Microbuthus satyrus, Lowe & Kovařík & Stockmann & Šťáhlavský, 2018

Lowe, Graeme, Kovařík, František, Stockmann, Mark & Šťáhlavský, František, 2018, Review of Microbuthus with description of M satyrus sp n (Scorpiones Buthidae) from Oman and Yemen, Euscorpius 263, pp. 1-22 : 4-20

Microbuthus satyrus sp. n.

( Figs. 1–31, 37, 40–45, 71, 77–78, 79, 83, 86, Table 1) http://zoobank.org/urn:lsid:zoobank.org:act:BFCF2

D4C-7E63-4070-AC45-87BAE0A3DAAF

TYPE LOCALITY AND TYPE REPOSITORY. Oman, Jabal Samhan , 17°11'10.1"N 54°56'34.3"E ( FKCP).

TYPE MATERIAL. Oman: Jabal Samhan , 17°11'10.1" N 54°56'34.3"E ( Fig. 79), 425 m a.s.l., 1♂ (holotype, No. 1381, Figs. 1–2, 5, 7, 9–13, 15–17, 22–31, 37, 40– 45, 79, 83), 1♂ 1♀ (paratypes, Figs. 3–4, 6, 8, 14, 18–21, 71), 29.X.2017, leg. M. Stockmann ( FKCP) . Yemen: Shabwah gov., S of An Nuqbah, Al Aram vill., 14° 13'48"N 047°04'59"E, 970 m a.s.l. ( Fig. 80), 22.X.2005, 1♀ (paratype), leg. D. Král ( FKCP) .

ETYMOLOGY. Named after deities in Greek mythology, the satyrs (σάτυρος, satyros).

DISTRIBUTION. Oman, Yemen.

DIAGNOSIS. Total length of adult 17 mm (males) to 22 mm (females); base color uniform coal black; legs, pedipalp femur and fingers yellow to orange, telson reddish brown; metasomal segments stout, L/W ratios: I 0.99–1.04 (♀ ♂), II 1.29–1.27 (♀ ♂), III 1.26–1.29 (♀ ♂), IV 1.29–1.20 (♀ ♂), V 1.32–1.24 (♀ ♂), segments IV–V swollen, ovate, wider than I–III, heavily sclerotized, ventral and lateral surfaces strongly convex, pitted, lacking macrosetae, dorsal surfaces widely, deeply excavated; posterior slope of dorsal surfaces with coarse granules; ventrolateral carinae on V distinct, with weak, blunt granules, extending only over posterior 1/4 of segment; telson widest at base, slightly tapering posteriorly; vesicle deepest near middle, ventral surface with scattered shallow depressions, ventromedian carina with 13–16 granules; legs III–IV with weak to moderate tibial spurs; basitarsi without bristle combs; pedipalps slender, femur L/W 3.53–3.8 (♀ ♂), patella L/W 3.57– 3.12 (♀ ♂); chela fixed finger with trichobothrium dt at 1/4 of length from tip; pectine teeth, male 13, females 10–11.

COLORATION ( Figs. 1–4). Base color uniform coal black, pedipalp manus brown to black; pedipalp femur and fingers pale yellow; pedipalp trochanter and patella orange to black, legs with basitarsi, telotarsi, and tibiae pale yellow; telson deep reddish brown, darker proximally; sternites III–V dirty yellow brown, VI dark brown, VII blackish brown.

CARAPACE ( Figs. 5–6). Strongly trapezoidal, W/L 1.39–1.26 (♀ ♂); anterior margin very weakly emarginate; surface densely, coarsely granular, with most carinae indistinct; anterior median carinae granular, divergent, becoming indistinct less than halfway towards anterior marginal shelf; median ocular tubercle prominent, raised, with large ocelli; 4 pairs of lateral eyes, including 2 major lower anterior ocelli, 2 minor lower and upper posterior ocelli; superciliary carinae indistinct, top of ocular tubercle flat, with curved row of granules above each eye; central median and posterior median carinae irregular, coarsely granulate, fused into reticular grid enclosing four smooth, oval depressions; posterior median furrow wide, shallow; sides of carapace steeply sloped, densely granular, lateral margins with finely granulated rim.

COXOSTERNAL AREA ( Figs. 7–8). All coxae granular, lacking macrosetae; coxa IV narrow, elongate, with moderately developed, granular marginal carinae; sternum type 1 (Soleglad & Fet, 2003), an irregular pentagon almost triangular in shape, coarsely granular, with deep, transverse postero-median sulcus; genital opercula smooth, with 2–4 lateral macrosetae; genital papillae present; genital opercula rounded triangular, lateral surfaces roughened; pectines with 3 marginal lamellae, 6 middle lamellae; fulcra bearing two short reddish setae; pectine teeth: males 13, females 10–11.

MESOSOMA ( Figs. 1–8). Pretergites with rough surface bearing scattered microgranules, posterior borders minutely granular; tergites I–VI densely granular, granulation fine on anterior half, coarse on posterior half; carinae indistinct on tergites I–III; tergites IV–VI with three short, longitudinal, granular carinae, very weak on IV, weak on V–VI; carinae confined to posterior half of tergites; tergites IV–VI with median and lateral carinae partially bridged by transverse row of granules; tergite VII with 5 broad, granular ridges; sternite III roughened medially, coarsely shagreened near lateral margins, densely micro-shagreened on anterolateral concavities above pectines; sternites IV–V densely shagreened, more coarsely along lateral margins, posteromedial area nearly smooth; sternite VI densely, coarsely shagreened, posteromedial area rugose-granulate, sternite VII with dense granulation; sternites IV–VI with weak median sulcus, VI with weak median carinae, VII without developed carinae; lateral margins of all sternites crenulate; sternites with posterior margins microdenticulate on III–V, granulate-crenulate on VI–V.

METASOMA ( Figs. 13–20, 37). Segments I–III with 6 weak to moderate granulate-crenulate carinae masked by heavy reticular granulation; intercarinal surfaces of I– III coarsely reticulo-granulate; segments IV–V wider and taller than I–III, strongly swollen, ventral and lateral surfaces convex, smooth, polished, with numerous shallow depressions or indentations, each giving rise to a very short, filiform microseta; dorsosubmedian and dorsolateral carinae indistinct, forming smooth blunt lips bordering dorsal anterior and posterior concavities; metasoma V with weak, ventrolateral carinae bearing weak, blunt granules, confined to posterior 1/4 of segment; other carinae of IV–V obsolete; anterior half of dorsal surface of segment IV forming deep, rugose, anteriorly sloped, V-shaped trough fitting the aculeus; posterior half a wide, granular, nearly flat slope; anterior part of dorsal surface of segment V smooth, irregular, strongly sloped, base concave with longitudinal striae; posterior slope of segment V deeply concave, smooth, with only faint median sulcus; lateral anal lobe smooth.

TELSON ( Figs. 13–20). Vesicle ovoid in dorsal view, slightly tapered, wider basally, narrowing distally; dorsal surface smooth, weakly convex; upper lateral surface with distinct curved, longitudinal sulcus; ventral surface smooth with scattered depressions anterolaterally, anterior part bearing prominent, serrulate ventromedian carina with 13–16 polished, widely transverse granules; ventromedian carina flanked on either side by weak furrow; ventral surface with scattered microsetae; subaculear tubercle absent; aculeus short, stout, bent sharply downward perpendicular to plane of dorsal surface of vesicle; tip of aculeus extending below deepest part of vesicle.

PEDIPALP ( Figs. 21–31). Femur, patella, and chela very slender, elongate. Femur: with 5 moderate, granular carinae; external carina broad, heavily granulated, encircling trichobothria e 1 and e 2; intercarinal surfaces roughened, shagreened to densely, finely granular. Patella: with 8 carinae; external carina strong, almost smooth; dorsoexternal, dorsomedian, dorsointernal, internal and ventrointernal carinae moderate, irregularly granulated; ventromedian and ventroexternal carinae weak, finely granulate; intercarinal surfaces roughened with dense, fine granulation dorsally, scattered fine granulation ventrally. Chela: manus with carinae weak or absent, weakly granulose; digital, dorsal secondary and dorsal marginal carinae weak to moderate on fixed finger, dorsal interior carina very weak on apical end of fixed finger, obsolete on more proximal fixed finger and manus; external surfaces roughened, with weak reticulate granulation, ventral and internal surfaces smoother; chela fingers elongate, smooth, vertically arched, leaving clear gap when closed; fixed finger deflected strongly upward at base, flanked externally and internally by rounded lobes bearing three elongate clavate microsetae; both fingers bearing enlarged, erect, spinoid denticles dividing dentate margins into weakly scalloped subrows of fine, widely spaced microdenticles; fixed finger with subdistal pair of denticles plus 3 additional enlarged denticles on margin; movable finger with subdistal pair of denticles and two enlarged denticles along margin, subrows with 5, 7–8, 10–11 microdenticles; fixed finger with external groove at base of enlarged terminal tooth, receiving terminal tooth of movable finger. Trichobothriotaxy: type Aβ, neobothriotaxic minorante ( Vachon, 1974, 1975), as diagnosed for the genus.

LEGS ( Figs. 9–12). Inferior carinae of femur and patella on all legs crenulate to serrate; intercarinal surfaces finely granular; legs III–IV with weak to moderate tibial spurs; inferior margins of basitarsi with two rows of 4–8 stout, spiniform setae; soles of telotarsi with two rows of several short setae; ungues short, stout; legs with two tarsal spurs, prolateral tarsal spurs basally bifurcate, retrolateral tarsal spurs simple.

CHELICERA. Dark reticulated pigmentation on movable finger, and distal dorsal and distal ventral areas of manus; fingers with normal buthid dentition ( Vachon, 1963; Sissom, 1990): movable finger with dorsal distal tine and ventral distal tine of equal size, dorsal margin with 4 denticles (subdistal, medial, two small basal), ventral margin with 2 denticles (large subdistal, small basal); fixed finger with large subdistal denticle and basal bicusp; ventral aspect of fixed finger with two prominent denticles, distal denticle slightly larger than proximal denticle; dense brush of long setae on ventral and internal aspect of fixed finger, and ventral aspect of movable finger.

HEMISPERMATOPHORE ( Figs. 42–45). Flagelliform; elongate, trunk ca. 6.6 times length of capsule; flagellum with short pars recta ca. 0.44 times length of trunk, with slight expansion along anterior margin, and longer, thinner, hyaline, cylindrical, gradually tapering pars reflecta; sperm hemiduct tripartite, with larger anterior and posterior lobes, and smaller acuminate median lobe; basal lobe horn-like, relatively tall, robust, a strongly curved hook with pointed tip, chord length about half length of median lobe.

AFFINITIES. The morphometrics of the metasomal segments, or their lack of dense pilosity, easily differentiate M. satyrus sp. n. from other Arabian Microbuthus ( M. gardneri and M. kristensenorum ), except for M. litoralis , which it more closely resembles (see Figs. 33– 37). However, M. litoralis has different colored legs and pedipalps, the pedipalp femur always being dark in M. litoralis ( Fig. 42) vs. yellow in M. satyrus sp. n. ( Figs. 1–4, 28–30). Also, metasoma V has weak ventrolateral carinae, with only blunt granules, confined to the posterior 1/4 of the segment in M. satyrus sp. n. ( Figs. 40– 41), whereas these carinae are strong, armed with sharp granules and extend over the posterior 1/3 of the segment in M. litoralis ( Figs. 38–39). Finally, the posterior ventral surface of metasoma V bears dense, strong granulation in M. litoralis , but is only weakly granulated to smooth M. satyrus sp. n.

Among the North African taxa: M. maroccanus differs from M. satyrus sp. n. in color, with orange legs and pedipalp fingers, and fuscous pedipalp femur ( Fig. 70), and in metasomal segment V being furnished with well developed ventrolateral carinae extending over at least the posterior 2/3 of the segment, that bear large dentate granules, and the segment has dense, strong granulation on its posterior ventral surface ( Fig. 32); M. flavorufus differs, e.g., in its light yellow to reddishyellow color of carapace and mesosoma, and in having more stout metasomal segments, ♀ L/W ratios of metasoma I –V: 0.81, 1.00, 1.06, 0.95, 1.25 (data from Lourenço & Duhem, 2007), respectively, vs. 0.97, 1.30, 1.27, 1.29, 1.32 in M. satyrus sp. n., and in particular, metasoma IV is wider than long in M. flavorufus , vs. longer than wide in M satyrus sp. n..

ECOLOGY. The type locality is in a rocky wadi with boulders, surrounded by mountains on the southern aspect of the Jabal Samhan escarpment facing the coastline. The area is well vegetated with small trees and shrubs, and the substrate is clay soil with many rocks. The climate in this area is mild year round, with temperatures around 30°C and a wet season corresponding to the Khareef monsoon from June to Sep- corresponds to posterior width), depth (D).

tember. Two males were located by UV detection a few hours after sunset. They were resting in ambush positions on rocks at the top of the cliffs close to the wadi. Other species of scorpion observed in the area were: Nebo whitei Vachon, 1980 , Hottentotta salei (Vachon, 1980) , Butheolus gallagheri Vachon, 1980 and Compsobuthus acutecarinatus (Simon, 1882) .

Selection of habitat and climatic conditions of M. satyrus sp. n. are similar to those of the geographically proximate species M. kristensenorum , which occurs in surrounding areas of coastal Dhofar ( Fig. 86). However, within Dhofar, M. satyrus is so far known only from the type locality at the base of the Jabal Samhan escarpment. It is possible that this area offers unique environmental conditions that favor this species. Another buthid scorpion, Leiurus heberti Lowe, Yagmur et Kovařík, 2014 , also appears to be localized to the region of Jabal Samhan, while a related species, L. haenggii Lowe, Yagmur et Kovařík, 2014 occurs in surrounding areas ( Lowe et al., 2014).

A highly gravid female was found during daytime at the base of a larger tree in a small hollow under a stone, where conditions were more humid and cooler. A few days later, the female gave birth to 7 young, which dispersed from the female mesosoma six days later. This contrasts with the smaller litter sizes reported for M. fagei (3–4 young; Lourenço, 2007), and M. litoralis (4 young; Lourenço, 2011a), but corresponds with our experience with wild caught specimens of other Microbuthus species kept in captivity. Various broods in the breeding stock of M.S. were observed, in which M. kristensenorum produced several litters with 3–10 newborns, and M. gardneri , two litters with 6 and 7 newborns.

Comparative Microbuthus material examined, including new records.

Microbuthus maroccanus Lourenço, 2002

Morocco: Tan-Tan Province , 8 km SE of Tarfaya , 27°53.61'N 012°52.52'W ( WGS84 ), 5.II.2005, 1♀ 2♂ 1im., leg. R. et H. Fouquè et S. Bečvář ( FKCP) . Western Sahara: 55 km S of Boujdour , 25°37.69'N 014 °

39.80'W ( WGS 84), 6.II.2005, 2♂ 7♀ ( Figs. 32, 70), leg. R. & H. Fouquè & S. Bečvář ( FKCP); 150 km S of Boujdour , 24°48,85'N 014°51.31'W ( WGS84 ) , 6–7.II. 2005, 1♀ 2ims, leg. R. et H. Fouquè et S. Bečvář ( FKCP); 20 km SE of El Aaiún , 27°03.04'N 013° 03.14'W ( WGS84 ) , 8–9.II.2005, 1♂ 1♀ leg. R. et H. Fouquè et S. Bečvář ( FKCP) .

Microbuthus gardneri Lowe, 2010 Oman: Jabal Bani Jabir , 22.813172°N 59.058884°

E, 1730 m a.s.l., 1♂ 1♀ ( Figs. 33, 46–49, 74, No. 1382),

31.X.2017, leg. M. Stockmann ( FKCP).

Microbuthus kristensenorum Lowe, 2010

Oman: Dhofar Province, Wadi Nashib , ca 20 km N of Salalah ( Fig. 81), 25–28.VIII.2007 , 1♀, leg. J. Horák ( FKCP); 1 km E of Qayrun Hariti , Dhofar, 17°15'48.7"N 54°05'15"E, 848 m a.s.l., 1♀, IX.2013, leg. T . Mazuch et P. Novák ( FKCP); Arlit , 16°49.73'N 53°19.65'E, 1036 m a.s.l., 23.III.2014 , 1♀ and 1♀ 1♂ reared from ♀ litter ( Figs. 34, 50–53, 73, No. 906), leg. D. Hoferek ( FKCP); W of Qairoon Hairitti , 17.256408°N 54.022032°E, 805 m a.s.l. , 1♂ 1♀ ( Figs. 35, 54–57, 72, No. 1380), 28.X. 2017, leg. M. Stockmann ( FKCP) .

Microbuthus litoralis (Pavesi, 1885)

Djibouti: Gulf of Aden, Tadjura Bay , 1♀ juv. (holotype of Microbuthus pusillus Kraepelin, 1898 ) ( ZMUH); Obok, II.1893, 2♀ ( ZMUH); northeast, 8.V. 2009, 1♀ 4juvs., leg. T. Anthony ( ZMUH No. A 23/11) .

Eritrea: S of Assab , IV.1982, 1♀ (neotype), leg. P. M. Brignoli ( ZMUH No. A 22/11) ; Assab , XII.1899, 1♂ ( ZMUH) ; Dese Island , 15°26'39.2"N 39°45'32.7"E, 8 m a.s.l. ( Fig. 82), 5–7.XI.2015 (Locality No. 15 EJ) , 1♀ im. ( Figs. 36, 38–39, 64–65), leg. T. Mazuch & F. Kovařík ( FKCP) . Yemen: Al Hudaydah gov., Wadi Zabid (E Zabid ), 14°09'N 043°31'E, 325 m a.s.l. ( Fig. 80), 22– 23.III.2007, 1♀ im., leg. D. Král ( FKCP) ; Ta’izz gov., N of Al Makha by road, 13°23'37"N 43°16'22"E, 5 m a.s.l., 28.X.2007, 1♀ 1♂ ( Figs. 66–69), leg. David Král (F KCP) .