Leiurus hebraeus ( Birula, 1908 )

Lowe, Graeme, Yağmur, Ersen Aydın & Kovařík, František, 2014, A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula, Euscorpius 191, pp. 1-129 : 73-79

Leiurus hebraeus ( Birula, 1908) stat. n.

( Figs. 60–71, 87F, 88F, 89F, 90F, 91G, 92K, 93I–J, 94D–E, 95, 98–99, Tabs. 3A, 5)

Buthus quinquestriatus hebraeus Birula, 1908: 124–129 .

REFERENCES

Scorpio occitanus: Audouin in Savigny, 1826: 173, pl. VIII, fig. 1 (in part); Audouin in Savigny, 1827: 410-411, pl. VIII, fig. 1 (in part).

Buthus quinquestriatus: Kraepelin, 1891: 58-60 , pl. I, fig. 7, pl. II, fig. 38 (in part); Pocock, 1891: 242– 243 (in part); Pavesi, 1895: 4; Kraepelin, 1899: 27- 28 (in part); Simon, 1892: 83 (?); Davydov, 1898: 41–43; Schenkel, 1932: 380; Werner, 1934: 269 (in part); Werner, 1935: 211; Bodenheimer, 1937: 235; Whittick, 1955; Whittick, 1970.

Buthus quinquestriatus hebraeus: Birula, 1910: 118 .

Buthus ( Buthus) 5- striatus hebraeus: Birula, 1910: 170 .

Buthus ( Buthus) quinquestriatus: Birula, 1917a: 23 , 213 (in part); Whittick, 1941: 43 (in part).

Leiurus quinquestriatus: Vachon, 1950a: 197 (in part); Stahnke, 1972: 130 (in part); Lamoral & Reynders, 1975: 509–510 (in part); Wahbeh, 1976: 89; Levy & Amitai, 1980: 47–48 (in part); Warburg et al., 1980: 206; Kinzelbach, 1984: 100, Fig 2; Kinzelbach, 1985: map II (in part); Amr et al., 1988: 373; El- Hennawy, 1988: 14–15, 18; El-Hennawy, 1992: 101, 125–126 (in part); Amr & El-Oran, 1994: 186; Kovařík, 1998: 112 (in part); Kabakibi et al., 1999: 80, fig. 1; Amr & Abu Baker, 2004: 237–238, 241; Yağmur et al., 2009: 2–3 (in part), fig. 10–11; Shehab et al., 2011: 335–337, fig. 2D, tabs. 1–2.

Leiurus quinquestriatus hebraeus: Vachon, 1966: 212 ; Levy, Amitai & Shulov, 1970: 231–242 (in part); Levy & Amitai, 1980: 48–53, fig. 47–51, map 3, Appendix (in part); El-Hennawy, 1992: 101, 126; Sissom, 1994: 23; Fet & Lowe, 2000: 156–157; Yağmur et al., 2009: 12; Khalil & Yağmur, 2010: 2.

Leiurus quinquestriatus voelschowi: Pohl, 1967: 209– 215 , figs. 1–4; Kovařík, 1997: 180 (in part, record from Jordan).

Leirus (sic) quinquestriatus hebraeus: Levy & Amitai, 1980 : Appendix.

TYPE MATERIAL. Lectotype adult ♀, 4 immature paralectoptypes, Jordan, Wadi ‘ Arrud ( ZISP 578 ) (not examined, Figs. 70–71).

OTHER MATERIAL EXAMINED. Israel: 3 ♂, 7 ♀, 15 juvs, Palaestina (ev. Jordanien) ( NHMB 17 a) ; 1 ♀, Palaestina (ev. Jordanien), Jericho , 31°30.6'N 35°16.8'E, IV.1927, leg. O. Wohlberedt ( NHMB 17 - Ia) ; 2 ♀, ca. 11 km S of Beersheba, Hadarom , 31°06.19'N 34°49.41'E, 350 m a.s.l., 15.IV.1983, leg. A.M. de Saint Michel ( MEB 034 ) ; 1 ♂, 2 juvs, between Nesher & Yagur, Haifa District, 32°45.22'N 35°03.69'E, 27 m a.s.l., 20.IX.1984, leg. M. R . Warburg ( MEB 197 ) ; 1 ♂, 1 ♀, Haifa area, 32°48'N 34°59'E, 300 m a.s.l., 30.XI.1984, leg. M. R . Warburg ( MEB 198 ) ; 1 ♂, 3 ♀, 3 juvs, Negev desert, Vadi Hazaz near Sede Boqer ( Haluqim Ridge), XI.- XII .2004, leg. J. Král ( FKCP); 1 ♂, 2 ♀, Negev desert, Sede Boqer , IX.2007, leg. J. Král and M. Forman ( FKCP) . Jordan: 1 ♂, 4 ♀, Madaba, 31°43'N 35°48'E, 764 m a.s.l., 22. III .1983, leg. M.A. Jafar ( MEB 346 ) ; 1 ♀, Tabaqat , 32°27'N 35°37'E, – 200 m a.s.l., 4. V .1995, leg. V. Sejva (GL) ; 1 ♀, Kurayyima Udoli , 32°16.75'N 35°35.87'E, 23. VII .2000 (GL). Syria: 4 ♂, Bosra, V .1994, leg. D. Modrý ( FKCP); 1 ♀, Seydmaya, V . 1994, leg. D. Modrý ( FKCP).

DIAGNOSIS (adults). Small to medium sized Leiurus , 58–77 mm in length, carapace L 6.5–9.3 mm; base color yellow to orange-brown, with variable fuscosity on carapace and tergites; metasoma V either clear or fuscous except for posterior end; carapace with area between anterior median carinae finely shagreened, with scattered medium to fine granules, area between posterior median carinae with shallow median furrow flanked by arcs of medium or fine granules; medial intercarinal surfaces of tergites II – III shagreened, with variable medium to fine granulation; posterior margin of coxa III smooth or with sparse fine granules; metasoma robust, metasoma II L/ W 1.38 –1.64, metasoma III L/ W 1.49 –1.69, metasoma IV L/ W 1.71 –2.03; ventromedian carinae of metasoma II and III with 14–19 denticles (46/48 carinae); metasoma V with enlarged subtriangular or lobate denticles on ventrolateral carinae; pedipalps relatively stout, patella L/W ♂ 2.81–2.95, ♀ 2.48–2.83; leg III patella L/D 3.13– 3.71; pectine teeth ♂ 33–37, ♀ 27–32; pectines medium sized, pectine L/ carapace L ♂ 1.14–1.29, ♀ 0.92–1.21, mid-pectine sensillar margin L/ metasoma I W ♂ 0.169 – 0.180, ♀ 0.100 –0.136; basal 1–2 pectine teeth of males overlap if anterior pectine margins aligned to posterior margins of coxae IV; pectine basal piece very lightly, finely shagreened, almost smooth in females, almost smooth or weakly granulated in males; leg III basitarsus with 7–12 retrosuperior setae; pedipalp chela fixed finger with trichobothrium db either distal or proximal to est; sternite VII with area between median carinae densely, finely microgranulated or shagreened; sternite carination: males, sternite III with median carinae moderate to strong, sternites IV– V with lateral carinae strong, median carinae weak to moderate; females, sternite III with median carinae weak or obsolete, sternites IV– V with lateral carinae weak to moderate, median carinae obsolete .

MEASUREMENTS.

Male, Israel ( NHMB 17 a) (mm). Total L 58.0; metasoma + telson L 36.0; carapace L 6.74, W 7.22, carapace preocular L 3.05; metasomal segments (L/ D /W) I 4.98/ 4.01/4.34, II 5.82 / 3.63/3.97, III 6.08 / 3.56/3.85, IV 6.92/ 3.44/3.59, V 7.67 / 2.89/3.49; telson L 7.00; vesicle L 3.85, D 2.78, W 2.87; pedipalp chela L 11.62, manus ventral L 3.51, manus W 1.93, manus D 2.14, fixed finger L 6.79, movable finger L 7.98; pedipalp femur L 5.98, W 2.03, patella L 7.36, W 2.55; pectine L 7.83, mid-pectine sensillar margin L 0.735; leg III femur L 6.87; leg III patella L 5.79, D 1.825; chela db–est (left/ right) distance 0.292 / 0.958; pectine teeth (left/ right) 35/ 35 .

Female, Israel ( NHMB 17 a) (mm). Total L 76.0; metasoma + telson L 44.0; carapace L 8.09, W 8.91, carapace preocular L 3.82; metasomal segments (L/ D /W) I 5.70/ 4.30/4.83, II 6.44 / 4.15/4.30, III 6.48 / 4.05/4.10, IV 7.53/ 3.82/3.84, V 9.06 / 3.49/4.06; telson L 8.41; vesicle L 4.53, D 3.28, W 3.49; pedipalp chela L 14.34, manus ventral L 4.28, manus W 2.41, manus D 2.56, fixed finger L 8.20, movable finger L 10.03; pedipalp femur L 6.88, W 2.23, patella L 8.32, W 3.05; pectine L 8.83, mid-pectine sensillar margin L 0.653; leg III femur L 8.15; leg III patella L 6.92, D 2.00; chela db–est (left/ right) 0.354 / 0.417; pectine teeth (left/ right) 30/ 30 .

DISTRIBUTION. Israel, Jordan, Syria, Lebanon.

REMARKS. Birula (1908) separated L. q. hebraeus from the nominotypic subspecies by 11 characters. We assessed these characters by comparing 20 adult L. hebraeus from Israel and Jordan with 26 adult L. quinquestriatus from Egypt and Sudan:

(1) weaker carapace carination and granulation, (2) granulated ocular tubercle, (3) more coarsely shagreened tergites, (4) non-granulated intercarinal surfaces on the metasoma: (1) – (4) were unreliable when granulation was visualized in fine detail by UV fluorescence;

(5) ventromedian carinae of metasoma II – III with denticles increasing in size posteriorly (vs. uniform in size): this was confirmed for L. hebraeus , but also observed in some L. quinquestriatus , and hence was not diagnostic;

(6) smaller number of denticles (15 – 17) on ventromedian carinae of metasoma III (vs. 18 – 27): this was well supported with no overlap, i.e. 46/46 carinae of L. quinquestriatus with 19 or more denticles, 24/24 carinae of L. hebraeus with 18 or fewer denticles;

(7) lower range of denticles (18 – 31) on ventrolateral carinae of metasoma V (vs. 23 – 38): due to the broad overlap of ranges, this was not diagnostic;

(8) more stout pedipalp and metasomal segments: this was supported by smaller morphometric ratios, pedipalp patella L/W and metasoma III – IV L/W;

(9) metasoma I only barely wider than metasoma II (vs. significantly wider): this was not diagnostic, due to significant overlap of the ratio metasoma I W/ metasoma II W ( L. quinquestriatus 1.10 – 1.21, L. hebraeus 1.09 – 1.14);

(10) telson vesicle longer than aculeus (chord length), vesicle width equal to aculeus length (vs. about the same length or shorter, and width <aculeus length): there was a trend for L. quinquestriatus to have a relatively longer aculeus and a shorter, more bulbous vesicle than L. hebraeus , but there was significant overlap in the ratio of vesicle L/ telson L ( L. quinquestriatus 0.48 – 0.56, L. hebraeus 0.52 – 0.61);

(11) pedipalp movable finger short, less than twice manus length, with 12 denticle subrows (vs. long, slender,> twice manus length, and 13 subrows): this was not diagnostic, because although the movable finger was on average relatively shorter in L. hebraeus vs. L. quinquestriatus , there was broad overlap in the ratio movable finger L (chord)/ manus ventral L, and movable fingers had either 12 or 13 subrows in both species.

Levy et al. (1970) accepted characters (5) and (10) of Birula (1908), and proposed 3 additional characters to differentiate L. q. hebraeus :

(12) posterior median area of carapace granulated, with paired granule arcs (vs. smooth or sparsely granulated): this difference was confirmed for most specimens, although some L. quinquestriatus have weak arcs of fine granulation;

(13) lateral flanks of tergite V with irregular granulation (vs. short rows of granules, resembling carinae): this rather subjective character was quite variable, and short granule rows could be discerned or not in both L. hebraeus and L. quinquestriatus ;

(14) weak or obsolete carinae on sternites (slightly more distinct on posterior segments) (vs. distinct, elevated strong carinae on sternites IV–VII): this difference applies to males, however females have weaker carinae and some L. quinquestriatus females can have weak to obsolete carination on sternites IV–V.

Thus, from the above list, only characters (6), (14) (males only), and perhaps (12), were potential diagnostic characters separating L. hebraeus from L. quinquestriatus . Four examined adult and subadult males from southern Sinai resembled L. quinquestriatus in having slender pedipalp and metasomal segments, and reduced granulation between the posterior median carinae of the carapace. However, they grouped with the northern populations from Israel and Jordan in other morphometrics (i.e. lower metasoma I W/ II W, lower pedipalp movable finger L/ manus ventral L, higher vesicle L/ telson L) and meristics (number of denticles on ventromedian carinae of metasoma II–III). Levy et al (1980) also noted that the southern populations of L. q. hebraeus had more slender appendages. Partial overlap of characters suggests that there could be two subspecies, L. q. quinquestriatus and L. q. hebraeus , with a hybrid zone in southern Sinai. However, there appears to be a physical barrier preventing gene flow between these populations because Leiurus is excluded from the sandy coastal plain connecting North Africa and the Sinai Peninsula ( Levy & Amitai, 1980). Genetic isolation is supported by divergent evolution of distinct polypeptide toxins in their venoms ( Smertenko et al., 2001) and different physicochemical profiles of venom proteins ( Nascimento et al., 2006). Here, we propose to diagnose the African and Middle Eastern populations as distinct species by a combination of morphological characters. The southern Sinai populations occur in a different habitat (i.e. rocky mountains), and may belong to other undescribed species.