Taxonomic revision of the genus Asellia (Chiroptera: Hipposideridae) with a description of a new species from southern Arabia Author Benda, Petr Department of Zoology, National Museum (Natural History), Václavské nám. 68, CZ- 11 o 79 Praha 1, Czech Republic & Department of Zoology, Faculty of Science, Charles University, Viničná 7, CZ- 128 44 Praha 2, Czech Republic & Corresponding author: E-mail: petr _ benda @ nm. cz petr_benda@nm.cz Author Vallo, Peter Institute of Vertebrate éiology, Academy of Sciences of the Czech Republic, v. v. i., Květná 8, CZ- 603 6 o érno, Czech Republic Author Reiter, Antonín South Moravian Museum in Znojmo, Přemyslovců 8, CZ- 669 4 o Znojmo, Czech Republic & Department of Zoology, Faculty of Science, Charles University, Viničná 7, CZ- 128 44 Praha 2, Czech Republic text Acta Chiropterologica 2011 2011-03-23 13 2 245 270 journal article 10.3161/150811011X624749 59fb62a3-ae40-4cc0-b1c5-d36baebe11b3 1733-5329 3943317 Asellia arabica sp. nov. Synonymy Asellia tridens (Geoffroy, 1813) : Pockock, 1935: 442; Harrison, 1964: 98; Harrison, 1980: 390; Kingdon, 1990: 37 ; Harrison and Bates, 1991: 55. Asellia tridens tridens (Geoffroy, 1813) : Harrison, 1957: 5; Kock, 1969: 129; Nader, 1990: 340; Al-Jumaily, 1998: 483. Asellia patrizii De Beaux, 1931 : Al-Jumaily, 2004: 60. Type material Holotype : adult ( NMP 92790 [S+A]), Hawf ( Al Mahra Prov. ), 14 October 2005 , leg. P. Benda. Paratypes (9): 3♂♂ , 2♀♀ ( NMP 92791 , 92792– 92794 [S+A], 92789 [A]), Hawf ( Al Mahra Prov. ), 14 and 15 October 2005 , leg. P. Benda ; – 2♂♂ , 2♀♀ ( NMP 92795 , 92796 , 92798 [S+A], 92797 [A]), Damqawt ( Al Mahra Prov. ), 16 October 2005 , leg. P. Benda. Type locality Republic of Yemen , Province of Al Mahra , oasis of Hawf (easternmost edge of the country), 16°39’N , 53°03’E , 410 ma .s.l . Description and diagnosis Small bat and medium-sized to small representative of the genus Asellia Gray, 1838 . It is in most respects very similar to the other species of the genus, including the structure of the nose leaf (Fig. 11). In body and skull size, A. arabica sp. nov. clearly differs from both the largest and the smallest species of the genus, A. tridens (Geoffroy, 1813) and A. patrizii De Beaux, 1931 , respectively, but it partly overlaps with A. italosomalica De Beaux, 1931 ( Table 1 , Figs. 2 and 4 ). Forearm length 43.1–46.5 mm , condylocanine length of skull 14.2–15.3 mm , length of the upper tooth- row 5.7–6.1 mm . Asellia arabica sp. nov. has a relatively narrow skull (width across the zygomatic arches 8.7–9.3 mm ; relative zygomatic width, LaZ/LCc, 0.589 –0.622 ) and a relatively narrow and short ros-trum (width of rostrum across canines 4.4–4.8 mm ; relative width of rostrum across canines, CC/LCc, 0.297 –0.324 ) in comparison to the other Asellia species ; the skull and rostrum of A. arabica sp. nov. are, on average, the narrowest, and the rostrum is the shortest, relatively speaking. The braincase is low (height of braincase 4.7–5.1 mm ; relative height of braincase, ANc/LCc, 0.320 –0.342 ) and the tympanic bullae are rather large (largest horizontal diameter of bulla 2.6–2.9 mm ; relative size of bulla, LBT/LCc 0.181 –0.214 ). FIG. 10. Portrait of A. arabica sp. nov. from Ain Jarziz, Dhofar, SW Oman (photo by A. Reiter) TABLE 3. Forearm and skull dimensions (in mm) of the examined holotype specimens of Asellia . See Abbreviations and Terminology for explanation of dimension abbreviations
tridens diluta patrizii italosomalica arabica sp. n.
Pa r amete r MNHN BMNH MSNG MSNG NMP
1986-1068 12.11.14.2. 31313 30942 92790
LAt 49.10 52.40 39.00 43.40 46.50
LC r 18.47 18.76 15.02 17.08 16.81
LOc 18.48 14.88 16.97 16.52
LCc 16.33 16.43 13.24 14.85 14.71
LaZ 10.28 8.12 9.12 8.87
LaI 2.12 2.16 1.78 2.37 2.17
LaInf 6.60 5.66 4.80 5.52 5.19
LaN 7.48 7.08 6.18 7.02 6.69
LaM 8.54 7.43 8.08 7.86
ANc 5.60 5.81 4.53 5.08 4.82
AC r 6.97 7.48 6.15 6.43
LBT 2.98 2.62 2.81
CC 5.42 5.06 3.94 5.06 4.62
PP 6.38 6.32 4.68 5.83 5.39
M3M3 7.50 7.47 5.54 6.62 6.53
CM3 6.83 6.84 5.33 6.19 6.02
M1M3 4.16 4.23 3.25 3.73 3.61
CP 3.03 2.89 2.17 2.67 2.51
LMd 12.33 9.89 11.49 10.93
ACo 3.67 2.81 3.57 3.34
I1M3 6.49 7.67 7.14
CM3 7.42 5.73 6.82 6.51
M1M3 4.95 3.75 4.52 4.09
CP4 2.66 2.46 2.82 2.34
The baculum of A. arabica sp. nov. is a tiny bone, 1.0– 1.3 mm long, with a very narrow distal epiphysis, 0.1–0.2 mm wide (<18% of the baculum length) and a rather narrow proximal epiphysis that is 0.3–0.4 mm wide (<30% of the baculum length) ( Fig. 6l–n ). The colouration of the dorsal pelage of A. arabica sp. nov. is beige or pale-brownish grey (Fig. 11) with yellowish or pale rusty tinges; the ventral pelage is somewhat paler than the dorsal colouration. The nose leaf is almost unpigmented to very pale brownish; often the anterior leaf is slightly pigmented while the posterior leaf and supplementary leaflets are unpigmented. The wing membranes are pale brownish-grey. Dimensions of the holotype : see Table 3 . Genetics: Asellia arabica sp. nov. showed unique base positions within the mitochondrial gene for cytochrome b (1140 bp) at 24 sites: 486, 636, 681, 696, 750 (A→G), 201, 507 (C→A), 1092 (C→G), 145, 282, 492, 504, 690, 915, 958, 963, 1020, 1044 (C→T), 981, 1080, 1093 (T→C), 510, 717 (A/T→C), and 498 (G/T→C). At two sites, 324 and 609, uniquely different positions of Ginstead of A (A→G, A→A/G, and A→G/A) were present in three known haplotypes (DF1, DF2, DF3; see Appendix II). Asellia arabica sp. nov. was found to share identical unique base positions within the gene for cytochrome b with A. tridens (Geoffroy, 1813) at 88 sites: 18, 27, 105, 153, 237, 330, 357, 466, 582, 697, 709, 720, 807, 870, 933, 936, 945, 996, 1038, 1057, 1119 (A), 45, 57, 81, 84, 114, 165, 195, 204, 297, 315, 321, 329, 435, 438, 465, 477, 480, 519, 546, 564, 567, 589, 592, 618, 621, 624, 648, 687, 693, 699, 708, 713, 774, 789, 804, 831, 834, 840, 852, 858, 882, 888, 894, 916, 1126 (C), 129, 178, 372, 476, 591 (G), 66, 121, 126, 246, 285, 288, 345, 351, 399, 474, 528, 792, 897, 906, 999, 1077, and 1122 (T); and with A. italosomalica De Beaux, 1931 at 16 sites: 123, 864, 909, 984, 1023, 1116 (A), 174, 441, 447, 555, 663, 921, 1110 (C), 1017 (G), 684, and 828 (T).
Mitochondrial sequence of holotype and all examined paratypes ( NMP 9278992791 , 92793 , 92794 , 9279692798 ) — complete sequence of the mitochondrial gene for cytochrome b (GenBank Accession Number JF439015 ; haplotype DF1 — Appendix II), 5’ end: atg acc aac atc cga aaa tcc cac cca cta ttc aaa att atc aac gac tca ttc atc gac cta cct gcc ccc tca agc atc tcc tcc tga tga aac ttt ggc tca cta cta ggc gta tgc tta gct gtg caa atc cta aca gga tta ttc cta gcc ata cac tac aca tcc gac aca gcc acc gcc ttc tat tcc gtc aca cac atc tgc cga gac gtt aat tat ggc tga atc cta cgc tac ctt cat gcc aac gga gca tcc ata ttc ttc atc tgc ctt ttt ctt cat gta ggc cga gga att tac tac ggc tcc tac aca ttc aca gaa aca tga aac att ggt att att cta ctt ttc gcc gtc atg gca aca gcc ttc ata ggc tac gtc ctt cca tga gga caa ata tcc ttc tga gga gca aca gtc atc acc aac ctc ctc tca gcc atc cca tac atc gga act agc ctc gta gag tga gtt tga ggc ggc ttt tca gtc gac aaa gcc acc ctc act cga ttc ttc gcc ctc cac ttc ctc ctc cca ttc atc atc gca gcc ata gta ata gtc cac ctg cta ttc cta cat gaa acg ggc tca aac aac ccc aca gga atc ccg tca gac ata gac ata atc ccc ttc cac cca tac tac acc atc aag gat atc ctt ggc ctg atc cta ata atc ata gca ctg cta tcc cta gtc cta ttc gca cca gac ctc ctg gga gac cca gac aac tac act ccc gca aac cca ctc aac act cca ccc cat atc aaa cca gag tga tat ttc cta ttt gcc tac gcc atc cta cgg tca atc cca aac aaa cta gga gga gta gta gcc ctc gtc ctc tca atc ctt atc cta gct gta atc cct cta ctc cac aca tca aaa caa cgc agc ata acc ttc cgc cca tta agt caa tgc tta ttc tga ctc cta gtc gcc gac cta ctt aca cta aca tga atc ggg ggt caa cct gta gaa cac cca ttc att att atc gga caa ata gcc tca atc cta tac ttt ctc atc atc cta gtg ctc ctc cca ctc gca agc atc gca gaa aat cac cta tta aaa tga aga. Derivatio nominis The name arabica (Arabian) reflects the area of occurrence of the new species, i.e. the southern part of Arabia. Distribution Coastal areas of southern Arabia stretching from the Hadramaut Province in south-eastern Yemen (ca. 49°E ) to the Dhofar Province in south-western Oman (ca. 55°E ), i.e., ca. 650 km of a narrow coastal strip. Received 22 July 2010 , accepted 23 March 2011 APPENDIX I List od the specimens examined in morphological analysis Asellia arabica sp. nov. Oman (6): 2 ♂♂ , 2 ♀♀ ( NMP 92721–92724 [S+A]), Ain Jarziz ( Dhofar Prov. ), 27 October 2009 , leg. P. Benda , A. Reiter and M. Uhrin ; – 1 ♀ ( NMP 92753 [A]), Ain Tabruq ( Dhofar Prov. ), 29 October 2009 , leg. A. Reiter ; – 1 ind. ( BMNH 34.8 .4.1 [S+B]), Cave of Sahaur , Qara Mts. , SE Arabia , date unlisted, leg. B. S. Thomas. Yemen (14): 2 ♂♂ , 2 ♀♀ ( NMP 92795 , 92796 , 92798 [S+A], 92797 [A]), Damqawt ( Al Mahra Prov. ), 16 October 2005 , leg. P. Benda ; – 4 ♂♂ , 2 ♀♀ ( NMP 92790 , 92791 , 92792–92794 [S+A], 92789 [A]), Hawf ( Al Mahra Prov. ), 14 and 15 October 2005 , leg. P. Benda ; – 1 ♂ ( BCSU 013 [B]), Ryan ( Hadramaut Prov. ), July 1995 , leg. A. Basmaidi ; – 2 ♂♂ , 1 ♀ ( BCSU 128 , 136 , 195 [S+B]), Shikhawi ( Hadramaut Prov. ), 12 March 2000 , leg. A. K. Nasher. Asellia italosomalica De Beaux, 1931 Somalia (16): 1 ♀ ( MSNG 32582 [S+B]), Bender -Cassim , January–February 1932 , leg. I. Zanetti ; – 1 ♀ ( MZUF 8277 [S+A]), Callis , 20 October 1973 , leg. Granchi and B. Lanza ; – 2 ♂♂ , 2 ♀♀ ( MSNG 12232 a–d [S+B], incl. the paratype of A. tridens italosomalica De Beaux, 1931 ), Dolo , May–July 1911 , leg. C. Citerni ; – 2 ♂♂ ( MZUF 9940 , 9941 [S+A]), Mahas , 12 February 1977 , leg. A. Simonetta ; – 1 ♀ ( MSNG 30942 [S+B], holotype of A. tridens italosomalica De Beaux, 1931 ), Oddur , 1929, leg. N. Mosconi Bronzi ; – 1 ♀ ( MZUF 9942 [S+A]), Pozzi di Mahas , 11 April 1977 , leg. A. Simonetta ; – 1 ♂ , 1 ♀ ( MZUF 6291 [S+B], 6305 [S+A]), Run , 15 and 18 August 1969, leg. B. Lanza ; – 2 ♂♂ , 2 ♀♀ ( MZUF 13099 , 13100 [S+A], 15728 , 15734 [S+A]), Showli Berdi , 15 March 1984 and 15 November 1985 , leg. L. Chellazi and Messana. Yemen , Socotra (15): 10 ♀♀ ( BCSU pb2718 , 2719 , 2721 , BMNH 54.1010 ., NMP 90571–90575 [S+A], BCSU pb2720 , NMP 90570 [A]), Kam , date unlisted, leg. G. B. Popov , 5 May 2004 , leg. P. Benda and A. Reiter ; – 4 ♂♂ ( BCSU pb2749 , NMP 90590 , 90592 [S+A], NMP 90591 [A]), Mazaaba , 14 May 2004 , leg. P. Benda and A. Reiter ; – 1 ♂ ( NMP 90579 [S+A]), Suq , 7 May 2004 , leg. P. Benda and A. Reiter. Asellia patrizii De Beaux, 1931 Eritrea (8): 3 ♂♂ , 1 ♀ ( MSNG 33203 , 33204 [S+B], 31315a , 31315b [S+A], incl. three paratypes of A. patrizii De Beaux, 1931 ), Assab , July 1893 , leg. G. Pestalozza , October 1906, leg. P. Felter; 1 ♂ , 1 ♀ ( MSNG 31313 , 31314 [S+A], incl. the holotype of A. patrizii De Beaux, 1931 ), Gaarre , Dancalia , December 1928 , leg. S. Patrizi ; 2 ♀♀ ( BMNH 70.2288 ., 70.2289 . [S]), Nocra Island , Dahlak Islands , 27 December 1969 , leg. M. J. Largen. Ethiopia (8): 1 ♂ , 1 ♀ ( SMF 44998 , 44999 [S]), “an der Bahnlinie westl. zw. Metahara und Lake Basaka , Awash River Gebiet ”, 11 September 1973 , leg. H. Rupp ; 1 ♀ ( SMF 52377 [S+B]), W of Metahara , Shoa , 28 December 1970 , ded. M. Largen; 2 ♂♂ ( MZUF 6546 , 6547 [S+B]), Awash NP , 4.5 km from Metahara , 12 April 1971 , leg. M. L. Azzaroli and B. Lanza ; 1 ♂ , 1 ♀ ( MZUF 6545 [A], 6548 [S+A]), Dint. Metahara , Mte. Fantalle , 14 April 1971 , leg. M. L. Azzaroli and B. Lanza ; 1 ♂ ( BMNH 70.475 . [S]), North bank of river, Awash River Valley , Awash NP, Shoa , 25 September 1968 , leg. P. Morris. Asellia tridens (Geoffroy, 1813) Afghanistan (10): 9 inds. ( ZFMK 96.022 , 96.023 [S]), cave nr Kuh-i-Duzd , nr Dilaram , 32°11’N , 63°27’E , 1965, leg. J. Niethammer and D. Meyer -Oehme ; – 1 ♀ ( SMF 39167 [S]), Ebn-e-Yamin , Farah , 2 September 1965 , leg. D. Meyer -Oehme . – Algeria (20): 9 ♂♂ , 2 ♀♀ ( ISEA 9523 , 9524 , 9527 , 9528 , 9530 , 9532 , 9533 , IVB A477 [S+B], ISEA 9534 [S], 9526 , 9531 [B]), Ain Ouarka , 20 May 1982 , leg. K. Kowalski , B. Rzebik-Kowalska and J. Gaisler ; – 2 ♂♂ , 1 ♀ ( ISEA 9522 , 9534 , 9535 [S+B]), Benni Abbes , 15 May 1981, 19 July 1983 , leg. K. Kowalski and B. Rzebik-Kowalska ; – 1 ind. ( BMNH 94.7.12.1. [S]), Biskra , date and collector unlisted ; – 1 ♂ ( ISEA 9536 [S+B]), Brezina , 23 July 1983 , leg. K. Kowalski and B. Rzebik-Kowalska ; – 3 ♂♂ , 1 ♀ ( BMNH 12.11.14.1. , 12.11.14.2 ., MSNG 33629a , b [S+B], incl. the holotype of A. tridens diluta Andersen, 1918 ), El Golea , 16 May 1912 , leg. W. Rotschild and E. Hartert , April 1938, leg. P. Laurent. Egypt (47): 2 ♀♀ ( MSNG 33202a , b [S+B]), Abu Simbel , 27 January 1904 , leg. E. A. D’Albertis ; – 8 ♂♂ , 2 ♀♀ ( NMP 92585–92592 [S+A], 92593 , 92594 [A]), Aswan , 24 January 2010 , leg. P. Benda , I. Horáček and R. Lučan ; – 2 ♂♂ ( IVB 22 , 27 [S+B]), Dandara , temple, 27 April 1969 , leg. J. Gaisler ; – 1 ♂ ( MNHN 1986-1068 [S+B]), Egypte [= Thebes], leg. E. Geoffroy Saint-Hilaire ( lectotype of Rhinolophus tridens Geoffroy, 1813 ); 4 ♂♂ , 2 ♀♀ ( MSNG 33201a–d , 6994 , 6995 [S+B]), Garbu Suan , in faccia ad Assuan , February 1907 , leg. E. A. D’Albertis ; – 4 ♂♂ ( MSNG 33200a–d [S+B]), Sakkarah ( Cairo ), May 1906 , leg. F. W. Innes Bey ; – 3 inds. ( NMP 90351 , 90352 , 90354 [S+A]), Shali , Siwa oasis , 12 April 2002 , leg. P. Munclinger and P. Nová ; – 17 ♂♂ , 2 inds. ( IVB 30 , 31 , 33–36 , 38–43 , 45–51 [S+B]), Thebes , tombs, 30 April 1969 , leg. J. Gaisler. Ethiopia : 3 inds. ( BMNH 70.472 . – 70.474 . [S]), from cave floor, Awash NP , Shoa , 28 September 1968 , leg. P. Morris. Gambia (3): 1 ♂ ( SMF 82480 [S]), Georgetown [=Janjangbureh], MacCarthy Island , 20 April 1995 , leg. D. Kock ; 1 ♂ ( SMF 92154 [S]), Janjangbureh , 22 January 2001 , leg. L. Barnett and C. Emms ; – 1 ♀ ( BMNH 49.476 . [S+B]), 10 mi SE of Kontaur , 24 August 1948 , leg. A. P. Buxton. Iran (26): 5 ♂♂ , 10 ♀♀ , 6 inds. ( NMP 48173–48187 [S+A], JOC unnumbered [S+Sk]), Choqa Zanbil , 15 October 1998 , leg. P. Benda , J. Obuch and M. Uhrin ; – 1 ♂ ( BMNH 77.824 . [S+B]), Chehar -Zanbil , Dezful , SW Iran , 19 May 1964 , leg. R. Szbokar ; 1 ind. ( BMNH 77.827 . [S+B]), Iranshahr , Baluchistan , 12 March 1975 , ded. E. Etemad ; – 1 ♂ ( BMNH 77.826 . [S+B]), Kahnuge , Sof Kerman , 3 March 1971 , ded. E. Etemad ; – 1 ♀ ( BMNH 77.825 . [S+B]), Rudan Minab , SW of Iranshah , South coast of Iran , 10 March 1966 , leg. E. Etemad ; – 1 ind. ( BMNH 6.1.2.1 . [S]), Seistan , date and collector unlisted . – Iraq (8): 8 ♀♀ ( NMW 21994 , 2199622002 [S+A]), Mosul , 5 December 1910 , leg. S. Hassoun. Israel (1): 1 ♀ ( SMF 18987 [S]), Jaffa , ded. Z. Lev . – Libya (4): 1 ♀ ( NMP 48317 [S+A]), Ghat , 3 October 1999 , leg. P. Benda ; – 1 ind. ( MSNG 54783 [A]), Serdeles ( Tripolitania ), March 1934 , leg. G. Scortecci ; – 1 ♀ ( MSNG 32180 [S+B]), Ubari , October 1932 , leg. L. Cipriani ; – 1 ♀ ( MSNG 33222 [S+B]), Zuila , September 1933 , leg. E. Zavattari. Mauritania (9): 1 ♀ ( MZUF 20732 [A]), Ouadane , 23 November 2002 , leg. P. Agnelli ; – 1 ♂ (NMP 93647 [S+A]), Tarjit , 21 October 2010 , leg. P. Benda , A. Reiter and M. Uhrin ; – 7 ♀♀ ( NMP 93639–93641 , 93643–93645 [S+A], 93642 [A]), Tin Labbé , 19 October 2010 , leg. P. Benda , A. Reiter and M. Uhrin. Morocco (3): 3 ♀♀ ( NMP pb38423844 [S+A]), Tassetift , 22 April 2008 , leg. P. Benda , J. Červený , A. Konečný and P. Vallo. Oman (21): 2 ♂♂ , 1 ♀ ( NMP 92627 , 92628 [S+A], 92629 [A]), Al Ghubrah , 18 October 2009 , leg. P. Benda , A. Reiter and M. Uhrin ; – 2 ♀♀ ( NMP 92761 , 92762 [S+A]), Al Mintirib , 1 November 2009 , leg. P. Benda , A. Reiter and M. Uhrin ; – 4 ♂♂ ( NMP 90770–92772 [S+A], 92773 [A]), Muqal , 1 November 2009 , leg. P. Benda , A. Reiter and M. Uhrin ; – 5 ♂♂ , 7 ♀♀ ( NMP 92683–92692 , 92694 [S+A], 92693 [A]), Shisr , 24 October 2009 , leg. P. Benda , A. Reiter and M. Uhrin. Pakistan (6): 1 ind. ( BMNH 85.8.1.368 . [S+B]), Mekran Coast , date unlisted, leg. Capt. C. J. Bringham ; – 1 ♂ ( BMNH 25.7.3.1 . [S+B]), Nawab Issakhans Tomb , Tatta , Sind, 22 October 1922 , leg. C. McCann ; – 4 ♀♀ ( BMNH 19.11.7.7 .– 19.11.7.10 . [S+B]), Panjgur , Baluchistan , 7– 13 June 1918 , leg. E. J. Hotson. Saudi Arabia (8): 2 ♂♂ , 1 ♀ ( BMNH 25.4.3.1 .– 25.4.3.3 . [S+B]), Hufuf , C. Arabia , 4– 12 December 1923 , leg. R. E. Cheesman ; – 1 ♂ ( BMNH 1950.33 . [S+B]), Jedda , 20 October 1949 , leg. A. C. Trott ; – 4 ♀♀ ( SMF 80445–80448 [S]), Massullamiyah Cliff , N of al-Jubail, al-Sinaiyah, Eastern Prov. , 28 May 1992 , D. Kock and I. A. Nader . – Senegal (4): 3 ♂♂ , 1 ♀ ( ZFMK 76.2976.31 [S+B], 76.28 [B]), Tatki , 20 km E Fété-Olé , 21 November 1975 , leg. W. Boehme et al . – Sudan (33): 2 ♂♂ , 2 ♀♀ ( ZFMK 97.00197.003 , 97.005 [S+Sk+B]), Djebel , ca. 30 km NW of Khartoum , 18 March 1986 , leg. J. Niethammer ; – 2 ♂♂ , 7 ♀♀ , 6 inds. ( IVB 1–9 [S+B], NMP 90344–90349 [S+A]), Jebel el Azraq , 18 February 1966 , leg. P. Štys ; – 4 ♂♂ ( NMP 93664–93666 [S+A], 93667 [A]), Dongola Alajuz , 7 December 2010 , leg. P. Benda and J. Šmíd ; – 3 ♂♂ , 3 ♀♀ ( NMP 93651–93653 , 93655–93657 [S+A]), Karima , Jebel Barkal , 6 December 2010 , leg. P. Benda and J. Šmíd ; – 1 ♂ ( NMP 93680 [S+A]), Masoud , Bayudah Desert , 13 December 2010 , leg. P. Benda and J. Šmíd ; – 1 ♂ , 2 ♀♀ ( NMP 93670 , 93671 [S+A], 93672 [A]), Quikkah , 9 December 2010 , leg. P. Benda and J. Šmíd. Syria (44): 1 ♂ , 15 ♀♀ ( NMP 48019–48021 , 48024–48027 [S+A], 48022 , 48023 [S+B], SMF 55478 [S+A], 7410574110 [S+Sk+B]), Halabiyyeh , 15– 17 August 1978 , leg. R. Kinzelbach , 1 June 1989, leg. D. Kock, 17 June 1998, leg. M. Andreas, P. Benda and M. Uhrin ; – 1 ♂ ( NMP 48813 [S+A]), Qala’at ar Rahba , 17 May 2001 , leg. M. Andreas , P. Benda , A. Reiter and D. Weinfurtová ; 27 ♂♂ ( SMF 60365 , 73414–73418 , 74099 [S+Sk+B], 734111 , 73419 [S+Sk], 7340773410 , 73412 , 73413 [S+A], NMP 47926 [A], 48037 , 48038 , 4804048047 [S+A], 48039 [S+B]), Tadmor , Afqa cave , 11 March 1979 , leg. R. Kinzelbach , 20 September 1988, leg. D. Kock, 17 May 1989, leg. D. Kock, 30 April 1995, leg. M. Kaftan, 23 June 1998, leg. M. Andreas, P. Benda and M. Uhrin. Tchad (2): 1 ♂ , 1 ♀ ( BMNH 64.240 ., 64.241 . [S+B]), Zouar , Tibesti , Sahara , 1735 m , 9 and 16 October 1955 , leg. I. G. Gibson. Tunisia (24): 24 ♂♂ ( IVB 1.2.751.2.98 [S+B]), El Hammá , nr. Tozeur , autumn 1973, leg. Pavlík. Yemen (19): 1 ♂ ( BCSU 312 [S+B]), Bayt Al-Faqech ( Al-Hodeida Prov. ), October 2002 , leg. A. As-Sabri ; – 2 ♂♂ ( MSNG 33196a , b [S+B]), Mokkha , 1891, leg. G. Pestalozza ; – 3 ♂♂ , 8 inds. ( BMNH 95.6.1.11 .– 95.6.1.14 ., 99.11.6.93 ., 99.11.6.94 ., 99.11. 6.97 . [S], BMNH 99.3.14.25 ., MSNG 33197a, b , 33198 [S+B]), Aden , 1883 , 1892 , 1895 , 1899 , leg. Ruspoli , V. Ragazzi , R. O. Grant , J. W. Yerbury , A. B. Percival and W. Dodson ; 1 ♂ ( MSNG 33199 [S+B]), near Aden , 1893, leg. A. Pogliani ; 2 ♀♀ , 1 ind. ( BMNH 95.6.1.50 ., 95.6.1.51 ., 99.11.6.12 . [S+B]), Lahej , 9 and 28 March 1895, 9 May 1899 , leg. J. W. Yerbury , A. B. Percival and W. Dodson ; – 1 ♂ ( BMNH 57.426 . [B]), Qatn , E. Aden Prot. , Arabia, 2000’, 5 June 1956 , leg. D. J. Greathead.