A review of the genus Leiurus Ehrenberg, 1828 (Scorpiones: Buthidae) with description of four new species from the Arabian Peninsula Author Lowe, Graeme Monell Chemical Senses Center, 3500 Market St, Philadelphia, USA Author Yağmur, Ersen Aydın Alaşehir Vocational School, Celal Bayar University, Manisa, Turkey Author Kovařík, František P. O. Box 27, CZ - 145 01, Praha 45, Czech Republic www.kovarex.com/scorpio text Euscorpius 2014 2014-12-31 191 1 129 https://doi.org/10.18590/euscorpius.2014.vol2014.iss191.1 journal article 152185 10.18590/euscorpius.2014.vol2014.iss191.1 a6381f1d-22e1-4629-b369-07f5ab1d7834 1536-9307 6505622 E467B3C0-D693-4EAF-B5F0-759D8C63FE35 Leiurus macroctenus sp. n. ( Figs. 12–22 , 58A , 59A–B , 87G , 88G , 89G , 90G , 91B , 92H–I , 93H , 95 , 98–100 , Tabs. 1 , 3B , 4 ) http://zoobank.org/urn:lsid:zoobank.org:act:55E2C0 48-986D-43A2-AF63-5874138435D6 REFERENCES Buthus quinquestriatus : Finnegan, 1932: 91–92 (in part; record from Wadi Ghabartan). Leiurus quinquestriatus : Gallagher, 1978: 36–37 ; Lowe, 1993: 2 ; Kovařík 1998: 112 (in part) ; Fet & Lowe, 2000: 155 (in part); Lowe, 2001: 176 ; Fet et. al., 2003: 3 ; Soleglad & Fet, 2003b: 149 , fig. B-1, B-2; Hendrixson 2006: 84–91 , fig. 18 (in part, record from Oman ). Leiurus quinquestriatus hebraeus : Kamenz & Prendini, 2008: 40 . Leiurus sp. : Lowe, 2010a: 23 ; Lowe, 2010b: 26 , 37. HOLOTYPE . Adult ♂ , Oman , Nejd Desert ; S of Thumrait , UV detection, fine silty soil, open plain with rock outcrops and small wadis, 17°30.76'N 54°02.76'E , 580 m a.s.l. , 16.X.1993 , 19:28 h, leg. G. Lowe , NHMB . PARATYPES . Oman : 2 ♂ , Thamarit , Dhofar , sand & rock (limestone), 17°38'N 54°01'E , 350 m a.s.l. , I.1980 , leg. J.N. Barnes & M.D. Gallagher MDG 5954 ( MNHN ) ; 2 ♀ , Thamarit , Dhofar , sand & rock, 17°38'N 54°01'E , 350 m a.s.l. , I.1980 , leg. J.N. Barnes & M.D. Gallagher MDG 5952, 5953 ( MNHN ) ; 1 immature ♀ , nr Thamarit , Dhofar , under object on gravel in wadi, 17°41'N 54°03'E , 440 m a.s.l. , 22. II .1980, leg. J.N. Barnes & M.D. Gallagher MDG 5973 ( MNHN ); 1 ♂ , nr Thamarit , Dhofar , 17°42'N 54°02'E , 440 m a.s.l. , 4. III .1980, leg. J.N. Barnes & M.D. Gallagher MDG 5972 ( MNHN ); 1 juv, nr Thamarit , Dhofar , on sand, 17°40'N 54°02'E , 450 m a.s.l. , 22. III .1980, leg. J.N. Barnes 50A ( MNHN ); 1 juv , nr Thamarit , Dhofar , under object on sand, 17°40'N 54°02'E , 450 m a.s.l. , 22.III .1980 , leg. J.N. Barnes 54 ( NHMB ); 1 juv , nr Thamarit , Dhofar , 17°40'N 54°02'E , 450 m a.s.l. , 22.III .1980 , leg. J.N. Barnes 52 ( NHMB ); 1 juv, nr Thamarit , Dhofar , under object on sand, 17°40'N 54°02'E , 450 m a.s.l. , 22. III .1980, leg. J.N. Barnes 37 ( MNHN ); 1 ♂ , nr Thamarit , Dhofar , under rock, 17°41'N 54°02'E , 450 m a.s.l. , 23.III .1980 , leg. J.N. Barnes ( NHMB ); 1 juv , nr Thamarit , Dhofar , under object on sand, 17°40'N 54°00'E , 450 m a.s.l. , 24.III .1980 , leg. J.N. Barnes 35 ( NHMB ); 1 ♀ , nr Thamarit , Dhofar , under object on sand, 17°40'N 54°00'E , 450 m a.s.l. , 24.III .1980 , leg. J.N. Barnes ( NHMB ); 1 ♂ , nr Thamarit , Dhofar , 17°41'N 54°03'E , 450 m a.s.l. , 24.III .1980 , leg. J.N. Barnes 33 ( NHMB ); 1 juv , nr Thamarit , Dhofar , on sand, 17°40'N 54°02'E , 450 m a.s.l. , 24.III .1980 , leg. J.N. Barnes 55, ( NHMB ); 1 juv , nr Thamarit , Dhofar , on gravel, 17°40'N 54°02'E , 450 m a.s.l. , 30.III .1980 , leg. J.N. Barnes 58 ( NHMB ); 1 ♂ , 3 mi. NW of Thamarit , under sack on sand, 17°43'N 54°00'E , 420 m a.s.l. , 8. III .1980, leg. M.D. Gallagher MDG 5971 ( MNHN ); 1 ♂ , 4 mi. NE of Thamarit , Dhofar , in soft sand under sack in green wadi, 17°42'N 54°03'E , 8. III .1980, leg. J.N. Barnes & M.D. Gallagher MDG 5975 ( MNHN ); 1 ♂ , nr Al Ashkara , 22°00'N 59°40'E , XII .1980, leg. A.B. Paltrinieri ( NHMB 17bq); 1 ♂ , Yalooni , Jiddat al Harasis , in house, sand on stony plateau, 19°57'N 57°07'E , 154 m a.s.l. , 22.IV.1982 , leg. M.D. Gallagher MDG 6428/1,2 ( ONHM ) ; 1 ♂ , Wadi Haytam , sandy depression in rocks, 19°11'N 57°00'E , 13.V .1983 , leg. M.D. Gallagher MDG 6770 ( NHMB ); 1 ♂ , Thamarit , under rock on soft sand, 17°42'N 53°59'E , 24. V .1983, leg. J.N. Barnes 126 ( BMNH ); 1 ♂ , Thamarit , under rock on soft sand, 17°42'N 53°59'E , 24. V .1983, leg. J.N. Barnes 125 ( ONHM ); 1 ♂ , Yalooni , Jiddat al Harasis , 19°57'N 57°06'E , 154 m a.s.l. , 17.XII .1984 , leg. M.D. Gallagher ( NHMB ); 1 juv , Qarhat Mu'ammar , 21°38'N 59°18'E , 130 m a.s.l. , 28.II .1986 , leg. W. Büttiker ( NHMB 17bt); 1 ♀ , Ra's al Hadd , on sand under object, 0.5 km from sea , 22°31.8'N 59°46'E , 5 m a.s.l. , 3.II .1988 , 10:00 h, leg. M.D. Gallagher MDG 8020 ( NHMB 17br); 1 juv , Huqf , central Oman , under rock on sand, 20°53'N 57°36'E , 15.III .1988 , 14:00 h, M.D. Gallagher MDG 8031 ( NHMB 17bs); 1 ♂ , between Bani bu Ali & Al Ashkharah , 3 km W Qumaylah turning, under rubbish on sand, 21°56'N 59°35'E , 17.I.1991 , leg. S.M. Farook & A.S. Gardner ( MNHN ) ; 1 ♂ , 1♀ , 3 km N of Khubah , 22°16'N 59°48'E , 11.II .1993 , leg. A.S. Gardner ( NHMB ); 6 ♂ , between Khabbah & Daffah , under stones on limestone plateau , 22°15'N 59°48'E , 29.VIII.1993 , leg. A.S. Gardner & S.M. Farook ( NHMB ); 6 ♂ , 4 ♀ , 5 juvs , S of Thumrait , Nejd Desert , UV detection, near or partly in burrows , fine silty soil, open plain rock outcrops & small wadis, 17°30.76'N 54°02.76'E , 580 m a.s.l. , 16.X.1993 , 19:28 h, leg. G. Lowe ( NHMB ); 4 ♂ , 4 ♀ , 4 juvs , S of Thumrait , Nejd Desert , UV detection, on soil & at mouths of burrows, fine silty soil, open plain rock outcrops & small wadis, 17°30.8'N 54°02.94'E , 580 m a.s.l. , 16.X.1993 , 20:57 h, leg. G. Lowe ( NHMB , ONHM 1 ♀, WDS 1 ♂); 1 ♂ , Salalah-Thumrait road , Nejd Desert , UV detection, on ground, hills near main road, 17°25.56'N 54°03.14'E , 600 m a.s.l. , 16.X.1993 , 21:49 h, leg. G. Lowe ( NHMB ); 1 juv , E of Thumrait , road to Marmul , UV detection, open desert plain, sparse vegetation, 17°38.99'N 54°08.29'E , 500 m a.s.l. , 19.X.1993 , 21:10 h, leg. G. Lowe ( NHMB ); 3 ♂ , 2 ♀ , E of Thumrait , road to Marmul , UV detection, open plain, low rocky hills, 17°38.33'N 54°07.77'E , 500 m a.s.l. , 19.X.1993 , 21:31 h, leg. G. Lowe ( ONHM ) ; 3 ♂ , 1 juv , S of Thumrait , Nejd Desert , UV detection, open alluvial plain with rock outcrops and small wadis, on fine soil & partially in burrows, 17°30.77'N 54°02.82'E , 600 m a.s.l. , 19.X. 1993 , 23:02 h, leg. G. Lowe ( NHMB ); 2 juvs , Yalooni , UV detection, on gravel plain with Prosopis woodland, 19°57'N 57°06'E , 150 m a.s.l. , 21.X.1993 , leg. M. Fisher ( NHMB ); 2 juvs , Yalooni , on sand, amongst fallen twigs, clear moonless sky, slight breeze , 19°56'N 57°05'E , 154 m a.s.l. , 15.III .1994 , 22:00 h, leg. M.D. Gallagher MDG 8564.3-4 ( NHMB ); 1 ♀ , 2 km N of Yalooni , Jiddat Al Harasis , under barrel with beetles on sand , 19°58'N 57°07'E , 154 m a.s.l. , 17.III .1994 , 13:30 h, leg. M.D. Gallagher MDG 8569 ( NHMB ); 1 juv , nr Ad Daffah, low dunes, below rocky escarpment , 22° 15'N 59°48'E , 24.XI.1994 , leg. A.S. Gardner ( NHMB ); 1 juv , Jabal Qara , hilltop, UV detection, 17°14.51'N 54°04.25'E , 800 m a.s.l. , 30.IV.1995 , leg. J. Dundon ( NHMB ) ; 1 ♂ , Jabal Qara , hilltop, UV detection, 17° 14.51'N 54°04.25'E , 800 m a.s.l. , 30.IV.1995 , leg. J. Dundon ( USNM ) ; 1 ♀ , 2 juvs , Masirah Island , north Masirah , east coast, low rock littered slopes, compact silty soil , UV detection, 20°34.48'N 58°55.36'E , 25 m a.s.l. , 20.IX.1995 , 19:50 h, leg. G. Lowe & M.D. Gallagher ( NHMB ); 1 ♂ , 1♀ , Masirah Island , north Masirah , east coast, UV detection near edge of wide wadi with small side wadis, from edges of slopes, 20°35.68'N 58°53.98'E , 20 m a.s.l. , 20.IX.1995 , 21:10 h, leg. G. Lowe & M.D. Gallagher ( NHMB ) ; 1 ♂ , 3 ♀ , Masirah Island , north Masirah , east coast, wadi with row of trees & flowing water close to beach with hermit crabs, UV detection on gravelly ground away from trees, 20°39.78'N 58°54.28'E , 5 m a.s.l. , 20.IX.1995 , 22:30 h, leg. G. Lowe & M.D. Gallagher ( MNHN ) ; 1 ♂ , 3 ♀ , 6 juvs, Masirah Island , north Masirah , east coast, UV detection on sand and gravel, by row of trees along wadi with flowing water, 20°40.14'N 58°54.4'E , 20.IX.1995 , 23:20 h, leg. G. Lowe & M.D. Gallagher ( GL ) ; 1 ♀ , 2 juvs, Masirah Island , north Masirah , east coast, UV detection, near beach, gravelly/ silty flat with shrubs, 20°41.01'N 58°54.63'E , 21.IX.1995 , 0:05 h, leg. G. Lowe & M.D. Gallagher ( BMNH ) ; 1 ♂ , 5 ♀ , 3 juvs , Masirah Island , south Masirah , base of Jabal Suwayr , seepage area with trees & standing water at base of Jabal rock and boulder slopes , UV detection on sand & gravel plain below not far from beach, 20°11.65'N 58°39.42'E , 20 m a.s.l. , 21.IX.1995 , 18:10 h, leg. G. Lowe & M.D. Gallagher ( NHMB ); 3 ♂ , 8 ♀ , 3 juvs , Masirah Island , south Masirah , Wadi Qitaar , UV detection on sand deposits along edges of wide sandy wadi, 20°13.78'N 58°38.4'E , 20 m a.s.l. , 21.IX.1995 , 21:20 h, leg. G. Lowe & M.D. Gallagher ( NHMB ); 6 ♂ , 11 ♀ , 5 juvs, Masirah Island , south Masirah , UV detection on sand, area with shrubs by low rocky hills, 20°16.05'N 58°38'E , 25 m a.s.l. , 21.IX.1995 , 22:30 h, leg. G. Lowe & M.D. Gallagher ( ONHM ) ; 2 ♂ , 3 ♀ , 5 juvs, Masirah Island , UV detection on sand in wide wadi system, densely vegetated with low shrubs, 20°23.48'N 58°41.32'E , 30 m a.s.l. , 21.IX.1995 , 23:50 h, leg. G. Lowe & M.D. Gallagher ( NHMB ) ; 1 ♂ , Masirah Island , north Masirah , west coast, SE of Dafiyat , under wood debris on ground, gravel flat, 20°35.28'N 58°50.76'E , 22.IX.1995 , 0:50 h, leg. G. Lowe & M.D. Gallagher ( BMNH ) ; 1 ♀ , nr Duqm , cliff with nearby wadi and trees, 19°36.06'N 57°37.29'E , 50 m a.s.l. , 23.IX.1995 , 18:50 h, leg. G. Lowe & M.D. Gallagher ( MZUF ) ; 1 ♂ , 3 juvs , nr Duqm , UV detection , 19°30.59'N 57°34.9'E , 95 m a.s.l., 23. IX.1995 , 18:40 h, leg. G. Lowe & M.D. Gallagher ( NHMB ); 2 ♂ , 1 juv, nr Duqm , UV detection, vegetated wadi, 19°33.53'N 57°37.38'E , 75 m a.s.l. , 23.IX.1995 , 23:00 h, leg. G. Lowe & M.D. Gallagher ( MNHN ); 1 ♂ , 1♀ , E. of Duqm , UV detection on ground , densely vegetated wadi, 19°39'N 57°40'E , 30 m a.s.l. , 23. IX.1995 , 23:50 h, leg. G. Lowe & M.D. Gallagher ( NHMB ); 11 ♂ , 8 ♀ , 10 juvs , N edge of Wadi Dirif , UV detection on sandy substrate , rocky knoll overlooking wadi and on open flats of wadi, 18°57.51'N 57°21.73'E , 20 m a.s.l. , 24.IX.1995 , 18:40 h, leg. G. Lowe & M.D. Gallagher ( NHMB ); 1 ♂ , 1♀ , Wadi Dirif , above beach zone, UV detection, fine silty soil by rock outcrops, 18°57.57'N 57°20.63'E , 5 m a.s.l. , 24.IX.1995 , 23:30 h, leg. G. Lowe & M.D. Gallagher ( MCZN ) ; 4 ♂ , 2 juvs, Wadi Dirif , W edge of wadi, UV detection on sand, 18°57.2'N 57°20.63'E , 10 m a.s.l. , 24.IX.1995 , 23:40 h, leg. G. Lowe & M.D. Gallagher ( BMNH ); 13 juvs, Wadi Dirif , E edge of wadi, UV detection, rock outcrops on sand, 18°57.1'N 57°20.71'E , 10 m a.s.l. , 25.IX.1995 , 0:25 h, leg. G. Lowe & M.D. Gallagher ( ONHM ) ; 6 ♂ , 6 ♀ , Wadi Dirif , E edge of wadi , UV detection on sand, with rocks, 18°57.1'N 57°20.71'E , 10 m a.s.l. , 25.IX.1995 , 0:25 h, leg. G. Lowe & M.D. Gallagher ( NHMB ); 1 juv , Wadi Dirif , dunes by road , UV detection on sand, 18°57.07'N 57°20.09'E , 20 m a.s.l. , 24.IX.1995 , 22:50 h, leg. G. Lowe & M.D. Gallagher ( NHMB ); 4 juvs , Wadi Shuwaymiyah , UV detection, base of cliff along southern edge of wadi, sand and loose rock along base, 17°54.9'N 55°34.63'E , 25.IX.1995 , 22:00 h, leg. G. Lowe & M.D. Gallagher ( NHMB ); 1 ♂ , 2 ♀ , coast W of Shuwaymiyah , UV detection on sand, wide sandy wadi along coast, gentle slopes up to cliffs on edges, 17°53.91'N 55°39.16'E , 5 m a.s.l. , 25.IX.1995 , 23:40 h, leg. G. Lowe & M.D. Gallagher ( FKCP ) ; 2 ♂ , 2 juvs, plateau between Shalim & Ash Shuwaymiyah , UV detection, open rocky windswept plateau with sparse shrubs ground with bare rock and patches of soil, strong wind, 18°02.44'N 55°36.91'E , 265 m a.s.l. , 26.IX.1995 , 1:32 h, leg. G. Lowe & M.D. Gallagher ( GL ) ; 1 ♂ , 2 ♀ , 2 juvs, Saiwan , Rudist outcrop, 20°42.94'N 57°36.96'E , 20. II .1996, leg. A.S. Gardner ( ONHM ); 1 ♂ , Marmul , 18°10.09'N 0°14.76'E ( FKCP ) ; 1 ♂ , 30 km N of Haima , near Haylat Harashif , dry level sand desert, in large open area, arid gravel desert, 20°11.43'N 56°33.58'E , 6. XII . 1996, leg. M.D. Gallagher MDG 8798 ( FKCP ); 1 ♂ , 3 ♀ , Khawr Dirif , on sand under small plants and rocks, 18°56.7'N 57°20.73'E , 5 m a.s.l. , 7. XII .1996, leg. I.D. Harrison & M.D. Gallagher MDG 8802 ( USNM ); 3 ♂ , 3 ♀ , 5 juvs , 20km N of Liqbi , Sahil al Jazr , sand and scrub , 18°24.17'N 56°33.95'E , 15 m a.s.l. , 8.XII .1996 , leg. I.D. Harrison & M.D. Gallagher MDG 8805 ( NHMB ); 11 ♂ , 1♀ , 1 juv, Thumrait , wadi area SE of Thumrait and golf course, 17°42'N 53°59'E , 11-12. II – 9.X.1997, leg. J.N. Barnes ( ONHM , EV 1 ♂ ) ; 1 ♂ , Huqf , on open sand between bushes at foot of low hills, three quarter moon, strong wind, 20°25.7'N 57°26.42'E , 18.IV.1997 , M.D. Gallagher MDG 8859 ( ONHM ) ; 3 ♂ , 2 ♀ , 4 juvs , Ra's Al Ghubbah , hiding under shrubs from three quarter moon, on sand over rock slope from beach, 20°07.95'N 57°49.77'E , 5 - 20 m a.s.l. , 19.IV.1997 , 19:00–21:00 h, leg. M.D. Gallagher & I.D. & M. Harrison MDG 8866 ( NHMB ) ; 8 ♂ , 2 ♀ , Duhai , low aeolianite hilltop & slope to sand, to edge of trees, 500 m 2 area, 21°06.26'N 58°22.29'E , 70 m a.s.l. , 21.X.1997 , leg. M.D. Gallagher & I.D. Harrison MDG 8889, ( ONHM 2 ♀ , GL ) ; 4 ♀ , 1 juv, NE of Ras Al Jibsh , slope of sand and scrub hummocks, between sea and rocky mountains, on rocks, sand and under scrub, 21°35.4'N 59°25.1'E , 100 m a.s.l. , 19. II .1998, leg. J. Peterson , I.D. Harrison & M.D. Gallagher MDG 8945 ( MZUF ); 6 ♂ , 1♀ , 2 juvs , N of Al Ashkharah , beach sand, scrub and rocks, most hiding in scrub from three quarter moon, 21°54.7'N 59°36.78'E , 2 m a.s.l. , 7.IV.1998 , leg. I.D. , S. & R. Harrison . & M.D. Gallagher ( NHMB ) ; 1 ♂ , E. of Hayma , on sparse vegetation, 19°46.05'N 56°35.77'E , 3.X.1998 , leg. I.D. Harrison ( NHMB ) ; 3 ♂ , 1♀ , 1 juv, between Al Ashkharah and A'Seelal , under small bushes, 21°55.4'N 59°37. 67'E , 21.XI.1998 , leg. I.D. Harrison ( MNHN ); 1 juv , Jiddat al Harasis nr Habab (site F/9), under rocks in road cut, 19°42.28'N 56°48.72'E , 250 m a.s.l. , 29.XII . 1998 , 14:45 h, leg. A. Winkler & B. Winkler ( NHMB ); 5 ♂ , 2 juvs , nr Al Ashkharah , on grass, on sandy plain between sea & mountains, 21°54'N 59°36'E , 15.IV.1999 , leg. I.D. & S.E. Harrison ( NHMB ) ; 1 ♀ , nr Ras Khaysat al Liiyakh , ca. 20 km N Ras Madrakah , Al Wusta region , sitting under small plant 30 cm × 30 cm , 19° 13.5'N 57°45'E , 21.IX.2000 , leg. R . Harrison & S. Harrison ( ONHM ) ; 1 ♂ , Al Hayl , by rock, in rock/ pebble strewn sand on low ridge overlooking wide sandy wadi, 20°42'N 58°15'E , 11.XI.2001 , leg. I.D. Harrison ( ONHM ) ; 2 ♂ , Wadi Abaytham , under rock on low ridge overlooking wadi, sparse vegetation on ridge, denser in wadi, 18°58'N 57°36'E , 12.XI.2001 , leg. I.D. Harrison ( MCZN ) ; 1 ♀ , S Duqm (site F/24) , UV detection, in small dunes, mouth of burrow, 19°26.75'N 57°35.78'E , 147 m a.s.l. , 26.XII .2001, 19:00–20:00 h, leg. A. Winkler ( NHMB ); 1 juv , S Duqm (site F/24) , UV detection, in small dunes, mouth of burrow, 19°26.75'N 57°35.78'E , 147 m a.s.l. , 27.XII .2001 , 04:00–05:00 h, leg. A. Winkler ( NHMB ); 1 ♀ subadult, 3 juvs, Al– Saadanat , 19°20.37'N 57°35.47'E , 97 m a.s.l. , 18. III . 2014, leg. D. Hoferek ( FKCP ); 1 ♂ , 1 ♀ , UV detection 20:00–22:00 h, Wadi Dirif , 18°56.51'N 57°20.73'E , 7 m a.s.l. 19. III .2014, leg. D. Hoferek ( FKCP ); 4 ♀ , UV detection 20:00–22:00 h, Shalím , 18°02.46'N 55°36. 93'E , 290 m a.s.l. , 20. III .2014, leg. D. Hoferek ( FKCP ). Figure 12: Leiurus macroctenus sp. n. , holotype male. Habitus, dorsal aspect. Thumrait, Oman. Figure 13: Leiurus macroctenus sp. n. , holotype male. Habitus, ventral aspect. Thumrait, Oman. Figure 14: Leiurus macroctenus sp. n. , holotype male. A. Carapace and tergites. B. Coxosternal area and sternites. Thumrait, Oman. Figure 15: Leiurus macroctenus sp. n. , holotype male. Right pedipalp. A. Femur, dorsal aspect. B. Patella, dorsal aspect. C. Patella, external aspect. D. Chela, ventral aspect. E. Chela, dorsal aspect. F. Fixed finger dentition. G. Movable finger dentition. H. Chela, external aspect. Thumrait, Oman. Figure 16: Leiurus macroctenus sp. n. , paratype female. Habitus, dorsal aspect. Wadi Qitaar, Masirah Island, Oman. Figure 17: Leiurus macroctenus sp. n. , paratype female. Habitus, ventral aspect. Wadi Qitaar, Masirah Island, Oman. Figure 18: Leiurus macroctenus sp. n. , paratype female. A. Carapace and tergites. B. Coxosternal area and sternites. Wadi Qitaar, Masirah Island, Oman. ETYMOLOGY. The name macroctenus (= ‘large comb’) refers to the large size of the pectine teeth in both sexes, which distinguishes this species from other members of the genus. DIAGNOSIS (adults). Medium to large Leiurus , 61–88 mm in length, carapace L 7.1–10.2 mm , base color yellow to pale orange, carapace with limited fuscosity on median interocular tubercle; metasoma V slightly fuscous to dark except for posterior end; carapace with area between anterior median carinae smooth or bearing sparse medium to fine granules, area between posterior median carinae with moderately deep median furrow flanked by lateral arcs of medium to fine granules; medial intercarinal surfaces of tergites II–III densely shagreened and granulated; posterior margin of coxa III smooth or with sparse, fine granules; metasoma robust to moderately slender, metasoma II L/ W 1.48 –1.82, metasoma III L/ W 1.59 –2.03, metasoma IV L/ W 1.87 –2.40; ventromedian carinae of metasoma II and III with 14–23 denticles (78/80 carinae); metasoma V with enlarged subtriangular or lobate denticles on ventrolateral carinae; pedipalps moderately slender, patella L/W ♂ 2.83– 3.64, ♀ 2.55–3.32; leg III patella L/D ♂ 3.43–4.48, ♀ 3.38–4.00; pectine teeth ♂ 33–43, ♀ 30–37; pectines large, broad, pectine L/ carapace L ♂ 1.15–1.52, ♀ 1.03–1.26, mid-pectine sensillar margin L/ metasoma I W ♂ 0.216 –0.254 , ♀ 0.156 –0.203 ; basal 3–6 pectine teeth of males overlap if anterior pectine margins aligned to posterior margins of coxae IV; pectine basal piece lightly, finely shagreened; leg III basitarsus with 15–26 retrosuperior setae; pedipalp chela fixed finger with trichobothrium db either distal or proximal to est ; sternite VII with area between median carinae smooth or slightly roughened anteriorly; sternite carination: males : sternite III with median carinae strong, sternites IV–V with lateral carinae moderate to strong, median carinae weak or obsolete, shagreened; females : sternite III with median carinae weak and finely granulated, or smooth and obsolete, sternites IV–V with lateral carinae weak or moderate, finely granulated, median carinae obsolete. COMPARISONS. L . macroctenus sp. n. is distinguished from most other species of Leiurus by its large pectine teeth, a distinction that is more pronounced when comparing females, and by strongly overlapping basal pectine teeth of males. The male of L. brachycentrus also has large pectine teeth, but the teeth of females are much smaller than those of female L . macroctenus . The number of retrosuperior setae in the basitarsal bristle combs of L . macroctenus is significantly higher than in other species, which is likely an adaptation to soft sandy substrates. DESCRIPTION ( holotype male). Coloration. Base color pale orange or yellow; melanic pigmentation confined to patches around median ocular tubercle and carinae of interocular triangle; ventromedian carinae of metasoma II–IV without pigmentation; metasoma V darkened except for distal portion of segment. Carapace. Subrectangular, W/L 0.97, with steep lateral flanks; top nearly flat, with median eyes strongly raised; interocular triangle convex laterally, depressed medially; anterior margin very slightly emarginate, minutely denticulate laterally with several fine median granules, bordered by row of coarse granules; 11 short macrosetae on anterior margin, carapace otherwise devoid of macrosetae; 5 lateral eyes (3 large, 2 small) on each side; carination : anterior median, central lateral, posterior median and posterior lateral carinae strongly developed, coarsely granular; supraocular section of superciliary carinae with weaker granulation (likely due to abrasion), postocular section coarsely granular; anterior median carinae not extending to anterior margin of carapace, separated from anterior marginal row of granules; central lateral and posterior median carinae fused into lyre configuration; central median carinae coarsely granular, bifurcated, posterior branch weakly curved; posterior lateral carinae curved with lateral extension, projecting past posterior margin of carapace; lateral ocular carinae moderate, with small, spaced granules; granulation : sparse patches of 11 medium to large granules on anterolateral corners of interocular triangle, 7–8 small and large granules in front of lateral ocular carinae; surface between anterior median carinae sparsely, finely shagreened with few isolated small granules; other intercarinal surfaces smooth or faintly microshagreened or with isolated small granules; posterior median furrow very shallow, nearly flat, with few fine granules or micro-shagreened along median line, flanked by lateral arcs of small to medium granules; posterior margin of carapace between posterior lateral carinae rimmed by regular row of small contiguous granules. Chelicera. Dorsal surface of manus smooth, glabrous, with 6 short, pale microsetae, 3 near apical margin, 3 subapical, each surrounded by granules; dorsointernal carina at base of fixed finger strong, well granulated, terminating anteriorly with prominent granules projecting over front of manus; single long macroseta in middle of dorsointernal carina; dorsal surface of movable finger smooth, with 4 pale microsetae; fingers with characteristic buthid dentition ( Vachon, 1963 ); movable finger dorsal margin with 5 teeth: dorsal distal tine, subdistal, median and 2 basal teeth fused in bicusp; ventral margin with 3 teeth: ventral distal tine, median and basal teeth; fixed finger margin with 4 teeth: distal tine, subdistal, median and basal teeth; ventral aspect of fixed finger with 2 teeth. Coxosternal area. Coxae smooth; coxal endite II with weak, finely granulated carina; coxae II–III with coarsely granular anterior carinae, distal margins bearing medium to coarse granules; proximal 1/3 of anterior carina of coxa III finely granulate; 3 long macrosetae along anterior carina of coxa II, 3 macrosetae along anterior carina of coxa III; anterior carina of coxa IV with regular small to medium granulation, with single proximal macroseta; posterior margin of coxa IV with very finely granulated carina on proximal half; sternum with scattered fine granules, subtriangular with slightly convex, nearly straight lateral margins, deep posteromedian longitudinal sulcus and pit with finely granulated posterior lip, two short macrosetae; genital opercula smooth with 4 short macrosetae. Pectines. Basal piece with concave anterior margin divided by small median groove, surface finely shagreened medially, almost smooth laterally; 4 short macrosetae on anterior margin, 1 posteromedial; pectines long, tips extending past proximal 4/5 of trochanter IV; pectines with 3 marginal lamellae, small accessory lamella distal to first marginal lamella, 9 middle lamellae, 38–39 teeth; marginal and middle lamellae with dense cover of short reddish macrosetae; fulcra with 3–6 setae; pectine teeth long; when anterior margins of left and right pectines aligned with posterior edges of coxae IV, basal 4 teeth overlap, without gap between basal middle lamellae, pectine teeth contact median carinae of sternite III; pectine teeth relatively large, mid-pectine sensillar margin L/ pectine L 0.096, mid-pectine sensillar margin L/ metasoma I W 0.231 . Figure 19: Leiurus macroctenus sp. n. , paratype female. Right pedipalp. A. Femur, dorsal aspect. B. Patella, dorsal aspect. C. Patella, external aspect. D. Chela, ventral aspect. E. Chela, dorsal aspect. F. Chela, external aspect. Wadi Qitaar, Masirah Island, Oman. Mesosoma. Tergites : pretergites smooth; tergites I– II with 5 granular carinae; median and inner lateral carinae linear with medium granules; outer lateral carinae aligned with posterior lateral carinae of carapace, strongly raised with enlarged posterior granules, hind ends with short lateral extensions, projecting past posterior margins of tergites; lateral flanks steeply sloped, with sparse granulation; medial intercarinal surfaces with dense, fine granulation or shagreened, with transverse anterior series of fine granules; tergites III–VI with 3 linear or slightly curved carinae with medium to large, erect granules; medial intercarinal surfaces densely shagreened, with few larger granules, transverse anterior series of fine granules becoming more diffuse in posterior tergites; heavier fine granulation on median patch in front of median carina; lateral surfaces moderately sloped, shagreened, with many medium size granules; tergite VII with 5 strong, coarsely granular carinae; inner and outer lateral carinae joined anteriorly by transverse granule rows; intercarinal surfaces smooth or very faintly shagreened, fine granulation on anteromedian and paired anterolateral patches; posterior margins of tergites I–VI rimmed with linear rows of medium sized granules; posterior margin of tergite VII with 3 small granules between inner lateral carinae; sternites : sternite III with moderately strong, broad, finely granulated median carinae; sternites IV–V with strong, finely granulated lateral carinae, very weak, almost obsolete densely shagreened median carinae; sternite VI with strong, coarsely granulated lateral carinae, moderate, finely granulated median carinae; sternite VII with strong, coarsely granulated median and lateral carinae; lateral margins of sternites IV–VII armed with regular denticulate granules; medial intercarinal surface of sternite III densely shagreened throughout, of IV–VI densely shagreened anteriorly; lateral intercarinal surfaces on IV–VI heavily shagreened anteriorly, lightly on post-spiracular area; setation : sternite III with 5–6 macrosetae on median carinae, 4–5 on external side of median carinae; sternites IV–VII with 2 paired macrosetae on median carinae, one pair in middle of sternite, other on posterior margin; lateral carinae on IV–VI with posterior marginal macrosetae, mid-carinal setae on IV; intercarinal macrosetae: one middle pair of lateral marginal setae on sternites IV–VI; paired mediolateral setae 3 on IV, 2 on V, 1 on VI; one pair of lateromarginal setae on VII; intercarinal posterior marginal macrosetae on III–VII: 4, 4, 2, 2, 0. Figure 20: Leiurus macroctenus sp. n. Metasoma. A. Male, lateral aspect. B. Male, ventral aspect. C. Female, lateral aspect. D. Female, ventral aspect. Male holotype, Thumrait, Oman. Female paratype, Wadi Qitaar, Masirah Island, Oman. Figure 21: Leiurus macroctenus sp. n. , holotype male. Tarsi. A–D. Left telotarsus and distal basitarsus, ventral aspect. A. Leg I. B. Leg II. C. Leg III. D. Leg IV. E. Right basitarsus III retrolateral aspect. Upper scale bar: A–D , lower scale bar: E . Thumrait, Oman. Hemispermatophore (topoparatype male). Trunk elongate, slender; flagellum long, filiform, pars recta 0.4 times length of trunk, pars reflecta half length of trunk; inner lobe a broad lamina, gently tapering apically; median and outer lobes narrower at base, sharply tapering to thin apical processes; inner lobe longer than median and outer lobes; outer lobe flexed at apex; basal lobe thin, curved hook; measurements: trunk L (to base of flagellum) 7.2 mm , pars recta 2.9 mm , inner lobe (from base of flagellum) 810  m, median lobe 410  m, outer lobe 530  m, basal lobe 200  m. Figure 22: Leiurus macroctenus sp. n. , paratype female. Trichobothrial map of pedipalp. A. Femur, dorsal aspect. B. Patella, dorsal aspect. C. Patella, external aspect. D. Chela, external aspect. E. Chela, ventral aspect. Wadi Qitaar, Masirah Island, Oman. Metasoma. Long, slender; total metasoma and telson L/ carapace L 5.4; carination : segment I with 10 complete carinae; segments II–III with 8 complete carinae, median lateral carinae restricted to posterior 0.25–0.38 of II, posterior 0.20 of III; metasoma IV with 8 carinae, V with 7 carinae; carinae on segments I–IV with crenulate or denticulate granulation; dorsosubmedian carinae moderate on I, moderate to weak on II– III, weak on IV; dorsolateral, ventrolateral and ventromedian carinae moderate on I, strong on II–IV; median lateral carinae moderate on I–III; ventromedian carinae on II–III with 16–19 granules, posterior granules larger; metasoma V with dorsolateral carinae very weak, faintly granulated on anterior 1/3, obsolete on posterior 2/3; ventrolateral carinae strong with dentate granules increasing in size posteriorly, larger subtriangular denticles alternating with series of smaller crenulations, ventrosubmedian carinae marked by series of small to medium dentate granules on anterior 2/3 of segment, ventromedian carina strong, armed with small to large dentate granules; lateral anal margin with 3 lobes, ventral lobe well separated from dorsal lobes, ventral anal margin with 13 irregular, narrow to wide transverse crenulations; intercarinal surfaces : segments I–IV smooth, segment V smooth dorsally, finely sparsely shagreened laterally and ventrally; setation : segments I– IV: ventromedian carinae with 3 macrosetae (one posterior marginal), ventrolateral carinae with 2 external macrosetae; metasoma V with 5 macrosetae on lateral surface (2 lateral anal), 4 pairs on ventral surface. Telson. Vesicle smooth, bulbous; ventral surface with scattered fine microsetae and several short macrosetae; aculeus slightly shorter than vesicle. Pedipalp. Femur : slender, L/ W 3.52 ; dorsoexternal, dorsointernal and ventrointernal carinae strong with coarse, closely spaced dentate granules; internal carina moderate with large dentate granules spaced well apart; external carina obsolete, smooth ridge with isolated medium granules; ventral surface smooth with isolated small proximal granules, external surface smooth, dorsal and internal surfaces weakly, sparsely shagreened; linear cluster of 17–20 accessory macrosetae on lower distal external surface; patella : slender, L/ W 3.20 ; dorsointernal carina moderate with coarse non-contiguous granulation; dorsomedian carina weak, almost smooth; dorsoexternal, external, ventroexternal carinae weak, smooth, ventromedian carina obsolete; ventrointernal carina weak, with well spaced medium granules and ventral patellar spur; internal carina moderate, with well spaced small to medium dentate granules and dorsal patellar spur; all intercarinal surfaces smooth; chela : slender, L/ W 6.34 , movable finger L/ manus ventral L 2.2; dorsal marginal and ventroexternal carinae weak, smooth, other carinae obsolete; all intercarinal surfaces smooth; manus and fixed finger with sparse short macrosetae; movable finger with numerous short macrosetae on ventral aspect, culminating in dense subapical brush; 13 primary denticle subrows on movable fingers, 12–13 on fixed fingers; 12 subrows flanked by internal and external accessory denticles. Trichobothriotaxy : orthobothriotaxic, type A  ( Vachon, 1974 ), db on fixed finger distal to est . Legs. Moderately long, slender, femur III L/ carapace L 1.13, patella III L/D 4.00; inferior carinae strongly denticulate on femur I–IV and patella I–III, almost smooth on patella IV; tibia III–IV with long spurs; retrolateral tarsal spurs simple, non-setose; prolateral tarsal spurs basally bifurcate, bearing 1–5 macrosetae; basitarsi I–III with well developed bristlecombs, at least as wide as basitarsal segment; basitarsus III setal counts (left/ right): retrosuperior 20/20, retroinferior 15/14 (including basal accessory seta), inferior 15/12; ventral surface of telotarsi with numerous, long fine macrosetae. Measurements of holotype male (mm). Total L 72.50; metasoma + telson L 45.50; carapace L 8.36, W 8.08 , carapace preocular L 3.72; metasomal segments (L/ W/ D) I 5.90/ 4.59/ 3.96, II 7.05/ 4.13/ 3.81, III 7.17/ 3.84/ 3.65, IV 7.87/ 3.55/ 3.32, V 9.09/ 3.66/ 3.02; telson L 8.20; vesicle L 4.40, W 3.22 , D 3.11; pedipalp chela L 15.02, manus ventral L 4.93, manus W 2.37 , manus D 2.84, fixed finger L 8.96, movable finger L 10.84; pedipalp femur L 7.47, W 2.12 , patella L 8.94, W 2.79 ; pectine L 11.09; leg III femur L 9.46; leg III patella L 7.91, D 1.978.

males	females
Thumrait	central coast	Masirah Island	Thumrait	central coast	Masirah Island
Pedipalp chela	6.19 – 7.11	5.87 – 6.31	6.01 – 6.78	5.35 – 6.57	5.13 – 5.80	5.27 – 6.13
L/ manus W	6.51 ± 0.29 (11)	6.13 ± 0.13 (9)	6.23 ± 0.22 (10)	5.95 ± 0.40 (13)	5.51 ± 0.26 (12)	5.56 ± 0.22 (16)
Patella III	3.86 – 4.48	3.43 – 4.10	3.58 – 4.00	3.66 – 4.00	3.45 – 3.88	2.12 – 2.54
L/D	4.06 ± 0.19 (9)	3.75 ± 0.22 (9)	3.83 ± 0.16 (10)	3.84 ± 0.11 (8)	3.63 ± 0.12 (12)	2.35 ± 0.12 (16)
Metasoma I	1.10 – 1.30	1.12 – 1.30	1.21 – 1.37	1.12 – 1.29	1.09 – 1.23	3.38 – 4.00
L/W	1.23 ± 0.06 (11)	1.20 ± 0.06 (8)	1.28 ± 0.05 (10)	1.22 ± 0.05 (13)	1.16 ± 0.04 (12)	3.70 ± 0.16 (16)
Metasoma II	1.57 – 1.72	1.50 – 1.70	1.57 – 1.79	1.52 – 1.73	1.49 – 1.62	1.15 – 1.32
L/W	1.66 ± 0.05 (11)	1.58 ± 0.07 (9)	1.67 ± 0.07 (10)	1.64 ± 0.06 (13)	1.56 ± 0.04 (12)	1.22 ± 0.04 (16)
Metasoma III	1.67 – 2.03	1.60 – 1.82	1.64 – 1.94	1.67 – 1.95	1.59 – 1.78	1.48 – 1.73
L/W	1.84 ± 0.11 (11)	1.70 ± 0.07 (9)	1.78 ± 0.08 (10)	1.79 ± 0.08 (13)	1.68 ± 0.06 (12)	1.62 ± 0.07 (16)
Metasoma IV	2.14 – 2.40	1.94 – 2.22	2.11 – 2.33	2.07 – 2.34	1.92 – 2.14	1.60 – 1.87
L/W	2.25 ± 0.07 (11)	2.07 ± 0.08 (9)	2.20 ± 0.07 (10)	2.18 ± 0.07 (13)	2.05 ± 0.07 (12)	1.75 ± 0.06 (16)
Metasoma V	2.40 – 2.67	2.19 – 2.49	2.32 – 2.74	2.15 – 2.63	2.20 – 2.39	1.87 – 2.30
L/W	2.50 ± 0.07 (11)	2.36 ± 0.10 (9)	2.53 ± 0.11 (10)	2.42 ± 0.13 (13)	2.27 ± 0.07 (12)	2.13 ± 0.11 (16)

Table 1: Variation in adult morphometric ratios among different populations of Leiurus macroctenus sp. n. in Oman (ranges, mean ± SD, sample sizes in parentheses). Higher values indicate more slender segments. Paratype female ( Thumrait ). Differs from male as follows: larger, more robust, wider body, carapace W/L 1.06; weaker carination on carapace, tergites and coxae; pectines narrower, shorter, tips extending to proximal 1/5 of trochanter IV, teeth smaller with shorter sensillar margins, mid-pectine sensillar margin L/ pectine L 0.084, mid-pectine sensillar margin L/ metasoma I W 0.189 ; 33–34 pectine teeth, basal pectine teeth nonoverlapping; sternite III with weak, narrow, finely granular median carinae; sternites IV–V with strong but narrower, more finely granulated lateral carinae, median carinae obsolete; sternites VI with narrower, finely granulated lateral carinae, weaker, narrower median carinae with regular small granules; VII with slightly stronger, granular carinae; intercarinal surfaces of sternites III–VI smooth medially, finely micro-shagreened anterolaterally; ventromedian carinae on metasoma II– III more strongly crenulate or dentate, with larger posterior denticles; pedipalp chela more robust, L/ W 5.35 . Measurements of paratype female (mm). Total L 84.00; metasoma + telson L 49.00; carapace L 9.70, W 10.29 , carapace preocular L 4.19; metasomal segments (L/ D/ W) I 6.27/ 4.54/ 5.17, II 7.56/ 4.44/ 4.58, III 7.79/ 4.28/ 4.37, IV 8.74/ 3.85/ 4.08, V 10.00/ 3.58/ 4.34; telson L 9.00; vesicle L 4.83, D 3.65, W 3.71 ; pedipalp chela L 17.18, manus ventral L 5.43, manus W 3.21 , manus D 3.85, fixed finger L 9.93, movable finger L 12.22; pedipalp femur L 8.54, W 2.59 , patella L 10.09, W 3.30 ; pectine L 11.71; leg III femur L 10.32; leg III patella L 8.60, D 2.19. Variation. Base color varied with location and substrate, being pale yellow in samples from Thumrait and Jiddat al Harasis, and light brown or orange in some coastal sites including Masirah Island. Morphometric variation was observed in subpopulations sampled from three separate regions: inland Nejd desert around Thumrait, central coast of Oman (Ras ad Duqm south to Ras Sawqirah) and Masirah Island. Variation in size was sexdependent: males from central coast were on average smaller with less size variation (carapace L 7.60 ± 0.22 mm ) than males from Thumrait (8.21 ± 0.50 mm ) or Masirah Island (7.78 ± 0.49 mm ); female mean sizes were similar for these populations, but sample variance was smallest in central coast and greatest in Thumrait populations. Variation in specific morphometric ratios among different populations is summarized in Table 1 . Samples from the central coast (Ras ad Duqm south to Ras Sawqirah) had on average less slender legs, pedipalp chelae and metasomal segments compared to samples from Thumrait and Masirah Island. In contrast, the morphometrics of Thumrait and Masirah Island populations were broadly overlapped. Overall morphometric variation is summarized in Table 4 . Sexual dimorphism : compared to females, the males had on average more slender pedipalps and larger pectine teeth ( Tab. 4B ). The number of denticles on ventromedian carinae of metasoma III was significantly lower in males (16.79 ± 1.61) than females (18.81 ± 1.47) (P = 0.00027, t test). Diagnostic characters : (1) leg III retrosuperior setae (bristle-comb): the number of setae exhibited a unimodal distribution with no significant difference between males and females; setal count = 19.96 ± 2.61 (mean ± SD, n = 100, sexes pooled), with 95/100 (95%) of basitarsi bearing 17 or more setae); (2) female pectine teeth: sensillar margins were relatively long over the length of the pectine, but shorter for the basal tooth; mid-pectine sensillar margin L/ carapace L 0.09 – 0.11 (0.10 ± 0.01, n = 28), basal-pectine sensillar margin L/ carapace L 0.03 – 0.08 (0.06 ± 0.01, n = 28). Basal teeth were also very different in proportions, with a longer trunk and shorter sensillar margin: mid-pectine trunk L/ sensillar margin L 0.31–0.49 (0.38 ± 0.04, n = 30), basal-pectine trunk L/ sensillar margin L 0.56–1.45 (0.96 ± 0.23, n = 28). Fig. 95B plots the range of covariation of the ratios of mid-pectine sensillar margin L/ carapace L, and midpectine sensillar margin L/ metasoma I W, showing the large morphometric difference from other Leiurus . The large pectine teeth are a more robust diagnostic character in females. In males, the enlargement of pectine teeth as a secondary sexual characteristic obscures the interspecies difference; (3) pedipalp fixed finger db was distal to est in 142/175 (81%), and proximal in 33/175 (19%) of fixed fingers ( Fig. 95A ). DISTRIBUTION. Known only from the central and north-eastern coast (including Masirah Island, reaching as far north as Ras Al Hadd), and the south-central plains (Jiddat al Harasis and Nejd Desert) of Oman . The elevation range of records is 2 – 800 m , with the lowest on the coast, and the highest in the Jabal Qara hills. ECOLOGY. These are arenicolous scorpions from coastal and adjacent inland alluvial plains and steppes in Oman . In the Nejd desert north of Jabal Qara, L. macroctenus was observed in burrows excavated in fine sedimentary soil around rock outcroppings. Individuals were often waiting in nocturnal ambush positions at the burrow entrance with carapace and pedipalps protruding, chela fingers open, while their mesosoma and metasoma were protected inside the burrow. In sandy wadis on Masirah Island, they were observed at night resting on open sand with tarsi spread in a wide circle, a characteristic behavior of psammophilous scorpions localizing moving prey by substrate vibrations detected by basitarsal slit sensillae ( Brownell, 1977 ; Brownell & Farley, 1979 ). Adaptation to sandy substrates is suggested by the well developed basitarsal bristle combs that are broader and denser than those of other Leiurus . On the mainland, L. macroctenus was absent from the Wahiba Sands and Rub’ al Khali. It could be excluded from those dunes by intraguild predation from the larger Apistobuthus , which is absent from Masirah Island. The Wahiba Sands also present an ecological barrier to the dispersal of other terrestrial fauna ( Munton 1988 ).