Butheolus harrisoni, Lowe, 2018

Lowe, Graeme, 2018, The genera Butheolus Simon, 1882 and Xenobuthus gen. nov. (Scorpiones: Buthidae) in Oman, Euscorpius 261, pp. 1-73 : 15-26

Butheolus harrisoni , sp. n.

( Figs. 67–162 , 333 , 338 , Tabs. 2–5 )

http://zoobank.org/urn:lsid:zoobank.org:act:CCE68 C77-AC8B-4D80-AECD-5EF8297554F

Butheolus gallagheri: Wood, 1993: 17 .

TYPE MATERIAL. Holotype ♂, Oman, Jabal Qara , Salalah-Thumrait road , UV detection, arid rocky wadi , edge of Nejd Desert , 17°17.26'N 54°05.36'E, 800 m a.s.l., 17.X.1993, 20:53 h, leg. G. Lowe ( NHMB).

PARATYPES. Oman: 3♂, 6♀, 1 juv ♂, same locality as holotype ( NHMB 3♂, 4♀; ONHM 1♂, 2♀) ; 1♂, Jabal Qara , 6.IX.1989, leg. M.D. Gallagher MDG 8146.2 ( ONHM 1351 ) ; 6♂, 7♀, 1 juv♂, 1 juv♀, Jabal Qara ; north slopes, Nejd, UV detection, rocky wadi & rocky slopes , 17°17.83'N 54°05.11'E, 800 m a.s.l., 16.X.1993, 22:38 h, leg. G. Lowe ( BMNH 1♂; MNHN 1♂; NHMB 3♂, 6♀; USNM 1♂, 1♀ ); 1♀, Jabal Qara , north slopes, Nejd Desert, UV detection, vegetated wadi bottom, chirping crickets, 17°17.01'N 54°05.81'E, 520 m a.s.l., 16.X.1993, 23:39 h, leg. G. Lowe ( MCZ) ; 1♀, Jabal Qara , Salalah-Thumrait road, UV detection, grassy/ rocky plateau with low hills and slopes, just south of police checkpoint, 17°15.98'N 54°04.88'E, 850 m a.s.l., 17.X.1993, 20:04 h, leg. G. Lowe ( MNHN) ; 4♂, 2♀, 2 juv ♂, 1 juv ♀, Jabal Qara , Salalah-Thumrait road , UV detection, wide rocky wadi, nr edge of wadi and rocky flats , 17°17.58'N 54°04.97'E, 800 m a.s.l., 17.X.1993, 21:45 h, leg. G. Lowe ( NHMB); 1♀, Jabal Qara , Nejd Desert, road to Ayun, UV detection, edge of wadi, major wadi below Ayun , fine silt and gravel in middle of wadi, rocky slopes and large rock formations on edges, day to dusk (few scorpions), 17°13.4'N 53°54.36'E, 600 m a.s.l., 20.X.1993, 18:36 h, leg. G. Lowe ( BMNH) ; 1♂, 2♀, 3 juv ♂, 1 juv ♀, Jabal Qamr , wadi between steep winding roads, UV detection, on gravel in wide wadi with boulders/ rounded rocks, 16°52.3'N 53°43.28'E, 40 m a.s.l., 27.IX.1995, 23:25 h, leg. G. Lowe, M.D. Gallagher (GL); 7♂, 2♀, 1 juv ♂, E of Jabal Qamr , along coastal wadi, UV detection on slope and gravel at base of slope, vegetated slopes with rocks, soil suitable for burrowing, 16°53.71'N 53°48.75'E, 15 m a.s.l., 28.IX. 1995, 00:30 h, leg. G. Lowe, M.D. Gallagher ( ONHM) ; 1 subadult ♂, Jabal Qamar , 2 km E of Ardit (Loc. F/10), windswept plateau with small rocky wadis, under rocks on slope, 16°50.7'N 53°23.69'E, 1060 m a.s.l., 1.I.1999, 11:00–16:30 h, leg. A. Winkler, B. Winkler ( ZSM) ; 1♀, Wadi Uyun (Loc. F/12), sand alluvium under big bush under stone, 17°14.39'N 53°53.99'E, 400 m a.s.l., 2.I.1999, 16:30 h, leg. A. Winkler ( ZSM) ; 1♀, Wadi Uyun (Loc. F/12), sand alluvium, under big bush under stone, 17°14.39'N 53°53.99'E, 400 m a.s.l., 3.I.1999, 10:00–12:00 h, leg. A. Winkler ( ZSM); 1 juv ♀, Jabal Qamar , 2 km E of Ardit (Loc. F/15 II), UV detection, windswept plateau with rocky wadis, between small shrubs, 16°50.7'N 53°23.69'E, 1060 m a.s.l., 31.I.2000, 16:00–21:00 h, leg. B. Winkler ( ZSM); 1 juv ♂, Jabal Qamar , 3 km E of Ardit (Loc. F/13), UV detection, windswept plateau with rocky wadis, under small stone, 16°50.71'N 53°23.72'E, 1076 m a.s.l., 19.XII.2001, 18:00–20:30 h, leg. A. Winkler ( ZSM) ; 1♂, 2♀, Jabal Qamar , 3 km E of Ardit (Loc. F/14), UV detection, windswept plateau with rocky wadis, under small stone, 16°50.87'N 53°23.77'E, 1067 m a.s.l., 19.XII.2001, 21:00–22:15 h, leg. A. Winkler ( ZSM) .

DIAGNOSIS. A member of the genus Butheolus differentiated as follows: small scorpions, adults 25–35 mm; bicolored, carapace and mesosoma moderate to dark brown but usually lighter on posterior tergites, metasomal segments IV–V and telson dark brown, metasoma I–II lighter brown to yellow, pedipalps and legs yellow except for fuscosity at base of chela fingers; carapace and tergites with mixture of fine and coarse granulation; pedipalp patella with dorsomedian carina complete; pedipalp chela with carinae weak or obsolete; ventral surface between ventromedian carinae of metasoma I with sparse, fine granulation or smooth in females; ventral and lateral intercarinal surfaces of metasoma II–V with moderately coarse granulation; metasoma and telson with sparse, short macrosetae; telson with subaculear tubercle small or absent, obtuse angle between posterior vesicle surface and aculeus base; pedipalp and metasoma moderately stout; pedipalp femur L/W ♂ 2.72–3.23, ♀ 2.29–2.47; pedipalp patella L/W ♂ 2.84–3.46, ♀ 2.33–2.64; pedipalp chela L/W ♂ 4.90–5.48, ♀ 4.07–4.67; metasoma I L/W ♂ 0.81–0.91, ♀ 0.74–0.83; metasoma IV L/D ♂ 1.43–1.69, ♀ 1.37– 1.54, metasoma V L/D ♂ 1.92–2.28, ♀ 1.86–2.10, telson L/D ♂ 2.33–2.73, ♀ 2.27–2.51; pectine teeth ♂ 17–21, ♀ 14–17; basitarsus III retrosuperior setae 6–10.

COMPARISONS. Most similar to B. gallagheri , from which it differs in the following characters: pale color of metasoma I–II, coarser granulation on carapace and tergites, complete dorsomedian carina on pedipalp patella, subaculear tubercle on telson vesicle small or indistinct; on average, higher numbers of setae in basitarsal bristlecombs ( Figs. 148–149 ), more robust pedipalp femur and patella ( Figs. 151, 157 ), wider and deeper pedipalp chela manus ( Fig. 136 ) relative to carapace.

ETYMOLOGY. A patronym in honor of Ian D. Harrison, who collected and contributed much material in support of studies on the scorpion fauna of Oman.

DESCRIPTION.

Based on holotype ♂, 32 ♂ and 29 ♀ paratypes.

Coloration ( Figs. 69–100 , 119–122 , 127–132 ). Bicolored or with variable fuscous pattern; carapace and tergites vary from dark brown to light yellowish-brown, tergite VII may be lighter than others; metasomal segments I–II range from yellow to light brown, IV–V dark brown, III more variable, ranging from bright yellow to a dark brown matching IV–V; telson dark reddish brown, not as dark as metasoma IV–V, vesicle with pale spots at bases of macrosetae and a pair of pale stripes on ventral surface; carapace slightly lighter in postocular, posterior-median triangular area, tergites with lighter lateral patches in transverse reticulated bands; dorsal aspect of chelicerae dark on fingers and distal margin of manus, yellow-brown reticulated on distal quarter 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, and dorsal femur and patella may occasionally have weak fuscous markings that leave pale spots at the bases of trichobothria; coxosternal area and ventral surface of mesosoma yellow, sternite VII may be light brownish-yellow.

Carapace ( Figs. 69 , 71 , 73 , 75 ). Strongly trapezoidal, W/L 0.78–0.97, posterior W/ anterior W 2.00 – 2.46; 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 8–10 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 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 to coarse granulation except for smooth patches on postocular plateau, posterior transverse and posterior marginal furrows; granulation fine on lateral flanks, coarse on interocular triangle, moderately coarse on posterior margin of carapace and borders of postocular triangular plateau; granulation weaker in females than males; superciliary carinae of females smooth, of males smooth above, granulated anteriorly and posteriorly.

Chelicerae ( Figs. 67–68 ). 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 2–3 pale microsetae.

Coxosternal area ( Figs. 70 , 72 , 74 , 76 ). Males. Coxa I coarsely granulated, endite smooth on anterior half, granulated on posterior half; coxa II coarsely, irregularly granulated, endite with granules concentrated along midline, surface smooth on medial and lateral margins; coxa III irregularly, coarsely granulated, granules concentrated along anterior margin; coxa IV with numerous coarse granules in anterior marginal band and posterior marginal row, central smooth strip in narrow anterior half of coxa, finely granulated in broad posterior half; distal anterior surface of coxae II–IV with reduced granulation or smooth; coxae I–III with scattered, mostly anterior macrosetae: coxa I 7–11, II 12–18, III 4–8; coxa IV typically with single macroseta on anterior proximal limit; sternum weakly granulated, subtriangular, with deep posteromedian pit, usually 2 macrosetae; genital opercula with fine granulation near anterior margins, otherwise smooth, with 3–7 macrosetae, posterolateral margins concave. Females. Coxa I with weak coarse granulation, smooth on anterior part of endites; coxa II with very weak granulation, almost smooth; coxa III smooth except for anterior marginal granulation; coxa IV smooth except for anterior and posterior marginal fine granulation; sternum smooth, with larger median pit; genital opercula smooth, elongate; setation as in male, but with longer setae.

Pectines ( Figs. 70 , 72 , 74 , 76 ). Basal piece with concave anterior margin and small median groove and pit, coarsely granulated in males, smooth anteriorly and finely shagreened posteriorly in females, bearing 2–8 macrosetae; pectines with 3 marginal lamellae, 5–7 middle lamellae, extending to proximal 1/5–1/3 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. 123–126 ). Flagelliform, trunk elongate, ca. 6 times length of capsule region; flagellum short with thicker pars recta and thinner pars reflecta; 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. 69–76 , 333 ). 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 weak; 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 or shagreened, laterally shagreened or finely granular; sternite VI with smooth or obsolete inner lateral carinae, smooth or 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, non-contiguous, closely spaced, digitate denticles; sternite VII densely, finely granular, with granulate median and lateral pairs of carinae confined to posterior 3/4–4/5 of sternite, only median pairs extending to posterior margin; all sternites with scattered, sparse macrosetae, sternite VII with 4 stereotypic isolated macrosetae on mid- to anterior external sides 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 smooth traces of outer lateral carinae on VI; posterior marginal denticles of sternites III–VI smaller than in males; sternite VII with two pairs of weakly granular or almost smooth carinae, weaker than carinae of males, with weaker dense, fine granulation or shagreened on median and lateral surfaces; mesosoma much wider in females than males.

Metasoma ( Figs. 69–70 , 73–74 , 127–132 ). Moderate in length with robust segments, metasoma + telson L/ carapace L ♂ 4.8–5.8, ♀ 4.7–5.5; carination: segments I–III with 10 complete carinae, IV with 4 complete carinae (dorsosubmedian and dorsolateral carinae usually visible only on anterior 1/2 of segment), V with 2 carinae (ventrolateral) strongly developed, dorsolateral carinae may be weakly developed and only visible in anterior part of segment; 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; females with coarser granulation of carinae than males, with segment I having weaker granulation on ventrolateral and ventral carinae; lateral anal margin with 2 blunt granules or lobes, ventral anal margin with 13–18 granules; intercarinal surfaces: segment I with dense, fine granulation on lateral surfaces, weakly shagreened on ventrolateral and ventral surfaces; II–V with dense, fine granulation on lateral, ventrolateral and ventral surfaces, V with some coarser granules on posterior 1/3 of ventral surface; 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, granulation less dense and coarser on all surfaces, segments I–II may have ventrolateral and ventral surfaces sparsely granulated or smooth; dorsal surfaces in female smooth except for few isolated small granules, usually on segments I – II; setation: scattered microsetae and sparse, short macrosetae present on all segments, macrosetae slightly longer in females; dorsal surfaces lack setation, except for metasoma V which may bear 2–4 macrosetae along dorsolateral margins.

Telson ( Figs. 69–70 , 73–74 , 127–132 ). Vesicle smooth dorsally, rugose and weakly granulated laterally and ventrally with small granules more apparent on anterior-ventral surface, ovoid with gentle posterior slope; subaculear tubercle very weak or indistinct; smooth lateral and ventrosubmedian longitudinal troughs weak; scattered microsetae and short macrosetae on lateral and ventral surfaces; rugose surfaces arise from development of shallow indentations at setal insertion points; aculeus stout, shorter than vesicle.

Pedipalps ( Figs. 77–118 ). Males ( Figs. 77–88 , 101– 103, 107–110, 113–115 ). Femur: L/ W 2.72 –3.23; dorsointernal, dorsoexternal and ventrointernal carinae strong, with uniform coarse granulation; external carina strong, weakly granular, with fine granules proximally, larger granules distally; internal carina moderate with medium to coarse granules; dorsal surface with fine and coarse granulation, internal surface densely finely granulated or shagreened, external surfaces weakly granulated or shagreened, nearly smooth in parts, ventral surface with sparse mix of medium and fine granules; 7–12 accessory macrosetae on distal external surface; patella: L/ W 2.84 –3.46; dorsointernal carina strong, granular; dorsomedian carina strong, granular, complete; dorsoexternal carina moderate, weakly granular; external carina moderate, smooth; ventroexternal and ventromedian carinae weak or obsolete, indicated by rows of fine granules; ventrointernal carina weak, indicated by coarse granules; internal carina moderate, finely granulated; dorsal, external and internal surfaces with sparse fine granulation or sparsely shagreened; ventral surface with very sparse fine granulation, or smooth; chela: slender, L/ W 4.90 –5.48, carinae obsolete with only weak, smooth traces of digital, exterior secondary and exterior marginal carinae discernible at base of manus; surface smooth with sparse macrosetae and microsetae; 5–8 primary denticle subrows on movable finger, 5–7 on fixed finger (including cases of fusion of proximal rows), subrows except proximal typically flanked by internal and external accessory denticles; 4–8 internal or external accessory denticles on movable finger, 4–7 on fixed finger. Females ( Figs. 89–100 , 104–106, 111–112, 116– 118 ). Femur: more robust than in males, L/ W 2.29 –2.47; dorsointernal and dorsoexternal carinae moderate, with uniform medium to coarse granules; external carina weak, smooth, distal half with sparse, coarse granules; ventrointernal carina moderate with uniform coarse granules; internal carina weak with irregular large granules; dorsal surface with moderate cover of mixed fine granules or microgranules, internal and ventral surfaces almost smooth with sparse weak, fine granules or weakly shagreened, external surfaces smooth; 6–9 accessory macrosetae on distal external surface, including linear series of 4–6 on ventral side of external carina; patella: more robust than in males, L/ W 2.33 –2.64; dorsomedian carina complete; dorsointernal, dorsomedian and dorsoexternal carinae weak, smooth, other carinae obsolete; all intercarinal surfaces smooth; chela: more robust than in males, L/ W 4.07 –4.67, all carinae obsolete, surface smooth with sparse macrosetae and microsetae; finger dentition similar to males, 6–8 primary denticle subrows on movable finger, 6–7 on fixed finger, 5–8 internal or external accessory denticles on movable finger, 5–7 on fixed finger. Trichobothriotaxy: orthobothriotaxic, type Aβ ( Vachon, 1974) ( Figs. 101– 118 ).

Legs (69–70, 73–74, 119–122). Males: Femur and patella I–IV with strongly serrate-denticulate inferior carinae, other carinae denticulate-granulate; prolateral surfaces of femur I–IV with dense, fine granulation, of patella I–IV weakly shagreened, nearly smooth; females: femur and patella I–IV with weakly serrate-crenulate inferior carinae, other carinae weakly granulate; prolateral surfaces of femur I–IV mostly smooth, sparsely shagreened or granulate on proximal parts, of patella I– IV nearly smooth; in both sexes, tibia III–IV with spurs; retrolateral tarsal spurs simple, prolateral tarsal spurs basally bifurcate; basitarsi I–III with 4–10 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 2 .

Variation. Color patterns were variable. The carapace and tergites ranged from a lighter dusky brown to darker brown. Metasoma III color ranged from light brown to dark brown in Jabal Qara specimens, and from yellow to light brown in Jabal Qamr specimens. At a single collection site in Jabal Qara, examples of both light and dark brown metasoma III were observed. Meristic and morphometric variation are summarized in Tabs. 3–5 , and Figs. 133–162 .

DISTRIBUTION. Recorded from inland sites in Jabal Qara mountains and coastal sites around Jabal Qamr, in the Dhofar Province of Oman ( Fig. 338 ).

ECOLOGY. The species occurs in mesic environments along the coast as a characteristic faunal element of lush woodland or grassland zones of south facing slopes of the Jabal Qara ( Sale, 1980). Collections were made in wadi bottoms among small shrubs. However, it can tolerate more xeric conditions, as there are records from semi-arid scrub zones of northern Jabal Qara near the edge of the Nejd Desert, and from the plateau of Jabal Qamr. Recorded elevation range was 15–1,076 m a.s.l. The substrate preference was rocky or gravelly wadis, and some were found by day in shallow scrapes under rocks. Scorpions that occurred together with B. harrisoni sp. n. were: Compsobuthus acutecarinatus ( Simon, 1882) , Hottentotta salei ( Vachon, 1980) , Leiurus haenggii Lowe et al., 2014 , Nebo whitei Vachon, 1980 , Microbuthus kristensenorum Lowe, 2010 , and Xenobuthus xanthus sp.n.. The two similar species, B. harrisoni and B. gallagheri , have so far not been recorded together at the same site (UV detection sampling at 14 separate sites for B. harrisoni , 3 for B. gallagheri ). The lack of overlap may reflect spatial partitioning according to local environmental factors and competitive exclusion in preferred habitats of each species.

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 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 69–70: Habitus of male Butheolus harrisoni sp. n. Fig. 69. Dorsal aspect. Fig. 70. Ventral aspect. Holotype male (♂), Oman, Jabal Qara, Salalah-Thumrait road, 17°17.26'N 54°5.36'E, 17.X.1993, leg. G. Lowe. Scale bar: 5 mm.

Figures 71–72: Prosoma and mesosoma of male Butheolus harrisoni sp. n. Fig. 71. Dorsal aspect, carapace and tergites. Fig. 72. Ventral aspect, coxosternal area and sternites. Holotype male. Scale bar: 1 mm.

Figures 73–74: Habitus of female Butheolus harrisoni sp. n. Fig. 73. Dorsal aspect. Fig. 74. Ventral aspect. Paratype female, same locality data as male in Figs. 67–68. Scale bar: 5 mm.

Figures 75–76: Prosoma and mesosoma of adult female Butheolus harrisoni sp. n. Fig. 75. Dorsal aspect, carapace and tergites. Fig. 76. Ventral aspect, coxosternal area and sternites. Paratype female, same locality data as male in Figs. 67–68. Scale bar: 1 mm.

Figures 77–100: Pedipalp of Butheolus harrisoni sp. n. Figs. 77–88. Holotype male (♂). Dorsal (77–79), external (80–82), ventral (83–85) and internal (86–88) aspect of femur (77, 80, 83, 86), patella (78, 81, 84, 87) and chela (79, 82, 85, 88). Figs. 89– 100. Paratype female (♀), same locality data as male in Figs. 67–68. Dorsal (89–91), external (92–94), ventral (95–97) and internal (98–100) aspect of femur (89, 92, 95, 98), patella (90, 93, 96, 99) and chela (91, 94, 97, 100). Scale bars: 1 mm.

Figures 101–118: Trichobothrial maps and chela finger dentition of Butheolus harrisoni sp. n. Holotype male (♂): dorsal (101), external (102) and ventral (103) aspect of chela; dorsal (107) and external (108) aspect of patella; dorsal (113), external (114) and internal (115) aspect of femur; dentition of movable finger (MF) (109) and fixed finger (FF) (110). Paratype female (♀), same locality data as male in Figs. 67–68: dorsal (104), external (105) and ventral (106) aspect of chela; dorsal (111) and external (112) aspect of patella; internal (116), dorsal (117) and external (118) aspect of femur. Scale bars: 1 mm (♂, ♂ dentition, ♀).

Figures 119–122: Leg tarsi and tibiae of male Butheolus harrisoni sp. n. Holotype male. Retrolateral aspect, left leg I (119), II (120), III (121), IV (122) tibia, basitarsus and telotarsus. White light illumination. Scale bar: 1 mm.

Figures 123–126: Hemispermatophore of Butheolus harrisoni sp. n. Fig. 123. Whole right hemispermatophore, convex aspect. Scale bar: 1 mm. Figs. 124–126. Capsule region of right hemispermatophore, anterior (124), convex (125) and posterior (126) aspects. In convex view (125), the capsule is compressed to show outlines of lobes. Paratype male, same locality data as male in Figs. 67–68. White light illumination. Scale bar: 200 µm.

Figures 127–132: Metasoma and telson of Butheolus harrisoni sp. n. Figs. 127–129. Holotype male (♂). Dorsal (127), right lateral (128) and ventral (129) aspects. Figs. 130–132. Paratype female (♀), same locality data as male in Figs. 67–68. Dorsal (130), right lateral (131) and ventral (132) aspects. Scale bars: 2 mm.

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).