 97–98, 102–103; Lourenço & Duhem, 2012: 122; Loria & Prendini, 2014, App. S1: 2, App. S2: 2; Lowe et al., 2014: 117.

Butheolus (Butheolus) gallagheri: El-Hennawy, 1992: 101 , 114; Kovařík, 1998b: 105.

TYPE MATERIAL. Holotype ♂, Oman, Wadi Rabkut (Raykhut), Jabal Samhan , Dhofar, 17°24'N 55°16'E, 75 rn a.s.l., under rocks on side of wadi, 17–18.X.77, leg. M. D. Gallagher, MDG4738 ( MNHN VA 2261 ) (examined).

OTHER MATERIAL EXAMINED. Oman: 1♂, Jabal Samhan , Dhofar, 17°07'N 54°42'E, 1280 m a.s.l., 7.IX. 1989, leg. M.D. Gallagher MDG 8149, ( ONHM 1350 ); 1♂, Khor Rori Beach, UV detection, rock outcrop, archaeological site, 17°02.1'N 54°26.22'E, 0 m a.s.l., 18.X.1993, 20:27 h, leg. G. Lowe, ( USNM); 2♀, Khor Rori Beach, UV detection, inland rocky hillock near beach, on rocky slopes, 17°02.21'N 54°26.19'E, 0 m a.s.l., 18.X.1993, 20:32 h, leg. G. Lowe, ( ONHM); 8♂, 12♀, 1 juv♀, main road above Khor Rori Beach , UV detection on ground, densely vegetated wadi, E of Taqah; warm and humid with many insects, 17°03.22'N 54°25.33'E, 50 m a.s.l., 18.X.1993, 21:24 h, leg. G. Lowe ( BMNH 1♂; FKCP 1♂, 1♀; GL 1♂, 1♀; MCZ 1♂; NHMB 2♂, 10♀; ONHM 1♂, 1♀); 2♂, 1♀, Khor Rori Beach , UV detection, densely vegetated wadi, on ground, 17°03.18'N 54°25.51'E, 22 m a.s.l., 24-XII- 2001, 02:25-04:45 h, leg. A. Winkler ( ZSM); 1♂, Salalah, leg. A. Ullrich (GL) .

DIAGNOSIS. A member of the genus Butheolus differentiated as follows: small scorpions, adults 22–32 mm; base color a uniform dark chocolate brown, with all metasomal segments and telson dark; pedipalps and legs yellow except for fuscosity at base of chela fingers; carapace and tergites uniformly, finely granular; pedipalp patella with dorsomedian carina incomplete, restricted to distal half of segment, weak or obsolete on proximal half; pedipalp chela with carinae weak or obsolete; ventral surface between ventromedian carinae of metasoma I with dense fine granulation in females; ventral and lateral intercarinal surfaces of metasoma II– V with fine granulation; metasoma and telson with sparse, short macrosetae; telson with moderate to small subaculear tubercle, obtuse angle between posterior vesicle surface and aculeus base; pedipalp and metasoma relatively slender: pedipalp femur L/W ♂ 3.00–3.38, ♀ 2.43–2.76; pedipalp patella L/W ♂ 3.20–3.65, ♀ 2.64– 2.84; pedipalp chela L/W ♂ 5.16–5.65, ♀ 4.24–4.79; metasoma I L/W ♂ 0.81–0.92, ♀ 0.76–0.81; metasoma IV L/D ♂ 1.44–1.60, ♀ 1.37–1.54, metasoma V L/D ♂ 1.96–2.12, ♀ 1.82–2.08, telson L/D ♂ 2.46–2.83, ♀ 2.33–2.56; pectine teeth ♂ 17–21, ♀ 14–17; basitarsus III retrosuperior setae 4–8.

REDESCRIPTION.

Based on 14 ♂, 16 ♀ (including holotype ♂).

Coloration ( Figs. 1–32 , 51–54 , 59–66 ). Base color a uniform dark brown on dorsal aspect of prosoma. mesosoma and all aspects of metasoma and telson; metasoma darker than mesosoma; carapace slightly lighter in postocular, posterior-median triangular area, tergites with lighter lateral patches in transverse reticulated bands; telson vesicle with pale spots at bases of macrosetae and a pair pale stripes on ventral surface; dorsal aspect of chelicerae dark on fingers and distal margin of manus, yellow-brown reticulated on distal half of manus; legs yellow except for reddish spots on distal external articular condyles; pedipalps yellow except for fuscous patch on distal chela manus at base of pedipalp fingers; ventral aspect of mesosoma yellow except for weak fuscosity on sternite VII.

Carapace ( Figs. 1 , 3 , 5 , 7 ). Strongly trapezoidal, W/L 0.79–0.92, posterior W/ anterior W 1.97 –2.38; lateral flanks steeply sloped; median ocular tubercle prominent; postocular area forming triangular posteromedial plateau with shallow transverse posterior marginal furrows; interocular triangle sloped downwards towards anterior margin; anterior margin with 9–11 macrosetae that are longer in females, carapace otherwise devoid of macrosetae; anterolateral margins with 5 pairs of lateral eyes: 3 major ocelli and 1 major or minor posterior ocellus below a granular ridge, 1 minor ocellus above granular ridge; whitish eyespot present below lateral eye cluster; all carinae of carapace obsolete except for superciliary carinae which may extend slightly anterior to median ocular tubercle; entire surface with dense fine granulation except for smooth patches on postocular plateau, posterior transverse and posterior marginal furrows; superciliary carinae may be smooth or weakly granulated, with more granules on posterior slopes; granulation may be coarser on lateral parts of interocular triangle, anterior margin of carapace, and margins of postocular triangular plateau area; males with more densely granulose interocular triangle.

Chelicerae ( Figs. 65–66 ). Dorsal surface of manus smooth, with two short, pale microsetae on apical margin, each with adjacent granules; dorsointernal carina strong, granulate, bearing one long, dark macroseta and one short, pale microseta; fingers robust with dentition typical of genus, movable finger dorsal margin with two large subdistal denticles and two small basal denticles, ventral margin with larger subdistal and smaller basal denticles, fixed finger with large subdistal denticle and proximal bicusp, two denticles on ventral surface; dorsal surface of movable finger smooth, with two pale microsetae.

Coxosternal area ( Figs. 2 , 4 , 6 , 8 ). Males. Coxa I finely granulated, endite smooth on anterior half, granulated on posterior half; coxa II densely, finely granulated, endite smooth on anterior third, granulated on posterior two thirds; coxa III densely, finely granulated, granules more coarse along anterior margin; coxa IV densely, finely granulated, anterior margin with band of coarser granules, posterior margin rimmed with linear series of granules; distal anterior surfaces of coxae II–IV smooth; coxae I–III with scattered, mostly anterior macrosetae: coxa I 5–9, II 8–12, III 3–5; coxa IV typically with single macroseta on anterior proximal limit; sternum weakly granulated, subtriangular, with deep posteromedian pit, usually 2 short macrosetae; genital opercula with fine granulation anteriorly, posteriorly smooth, with 2–5 macrosetae, posterolateral margins concave. Females. Coxae smooth except for weak granulation on medial endites of coxa I, anterior margins of coxa III–IV, and row of moderate granulation on anterior 2/3 of posterior margin of coxa IV; setation as in males; sternum smooth, with larger median pit; genital opercula smooth, elongate, with 4–8 macrosetae.

Pectines ( Figs. 2 , 4 , 6 , 8 ). Basal piece with concave anterior margin and small median groove and pit, finely granulated in males, smooth anteriorly and finely granulated posteriorly in females, bearing 5–8 macrosetae; pectines with 3 marginal lamellae, 5–7 middle lamellae, extending to proximal 1/4 of trochanter IV in males, distal end of coxa IV in females; teeth longer in males than females; marginal and middle lamellae with moderate cover of short macrosetae; fulcra with 2–4 setae.

Hemispermatophore ( Figs. 55–58 , 330 ). Flagelliform, trunk elongate, ca. 4.7 times length of capsule region; flagellum short; sperm hemiduct tripartite, posterior lobe large, laminate, median lobe small, acuminate, anterior lobe of intermediate length, tapered; posterior margin of median lobe overhanging posterior lobe, the two lobes fused along a median lobe carina; basal lobe a prominent, narrow, hook-like projection.

Mesosoma ( Figs. 1–8 , 332 ). Tergites: pretergites smooth, with sinuous, finely corrugated posterior margins; tergites densely, finely granulated, with smoother transverse lateral strips on tergites IV–VI; tergites I–II without carinae or with trace of weak median carina, III with weak median carina and weak or obsolete lateral carinae, IV–VI weakly tricarinate with median and paired lateral carinae, VII with 5 carinae, the median carina a weak granulated hump; all carinae finely granular, confined to posterior half of tergites, lateral carinae anteriorly divergent; tergite granulation and carination weaker in females than males; all tergites lacking macrosetae; sternites: males: sternites III–V lacking carinae, medially smooth, laterally shagreened or finely granular; sternite VI with smooth or obsolete inner lateral carinae, weakly granulated outer lateral carinae on posterior half of sternite; sternites IV–VI with wide, posteromedian smooth patch; posterior margins of sternites III–VI with fringe of numerous, small, noncontiguous, closely spaced, digitate denticles; sternite VII densely, finely granular, with granulate median and lateral pairs of carinae confined to posterior 2/3 of sternite, only median pairs extending to posterior margin; all sternites with scattered, sparse macrosetae, sternite VII with 4 stereotypic isolated macrosetae near outer anterior ends of carinae; females: sternite III smooth medially, shagreened laterally in areas covered by pectines, sternites IV–VI smooth; carinae absent on sternites III–V, only trace of lateral carinae on VI; posterior marginal denticles of sternites III–VI smaller than in males; sternite VII with two pairs of carinae, weaker than carinae of males, with weaker dense, fine granulation on lateral surfaces; mesosoma much wider in females than males.

Metasoma ( Figs. 1–2 , 5–6 , 59–64 ). Moderate in length with robust segments, metasoma + telson L/ carapace L ♂ 5.3–5.7, ♀ 4.8–5.3; carination: segments I–III with 10 complete carinae, IV with 4 complete carinae (dorsosubmedian and dorsolateral carinae visible only on anterior 1/5 of segment), V with 2 carinae (ventrolateral); carinae on segments I–IV uniformly granulate, ventrolateral carinae on V with smaller granules in anterior half, larger granules in posterior half; ventromedian carina on V obsolete, a trace indicated by linear series of granules; lateral anal margin with 2 blunt granules or lobes, ventral anal margin with 10–14 granules; intercarinal surfaces: dense, fine granulation on dorsolateral, lateral, ventrolateral and ventral surfaces of all segments; some coarser granules on posterior 1/3 of ventral surface of V; in males, dorsal surfaces sparsely, finely granulated on I–IV and anterior half of V, smooth on posterior slope of IV and posterior half of V in trough accommodating telson; in females, dorsal surfaces smooth except for a few isolated small granules (mostly on I–II); setation: scattered microsetae and sparse, short macrosetae present on all segments, setae slightly longer in females; dorsal surfaces lack setation, except for metasoma V which may bear 2–4 macrosetae along dorsolateral margins.

Telson ( Figs. 1–2 , 5–6 , 59–64 ). Vesicle smooth dorsally, strongly granulated laterally and ventrally, ovoid, with steep posterior slope due to development of weak subaculear tubercle; granules on lateral and ventral surfaces arranged in longitudinal bands, separated by smooth lateral and ventrosubmedian troughs; scattered microsetae and short macrosetae on lateral and ventral surfaces; vesicle granulation weaker in female, posterior areas almost smooth, with shallow indentations at setal insertion points; aculeus stout, shorter than vesicle.

Pedipalps ( Figs. 9–50 ). Males ( Figs. 9–20 , 33–35, 39–42, 45–47 ). Femur: L/ W 3.00 –3.38; dorsointernal carina moderate, with uniform medium-size granules; dorsoexternal carina strong with coarse granules; external carina moderate, nearly smooth with fine granules proximally, larger conical granules distally; ventrointernal carina moderate, with uniform medium-size granules; internal carina weak, with irregular fine and coarse granules; dorsal and internal surfaces with dense, fine granulation, ventral surface with sparse fine granulation or shagreened, external surfaces mostly smooth, with scattered fine granules near carinae; 3–6 accessory macrosetae on distal external surface; patella: L/ W 3.20 –3.65; dorsointernal carina moderate, finely granular; dorsomedian carina moderate, finely granular, continuous only in distal half of segment, residual trace at proximal end of segment indicated by broken granule row; dorsoexternal carina moderate, weakly granular or smooth; external carina weak, broad, smooth; internal carina moderate, finely granulated; other carinae obsolete; dorsal, external and ventral surfaces smooth or slightly roughened, without granulation; internal surface with weak, fine granulation; chela: slender, L/ W 5.16 – 5.65, all carinae obsolete, surface smooth with sparse macrosetae and microsetae; 5–7 primary denticle subrows on movable and fixed fingers (including cases of fusion of proximal rows), subrows except proximal typically flanked by internal and external accessory denticles; 5–7 internal or external accessory denticles on movable finger, 4–6 on fixed finger. Females ( Figs. 21– 32 , 36–38, 43–44, 48–50 ). Femur: more robust than in males, L/ W 2.43 –2.76; dorsointernal carina weak to moderate, with uniform medium-size granules; dorsoexternal carina moderate with coarse granules; external carina weak, smooth, with a few distal granules; ventrointernal carina weak, with uniform small granules; internal carina weak, broad, smooth; dorsal surface with sparse, fine granulation mostly in medial area, ventral surface smooth except for narrow medial band of weak, fine granules in proximal 1/3 of segment, external and internal surfaces smooth; 5–8 accessory macrosetae on distal external surface, including linear series of 3–5 on ventral side of external carina; patella: more robust than in males, L/ W 2.64 –2.84; dorsomedian carina weak, smooth, present only on distal half of segment; dorsointernal, dorsoexternal, external and internal carinae weak, smooth; other carinae obsolete; all intercarinal surfaces smooth; chela: more robust than in males, L/ W 4.24 –4.79, all carinae obsolete, surface smooth with sparse macrosetae and microsetae; finger dentition as in male. Trichobothriotaxy: orthobothriotaxic, type Aβ ( Vachon, 1974) ( Figs. 33–50 )

Legs ( Figs. 1–2 , 5–6 , 51–54 ). Femur and patella I– IV with finely denticulate inferior carinae, other carinae finely granulate; prolateral surfaces of femur I–IV with dense, fine granulation, of patella II–IV with sparse fine granulation, on both segments slightly weaker in females than males; tibia III–IV with spurs; retrolateral tarsal spurs simple, prolateral tarsal spurs basally bifurcate; basitarsi I–III with 4–8 long retrosuperior macrosetae arranged in bristle-combs; ventral surface of telotarsi with dual rows of short, fine macrosetae (occasionally reduced to single row on telotarsus I); tarsal ungues stout.

Measurements. See Table 1 .

Variation. Color patterns were stable in the sampled populations. Meristic and morphometric variation are summarized in Tabs. 3–5 , and Figs. 133–162 .

DISTRIBUTION. Known only from coastal sites around Jabal Samhan or Jabal Qara, in the Dhofar Province of Oman ( Fig. 338 ).

ECOLOGY. Most examined material was collected from low elevations (<100 m a.s.l.), but one record was at 1,280 m a.s.l. on the plateau atop the Jabal Samhan escarpment. The species occurs in mesic or humid environments along the coast, in densely vegetated rocky wadis with gravel and sandy soil. Scorpions that occurred together with B. gallagheri were: Hottentotta salei ( Vachon, 1980) and Leiurus haenggii Lowe et al., 2014 .

Figures

Figure 338. Map of records in the Arabian Peninsula of Butheolus and Xenobuthus gen. n. Symbols show locations of collection records of species examined in this study. B. gallagheri: black circles; B. harrisoni: orange circles; B. thalassinus: cyan circles; B. villosus: light green circle; X. anthracinus: magenta squares; X. xanthus: yellow squares. Locality data are from specimens examined, except for the type locality and two additional published records of B. thalassinus. The indefinite type locality of X. anthracinus (Hadramaut “by the way”; Pocock, 1895) is not plotted. Map scale: 150 km.

Figures 1–2: Habitus of male Butheolus gallagheri Vachon, 1980. Fig. 1. Dorsal aspect. Fig. 2. Ventral aspect. Oman, wadi above Khor Rori Beach, E. of Taqah, 17°3.2'N 54°25.33'E, 18.X.1993, leg. G. Lowe. Scale bar: 5 mm.

Figures 3–4: Prosoma and mesosoma of male Butheolus gallagheri Vachon, 1980. Fig. 3. Dorsal aspect, carapace and tergites. Same locality data as male in Figs. 1–2. Fig. 4. Ventral aspect, coxosternal area and sternites. Oman, Salalah, leg. A. Ullrich. Scale bar: 1 mm.

Figures 5–6: Habitus of female Butheolus gallagheri Vachon, 1980. Fig. 5. Dorsal aspect. Fig. 6. Ventral aspect. Same locality data as male in Figs. 1–2. Scale bar: 5 mm.

Figures 7–8: Prosoma and mesosoma of adult female Butheolus gallagheri Vachon, 1980. Fig. 7. Dorsal aspect, carapace and tergites. Fig. 8. Ventral aspect, coxosternal area and sternites. Same locality data as male in Figs. 1–2. Scale bar: 1 mm.

Figures 9–32: Pedipalp of Butheolus gallagheri Vachon, 1980. Figs. 9–20. Male (♂). Dorsal (9–11), external (12–14), ventral (15–17) and internal (18–20) aspect of femur (9, 12, 15, 18), patella (10, 13, 16, 19) and chela (11, 14, 17, 20). Figs. 21–32. Female (♀). Dorsal (21–23), external (24–26), ventral (27–29) and internal (30–32) aspect of femur (21, 24, 27, 30), patella (22, 25, 28, 31) and chela (23, 26, 29, 32). Both sexes with same locality data as male in Figs. 1–2. Scale bars: 1 mm.

Figures 33–50: Trichobothrial maps and chela finger dentition of Butheolus gallagheri Vachon, 1980. Male (♂): dorsal (33), external (34) and ventral (35) aspect of chela; dorsal (39) and external (40) aspect of patella; dorsal (45), external (46) and internal (47) aspect of femur; dentition of movable finger (MF) (41) and fixed finger (FF) (42). Female (♀): dorsal (36), external (37) and ventral (38) aspect of chela; dorsal (43) and external (44) aspect of patella; internal (48), dorsal (49) and external (50) aspect of femur. Same locality data as male in Figs. 1–2. Scale bars: 1 mm (♂, ♂ dentition, ♀).

Figures 51–54: Leg tarsi, tibiae and patellae of male Butheolus gallagheri Vachon, 1980. Retrolateral aspect, left leg I tibia, basitarsus and telotarsus (51), left legs II–IV, patella, tibia, basitarsus and telotarsus (52–54 respectively). Same locality data as male in Figs. 1–2. White light illumination. Scale bar: 1 mm.

Figures 55–58: Hemispermatophore of Butheolus gallagheri Vachon, 1980. Fig. 55. Whole right hemispermatophore, convex aspect. Scale bar: 1 mm. Figs. 56–58. Capsule region and flagellum of right hemispermatophore, anterior (56), convex (57) and posterior (58) aspects. In convex view (57), the capsule is compressed to show outlines of lobes. Same locality data as male in Figs. 1–2. White light illumination. Scale bar: 200 µm.

Figures 59–64: Metasoma and telson of Butheolus gallagheri Vachon, 1980. Figs. 59–61. Male (♂). Dorsal (59), right lateral (60) and ventral (61) aspects. Figs. 62–64. Female (♀). Dorsal (62), right lateral (63) and ventral (64) aspects. Same locality data as male in Figs. 1–2. Scale bars: 2 mm.

Figures 65–68: Chelicerae of male Butheolus gallagheri Vachon, 1980 and B. harrisoni sp. n. Figs. 65–66. B. gallagheri, right chelicera, dorsal (65) and ventral (66) aspect. Same locality data as male in Figs. 1–2. Figs. 67–68. B. harrisoni sp. n., right chelicera, dorsal (67) and ventral (68) aspect. Paratype male, Oman, Jabal Qara, north slopes, Nejd desert, 17°17.83'N 54°5.11'E, 16.X.1993, leg. G. Lowe. Scale bar: 500 µm.

Figures 133–147: Scatter plots showing variation in morphometric ratios of Butheolus gallagheri Vachon, 1980 and B. harrisoni sp. n. Selected measurements normalized to carapace length (CL). Each symbol represents one specimen. B. gallagheri: black symbols; B. harrisoni: orange symbols. Males: filled circles, females: open circles. Abbreviations: L, length; W, width, WA anterior width, WP posterior width, D depth.

Figures 148–150: Histograms showing variation in meristic counts of basitarsal setation and pectine teeth for Butheolus gallagheri Vachon, 1980 and B. harrisoni sp. n. Figs. 148–149. Distributions of number of retrosuperior macrosetae (“bristlecomb” setae) on basitarsi I–III for males (148) and females (149). Fig. 150. Distributions of number of pectine teeth in male (♂) and female (♀) combs. In all plots: B. gallagheri, black bars; B. harrisoni, orange bars. Ordinates are numbers of tarsi or combs, pooling data from left and right appendages of all specimens examined.

Figures 151–162: Scatter plots showing variation in morphometric ratios of species of Butheolus and Xenobuthus gen. n. Figs. 151–156. Selected ratios of males (♂). B. gallagheri: black filled symbols; B. harrisoni: orange filled symbols; B. thalassinus: cyan circles; X. anthracinus: magenta circles; X. xanthus: yellow circles. Figs. 157–162. Selected ratios of females (♀). B. gallagheri: black open symbols; B. harrisoni: orange open symbols; B. villosus: light green circles; X. anthracinus: magenta circles; X. xanthus: yellow circles. Abbreviations: L, length; W, width (carapace W = posterior width), D depth.

Figures 330–337: Diagnostic characters of genera Butheolus and Xenobuthus gen. n.. Figs. 330–331. Hemispermatophore basal lobes of B. gallagheri (330) and X. anthracinus (331); axial views of right (330) and mirrored left (331) hemispermatophore. Oriented with convex side up, anterior margin left and posterior margin right. Abbreviations: b, basal lobe; m, median lobe. Scale bars: 50 µm. Figs. 332–337. Dentition of posterior margin of sternite IV in male B. gallagheri (332), B. harrisoni sp. n. (333), X. anthracinus (334), X. xanthus sp. n. (335), Hottentotta jayakari (Pocock, 1895) (336) and H. pellucidus Lowe, 2010 (337). UV fluorescence. Scale bars: 100 µm. Locality data: 330 and 332, as in Tab. 1; 331 and 334, as in Tab. 7; 333, paratype male, same locality as holotype; 335, holotype; 336, Oman, Rte 13, ca. 5.9 km E junction Wadi Mistal Rd, 23°21.33'N 57°38.21'E, 460 m a.s.l., 26.IX.1994, leg. G, Lowe, M.D. Gallagher; 337, Oman, Jabal Bani Jabir, IX.1995, 1800 m a.s.l., leg A. Francois, C. DeLise (ONHM).