Vachoniolus gallagheri, Lowe, 2010

Lowe, Graeme, 2010, The genus Vachoniolus (Scorpiones: Buthidae) in Oman, Euscorpius 100 (100), pp. 1-37 : 10-17

Vachoniolus gallagheri , sp. nov.

Figs. 30–56 , 91–93 , 102–103 , 106–111 , 146–148 , Tab. 1

Holotype: adult ♂, Oman: wadi N of Adam , near main road, UV detection, sandy wadi, on sandy plain, 22°31.16'N 57°31.04'E, 420 m a.s.l., 17 September 1995, 22:20 h, leg. G. Lowe, NHMB.

Paratypes: Oman: 1 juvenile, near Qarn Nihaydah , under stone, 21°15'N 56°55'E, 8 October 1981, leg. M.D. Gallagher MDG 6280, GL ; 1 ♂, 1 ♀, 1 juvenile, ca. 57 km S of Hafit, pitfall No. 11, sand with little vegetation except Prosopis and Acacia , 23°29'N 55°52'E, 200 m a.s.l., trap collected 1 March 1994, leg. M.D. Gallagher & B.J. Tigar MDG 8592, NHMB ; 2 ♂, 6 ♀, 2 juveniles, sandy wadi W of Ghabah , UV detection, low sand dunes with scrub, adjacent to sabkha, scorpions walking rapidly on sand in wind, near bases of bushes, 21°23.89'N 57°09.56'E, 185 m a.s.l., 5 October 1994, leg. G. Lowe & M.D. Gallagher, NHMB, GL (1 ♂) , FKCP (1 ♀) ; 1 ♂, plain S of Adam , UV detection, open ground, silty gravel plain, 22°06.16'N 57°26.96'E, 250 m a.s.l., 16 September 1995, 19:00– 20:00 h, leg. G. Lowe & J. Dundon, ONHM ; 2 ♂, 1 juvenile, between Ghabah & Adam, UV detection on sand, sandy wadi on open gravel plain with trees and bushes, large solifuges common, 22°09.54'N 57°30.16'E, 250 m a.s.l., 16 September 1995, 22:00–23:00 h, leg. G. Lowe & J. Dundon, NHMB ; 4 ♂, 1 ♀, sandy wadi S of Nizwa roundabout, near main road, UV detection, 22°44.41'N 57°33.37'E, 380 m a.s.l., 17 September 1995, 20:30 h, leg. G. Lowe, NHMB, FKCP (1 ♂) ; 2 ♂, same locality as holotype, NHMB ; 1 ♂, Ghabah North, UV detection, active on ground, gravelly and sandy wadi, firm substrate, area of small wadis with shrubs, 21°23.63'N 57°09.64'E, 100 m a.s.l., 29 September 1995, leg. G. Lowe & M.D. Gallagher, MNHN ; 1 ♀, Saiwan , rudist outcrop, 20°42.94'N 57°36.96'E, 20 February 1996, leg. A.S. Gardner, ONHM ; 1 juvenile, Ghabah North area , S of Adam, in a sandy wadi, 21°45.74'N 57°17.31'E, 150 m a.s.l., 19 March 1996, 20:30–21:15 h, leg. J. Dundon 117, NHMB ; 1 ♂, 15 km S of Barzaman , soft sand in grassy hollow, 22°10'N 58°03'E, 250 m a.s.l., 20 October 1997, leg. M.D. Gallagher & I.D. Harrison MDG 8887, BMNH .

Diagnosis. Small to medium sized Vachoniolus , adults 35–50 mm, uniform pale yellow without melanic pigmentation on metasoma and telson ( Figs. 30–31, 34–35 ); tergites smooth in females, finely shagreened in males, with coarser granulation along posterior margins; positions of obsolete lateral carinae of tergites III–VI marked posteriorly by single granule; metasomal segments relatively stout ( Figs. 45–46 ), L/W ratios: I 1.25–1.33, II 1.48–1.65, III 1.60–1.85, IV 2.00–2.31, V 2.22–2.64; metasoma I–II with dorsolateral surfaces smooth; dorsosubmedian carinae on metasoma II–III weak, with widely spaced granules separated by two or more granule lengths; ventrosubmedian carinae of metasoma II–III crenulate to dentate, armed with moderate to large, closely spaced dentate granules, 4–8 granules on posterior half of carina; telson with moderately long aculeus, vesicle L/ telson L 0.45–0.52; pedipalp femur with trichobothrium d 5 usually distal to e 2, distance ratio (d 5 – e 2)/(e 2 – e 1) median value 0.25 ( Figs. 49 , 111 ); pedipalp patella neobothriotaxic with 8 trichobothria on external surface ( Figs. 50–51 ); pedipalp chela manus smooth in females, finely granulated on internal, dorsal and dorsolateral surface in males; tibial spurs present or absent on legs III–IV.

Etymology. The specific epithet is a patronym honoring Michael D. Gallagher for his many enduring contributions to the natural history of Oman, including collecting a large part of the materials for the author’s study of the scorpion fauna of Oman.

Comparisons. V. batinahensis sp. nov., differs as follows: larger average size, 2–3 granules on lateral carinae of tergites III–VI; weaker dentition on ventrosubmedian carinae of metasoma II–III; pedipalp patella with 7 trichobothria on external surface; pedipalp chela manus smooth; tibial spurs always present on legs III– IV; V. globimanus differs as follows: larger average size, males with shagreened tergites, finer and more dense dentition on metasomal carinae, more slender metasoma, male pedipalp chela manus smooth except for dorsointernal margin; male V. iranus Navidpour et. al., 2008, differs as follows: melanic pigmentation on carapace, pedipalp femur and patella; 2–3 granules on lateral carinae of tergites III–VI; coarsely dentate-granulate dorsosubmedian carinae on metasoma II–III; granulate dorsal and dorsolateral surfaces on metasoma I–V; enlarged bicuspid granule on mid-internal surface of pedipalp femur; more robust dorsointernal carina on pedipalp patella.

Description (holotype male unless otherwise specified).

Coloration ( Figs. 30–31 ). Body and appendages uniform pale yellow; melanic pigment on carapace under median and lateral eyes; denticles on fingers of chelicerae and pedipalp chelae, articular condyles of pedipalp movable finger, and telson aculeus castaneous.

Carapace ( Fig. 38 ). Subrectangular, 1.12 times wider than long, anterior inter-ocular width 0.47 times posterior width; surface convex with lateral flanks moderately sloped; anterior margin with weak median convexity, posterior margin nearly straight; moderately coarse to fine granulation of variable density on most of carapace; coarse granules on anterior inter-ocular area between lateral eyes, inner margins of lateral eyes, along anterior margin of carapace, and behind median eyes; anterior lateral, lateral and posterior areas around median ocular tubercle smooth, lustrous; lateral flanks of carapace with medium to fine granulation; posterior median sulcus finely shagreened, adjacent lateral surfaces smooth; median posterior margin of carapace bordered by strip of dense, fine granulation; median eyes large, ocular tubercle prominent, wide, inter-ocular distance equal to two ocular diameters; 4 lateral eyes on each side, anterior 3 ocelli equal in size, 4 th ocellus reduced; single median pair of short reddish macrosetae on anterior margin, carapace otherwise devoid of macrosetae.

Chelicera (paratype male) ( Figs. 39–40 ). Dorsal surface of manus smooth, convex, lustrous with dense subapical transverse band of small granules; dorsointernal carina at base of fixed finger with larger dentate granules; chaetotaxy: two short pale macrosetae on apical margin of manus; long reddish macroseta on dorsointernal carina; movable finger smooth, without macrosetae, ventral surface of manus smooth, with numerous long, fine microsetae on middle and internal surfaces, more dense on interior aspect, extending dorsally to base of fixed finger; ventral surfaces of fixed and movable fingers with dense brush of microsetae; dentition: fingers with normal buthid dentition (Vachon, 1963; Sissom, 1990); fixed finger with large distal tine, smaller subdistal denticle and large proximal bicusp; two large denticles on ventral surface, proximal denticle smaller than distal; movable finger with large dorsal and ventral distal tines; dorsal margin of movable finger with two large triangulate denticles and two small proximal denticles; ventral margin armed with two robust denticles, proximal smaller than distal.

Coxosternal area ( Figs. 31, 33 ). Coxal surfaces mostly smooth, lustrous, with scattered fine granulation; coxa I with coarsely granular endite and anteromedian carina, finely granulate distal margin; coxa II with coarsely granular anterior carina, finely granular endite and distal margin; coxae III–IV with anterior carina weak, with medium to fine granules; coxa III with sparse small granules on surface, distal margin finely denticulate; coxa IV surface and distal margin smooth, posterior margin finely shagreened in basal half; anterior margin with 3 macrosetae on coxa II, 3–4 macrosetae on coxa III; coxa IV with single basal macroseta; sternum subpentagonal with deep postero-median excavation, sparsely shagreened, bearing 2 macrosetae; genital opercula smooth, with 3 macrosetae; genital papillae present.

Pectines ( Figs. 31, 33 , 56 ). Basal piece smooth, rectangular, with deep anterior median notch; pectines moderately long, tips extending to basal third of trochanter IV; pectines with 3 marginal lamellae, 7–8 middle lamellae, 18–19 teeth; pectine teeth elongate, nearly uniform in size; marginal lamellae, middle lamellae and fulcra with numerous short, reddish macrosetae; fulcra with 7–11 setae; basal teeth not overlapping when anterior margins of pectines align with posterior margins of coxae IV.

Mesosoma ( Figs. 30–33 ). Tergites: pretergites smooth, lustrous; median carina obsolete on tergite I, very weak on II–IV, weak on V–VI; tergites I–VI with lateral carinae obsolete, marked near posterior margin by single enlarged granule; tergite VII with weak median hump, inner lateral carinae very weak, marked by row of granules, outer lateral carinae obsolete; posterior margins of all tergites nearly straight, smooth; tergites I–VI densely, finely, uniformly shagreened, with series of small granules along posterior margins; tergite V finely shagreened or smooth medially, sparsely granulated laterally; all tergites devoid of macrosetae, lateral margins microdenticulate. Sternites: sternites III–VI smooth, lustrous, lacking carinae; sternite III with anterior lateral areas finely shagreened; sternite VII with weak, smooth, nearly obsolete paired median carinae and weak, smooth to faintly granulose paired lateral carinae, intercarinal surfaces smooth; sternites III–VI with postero-medial margins smooth, postero-lateral and lateral margins finely denticulate; sternite VII with nearly smooth posterior margin, denticulate lateral margins. Chaetotaxy: sternite III with 7 short macrosetae in medial area, IV–VI with paired medial macrosetae, VII with one pair of macrosetae on median carinae, one pair on lateral carinae, one pair on anterior lateral surface; posterior margins of sternites with short to medium length setae, III–VI with 2–3 larger posterior lateral setae, mixed with 12, 12, 19 and 11 smaller setae, arranged nearly uniformly along posterior margins with medial gap; lateral margins of sternites with well spaced macrosetae (left–right): III 2–2, IV 4–4, V 6–5, VI 4–3, VII 1–2.

Hemispermatophore (paratype males) ( Figs. 41–42 ). Flagelliform; trunk elongate, slender; flagellum long, filiform, proportions variable; in 3 males, pars recta 0.53, 0.71 and 0.27 times length of trunk, pars reflecta 0.65, 0.47 and 0.80 times length of trunk, respectively; inner lobe a broad lamina, gently tapering with blunt apex; median and outer lobes shorter, laminate, almost fully (1 case) or fully fused (2 cases); median lobe with outer margin curved, tapering to fine apical process, outer lobe linear, apically truncate; basal lobe a short, rounded, mediolaterally compressed knob; measurements of paratype (from N of Adam, partially fused lobes): trunk length (to base of flagellum) 3.5 mm, pars recta 2.5 mm, pars reflecta 1.7 mm, inner lobe (from base of flagellum) 900 µm, median lobe (from juncture with inner lobe) 305 µm, outer lobe from juncture with median lobe 120 µm, from median lobe juncture with inner lobe 440 µm, basal lobe 60 µm.

Metasoma ( Figs. 45–46 ). Long, slender, total length plus telson length 6.1 times carapace length. Carination: metasoma I with 10 complete carinae; metasoma II–III with 8 complete carinae, median lateral carinae confined to posterior 1/2 of segment II, posterior 1/3 of segment III; segment IV with 4 carinae, segment V with 3 carinae; dorsosubmedian carinae weak with small, separated granules on segments I–III, obsolete on IV; dorsolateral carinae moderate on segments I–III with large dentate granules, weak with smaller dentate granules on IV, obsolete on V; median lateral carinae weak with small granules on segment I, weak to obsolete on II–III with smaller anterior granules, larger posterior granules; ventrolateral carinae on segment I moderate, smooth anteriorly, coarsely granulate posteriorly, on II strong with coarse granules, on III strong, coarsely granulate-crenulate with larger posterior granules, on IV strong, coarsely granulate-crenulate, on V strong, armed with small, medium and large dentate granules; ventrosubmedian carinae on segment I weak, granulose posteriorly, obsolete anteriorly, on II moderate to strong, coarsely granular, with larger posterior granules, on III strong, coarsely granule-crenulate with enlarged posterior granules, on IV obsolete, positions marked by scattered granules; posterior half of ventrosubmedian carina with 7–8 enlarged dentate granules on II, 6 on III; metasoma V with ventrosubmedian carinae obsolete, marked by series of enlarged dentate granules on anterior 3/4 of segment, ventromedian carina moderate, studded with small to large dentate granules; 5 denticles on ventral anal arc; lateral anal lobes split by single incision into broad dorsal lobe and dentate ventral lobe. Granulation: dorsal surfaces smooth on segments I–IV, smooth with scattered small granules on V, concave on I–III, convex on IV–V; dorsolateral surfaces smooth on segments I–III, with few small granules on I, finely granulate on IV–V; lateral surfaces smooth on I–III, finely granulate IV–V; ventrolateral and ventral surfaces smooth on I–II, lightly shagreened on III, densely shagreened on IV–V. Chaetotaxy: carinal macrosetae (left–right) dorsosubmedian: I, 0–0, II 1–0, III 2–2, IV 2–2, dorsolateral: I, 1–0, II 1–2, III 1–2, IV 2–2; ventrolateral: I 2–2, II 2–2, III 2–2, IV 2–2; ventrosubmedian: I 2–2, II 2–2, III 2–2, IV 2–2, V 3–3; intercarinal macrosetae: metasoma V 12–13 lateral setae, one postero-lateral pair on ventral surface.

Telson ( Figs. 45–46 ). Vesicle slim, width 0.69 times metasoma V width, dorsal surface smooth, lateral surface smooth with few small granules, ventral surface studded with medium to large granules, bearing numerous medium length reddish macrosetae: 1–2 on lateral surfaces, 14 on ventral surface, 2 on pedicel; aculeus slender, as long as vesicle.

Pedipalp ( Figs. 47–51 , 91–93 , 102 ). Femur ( Figs. 49 , 9 1 ): short, stout, 0.73 times carapace length, 2.42 times longer than wide; dorsoexternal, dorsointernal and ventrointernal carinae strong, with robust dentate granules; external surface smooth; dorsal surface flat, mostly smooth with scattered small granules and proximal cluster of 3–4 medium granules, ventral surface smooth, proximally concave, distally convex, with few small granules; internal surface with irregular small and large conical granules, internal carina obsolete; dorsoexternal carina with 1 distal macroseta, external surface with (left–right) 0–1 basal, 8–12 distal macrosetae. Patella ( Figs. 50–51 , 92 ): short, stout, 0.87 times carapace length, 1.95 times longer than wide; dorsointernal carina strong, granulose; ventrointernal carina strong, with large dentate granules; internal carina weak, marked by contiguous medium to coarse granules; other carinae obsolete; dorsal surface smooth with few fine granules, external surface smooth, ventral surface smooth with sparse fine granules along internal margin; ventrointernal surface smooth with several fine granules, dorsointernal surface finely granular. Chela ( Figs. 47– 48 , 93 ): relatively short, 1.37 times carapace length; manus extremely swollen, subglobose, width 0.65 times carapace length, fingers very short, movable finger 0.79 times ventral manus length; manus with dorsal surface densely, finely shagreened, external surface smooth to sparsely shagreened, ventroexternal surface densely, finely shagreened, ventral and ventrointernal surfaces smooth except for lightly shagreened proximal ventrointernal patch, internal surface sparsely to moderately shagreened with several small granules; all carinae obsolete; fixed and movable fingers smooth; dentate margins of finger straight, without proximal scalloping; closure of fingers leaves linear proximal gap, with tips of fingers apically divergent; ventral aspect of movable finger angled upwards near base; manus largely bare, with few short macrosetae; movable finger with numerous short macrosetae on ventral aspect, more dense apically; 7 primary denticle subrows on fixed and movable fingers; fixed fingers with 7 internal accessory denticles, 7–8 external accessory denticles; movable fingers with 7 internal accessory denticles, 6 and 8 external accessory denticles, tip of movable finger with 4 subdistal denticles ( Fig. 102 ). Trichobothrial pattern: neobothriotaxic, type Aβ (Vachon, 1974, 1975), with supernumerary esb 3 on patella; trichobothrium d 5 located slightly distal to e 2; femur d 2, patella d 2, chela Eb 3, Esb and esb petite; db on fixed finger level with esb, at base of fixed finger; eb on distal manus; dt at middle of fixed finger, approximately midway between est and et.

Legs ( Figs. 30–33 , 43–44 ). Leg I short with stout segments; leg II moderately slender; legs III–IV very slender; ventral carinae dentate on femur I, finely denticulate on femora II–III, weak, crenulate-denticulate on femur IV; tibial spur present on right leg III, minute and vestigial on left leg III, present on both legs IV; retrolateral pedal spurs on legs I–IV simple, without setae; prolateral pedal spurs basally bifurcate on legs I – III, basally trifurcate on leg IV, with (left–right) 1–1, 0– 1, 5–6, 5–4 setae; tibiae I–III with retrosuperior bristle-combs ( Fig. 43 ) with (left–right) 6–6, 8–8, 13–11 setae; basitarsi I–III with bristle-combs ( Fig. 43 ), retrosuperior series developed on I–III with (left–right) 9–8, 12–9, 17–16 setae; retroinferior series developed on II–III with 5–5, 11–10 setae; ventral surfaces of telotarsi with short fine macrosetae on legs I–II, medium length macrosetae on legs II–III ( Fig. 17 ), medium to long, fine macrosetae on legs III–IV.

Measurements of holotype male (mm). Total L 49.30; metasoma and telson L 33.00; carapace L 5.40, anterior W 2.84, posterior W 6.04, carapace preocular L 2.24; metasomal segments (L/W/D) I 4.47/3.44/2.77, II 5.18/3.35/2.92, III 5.37/3.35/2.91, IV 6.24/2.92/2.75, V 6.88/2.79/2.55; telson L 5.33; vesicle L 2.75, W 1.93, D 1.81; pedipalp chela L 7.40, chela manus ventral L 4.64, chela manus W 3.48, D 3.78, fixed finger L 2.24, movable finger L 3.65; pedipalp femur L 3.96, W 1.63, patella L 4.69, W 2.41; pectine L 5.36, leg III patella L 4.64, W 1.50.

Measurements of paratype female (adult from N of Adam) (mm). Total L 47.00; metasoma and telson L 26.60; carapace L 5.03, anterior W 2.73, posterior W 5.68, carapace preocular L 2.24; metasomal segments (L/W/D) I 3.61/2.87/2.41, II 4.18/2.64/2.49, III 4.26/ 2.61/2.59, IV 4.99/2.41/2.28, V 5.62/2.54/2.15; telson L 4.99; vesicle L 2.41, W 1.67, D 1.72; pedipalp chela L 5.93, chela manus ventral L 2.54, chela manus W 1.46, D 1.79, fixed finger L 2.58, movable finger L 3.61; pedipalp femur L 3.18, W 1.31, patella L 4.13, W 1.72; pectine L 3.10, leg III patella L 3.78, W 1.38.

Variation. Biometric data were obtained from 17 adults or sub-adults (14 males, 3 females). The body length was 34–50 mm (mean ± SD 41.8 ± 4.3 mm) and carapace length 3.96–6.79 mm (5.27 ± 0.64 mm). Pedipalp finger primary denticle subrows for adults: of 34 fixed fingers, there were 2 fingers with 6 subrows, 22 with 7, and 10 with 8; of 34 movable fingers, there was 1 finger with 5 subrows, 6 with 6, 24 with 7, and 3 with 8. The number of dentate granules on the posterior half of the ventrosubmedian carina of metasoma II and III: on II, 2 carinae with 5 granules, 13 with 6, 15 with 7, 4 with 8; on III, 5 carinae with 4 granules, 18 with 5, 11 with 6. Tibial spurs were either present or absent, and were absent more frequently on leg III than leg IV; of 30 leg III tibiae, spurs were present in 19 (63.3 %), and absent in 11 (36.7 %); of 29 leg IV tibiae, spurs were present in 25 (86.2 %), and absent in 4 (13.8 %). Variation in morphometrics is summarized in Figs. 106– 111 , Tab. 1. Trichobothrial distance ratio (d 5 – e 2)/(e 2 – e 1)> 0 (i.e. d 5 distal to e 2) for 27/28 femora (96.4 %).

Sexual dimorphism: females differed from males as follows: carapace smooth, lustrous with granulation weak to obsolete; tergites smooth, lustrous; pedipalp femur dorsally smooth, lustrous with only a few granules, internal surface mostly smooth with much smaller granules; pedipalp patella smooth, lustrous, with dorsointernal carina weak, finely granulated, ventrointernal carina weak with moderate to small granules ( Fig. 54 ); pedipalp chela manus not strongly swollen, smooth, lustrous ( Figs. 52–53 ); metasoma II–III with more prominent denticles on posterior half of ventrolateral and ventrosubmedian carinae; metasoma IV not as heavily granulated on lateral and ventral surfaces, with carinae weaker, more finely granular; ventrolateral and ventrosubmedian carinae on metasoma I moderate, almost smooth anteriorly, crenulate–granulate posteriorly; on II–III strong, crenulate–granulate anteriorly, dentate-granulate posteriorly; segment IV with ventrolateral carinae moderate, weakly granular; ventrosubmedian carinae obsolete; pectines much shorter ( Fig. 55 ), terminating well short of distal ends of coxae IV ( Figs. 35, 37 ). Pectine teeth: males 14–19 (of 28 combs from n = 14 males: 1 comb with 14 teeth, 4 combs with 15 teeth, 6 with 16, 7 with 17, 8 with 18, 2 with 19), females 9–14 (of 6 combs from n = 3 females: 1 comb with 9 teeth, 1 with 11, 2 with 12, 2 with 14). There was significant sexual dimorphism in morphometrics ( Table 1): females tended to have shorter pedipalp femur, patella, chela and metasoma I–V relative to carapace length, more slender pedipalp patella, and pedipalp chela movable finger longer relative to manus.

Juveniles: differed from adults as follows: dorsoexternal, dorsointernal and ventrointernal carinae on pedipalp femur weak to obsolete, with small granules; carinae on patella obsolete; chela manus not swollen in juvenile males; fewer macrosetae on the tibiae, basitarsi and telotarsi of all legs. In smallest juveniles (3 specimens, carapace lengths 1.53 mm, 1.59 mm, 1.65 mm) the tibia, basitarsus and telotarsus of legs I–III lacked macrosetae and bristle combs, and series of elongate spiniform processes (spine combs) extended from the retrosuperior margins of these leg segments (1– 2 processes on tibia, 6–7 on basitarsi, 3–5 on telotarsi). Spines consisted of a short, broad basal portion and a long, fine, needle-like distal portion.

Distribution. This species is known only from arid plains of north-central Oman, south or south-west of the Al Hajar mountains, at elevations of 100–420 m a.s.l.

Ecology. All specimens were collected from sandy soils on arid, alluvial fans and wadis in the hot interior desert of north-central Oman. The development of bristle combs on the tibia and basitarsus of legs I–III is consistent with psammophilous, arenicolous specialization (Polis, 1990; Fet, Polis & Sissom, 1998; Prendini, 2001a). Compared to the more ultrapsammophilous V. globimanus , this species was found on more compacted sandy substrates. Burrowing in firmer soils may be correlated with the more robust development of the pedipalps and metasomal segments ( Figs. 106–111 ), heavier carinae on the metasoma and pedipalp patella, and less elongate tarsal ungues ( Fig. 43–44 ).

Figures

Figures 30–37: Vachoniolus gallagheri, sp. nov., holotype male (30–33) and paratype female from N of Adam (34–37). Habitus viewed under reflected white light (30–31, 34–35) or UV fluorescence (32–33, 36–37). 30, 32, 34, 36. Dorsal aspect. 31, 33, 35, 37. Ventral aspect. Scale bars: 10 mm for both male and female.

Figures 38–46: Vachoniolus gallagheri, sp. nov. 38, 43–46, holotype male; 39–42, paratype male (same locality as holotype). 38. Carapace, dorsal aspect. 39. Right chelicera, dorsal aspect. 40. Right chelicera, ventral aspect. 41–42. Right hemispermatophore, lobes at base of flagellum. 41, convex aspect; 42 outer aspect. 43. Right basitarsus and telotarsus III, retrolateral aspect. 44. Right telotarsus III, ventral aspect. 45–46. Metasoma and telson. 45. Right lateral aspect. 46. Ventral aspect. Scale bar: 38: 3.0 mm; 39–40, 44: 1.5 mm; 41–42: 1.0 mm; 43: 2.1 mm; 45–46: 4.25 mm.

Figures 47–56: Vachoniolus gallagheri, sp. nov. 47–51, 56, holotype male; 52–55, paratype female (S of Nizwa roundabout). 47. Right pedipalp chela of male, external aspect. 48. Right pedipalp chela of male, ventral aspect. 49. Right pedipalp femur of male, dorsal aspect. 50. Right pedipalp patella of male, dorsal aspect. 51. Right pedipalp patella of male, external aspect. 52. Right pedipalp chela of female, external aspect. 53. Right pedipalp chela of female, ventral aspect. 54. Right pedipalp patella of female, dorsal aspect. 55. Left pectine, female, ventral aspect. 56. Left pectine, male, ventral aspect. Scale bar: 47–48, 55–56: 3.0 mm; 49–54, 2.0 mm. Trichobothrial notation after Vachon (1974).

Figures 88–99: Pedipalp segments of Vachoniolus, species, viewed under UV fluorescence. 88–90, V. batinahensis, sp. nov., holotype male; 91–93, V. gallagheri, sp. nov., holotype male; 94–96, V. globimanus, male from Barik; 97–99, V. iranus, male from Iran. 88, 91, 94, 97: pedipalp femur, dorsal aspect; 89, 92, 95, 98: pedipalp patella, dorsal aspect; 90, 93, 96, 99: pedipalp chela, external aspect. Scale bar: 88–89, 99: 2.0 mm; 90: 2.4 mm; 91–92: 1.7 mm; 93, 96: 2.7 mm; 94–95: 2.1 mm; 97–98: 1.3 mm.

Figures 100–105: Pedipalp finger dentition of Vachoniolus species in Oman. Each figure shows a sketch of the denticle patterns on the dentate margins of right fixed finger (left drawing), and right movable finger (right drawing). Position of internal fixed finger trichobothrium it is indicated. 100. V. batinahensis, sp. nov., holotype male. 101. V. batinahensis, sp. nov., paratype female (al Abyad). 102. V. gallagheri, sp. nov., holotype male. 103. V. gallagheri, sp. nov., paratype female (N of Adam). 104. V. globimanus, female (Marsawdad). 105. V. globimanus, male (Shigag).

Figures 106–111: Graphical illustration of variation in selected morphometric ratios of Vachoniolus. 106. Scatter plot of male variation in slenderness of metasoma I and relative length of pectine (metasoma I L/W vs. pectine L/ carapace L). 107. Scatter plot of variation in relative length of metasoma IV and slenderness of telson (metasoma IV L/ carapace L vs. telson L/W). 108. Scatter plot of male variation in slenderness of metasoma IV and relative proportion of pedipalp chela fixed finger (metasoma IV L/W vs. pedipalp chela fixed finger L/ manus L). 109. Scatter plot of male variation in slenderness of metasoma V and pedipalp femur (metasoma v L/W vs. pedipalp femur L/W). 110. Scatter plot of male variation in slenderness of pedipalp femur and patella (pedipalp femur L/W vs. pedipalp patella L/W). 111. Histogram plots showing distributions of the trichobothrial distance ratio (d5–e2)/(e2–e1) (data from both sexes). Vertical line is crossover point for d5 relative to e2. Note: each histogram bar represents the number of cases with ratio ≤ right abscissa tick mark; for V. globimanus, all data ≤ 0 are actually equal to 0 (i.e. no cases of d5 proximal to e2). The ratio distribution of V. batinahensis was significantly different from that of V. gallagheri (P = 0.0001), and the latter was also significantly different from that of V. globimanus (P = 4·10-6) (Mann-Whitney test). Color codes of all plots: green: V. batinahensis; yellow: V. gallagheri; magenta: V. globimanus; blue: V. iranus. Data in 106–110 from adults, in 111 from all ages.

Figures 146–148: Maps showing collection sites of Vachoniolus material examined in this study. 146. Records plotted on geopolitical map. Arrows: type localities for V. batinahensis and V. gallagheri, and approximate type locality of V. minipectenibus (= V. globimanus). 147. Records plotted on color topographic map with shaded relief. 148. Records plotted on terrain map obtained by satellite imagery. Green circles: V. batinahensis, sp. nov.; yellow circles: V. gallagheri, sp. nov.; magenta circles: V. globimanus.

Figures 9–17: Vachoniolus batinahensis, sp. nov. 9, 14–17, holotype male; 10–13, paratype male (same locality as holotype). 9. Carapace, dorsal aspect. 10. Right chelicera, dorsal aspect. 11. Right chelicera, ventral aspect. 12–13. Right hemispermatophore, lobes at base of flagellum. 12, convex aspect; 13 outer aspect. 14–15. Metasoma and telson. 14. Right lateral aspect. 15. Ventral aspect. 16. Right basitarsus and telotarsus III, retrolateral aspect. 17. Right telotarsus III, ventral aspect. Scale bar: 9, 16: 3 mm; 10, 11: 1.7 mm; 12, 13: 1 mm; 14, 15: 5.7 mm; 17: 1.5 mm.