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<mods:title id="5C6BAB842FF081703114BF2B0BA3BFA4">Notes on Compsobuthus: redescription of C. arabicus Levy et al., 1973 from Arabia, and description of two new species from North Africa (Scorpiones: Buthidae)</mods:title>
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captionTargetPageId-5="10" captionTargetPageId-6="11" captionTargetPageId-7="12" captionTargetPageId-8="13" captionTargetPageId-9="14" captionText-0="Figures 1–9: Compsobuthus arabicus Levy, Amitai &amp; Shulov, 1973, holotype female from Daugha, Saudi Arabia. Figures 1–2. Habitus in dorsal (1) and ventral (2) views. Scale bar: 10 mm. Figures 3–5. Metasoma and telson in lateral (3), dorsal (4) and ventral (5) views. Scale bar: 5 mm. Figure 6. Telson in lateral view. Figure 7. Carapace and tergites I–IV. Figure 8. Posterior coxosternal area, sternum, genital opercula, pectines and sternites III–IV. Figure 9. Right legs III–IV, tarsal segments, retrolateral aspect." captionText-1="Figures 10–17: Compsobuthus arabicus, holotype female, right pedipalp. Figures 10–12. Chela in dorsal (10), external (11) and ventrointernal (12) views. Figures 13–15. Patella in dorsal (13), external (14) and ventral (15) views. Figure 16. Trochanter and femur, dorsal view. Figure 17. Movable finger, dorsal view showing dentition. White circles: trichobothrial positions (Figures 10–14, 16)." captionText-10="Figures 69–77: Compsobuthus arabicus. Figure 69. Carapace and tergites I–II. Figures 70–71. Right chelicera, dorsal (70) and ventral (71) views. Figures 72–73. Right hemispermatophore, capsule region, convex (72) and anterior (73) views. Figures 74–75. Metasoma and telson, lateral (74) and ventral (75) aspects. Figures 76–77. Right leg III, telotarsus and basitarsus, retrolateral view (76), and telotarsus, ventral view (77). Scale bar: 5 mm (69); 1 mm (70–71); 0.68 mm (72–73); 3 mm (74–75); 1.175 mm (76); 0.84 mm (77). Males from: 55 km NW of Ibra (23°36.5'N 56°05.33'E, 22.XI.1995) (69, 74–75); W of Ghabah (21°23.89'N 57°09.56'E, 5.X.1994) (70–71, 76–77); 15 km NNE of Fasad (18°45.2'N 53°08.9'E, 29-30.I.1998) (72–73)." captionText-11="Figures 78–84: Compsobuthus arabicus. Figures 78–82. Right pedipalp, femur dorsal (78), patella dorsal (79), external (80); chela external (81), ventral (82). Figures 83–84. Left pectine, male (83), female (84). Scale bar: 1 mm. Male from 55 km NW of Ibra (23°36.5'N 56°05.33'E, 22.XI.1995) (78–83); female from 10 km N of Adam (22°29.66'N 57°33'E, 23.X.1995) (84)." captionText-12="Figures 85–93: Scatter plots showing variation in male morphometric ratios of Compsobuthus arabicus, and C. polisi Lowe, 2001. Metasoma I L/W vs. metasoma II L/W (85), metasoma III L/W vs. metasoma IV L/W (86), metasoma V L/D vs. metasoma V L/W (87), metasoma I L/D vs. metasoma II L/D (88), metasoma III L/D vs. metasoma IV L/D (89), pedipalp femur L/ carapace L vs. Pectine L/ Metasoma V W (90), pedipalp femur L/W vs. pedipalp patella L/W (91), pedipalp movable finger L/ manus ventral L vs. pedipalp chela L/W (92), and metasoma III L/W vs. metasoma V W/D (93). Black circles: C. arabicus samples from Oman; green circles: C. arabicus samples from United Arab Emirates; yellow circles: C. polisi samples (all from Oman)." captionText-13="Figures 94–102: Scatter plots showing variation in female morphometric ratios of Compsobuthus arabicus, and C. polisi. Metasoma I L/W vs. metasoma II L/W (94), metasoma III L/W vs. metasoma IV L/W (95), metasoma V L/D vs. metasoma V L/W (96), metasoma I L/D vs. metasoma II L/D (97), metasoma III L/D vs. metasoma IV L/D (98), pedipalp femur L/ carapace L vs. Pectine L/ Metasoma V W (99), pedipalp femur L/W vs. pedipalp patella L/W (100), pedipalp movable finger L/ manus ventral L vs. pedipalp chela L/W (101), and metasoma III L/W vs. metasoma V W/D (102).. Black circles: C. arabicus samples from Oman; red circle: C. arabicus holotype from Daugha, Saudi Arabia; yellow circles: C. polisi samples (all from Oman)." captionText-14="Figures 103–104. Compsobuthus arabicus, in vivo habitus of females from Oman, W of Wahiba Sands (103), and SW of Wahiba (104)." captionText-15="Figures 105–106. Compsobuthus arabicus, female from Oman, W of Wahiba sands, 22.02990°N 58.18490°E with newborns (105) and juveniles after first ecdysis (106)." captionText-16="Figures 107–108. Compsobuthus arabicus, localities. Figure 107. Oman, W of Wahiba sands. Figure 108. Oman, E of Aydam." captionText-17="Figure 109. Map showing records of Compsobuthus arabicus Levy, Amitai &amp; Shulov, 1973, in the southeastern Arabian Peninsula. The holotype and paratype localities are indicated by arrows. Plotted records were confirmed by examination of materials, with the exception of two of the three paratype localities of Thesiger." captionText-18="Figures 110–113. Compsobuthus turieli sp. n. Figures 110–111. Male holotype in dorsal (110) and ventral (111) views. Figures 112–113. Female paratype from Morocco, North of Zag in dorsal (112) and ventral (113) aspects. Scale bar: 10 mm." captionText-19="Figures 114–122: Compsobuthus turieli sp. n. Figures 114–117, 121–122. Male holotype, telson lateral (114), metasoma and telson lateral (115), dorsal (116), and ventral (117) views. Right chelicera dorsal (121) and ventral (122). Figures 118–120. Female paratype from Morocco, North of Zag, metasoma and telson lateral (118), dorsal (119), and ventral (120) views. Scale bar: 10 mm (115–120)." captionText-2="Figures 18–22: Compsobuthus arabicus, from Oman, habitus. Figures 18–19. Male from SW of Wahiba, 21°19.35'N 58°15.55'E, dorsal (18) and ventral (19) views. Figures 20–21. Female from Dhofar Province, Wadi Qitbit, 19°9'20&quot;N 54°30'27&quot;E. Scale bar: 10 mm." captionText-20="Figures 123–134: Compsobuthus turieli sp. n. Figures 123–126: Mesosoma. Figures 123, 126. Male holotype, carapace and tergites I–IV (123) and posterior coxosternal area and sternites (126). Figures 124–125. Female paratype from Morocco, North of Zag, carapace and tergites I–IV (124) and posterior coxosternal area and sternite III (125). Figures 127–134: Left legs, retrolateral aspects. Figures 127–130. Male holotype. Figures 131–134. Female paratype from Morocco, North of Zag." captionText-21="Figures 135–155: Compsobuthus turieli sp. n., pedipalps. Figures 135–144. Male holotype, chela dorsal (135), external (136) and ventrointernal (137), patella dorsal (138), external (139) and ventral (140), trochanter and femur internal (141), dorsal (142) views, and movable (143) and fixed (144) fingers dentition.Figures 145–155. Female paratype from Morocco, North of Zag, chela dorsal (145), external (146) and ventrointernal (147), patella dorsal (148), external (149) and ventral (150), trochanter and femur internal (151), ventral (152), and dorsal (153) views, and movable (154) and fixed (155) finger dentition. Trichobothrial pattern is indicated in Figures 135–139, 141–142 (white circles)." captionText-22="Figures 156–157. Compsobuthus turieli sp. n. Figure 156. Locality, Morocco, North of Zag. Figure 157. Female paratype from this locality with newborns." captionText-23="Figures 158–159. Compsobuthus turieli sp. n., juvenile (158) and female (159) paratypes from Morocco, North of Zag in vivo habitus." captionText-24="Figures 160–167: Hemispermatophores. Figures 160–163. Compsobuthus turieli sp. n., male holotype, whole hemispermatophore (160), capsule in convex compressed (161), anterior (162), and posterior (163) views. Figures 164–167. C. ullrichi sp. n., male holotype, whole hemispermatophore (164), capsule in posterior (165), convex compressed (166), and anterior (167) views. Scale bars: 1 mm (160, 164), 200 μm (161–163, 165–167)." captionText-25="Figures 168–171: Figures 168–169. Compsobuthus ullrichi sp. n., male holotype in dorsal (168) and ventral (169) views. Figures 170–171. C. levyi, male holotype in dorsal (170) and ventral (171) views. Scale bar: 10 mm." captionText-26="Figures 172–179: Figures 172, 174–176. Compsobuthus levyi, male holotype, telson lateral (172), metasoma and telson lateral (174), dorsal (175), and ventral (176) views. Figures 173, 177–179. C. ullrichi sp. n., male holotype, telson lateral (173), metasoma and telson lateral (177), dorsal (178), and ventral (179) views. Scale bar: 10 mm (174–179)." captionText-27="Figures 180–189: Figures 180–181, 184–187. Compsobuthus ullrichi sp. n., male holotype, carapace and tergites I–IV (180), posterior coxosternal area and sternites (181), and left legs I–IV, retrolateral aspects (184–187). Figures 182–183, 188–189. C. levyi, male holotype, carapace and tergites I–III (182), posterior coxosternal area and sternites (183), and left legs III–IV, retrolateral aspects (188–189)." captionText-28="Figures 190–209: Pedipalps. Figures 190–200. Compsobuthus ullrichi sp. n., male holotype, chela dorsal (190), external (191) and ventrointernal (192), patella dorsal (193), external (194) and ventral (195), trochanter and femur dorsointernal (196), dorsal (197), and ventral (198) views, and movable (199) and fixed (200) fingers dentition. Figures 201–209. C. levyi, male holotype, chela dorsal (201), external (202) and ventrointernal (203), patella dorsal (204), external (205) and ventral (206), trochanter and femur dorsal (207) views, and movable (208) and fixed (209) finger dentition. Trichobothrial pattern is indicated in Figures 190–194, 196–197, 201–205, 207 (white circles)." captionText-29="Figures 210–224: Mitotic metaphases (210, 213, 216, 219, 222), male postpachytenes (211, 214, 217, 220, 223), and ideograms (212, 215, 218, 221, 224) (y axis: % of the diploid chromosome length, dark grey marks: chromosomes in multivalent association) of Compsobuthus species from Arabia and North Africa. Figure 210. Female of C. acutecarinatus (2n=22). Figures 211–212. Male of C. acutecarinatus (2n=22, 11II). Figures 213–215. Male of C. acutecarinatus (2n=22, 9II+CIV). Figures 216–218. Male of C. arabicus (2n=22, 9II+CIV). Figures 219–221. Male of C. maindroni (2n=22, 11II). Figures 222–224. Male of C. ullrichi sp. n. (2n=22, 9II+CIV).Arrows show chromosomes in multivalent association during postpachytene (214, 217, 223). Scale bar: 10 μm" captionText-3="Figures 22–37: Figures 22–27: Compsobuthus arabicus, prosoma and mesosoma. Figures 22–23. Male, SW of Wahiba, carapace and tergites I–IV (22), posterior coxosternal area and sternites (23). Figures 24–25. Female, Wadi Qitbit, carapace and tergites I–III (24), posterior coxosternal area and sternite III (25). Figure 26. Female, Oman, W of Wahiba sands (22.02990°N 58.18490°E), posterior coxosternal area and sternites. Figure 27. Female, SW of Wahiba (21°19.35’N 58°15.55’E, XI.2017), posterior coxosternal area and sternites III–IV.Figures 28–29. C. polisi, female paratype from Oman, Wadi Dirif, carapace and tergites I–II (28), posterior coxosternal area and sternite III (29). Figures 30–37: C. arabicus, legs, tibiae and tarsi, retrolateral aspects. Figures 30–33. Male, right legs I–IV. Figures 34–35. Female, SW of Wahiba, left legs III–IV. Figures 36–37. Female, Wadi Qitbit, left legs III–IV." captionText-4="Figures 38–54: Compsobuthus arabicus, pedipalps. Figures 38–45. Male, SW of Wahiba, chela dorsal (38), external (39) and ventral (40), patella dorsal (41), external (42) and ventral (43), trochanter and femur internal (44), dorsal (45), and ventral (46) views. Figures 47–54. Female, SW of Wahiba, chela dorsal (47), external (48) and ventrointernal (49), patella dorsal (50), external (51) and ventral (52), trochanter and femur dorsointernal (53) and dorsal (54) views. White circles: trichobothrial positions (Figures 38–42, 44–45)." captionText-5="Figures 55–64: Figures 55–63. Compsobuthus arabicus. Figures 55–57. Male, SW of Wahiba, metasoma and telson lateral (55), ventral (56), and dorsal (57) views. Figures 58–60. Female, SW of Wahiba, metasoma and telson lateral (58), ventral (59), and dorsal (60) views. Figures 61–63. Female, Wadi Qitbit, metasoma and telson lateral (61), ventral (62), and dorsal (63) views. Figure 64. C. polisi Lowe, 2001, female paratype from Oman, Wadi Dirif, metasoma and telson lateral view. Scale bar: 10 mm." captionText-6="Figure 65. Compsobuthus arabicus. Male, Wahiba Sands (22°10.68'N 58°51.16'E, 7.X.1993), dorsal habitus. Color image with cuticular surface detail highlighted digitally by UV fluorescence. Scale bar: 5 mm." captionText-7="Figure 66. Compsobuthus arabicus. Male, Wahiba Sands (22°10.68'N 58°51.16'E, 7.X.1993), ventral habitus. Color image with cuticular surface detail highlighted digitally by UV fluorescence. Scale bar: 5 mm." captionText-8="Figure 67. Compsobuthus arabicus. Female, 30 km S of Adam (22°05.9'N 57°31.12'E, 19.III.1996), dorsal habitus. Color image with cuticular surface detail highlighted digitally by UV fluorescence. Scale bar: 5 mm." captionText-9="Figure 68. Compsobuthus arabicus. Female, 30 km S of Adam (22°05.9'N 57°31.12'E, 19.III.1996), ventral habitus.Color image with cuticular surface detail highlighted digitally by UV fluorescence. Scale bar: 5 mm." figureDoi-0="http://doi.org/10.5281/zenodo.5741451" figureDoi-1="http://doi.org/10.5281/zenodo.5741453" figureDoi-10="http://doi.org/10.5281/zenodo.5741471" figureDoi-11="http://doi.org/10.5281/zenodo.5741473" figureDoi-12="http://doi.org/10.5281/zenodo.5741475" figureDoi-13="http://doi.org/10.5281/zenodo.5741477" figureDoi-14="http://doi.org/10.5281/zenodo.5741479" figureDoi-15="http://doi.org/10.5281/zenodo.5741481" figureDoi-16="http://doi.org/10.5281/zenodo.5741485" figureDoi-17="http://doi.org/10.5281/zenodo.5741487" figureDoi-18="http://doi.org/10.5281/zenodo.5741489" figureDoi-19="http://doi.org/10.5281/zenodo.5741493" figureDoi-2="http://doi.org/10.5281/zenodo.5741455" figureDoi-20="http://doi.org/10.5281/zenodo.5741495" figureDoi-21="http://doi.org/10.5281/zenodo.5741497" figureDoi-22="http://doi.org/10.5281/zenodo.5741499" figureDoi-23="http://doi.org/10.5281/zenodo.5741501" figureDoi-24="http://doi.org/10.5281/zenodo.5741503" figureDoi-25="http://doi.org/10.5281/zenodo.5741505" figureDoi-26="http://doi.org/10.5281/zenodo.5741507" figureDoi-27="http://doi.org/10.5281/zenodo.5741509" figureDoi-28="http://doi.org/10.5281/zenodo.5741511" figureDoi-29="http://doi.org/10.5281/zenodo.6544482" figureDoi-3="http://doi.org/10.5281/zenodo.5741457" figureDoi-4="http://doi.org/10.5281/zenodo.5741459" figureDoi-5="http://doi.org/10.5281/zenodo.5741461" figureDoi-6="http://doi.org/10.5281/zenodo.5741463" figureDoi-7="http://doi.org/10.5281/zenodo.5741465" figureDoi-8="http://doi.org/10.5281/zenodo.5741467" figureDoi-9="http://doi.org/10.5281/zenodo.5741469" httpUri-0="https://zenodo.org/record/5741451/files/figure.png" httpUri-1="https://zenodo.org/record/5741453/files/figure.png" httpUri-10="https://zenodo.org/record/5741471/files/figure.png" httpUri-11="https://zenodo.org/record/5741473/files/figure.png" httpUri-12="https://zenodo.org/record/5741475/files/figure.png" httpUri-13="https://zenodo.org/record/5741477/files/figure.png" httpUri-14="https://zenodo.org/record/5741479/files/figure.png" httpUri-15="https://zenodo.org/record/5741481/files/figure.png" httpUri-16="https://zenodo.org/record/5741485/files/figure.png" httpUri-17="https://zenodo.org/record/5741487/files/figure.png" httpUri-18="https://zenodo.org/record/5741489/files/figure.png" httpUri-19="https://zenodo.org/record/5741493/files/figure.png" httpUri-2="https://zenodo.org/record/5741455/files/figure.png" httpUri-20="https://zenodo.org/record/5741495/files/figure.png" httpUri-21="https://zenodo.org/record/5741497/files/figure.png" httpUri-22="https://zenodo.org/record/5741499/files/figure.png" httpUri-23="https://zenodo.org/record/5741501/files/figure.png" httpUri-24="https://zenodo.org/record/5741503/files/figure.png" httpUri-25="https://zenodo.org/record/5741505/files/figure.png" httpUri-26="https://zenodo.org/record/5741507/files/figure.png" httpUri-27="https://zenodo.org/record/5741509/files/figure.png" httpUri-28="https://zenodo.org/record/5741511/files/figure.png" httpUri-29="https://zenodo.org/record/6544482/files/figure.png" httpUri-3="https://zenodo.org/record/5741457/files/figure.png" httpUri-4="https://zenodo.org/record/5741459/files/figure.png" httpUri-5="https://zenodo.org/record/5741461/files/figure.png" httpUri-6="https://zenodo.org/record/5741463/files/figure.png" httpUri-7="https://zenodo.org/record/5741465/files/figure.png" httpUri-8="https://zenodo.org/record/5741467/files/figure.png" httpUri-9="https://zenodo.org/record/5741469/files/figure.png" pageId="4" pageNumber="3">Figures 1–224</figureCitation>
, 
<tableCitation id="F41834D6FFA4107696B2A5125E19FE56" box="[467,584,369,393]" captionStart-0="Table 1" captionStart-1="Table 2" captionStart-2="Table 3" captionStart-3="Table 4" captionStartId-0="6.[113,169,998,1019]" captionStartId-1="15.[113,169,1123,1144]" captionStartId-2="34.[116,172,986,1007]" captionStartId-3="39.[117,173,697,718]" captionTargetId-0="graphics-45@6.[115,1476,237,975]" captionTargetId-1="graphics-22@15.[115,1471,234,1103]" captionTargetId-2="graphics-458@34.[119,1471,235,973]" captionTargetId-3="graphics-629@39.[118,1476,236,680]" captionTargetPageId-0="6" captionTargetPageId-1="15" captionTargetPageId-2="34" captionTargetPageId-3="39" captionText-0="Table 1. Comparative measurements of adults of Compsobuthus arabicus and C. turieli sp. n. Abbreviations: length (L), width (W, in carapace it corresponds to posterior width), depth (D)." captionText-1="Table 2. Compsobuthus arabicus, morphometric variation.Abbreviations: length (L), width (W), depth (D)." captionText-2="Table 3. Comparative measurements of adults of Compsobuthus ullrichi sp. n., C. levyi and C. kabateki. Abbreviations: length (L), width (W, in carapace it corresponds to posterior width), depth (D)." captionText-3="Table 4. The diploid number, the postpachytene configuration, the percentage of the total chromosome length of the diploid set (% TCL) and the origin of Compsobuthus species cytogenetically investigated. Specimens karyotyped: C. acutecarinatus (Simon, 1882): Oman, Dhalkut beach, 16.6989°N 53.2214°E, 1♂; E of Aydam, 17.8887°N 53.06640°E, 1♂; W of Qairoon Hairitti,17.2564°N 54.0220°E, 2♂; WadiAsh Shuwaymiyyah, 17.9293°N 55.5510°E, 1♂1♀. C. arabicus Levy, Amitai &amp; Shulov, 1973: Oman, Al Wasil, 22.4914°N 58.7151°E, 2♂; Jabal Bani Jabir, 22.8132°N 59.0589°E, 1♂; E of Aydam, 17.8887°N 53.06640°E, 1♂; W of Wahiba, 22.0299°N 58.1849°E, 1♂. C. maindroni (Kraepelin, 1900): Oman, Khatm Ha‘ir Asfar, 24.7750°N 55.9146°E, 2♂; Say Plateau, 23.1185°N 57.6311°E, 2♂. C. ullrichi sp. n., 1♂ holotype." httpUri-0="http://table.plazi.org/id/EDE551E5FFA610749710A7855E70FBC6" httpUri-1="http://table.plazi.org/id/EDE551E5FFAF107D9710A000582FFBA6" httpUri-2="http://table.plazi.org/id/EDE551E5FF8210509715A7B95EC7FBD2" httpUri-3="http://table.plazi.org/id/EDE551E5FF8710559714A6DA5F1EFCA3" pageId="4" pageNumber="3" tableUuid-0="EDE551E5FFA610749710A7855E70FBC6" tableUuid-1="EDE551E5FFAF107D9710A000582FFBA6" tableUuid-2="EDE551E5FF8210509715A7B95EC7FBD2" tableUuid-3="EDE551E5FF8710559714A6DA5F1EFCA3">Tables 1–4</tableCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="F18052E6FFA410769710A5CD5EA1FDB9" pageId="4" pageNumber="3" type="reference_group">
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<taxonomicName id="7E9A7AEEFFA410769710A5CD5E36FE19" ID-CoL="3SVP" authority="Vachon, 1949: 93" authorityName="Vachon" authorityPageNumber="93" authorityYear="1949" box="[113,615,430,454]" class="Arachnida" family="Buthidae" genus="Compsobuthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="genus">
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<emphasis id="8BEEDD7FFFA410769710A5CD5D41FE19" box="[113,272,430,454]" italics="true" pageId="4" pageNumber="3">Compsobuthus</emphasis>
<bibCitation id="6CB00901FFA410769678A5CD5D8AFE19" author="VACHON, M." box="[281,475,430,454]" page="93" pageId="4" pageNumber="3" year="1949">Vachon, 1949: 93</bibCitation>
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(
<treatmentCitation id="383B277CFFA41076968DA5CD5E0EFE19" author="VACHON, M." box="[492,607,430,454]" page="213" pageId="4" pageNumber="3" year="1952">
<bibRefCitation id="DD0B7C9CFFA41076968DA5CD5E0EFE19" author="VACHON, M." bookContentInfo="1 - 482" box="[492,607,430,454]" journalOrPublisher="Institut Pasteur d'Algerie, Alger" pageId="4" pageNumber="3" pagination="46 - 104" part="29" refId="ref16361" refString="VACHON, M. 1952. Etudes sur les scorpions. Institut Pasteur d'Algerie, Alger, 1 - 482. (published 1948 - 1951 in Archives de l'Institut Pasteur d'Algerie, 1948, 26: 25 - 90, 162 - 208, 288 - 316, 441 - 481. 1949, 27: 66 - 100, 134 - 169, 281 - 288, 334 - 396. 1950, 28: 152 - 216, 383 - 413. 1951, 29: 46 - 104)." title="Etudes sur les scorpions" type="book chapter" year="1952">1952: 213</bibRefCitation>
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)
</taxonomicName>
; 
<treatmentCitation id="383B277CFFA410769514A5CD5D5FFE39" author="FET, V. &amp; G. LOWE" page="124" pageId="4" pageNumber="3" year="2000">
<bibRefCitation id="DD0B7C9CFFA410769514A5CD5D5FFE39" author="Fet &amp; Lowe" bookContentInfo="689 pp." editor="Fet, V. &amp; W. D. Sissom &amp; G. Lowe &amp; M. E. Braunwalder" firstAuthor="Fet" journalOrPublisher="New York: The New York Entomological Society" page="124" pageId="4" pageNumber="3" pagination="54 - 286" refId="ref14898" refString="FET, V. &amp; G. LOWE. 2000. Family Buthidae C. L. Koch, 1837. Pp. 54 - 286 in Fet, V., W. D. Sissom, G. Lowe &amp; M. E. Braunwalder. Catalog of the Scorpions of the World (1758 - 1998). New York: The New York Entomological Society, 689 pp." title="Family Buthidae C. L. Koch, 1837" type="book chapter" volumeTitle="Catalog of the Scorpions of the World (1758 - 1998)" year="2000">Fet &amp; Lowe, 2000: 124</bibRefCitation>
</treatmentCitation>
(complete reference list until 1998); 
<treatmentCitation id="383B277CFFA4107695C4A5AD5D5BFDD9" author="KOVARIK, F." page="88" pageId="4" pageNumber="3" year="2003">
<bibRefCitation id="DD0B7C9CFFA4107695C4A5AD5D5BFDD9" author="KovarIk" firstAuthor="KovarIk" page="88" pageId="4" pageNumber="3" pagination="87 - 112" refId="ref15081" refString="KOVARIK, F. 2003. Eight new species of Compsobuthus Vachon, 1949 from Africa and Asia (Scorpiones: Buthidae). Serket, 8 (3): 87 - 112." type="journal article" year="2003">KovařÍk, 2003: 88</bibRefCitation>
</treatmentCitation>
(in part); 
<treatmentCitation id="383B277CFFA4107696F4A58D5E3EFDD9" author="KOVARIK, F." box="[405,623,494,520]" page="31" pageId="4" pageNumber="3" year="2009">
<bibRefCitation id="DD0B7C9CFFA4107696F4A58D5E3EFDD9" author="KovarIk" bookContentInfo="170 pp." box="[405,623,494,520]" firstAuthor="KovarIk" journalOrPublisher="Prague: Clairon Production" page="31" pageId="4" pageNumber="3" refId="ref15116" refString="KOVARIK, F. 2009. Illustrated catalog of scorpions. Part I. Introductory remarks; keys to families and genera; subfamily Scorpioninae with keys to Heterometrus and Pandinus species. Prague: Clairon Production, 170 pp." title="Illustrated catalog of scorpions. Part I. Introductory remarks; keys to families and genera; subfamily Scorpioninae with keys to Heterometrus and Pandinus species." type="book" year="2009">KovařÍk, 2009: 31</bibRefCitation>
</treatmentCitation>
; 
<treatmentCitation id="383B277CFFA4107695E7A58D5E19FDF9" author="KOVARIK, F. &amp; A. A. OJANGUREN-AFFILASTRO" page="145–158" pageId="4" pageNumber="3" year="2003">
<bibRefCitation id="DD0B7C9CFFA4107695E7A58D5E19FDF9" author="KovarIk &amp; Ojanguren-Affilastro" bookContentInfo="400 pp." firstAuthor="KovarIk" journalOrPublisher="Prague: Clairon Production" page="145 - 158" pageId="4" pageNumber="3" refId="ref15488" refString="KOVARIK, F. &amp; A. A. OJANGUREN-AFFILASTRO. 2013. Illustrated catalog of scorpions. Part II. Bothriuridae; Chaerilidae; Buthidae I. Genera Compsobuthus, Hottentotta, Isometrus, Lychas, and Sassanidotus. Prague: Clairon Production, 400 pp." title="Illustrated catalog of scorpions. Part II. Bothriuridae; Chaerilidae; Buthidae I. Genera Compsobuthus, Hottentotta, Isometrus, Lychas, and Sassanidotus" type="book" year="2013">KovařÍk &amp; Ojanguren-Affilastro, 2013: 145–158</bibRefCitation>
</treatmentCitation>
, figs. 777–941; 
<treatmentCitation id="383B277CFFA4107697C0A64D5D9CFD99" author="KOVARIK, F., G. LOWE, J. PLISKOVA &amp; F. STAHLAVSKY" box="[161,461,558,584]" httpUri="http://treatment.plazi.org/id/BD662E6FFF89FFDEFE5E85AA8538FBF2" page="1-21" pageId="4" pageNumber="3" year="2016">
<bibRefCitation id="DD0B7C9CFFA4107697C0A64D5D9CFD99" author="KovarIk" box="[161,461,558,584]" etAl="et al." firstAuthor="KovarIk" page="1 - 21" pageId="4" pageNumber="3" pagination="1 - 21" refId="ref15356" refString="KOVARIK, F., G. LOWE, J. PLISKOVA &amp; F. STAHLAVSKY. 2016 a. Scorpions of the Horn of Africa (Arachnida, Scorpiones). Part VI. Compsobuthus Vachon, 1949 (Buthidae) with description of C. eritreaensis sp. n. Euscorpius, 226: 1 - 21." type="journal article" year="2016" yearSuffix="a">KovařÍk et al., 2016a: 1–21</bibRefCitation>
</treatmentCitation>
, figs. 1–77; 
<treatmentCitation id="383B277CFFA410769535A64D5C86FDB9" ID-DOI="http://doi.org/10.5281/zenodo.5741277" ID-Zenodo-Dep="5741277" author="KOVARIK, F." httpUri="http://treatment.plazi.org/id/CD1987EBFF88FFDEFEC3F955FD0BB9F0" page="1-11" pageId="4" pageNumber="3" year="2018">
<bibRefCitation id="DD0B7C9CFFA410769535A64D5C86FDB9" author="KovarIk" firstAuthor="KovarIk" page="1 - 11" pageId="4" pageNumber="3" pagination="1 - 11" refId="ref15194" refString="KOVARIK, F. 2018 a. Scorpions of the Horn of Africa (Arachnida, Scorpiones). Part XVI. Compsobuthus maidensis sp. n. (Buthidae) from Somalia. Euscorpius, 260: 1 - 11." type="journal article" year="2018" yearSuffix="a">KovařÍk, 2018a: 1–11</bibRefCitation>
</treatmentCitation>
, figs. 1–39, tab. 1; 
<treatmentCitation id="383B277CFFA4107696C2A62D5ED2FDB9" ID-DOI="http://doi.org/10.5281/zenodo.5741277" ID-Zenodo-Dep="5741277" author="KOVARIK, F." box="[419,643,590,616]" httpUri="http://treatment.plazi.org/id/CD1987EBFF88FFDEFEC3F955FD0BB9F0" page="4-6" pageId="4" pageNumber="3" year="2018">
<bibRefCitation id="DD0B7C9CFFA4107696C2A62D5ED2FDB9" author="KovarIk" box="[419,643,590,616]" firstAuthor="KovarIk" page="4 - 6" pageId="4" pageNumber="3" pagination="1 - 12" refId="ref15236" refString="KOVARIK, F. 2018 b. Notes on the genera Buthacus, Compsobuthus, and Lanzatus with several synonymies and corrections of published characters (Scorpiones: Buthidae). Euscorpius, 269: 1 - 12." type="journal article" year="2018" yearSuffix="b">KovařÍk, 2018b: 4–6</bibRefCitation>
</treatmentCitation>
, figs. 1–9.
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="F18052E6FFA410769710A6ED5EDFFD79" box="[113,654,654,678]" pageId="4" pageNumber="3" type="type_taxon">
<paragraph id="B925016DFFA410769710A6ED5EDFFD79" blockId="4.[113,654,654,678]" box="[113,654,654,678]" pageId="4" pageNumber="3">
<typeStatus id="6621BFCFFFA410769710A6ED5CF6FD7A" box="[113,167,654,677]" pageId="4" pageNumber="3">TYPE</typeStatus>
SPECIES. 
<taxonomicName id="7E9A7AEEFFA410769669A6ED5EDBFD79" authority="Simon, 1882" authorityName="Simon" authorityYear="1882" box="[264,650,654,678]" class="Arachnida" family="Buthidae" genus="Buthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="acutecarinatus">
<emphasis id="8BEEDD7FFFA410769669A6ED5DA9FD79" box="[264,504,654,678]" italics="true" pageId="4" pageNumber="3">Buthus acutecarinatus</emphasis>
Simon, 1882
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="F18052E6FFA410769710A6AD5EE0FA19" pageId="4" pageNumber="3" type="diagnosis">
<paragraph id="B925016DFFA410769710A6AD5EE0FA19" blockId="4.[113,773,718,1478]" pageId="4" pageNumber="3">
DIAGNOSIS. Small to medium-sized buthid scorpions, total length 
<quantity id="7E62AC88FFA4107697A0A68D5D6FFCDA" box="[193,318,750,774]" metricMagnitude="-2" metricUnit="m" metricValue="3.75" metricValueMax="5.5" metricValueMin="2.0" pageId="4" pageNumber="3" unit="mm" value="37.5" valueMax="55.0" valueMin="20.0">20–55 mm</quantity>
. Carapace with distinct carinae, central lateral and posterior lateral carinae connected to form continuous linear series of granules, projecting beyond posterior margin as distinct spiniform processes. Carapace in lateral view with entire dorsal surface horizontal, 5 pairs of lateral eyes. Cheliceral fixed finger with two ventral denticles. Sternum 
<typeStatus id="6621BFCFFFA410769637A7CD5DD4FC19" box="[342,389,942,966]" pageId="4" pageNumber="3">type</typeStatus>
1, sub-triangular. Pectines with fulcra, pectine teeth number 9–34. Hemispermatophore flagelliform, capsule in ‘3+1’-lobe configuration (‘ 
<taxonomicName id="7E9A7AEEFFA4107695D0A78D5EADFBD9" authorityName="Leach" authorityYear="1815" box="[689,764,1006,1030]" class="Arachnida" family="Buthidae" genus="Buthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="8BEEDD7FFFA4107695D0A78D5EADFBD9" box="[689,764,1006,1030]" italics="true" pageId="4" pageNumber="3">Buthus</emphasis>
</taxonomicName>
’ group; 
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), with 3 sperm hemiduct lobes well separated from flagellum, basal lobe a well-developed acuminate hook. Tergites I–VI with three carinae projecting beyond posterior margins as distinct spiniform processes. Sternites III–VI with slit-like spiracles. Orthobothriotaxic 
<typeStatus id="6621BFCFFFA410769710A0CD5CF1FB19" box="[113,160,1198,1222]" pageId="4" pageNumber="3">type</typeStatus>
A. Pedipalp femur with dorsal trichobothria arranged in 
<emphasis id="8BEEDD7FFFA4107697F0A0AE5CCFFB3A" bold="true" box="[145,158,1229,1253]" italics="true" pageId="4" pageNumber="3">β</emphasis>
-configuration. Pedipalp patella with trichobothrium 
<emphasis id="8BEEDD7FFFA41076958FA0AD5EAAFB39" box="[750,763,1230,1254]" italics="true" pageId="4" pageNumber="3">d</emphasis>
<subScript id="251E0328FFA41076959AA0BD5F52FB33" attach="left" box="[763,771,1246,1260]" fontSize="6" pageId="4" pageNumber="3">3</subScript>
internal to dorsomedian carina; chela with 
<emphasis id="8BEEDD7FFFA410769536A08D5E23FAD9" box="[599,626,1262,1286]" italics="true" pageId="4" pageNumber="3">db</emphasis>
basal to 
<emphasis id="8BEEDD7FFFA4107695BFA08C5EADFAD9" box="[734,764,1263,1286]" italics="true" pageId="4" pageNumber="3">est</emphasis>
, 
<emphasis id="8BEEDD7FFFA410769710A16D5CDBFAF9" box="[113,138,1294,1318]" italics="true" pageId="4" pageNumber="3">eb</emphasis>
located on fixed finger; chela manus with 
<emphasis id="8BEEDD7FFFA4107695EFA16D5EFDFAF9" box="[654,684,1294,1318]" italics="true" pageId="4" pageNumber="3">Eb</emphasis>
<subScript id="251E0328FFA4107695CDA17D5E95FAF3" attach="left" box="[684,708,1310,1324]" fontSize="6" pageId="4" pageNumber="3">1–3</subScript>
triad angled proximally (
<emphasis id="8BEEDD7FFFA41076963FA14D5D3BFA99" bold="true" box="[350,362,1326,1350]" italics="true" pageId="4" pageNumber="3">δ</emphasis>
-configuration) or almost collinear (
<emphasis id="8BEEDD7FFFA41076971AA12E5CD6FABA" bold="true" box="[123,135,1357,1381]" italics="true" pageId="4" pageNumber="3">λ</emphasis>
-configuration). Dentate margin of pedipalp chela movable finger with distinct granules divided into 8–14 linear rows, with 4 terminal granules and one basal terminal granule. Tibial spurs present on third and fourth pairs of legs.
</paragraph>
</subSubSection>
<subSubSection id="F18052E6FFA410769710A18D5815FD5C" pageId="4" pageNumber="3" type="discussion">
<paragraph id="B925016DFFA410769710A18D5815FD5C" blockId="4.[113,771,1518,2022]" lastBlockId="4.[816,1475,235,643]" pageId="4" pageNumber="3">
REMARKS ON KARYOTYPES. We analyzed karyotypes of four 
<taxonomicName id="7E9A7AEEFFA410769710A26D5D33F9F9" box="[113,354,1550,1574]" class="Arachnida" family="Buthidae" genus="Compsobuthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="8BEEDD7FFFA410769710A26D5D5CF9F9" box="[113,269,1550,1574]" italics="true" pageId="4" pageNumber="3">Compsobuthus</emphasis>
species
</taxonomicName>
(
<tableCitation id="F41834D6FFA410769613A26D5D99F9F9" box="[370,456,1550,1574]" captionStart="Table 4" captionStartId="39.[117,173,697,718]" captionTargetId="graphics-629@39.[118,1476,236,680]" captionTargetPageId="39" captionText="Table 4. The diploid number, the postpachytene configuration, the percentage of the total chromosome length of the diploid set (% TCL) and the origin of Compsobuthus species cytogenetically investigated. Specimens karyotyped: C. acutecarinatus (Simon, 1882): Oman, Dhalkut beach, 16.6989°N 53.2214°E, 1♂; E of Aydam, 17.8887°N 53.06640°E, 1♂; W of Qairoon Hairitti,17.2564°N 54.0220°E, 2♂; WadiAsh Shuwaymiyyah, 17.9293°N 55.5510°E, 1♂1♀. C. arabicus Levy, Amitai &amp; Shulov, 1973: Oman, Al Wasil, 22.4914°N 58.7151°E, 2♂; Jabal Bani Jabir, 22.8132°N 59.0589°E, 1♂; E of Aydam, 17.8887°N 53.06640°E, 1♂; W of Wahiba, 22.0299°N 58.1849°E, 1♂. C. maindroni (Kraepelin, 1900): Oman, Khatm Ha‘ir Asfar, 24.7750°N 55.9146°E, 2♂; Say Plateau, 23.1185°N 57.6311°E, 2♂. C. ullrichi sp. n., 1♂ holotype." httpUri="http://table.plazi.org/id/EDE551E5FF8710559714A6DA5F1EFCA3" pageId="4" pageNumber="3" tableUuid="EDE551E5FF8710559714A6DA5F1EFCA3">Table 4</tableCitation>
). All examined specimens of 
<taxonomicName id="7E9A7AEEFFA410769710A24D5D7FF999" authorityName="Simon" authorityYear="1882" box="[113,302,1582,1606]" class="Arachnida" family="Buthidae" genus="Buthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="acutecarinatus">
<emphasis id="8BEEDD7FFFA410769710A24D5D7FF999" box="[113,302,1582,1606]" italics="true" pageId="4" pageNumber="3">C. acutecarinatus</emphasis>
</taxonomicName>
, 
<taxonomicName id="7E9A7AEEFFA41076965EA24D5DEDF999" authorityName="Levy, Amitai &amp; Shulov" authorityYear="1973" box="[319,444,1582,1606]" class="Arachnida" family="Buthidae" genus="Compsobuthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="arabicus">
<emphasis id="8BEEDD7FFFA41076965EA24D5DEDF999" box="[319,444,1582,1606]" italics="true" pageId="4" pageNumber="3">C. arabicus</emphasis>
</taxonomicName>
, 
<taxonomicName id="7E9A7AEEFFA4107696ACA24D5E0AF999" baseAuthorityName="Kraepelin" baseAuthorityYear="1901" box="[461,603,1582,1606]" class="Arachnida" family="Buthidae" genus="Compsobuthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="maindroni">
<emphasis id="8BEEDD7FFFA4107696ACA24D5E0AF999" box="[461,603,1582,1606]" italics="true" pageId="4" pageNumber="3">C. maindroni</emphasis>
</taxonomicName>
and 
<taxonomicName id="7E9A7AEEFFA4107695F5A24D5F52F999" authority="Kovařík &amp; Lowe &amp; Stockmann &amp; Šťáhlavský, 2020" authorityName="Kovařík &amp; Lowe &amp; Stockmann &amp; Šťáhlavský" authorityYear="2020" box="[660,771,1582,1606]" class="Arachnida" family="Buthidae" genus="Compsobuthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="ullrichi" status="sp. nov.">
<emphasis id="8BEEDD7FFFA4107695F5A24D5F52F999" box="[660,771,1582,1606]" italics="true" pageId="4" pageNumber="3">C. ullrichi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="90DD6004FFA410769710A22D5CFFF9BA" box="[113,174,1614,1638]" pageId="4" pageNumber="3" rank="species">
<emphasis id="8BEEDD7FFFA410769710A22D5CDBF9B9" bold="true" box="[113,138,1614,1638]" pageId="4" pageNumber="3">sp</emphasis>
. 
<emphasis id="8BEEDD7FFFA4107697F9A22D5CF6F9B9" bold="true" box="[152,167,1614,1638]" pageId="4" pageNumber="3">n</emphasis>
.
</taxonomicNameLabel>
possess 2n=22 (
<figureCitation id="21A11DE8FFA41076963CA22D5EC6F9B9" box="[349,663,1614,1638]" captionStart="Figures 210–224" captionStartId="38.[116,193,1874,1896]" captionTargetBox="[113,1474,234,1856]" captionTargetId="figure-15@38.[113,1474,234,1862]" captionText="Figures 210–224: Mitotic metaphases (210, 213, 216, 219, 222), male postpachytenes (211, 214, 217, 220, 223), and ideograms (212, 215, 218, 221, 224) (y axis: % of the diploid chromosome length, dark grey marks: chromosomes in multivalent association) of Compsobuthus species from Arabia and North Africa. Figure 210. Female of C. acutecarinatus (2n=22). Figures 211–212. Male of C. acutecarinatus (2n=22, 11II). Figures 213–215. Male of C. acutecarinatus (2n=22, 9II+CIV). Figures 216–218. Male of C. arabicus (2n=22, 9II+CIV). Figures 219–221. Male of C. maindroni (2n=22, 11II). Figures 222–224. Male of C. ullrichi sp. n. (2n=22, 9II+CIV).Arrows show chromosomes in multivalent association during postpachytene (214, 217, 223). Scale bar: 10 μm" figureDoi="http://doi.org/10.5281/zenodo.6544482" httpUri="https://zenodo.org/record/6544482/files/figure.png" pageId="4" pageNumber="3">Figs. 210, 213, 216, 219, 222</figureCitation>
) and their chromosomes gradually decrease in length (
<figureCitation id="21A11DE8FFA41076953BA20D5D52F979" captionStart="Figures 210–224" captionStartId="38.[116,193,1874,1896]" captionTargetBox="[113,1474,234,1856]" captionTargetId="figure-15@38.[113,1474,234,1862]" captionText="Figures 210–224: Mitotic metaphases (210, 213, 216, 219, 222), male postpachytenes (211, 214, 217, 220, 223), and ideograms (212, 215, 218, 221, 224) (y axis: % of the diploid chromosome length, dark grey marks: chromosomes in multivalent association) of Compsobuthus species from Arabia and North Africa. Figure 210. Female of C. acutecarinatus (2n=22). Figures 211–212. Male of C. acutecarinatus (2n=22, 11II). Figures 213–215. Male of C. acutecarinatus (2n=22, 9II+CIV). Figures 216–218. Male of C. arabicus (2n=22, 9II+CIV). Figures 219–221. Male of C. maindroni (2n=22, 11II). Figures 222–224. Male of C. ullrichi sp. n. (2n=22, 9II+CIV).Arrows show chromosomes in multivalent association during postpachytene (214, 217, 223). Scale bar: 10 μm" figureDoi="http://doi.org/10.5281/zenodo.6544482" httpUri="https://zenodo.org/record/6544482/files/figure.png" pageId="4" pageNumber="3">Figs. 212, 215, 218, 221, 224</figureCitation>
, 
<tableCitation id="F41834D6FFA41076966DA2ED5D0AF979" box="[268,347,1678,1702]" captionStart="Table 4" captionStartId="39.[117,173,697,718]" captionTargetId="graphics-629@39.[118,1476,236,680]" captionTargetPageId="39" captionText="Table 4. The diploid number, the postpachytene configuration, the percentage of the total chromosome length of the diploid set (% TCL) and the origin of Compsobuthus species cytogenetically investigated. Specimens karyotyped: C. acutecarinatus (Simon, 1882): Oman, Dhalkut beach, 16.6989°N 53.2214°E, 1♂; E of Aydam, 17.8887°N 53.06640°E, 1♂; W of Qairoon Hairitti,17.2564°N 54.0220°E, 2♂; WadiAsh Shuwaymiyyah, 17.9293°N 55.5510°E, 1♂1♀. C. arabicus Levy, Amitai &amp; Shulov, 1973: Oman, Al Wasil, 22.4914°N 58.7151°E, 2♂; Jabal Bani Jabir, 22.8132°N 59.0589°E, 1♂; E of Aydam, 17.8887°N 53.06640°E, 1♂; W of Wahiba, 22.0299°N 58.1849°E, 1♂. C. maindroni (Kraepelin, 1900): Oman, Khatm Ha‘ir Asfar, 24.7750°N 55.9146°E, 2♂; Say Plateau, 23.1185°N 57.6311°E, 2♂. C. ullrichi sp. n., 1♂ holotype." httpUri="http://table.plazi.org/id/EDE551E5FF8710559714A6DA5F1EFCA3" pageId="4" pageNumber="3" tableUuid="EDE551E5FF8710559714A6DA5F1EFCA3">Table 4</tableCitation>
). This number of chromosomes was also previously found in 
<taxonomicName id="7E9A7AEEFFA410769625A2CD5D7DF939" authority="KovarIk, Lowe" authorityName="KovarIk, Lowe" class="Arachnida" family="Buthidae" genus="Compsobuthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="eritreaensis">
<emphasis id="8BEEDD7FFFA410769625A2CD5D8CF919" box="[324,477,1710,1734]" italics="true" pageId="4" pageNumber="3">C. eritreaensis</emphasis>
<bibRefCitation id="DD0B7C9CFFA410769682A2CD5D7DF939" author="KovarIk, Lowe, PlIskovA &amp; StAhlavsky" firstAuthor="KovarIk" journalOrPublisher="Euscorpius" pageId="4" pageNumber="3" pagination="1 - 21" part="226" refId="ref15356" refString="KOVARIK, F., G. LOWE, J. PLISKOVA &amp; F. STAHLAVSKY. 2016 a. Scorpions of the Horn of Africa (Arachnida, Scorpiones). Part VI. Compsobuthus Vachon, 1949 (Buthidae) with description of C. eritreaensis sp. n. Euscorpius, 226: 1 - 21." title="Scorpions of the Horn of Africa (Arachnida, Scorpiones). Part VI. Compsobuthus Vachon, 1949 (Buthidae) with description of C. eritreaensis sp. n." type="journal article" year="2016">KovařÍk, Lowe, PlÍškovÁ &amp; ŠťÁhlavský, 2016</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="DD0B7C9CFFA41076965AA2AD5E70F939" author="KovarIk" box="[315,545,1742,1768]" etAl="et al." firstAuthor="KovarIk" pageId="4" pageNumber="3" pagination="1 - 21" refId="ref15356" refString="KOVARIK, F., G. LOWE, J. PLISKOVA &amp; F. STAHLAVSKY. 2016 a. Scorpions of the Horn of Africa (Arachnida, Scorpiones). Part VI. Compsobuthus Vachon, 1949 (Buthidae) with description of C. eritreaensis sp. n. Euscorpius, 226: 1 - 21." type="journal article" year="2016" yearSuffix="a">KovařÍk et al., 2016a</bibRefCitation>
) and 
<taxonomicName id="7E9A7AEEFFA410769501A2AD5E49F8D9" authority="(Birula, 1905) (StAhlavsky et al., 2014)" authorityName="StAhlavsky et al." authorityYear="2014" baseAuthorityName="Birula" baseAuthorityYear="1905" class="Arachnida" family="Buthidae" genus="Compsobuthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="matthiesseni">
<emphasis id="8BEEDD7FFFA410769501A2AD5F52F939" box="[608,771,1742,1766]" italics="true" pageId="4" pageNumber="3">C. matthiesseni</emphasis>
(Birula, 1905) (
<bibRefCitation id="DD0B7C9CFFA410769677A28D5E41F8D9" author="StAhlavsky" box="[278,528,1774,1800]" etAl="et al." firstAuthor="StAhlavsky" pageId="4" pageNumber="3" pagination="355 - 358" refId="ref16198" refString="STAHLAVSKY, F., H. KOC &amp; E. A. YAGMUR. 2014. The first record of karyotypes in Leiurus abdullahbayrami and Compsobuthus matthiesseni (Scorpiones: Buthidae) from Turkey. Northwestern Journal of Zoology, 10 (2): 355 - 358." type="journal article" year="2014">ŠťÁhlavský et al., 2014</bibRefCitation>
)
</taxonomicName>
(
<tableCitation id="F41834D6FFA410769546A28D5E2AF8D9" box="[551,635,1774,1798]" captionStart="Table 4" captionStartId="39.[117,173,697,718]" captionTargetId="graphics-629@39.[118,1476,236,680]" captionTargetPageId="39" captionText="Table 4. The diploid number, the postpachytene configuration, the percentage of the total chromosome length of the diploid set (% TCL) and the origin of Compsobuthus species cytogenetically investigated. Specimens karyotyped: C. acutecarinatus (Simon, 1882): Oman, Dhalkut beach, 16.6989°N 53.2214°E, 1♂; E of Aydam, 17.8887°N 53.06640°E, 1♂; W of Qairoon Hairitti,17.2564°N 54.0220°E, 2♂; WadiAsh Shuwaymiyyah, 17.9293°N 55.5510°E, 1♂1♀. C. arabicus Levy, Amitai &amp; Shulov, 1973: Oman, Al Wasil, 22.4914°N 58.7151°E, 2♂; Jabal Bani Jabir, 22.8132°N 59.0589°E, 1♂; E of Aydam, 17.8887°N 53.06640°E, 1♂; W of Wahiba, 22.0299°N 58.1849°E, 1♂. C. maindroni (Kraepelin, 1900): Oman, Khatm Ha‘ir Asfar, 24.7750°N 55.9146°E, 2♂; Say Plateau, 23.1185°N 57.6311°E, 2♂. C. ullrichi sp. n., 1♂ holotype." httpUri="http://table.plazi.org/id/EDE551E5FF8710559714A6DA5F1EFCA3" pageId="4" pageNumber="3" tableUuid="EDE551E5FF8710559714A6DA5F1EFCA3">Table 4</tableCitation>
). This genus seems to have a conserved diploid number, similar to the genus 
<taxonomicName id="7E9A7AEEFFA4107697D7A34D5D6BF899" authorityName="Ehrenberg" authorityYear="1828" box="[182,314,1838,1862]" class="Arachnida" family="Buthidae" genus="Androctonus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="8BEEDD7FFFA4107697D7A34D5D6BF899" box="[182,314,1838,1862]" italics="true" pageId="4" pageNumber="3">Androctonus</emphasis>
</taxonomicName>
, which typically has 2n= 
<quantity id="7E62AC88FFA410769527A34D5E2DF899" box="[582,636,1838,1862]" metricMagnitude="-1" metricUnit="m" metricValue="6.096" pageId="4" pageNumber="3" unit="in" value="24.0">24 in</quantity>
all analyzed species (
<bibRefCitation id="DD0B7C9CFFA4107697A8A32D5DDFF8B9" author="SadIlek" box="[201,398,1870,1896]" etAl="et al." firstAuthor="SadIlek" pageId="4" pageNumber="3" pagination="69 - 76" refId="ref16022" refString="SADILEK, D., P. NGUYEN, H. KOC, F. KOVARIK, E. A. YAGMUR &amp; F. STAHLAVSKY. 2015. Molecular cytogenetics of Androctonus scorpions: an oasis of calm in the turbulent karyotype evolution of the diverse family Buthidae. Biological Journal of the Linnean Society, 115: 69 - 76." type="journal article" year="2015">SadÍlek et al. 2015</bibRefCitation>
). The chromosomes of all examined 
<taxonomicName id="7E9A7AEEFFA410769710A30D5D0FF859" box="[113,350,1902,1926]" class="Arachnida" family="Buthidae" genus="Compsobuthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="8BEEDD7FFFA410769710A30D5D5CF859" box="[113,269,1902,1926]" italics="true" pageId="4" pageNumber="3">Compsobuthus</emphasis>
species
</taxonomicName>
also exhibit typical buthid features such as holocentric organization and achiasmatic meiosis in males (e. g. 
<bibRefCitation id="DD0B7C9CFFA4107697EAA3CD5D03F819" author="Mattos" box="[139,338,1966,1990]" etAl="et al." firstAuthor="Mattos" pageId="4" pageNumber="3" pagination="121 - 136" refId="ref15967" refString="MATTOS, V. F., D. M. CELLA, L. S. CARVALHO, D. M. CANDIDO &amp; M. C. SCHNEIDER. 2013. High chromosome variability and the presence of multivalent associations in buthid scorpions. Chromosome Research, 21: 121 - 136." type="journal article" year="2013">Mattos et al., 2013</bibRefCitation>
). We observed only bivalents in 
<specimenCount id="AF9CCAE4FFA4107695C5A3CD5F52F819" box="[676,771,1966,1990]" pageId="4" pageNumber="3" type="male">one male</specimenCount>
of 
<taxonomicName id="7E9A7AEEFFA4107697F0A3AD5D1FF839" authorityName="Simon" authorityYear="1882" box="[145,334,1998,2022]" class="Arachnida" family="Buthidae" genus="Buthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="acutecarinatus">
<emphasis id="8BEEDD7FFFA4107697F0A3AD5D1FF839" box="[145,334,1998,2022]" italics="true" pageId="4" pageNumber="3">C. acutecarinatus</emphasis>
</taxonomicName>
from the locality W of Qairoon Hairitti (
<figureCitation id="21A11DE8FFA410769459A4885FC2FEDC" box="[824,915,235,259]" captionStart="Figures 210–224" captionStartId="38.[116,193,1874,1896]" captionTargetBox="[113,1474,234,1856]" captionTargetId="figure-15@38.[113,1474,234,1862]" captionText="Figures 210–224: Mitotic metaphases (210, 213, 216, 219, 222), male postpachytenes (211, 214, 217, 220, 223), and ideograms (212, 215, 218, 221, 224) (y axis: % of the diploid chromosome length, dark grey marks: chromosomes in multivalent association) of Compsobuthus species from Arabia and North Africa. Figure 210. Female of C. acutecarinatus (2n=22). Figures 211–212. Male of C. acutecarinatus (2n=22, 11II). Figures 213–215. Male of C. acutecarinatus (2n=22, 9II+CIV). Figures 216–218. Male of C. arabicus (2n=22, 9II+CIV). Figures 219–221. Male of C. maindroni (2n=22, 11II). Figures 222–224. Male of C. ullrichi sp. n. (2n=22, 9II+CIV).Arrows show chromosomes in multivalent association during postpachytene (214, 217, 223). Scale bar: 10 μm" figureDoi="http://doi.org/10.5281/zenodo.6544482" httpUri="https://zenodo.org/record/6544482/files/figure.png" pageId="4" pageNumber="3">Fig. 211</figureCitation>
) and in all males of 
<taxonomicName id="7E9A7AEEFFA410769314A4885953FEDC" baseAuthorityName="Kraepelin" baseAuthorityYear="1901" box="[1141,1282,235,259]" class="Arachnida" family="Buthidae" genus="Compsobuthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="maindroni">
<emphasis id="8BEEDD7FFFA410769314A4885953FEDC" box="[1141,1282,235,259]" italics="true" pageId="4" pageNumber="3">C. maindroni</emphasis>
</taxonomicName>
(
<figureCitation id="21A11DE8FFA410769273A488593FFEDC" box="[1298,1390,235,259]" captionStart="Figures 210–224" captionStartId="38.[116,193,1874,1896]" captionTargetBox="[113,1474,234,1856]" captionTargetId="figure-15@38.[113,1474,234,1862]" captionText="Figures 210–224: Mitotic metaphases (210, 213, 216, 219, 222), male postpachytenes (211, 214, 217, 220, 223), and ideograms (212, 215, 218, 221, 224) (y axis: % of the diploid chromosome length, dark grey marks: chromosomes in multivalent association) of Compsobuthus species from Arabia and North Africa. Figure 210. Female of C. acutecarinatus (2n=22). Figures 211–212. Male of C. acutecarinatus (2n=22, 11II). Figures 213–215. Male of C. acutecarinatus (2n=22, 9II+CIV). Figures 216–218. Male of C. arabicus (2n=22, 9II+CIV). Figures 219–221. Male of C. maindroni (2n=22, 11II). Figures 222–224. Male of C. ullrichi sp. n. (2n=22, 9II+CIV).Arrows show chromosomes in multivalent association during postpachytene (214, 217, 223). Scale bar: 10 μm" figureDoi="http://doi.org/10.5281/zenodo.6544482" httpUri="https://zenodo.org/record/6544482/files/figure.png" pageId="4" pageNumber="3">Fig. 220</figureCitation>
) during pachytene and postpachytene. However, we found also one quadrivalent in another 
<specimenCount id="AF9CCAE4FFA410769353A54858F1FE9C" box="[1074,1184,299,323]" pageId="4" pageNumber="3" type="male">two males</specimenCount>
of 
<taxonomicName id="7E9A7AEEFFA4107693A8A54859D4FE9D" authorityName="Simon" authorityYear="1882" box="[1225,1413,299,322]" class="Arachnida" family="Buthidae" genus="Buthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="acutecarinatus">
<emphasis id="8BEEDD7FFFA4107693A8A54859D4FE9D" box="[1225,1413,299,322]" italics="true" pageId="4" pageNumber="3">C. acutecarinatus</emphasis>
</taxonomicName>
from the localities Dhalkut beach and Wadi Ash Shuwaymiyyah (
<figureCitation id="21A11DE8FFA410769459A5085FC6FE5C" box="[824,919,363,387]" captionStart="Figures 210–224" captionStartId="38.[116,193,1874,1896]" captionTargetBox="[113,1474,234,1856]" captionTargetId="figure-15@38.[113,1474,234,1862]" captionText="Figures 210–224: Mitotic metaphases (210, 213, 216, 219, 222), male postpachytenes (211, 214, 217, 220, 223), and ideograms (212, 215, 218, 221, 224) (y axis: % of the diploid chromosome length, dark grey marks: chromosomes in multivalent association) of Compsobuthus species from Arabia and North Africa. Figure 210. Female of C. acutecarinatus (2n=22). Figures 211–212. Male of C. acutecarinatus (2n=22, 11II). Figures 213–215. Male of C. acutecarinatus (2n=22, 9II+CIV). Figures 216–218. Male of C. arabicus (2n=22, 9II+CIV). Figures 219–221. Male of C. maindroni (2n=22, 11II). Figures 222–224. Male of C. ullrichi sp. n. (2n=22, 9II+CIV).Arrows show chromosomes in multivalent association during postpachytene (214, 217, 223). Scale bar: 10 μm" figureDoi="http://doi.org/10.5281/zenodo.6544482" httpUri="https://zenodo.org/record/6544482/files/figure.png" pageId="4" pageNumber="3">Fig. 214</figureCitation>
), in 
<specimenCount id="AF9CCAE4FFA4107694ABA5085869FE5C" box="[970,1080,363,387]" pageId="4" pageNumber="3" type="male">two males</specimenCount>
of 
<taxonomicName id="7E9A7AEEFFA410769300A508588FFE5C" authorityName="Levy, Amitai &amp; Shulov" authorityYear="1973" box="[1121,1246,363,387]" class="Arachnida" family="Buthidae" genus="Compsobuthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="arabicus">
<emphasis id="8BEEDD7FFFA410769300A508588FFE5C" box="[1121,1246,363,387]" italics="true" pageId="4" pageNumber="3">C. arabicus</emphasis>
</taxonomicName>
from localities Jabal Bani Jabir and E of Wahiba (
<figureCitation id="21A11DE8FFA41076933AA5E858E5FE7C" box="[1115,1204,395,419]" captionStart="Figures 210–224" captionStartId="38.[116,193,1874,1896]" captionTargetBox="[113,1474,234,1856]" captionTargetId="figure-15@38.[113,1474,234,1862]" captionText="Figures 210–224: Mitotic metaphases (210, 213, 216, 219, 222), male postpachytenes (211, 214, 217, 220, 223), and ideograms (212, 215, 218, 221, 224) (y axis: % of the diploid chromosome length, dark grey marks: chromosomes in multivalent association) of Compsobuthus species from Arabia and North Africa. Figure 210. Female of C. acutecarinatus (2n=22). Figures 211–212. Male of C. acutecarinatus (2n=22, 11II). Figures 213–215. Male of C. acutecarinatus (2n=22, 9II+CIV). Figures 216–218. Male of C. arabicus (2n=22, 9II+CIV). Figures 219–221. Male of C. maindroni (2n=22, 11II). Figures 222–224. Male of C. ullrichi sp. n. (2n=22, 9II+CIV).Arrows show chromosomes in multivalent association during postpachytene (214, 217, 223). Scale bar: 10 μm" figureDoi="http://doi.org/10.5281/zenodo.6544482" httpUri="https://zenodo.org/record/6544482/files/figure.png" pageId="4" pageNumber="3">Fig. 217</figureCitation>
) and in the male hototype of 
<taxonomicName id="7E9A7AEEFFA410769430A5C85F91FE1C" authority="Kovařík &amp; Lowe &amp; Stockmann &amp; Šťáhlavský, 2020" authorityName="Kovařík &amp; Lowe &amp; Stockmann &amp; Šťáhlavský" authorityYear="2020" box="[849,960,427,451]" class="Arachnida" family="Buthidae" genus="Compsobuthus" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="ullrichi" status="sp. nov.">
<emphasis id="8BEEDD7FFFA410769430A5C85F91FE1C" box="[849,960,427,451]" italics="true" pageId="4" pageNumber="3">C. ullrichi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="90DD6004FFA4107694ABA5C85858FE1D" box="[970,1033,427,451]" pageId="4" pageNumber="3" rank="species">
<emphasis id="8BEEDD7FFFA4107694ABA5C85FB2FE1C" bold="true" box="[970,995,427,451]" pageId="4" pageNumber="3">sp</emphasis>
. 
<emphasis id="8BEEDD7FFFA410769495A5C85852FE1C" bold="true" box="[1012,1027,427,451]" pageId="4" pageNumber="3">n</emphasis>
.
</taxonomicNameLabel>
(
<figureCitation id="21A11DE8FFA41076937AA5C85828FE1C" box="[1051,1145,427,451]" captionStart="Figures 210–224" captionStartId="38.[116,193,1874,1896]" captionTargetBox="[113,1474,234,1856]" captionTargetId="figure-15@38.[113,1474,234,1862]" captionText="Figures 210–224: Mitotic metaphases (210, 213, 216, 219, 222), male postpachytenes (211, 214, 217, 220, 223), and ideograms (212, 215, 218, 221, 224) (y axis: % of the diploid chromosome length, dark grey marks: chromosomes in multivalent association) of Compsobuthus species from Arabia and North Africa. Figure 210. Female of C. acutecarinatus (2n=22). Figures 211–212. Male of C. acutecarinatus (2n=22, 11II). Figures 213–215. Male of C. acutecarinatus (2n=22, 9II+CIV). Figures 216–218. Male of C. arabicus (2n=22, 9II+CIV). Figures 219–221. Male of C. maindroni (2n=22, 11II). Figures 222–224. Male of C. ullrichi sp. n. (2n=22, 9II+CIV).Arrows show chromosomes in multivalent association during postpachytene (214, 217, 223). Scale bar: 10 μm" figureDoi="http://doi.org/10.5281/zenodo.6544482" httpUri="https://zenodo.org/record/6544482/files/figure.png" pageId="4" pageNumber="3">Fig. 223</figureCitation>
) during meiosis. Interestingly, the lengths of the chromosomes that form these multivalent associations are different among species (
<figureCitation id="21A11DE8FFA410769387A58859E6FDDC" box="[1254,1463,491,515]" captionStart="Figures 210–224" captionStartId="38.[116,193,1874,1896]" captionTargetBox="[113,1474,234,1856]" captionTargetId="figure-15@38.[113,1474,234,1862]" captionText="Figures 210–224: Mitotic metaphases (210, 213, 216, 219, 222), male postpachytenes (211, 214, 217, 220, 223), and ideograms (212, 215, 218, 221, 224) (y axis: % of the diploid chromosome length, dark grey marks: chromosomes in multivalent association) of Compsobuthus species from Arabia and North Africa. Figure 210. Female of C. acutecarinatus (2n=22). Figures 211–212. Male of C. acutecarinatus (2n=22, 11II). Figures 213–215. Male of C. acutecarinatus (2n=22, 9II+CIV). Figures 216–218. Male of C. arabicus (2n=22, 9II+CIV). Figures 219–221. Male of C. maindroni (2n=22, 11II). Figures 222–224. Male of C. ullrichi sp. n. (2n=22, 9II+CIV).Arrows show chromosomes in multivalent association during postpachytene (214, 217, 223). Scale bar: 10 μm" figureDoi="http://doi.org/10.5281/zenodo.6544482" httpUri="https://zenodo.org/record/6544482/files/figure.png" pageId="4" pageNumber="3">Figs. 215, 221, 224</figureCitation>
). This fact attests to the independent origin of these quadrivalents via reciprocal translocations that are very frequent chromosomal rearrangements within buthid scorpions (e. g. 
<bibRefCitation id="DD0B7C9CFFA41076926CA6285F39FD5C" author="StAhlavsky" etAl="et al." firstAuthor="StAhlavsky" pageId="4" pageNumber="3" pagination="355 - 358" refId="ref16198" refString="STAHLAVSKY, F., H. KOC &amp; E. A. YAGMUR. 2014. The first record of karyotypes in Leiurus abdullahbayrami and Compsobuthus matthiesseni (Scorpiones: Buthidae) from Turkey. Northwestern Journal of Zoology, 10 (2): 355 - 358." type="journal article" year="2014">ŠťÁhlavský et al., 2014</bibRefCitation>
; 
<bibRefCitation id="DD0B7C9CFFA410769413A6085868FD5C" author="Mattos" box="[882,1081,619,643]" etAl="et al." firstAuthor="Mattos" pageId="4" pageNumber="3" pagination="0192070" refId="ref15911" refString="MATTOS, V. F., L. S. CARVALHO, M. A. CARVALHO &amp; M. C. SCHNEIDER. 2018. Insights into the origin of the high variability of multivalent meiotic associations in holocentric chromosomes of Tityus (Archaeotityus) scorpions. PloS ONE 13, e 0192070." type="journal article" year="2018">Mattos et al., 2018</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>