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<mods:title id="443294A1665ECDD38869E3FE5A9EC466">Scorpions of the Horn of Africa (Arachnida Scorpiones) Part XXIII Buthus (Buthidae), with description of two new species</mods:title>
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captionTargetPageId-8="13" captionTargetPageId-9="14" captionText-0="Figures 14: Buthus awashensis. Figures 23. Male paratopotype in dorsal (1) and ventral (2) views. Figures 34. Female paratopotype in dorsal (3) and ventral (4) views. Scale bars: 10 mm." captionText-1="Figures 56. Buthus berberensis, male holotype in dorsal (5) and ventral (6) views." captionText-10="Figures 7489: Buthus pococki sp. n., pedipalp segments. Figures 7483. Male holotype. Chela dorsal (74), external (75) and ventral (76) views. Patella dorsal (77), external (78) and ventral (79) views. Trochanter and femur dorsal (80) and ventral (81) views. Movable (82) and fixed (83) finger dentition. Trichobothrial pattern is indicated by white circles (7478, 80). Figures 8489. Female paratopotype. Chela dorsal (84), external (85) and ventral (86) views. Patella dorsal (87), external (88) and ventral (89) views." captionText-11="Figures 9099: Buthus pococki sp. n. Figures 90, 92, 9497. Male holotype, carapace and tergites IIII (90), coxosternal area and sternites (92), left legs IIV, retrolateral aspect (9497 respectively). Figures 91, 93, 9899. Female paratopotype, carapace and tergites IIII (91), coxosternal area and sternite III (93), and right chelicera in dorsal (98) and ventral (99) views." captionText-12="Figures 100101: Localities. Figure 100. Buthus pococki sp. n., type locality (19SC). Figure 101. B. zeylensis, locality 19SH." captionText-13="Figures 102105: Buthus somalilandus sp. n. Figures 102103. Male in dorsal (102) and ventral (103) views. Figures 104105. Female in dorsal (104) and ventral (105) views. Scale bar: 10 mm." captionText-14="Figures 106113: Buthus somalilandus sp. n. Figures 106, 108110. Male, telson lateral (106), and metasoma and telson lateral (108), dorsal (109), and ventral (111) views. Figures 107, 111113. Female, telson lateral (107), and metasoma and telson lateral (111), dorsal (112), and ventral (113) views. Scale bar: 10 mm (108113)." captionText-15="Figures 114131: Buthus somalilandus sp. n. Figures 114123. Male. Chela dorsal (114), external (115) and ventral (116) views. Patella dorsal (117), external (118) and ventral (119) views. Trochanter and femur dorsal (120) and femur ventral (121) views. Movable (122) and fixed (123) finger dentition. Trichobothrial pattern is indicated by white circles (114118, 120). Figures 124129. Female. Chela dorsal (124), external (125) and ventral (126) views. Patella dorsal (127), external (128) and ventral (129) views. Trochanter and femur dorsal (130) and ventral (131) views." captionText-16="Figures 132139: Buthus somalilandus sp. n. Figures 132, 136139. Male holotype, carapace and tergites IIV (132), coxosternal area and sternites (134), left legs IIV, retrolateral aspect (136139 respectively). Figures 133134. Female paratopotype, carapace and tergites IIII (133), coxosternal area and sternites IIIVI (134)." captionText-17="Figures 140143: Buthus zeylensis from locality 19SH. Figures 140141. Male in dorsal (140) and ventral (141) views. Figures 142143. Female in dorsal (142) and ventral (143) views. Scale bar: 10 mm." captionText-18="Figures 144151: Buthus zeylensis from locality 19SH. Figures 144, 146148. Male, telson lateral (144), and metasoma and telson lateral (146), dorsal (147), and ventral (148) views. Figures 145, 149151. Female, telson lateral (145), and metasoma and telson lateral (149), dorsal (150), and ventral (151) views. Scale bar: 10 mm (146151)." captionText-19="Figures 152167: Buthus zeylensis from locality 19SH, pedipalp segments. Figures 152161. Male. Chela dorsal (152), external (153) and ventral (154) views. Patella dorsal (155), external (156) and ventral (157) views. Trochanter and femur dorsal (158) and ventral (159) views. Movable (160) and fixed (161) finger dentition. Trichobothrial pattern is indicated by white circles (152156, 158). Figures 162167. Female. Chela dorsal (162), external (163) and ventral (164) views. Patella dorsal (165), external (166) and ventral (167) views." captionText-2="Figures 715. Buthus berberensis, male holotype, carapace and tergites IIV (7), coxosternal area and sternites (8), left legs IIIV, retrolateral aspect (911 respectively), metasoma and telson lateral (12), and metasoma and telson lateral (13), dorsal (14), and ventral (15) views. Scale bar: 10 mm (1315)." captionText-20="Figures 168175: Buthus zeylensis from locality 19SH. Figures 168, 170, 172175. Male, carapace and tergites IIV (168), coxosternal area and sternites (170), left legs IIV, retrolateral aspect (172175 respectively). Figures 169, 171. Female, carapace and tergites IIII (169), coxosternal area and sternites IIIVI (171)." captionText-21="Figures 176190: Comparison of pdipalp chela (176, 178, 180, 182, 184), pedipalp patella (177, 179, 181, 183, 185), and telson (186190) of males of Buthus. Figures 176177, 186. B. awashensis, holotype. Figures 178179, 187. B. berberensis from locality 19SO. Figures 180181, 188. B. pococki sp. n., holotype. Figures 182183, 189. B. somalilandus sp. n., holotype. Figures 184185, 190. B. zeylensis from locality 19SH." captionText-22="Figures 191194: Buthus, in vivo habitus. Figure 191. B. awashensis, female paratype. Figure 192. B. pococki sp. n., male paratype. Figure 193. B. somalilandus sp. n., female, paratype. Figure 194. B. zeylensis from locality 19SH." captionText-23="Figures 195209: Male mitotic metaphases (195, 198, 201, 204, 207), postpachytenes (196, 199, 202, 205, 208) and ideograms (197, 200, 203, 206, 206) (y axis - % of the diploid chromosome length, dark grey marks chromosomes in multivalent association) of Buthus species from the Horn of Africa. Figures 195197. B. awashensis, topotype from locality 12EX (2n=22, 6II+CX), Figures 198200. B. berberensis from locality 19SO (2n=21, 9II+CIII), Figures 201203. B. pococki sp. n., paratopotype (2n=22, 11II), Figures 204206. B. pococki sp. n., paratype from locality 19SN (2n=22, 11II), Figures 207209. B. zeylensis from locality 19SH (2n=22, 11II). Arrows show chromosomes in multivalent association during postpachytene. Asterisks show extra-large chromosome from multivalents. Arrowheads indicate large chromosome pair. Scale bar: 10 μm (195196, 198199, 201202, 204205, 207208)." captionText-24="Figures 210211. Distribution of genus Buthus in Ethiopia and Somaliland (210) and type locality of B. somalilandus sp. n. (211)." captionText-3="Figures 1624: Buthus berberensis. Figures 1623. Male holotype, pedipalp segments. Chela dorsal (16), external (17) and ventral (18) views. Patella dorsal (19), external (20) and ventral (21) views. Trochanter and femur dorsal (22) and ventral (23) views. Trichobothrial pattern is indicated by white circles (1620, 22). Figure 24. Male from locality 19SO, in vivo habitus." captionText-4="Figures 2528: Buthus berberensis from locality 19SO. Figures 2526. Male in dorsal (25) and ventral (26) views. Figures 2728. Female in dorsal (27) and ventral (28) views. Scale bar: 10 mm." captionText-5="Figures 2936: Buthus berberensis from locality 19SO. Figures 29, 3133. Male, telson lateral (29), and metasoma and telson lateral (31), dorsal (32), and ventral (33) views. Figures 30, 3436. Female, telson lateral (30), and metasoma and telson lateral (34), dorsal (35), and ventral (36) views. Scale bars: 10 mm (3133, 3436)." captionText-6="Figures 3753: Buthus berberensis from locality 19SO, pedipalp segments. Figures 3746. Male. Chela dorsal (37), external (38) and ventral (39) views. Patella dorsal (40), external (41) and ventral (42) views.Trochanter and femur dorsal (43) and ventral (44) views. Movable (45) and fixed (46) finger dentition.Trichobothrial pattern is indicated by white circles (3741, 43). Figures 4753. Female. Chela dorsal (47), external (48) and ventral (49) views. Patella dorsal (50), external (51) and ventral (52) views. Trochanter and femur dorsal (53)." captionText-7="Figures 5461: Buthus berberensis from locality 19SO. Figures 54, 56, 5861. Male, carapace and tergites IIII (54), coxosternal area and sternites (56), left legs IIV, retrolateral aspect (5861 respectively). Figures 55, 57. Female, carapace and tergites IIV (55), coxosternal area and sternites IIIV (57)." captionText-8="Figures 6265: Buthus pococki sp. n. Figures 6263. Male holotype in dorsal (62) and ventral (63) views. Figures 6465. Female paratopotype in dorsal (64) and ventral (65) views. Scale bar: 10 mm." captionText-9="Figures 6673: Buthus pococki sp. n. Figures 66, 6870. Male holotype, telson lateral (66), and metasoma and telson lateral (68), dorsal (69), and ventral (70) views. Figures 67, 7173. Female paratopotype, telson lateral (67), and metasoma and telson lateral (71), dorsal (72), and ventral (73) views. Scale bars: 10 mm (6870, 7173)." figureDoi-0="http://doi.org/10.5281/zenodo.4648383" figureDoi-1="http://doi.org/10.5281/zenodo.4648385" figureDoi-10="http://doi.org/10.5281/zenodo.4648406" figureDoi-11="http://doi.org/10.5281/zenodo.4648408" figureDoi-12="http://doi.org/10.5281/zenodo.4648410" figureDoi-13="http://doi.org/10.5281/zenodo.4648412" figureDoi-14="http://doi.org/10.5281/zenodo.4648414" figureDoi-15="http://doi.org/10.5281/zenodo.4648416" figureDoi-16="http://doi.org/10.5281/zenodo.4648418" figureDoi-17="http://doi.org/10.5281/zenodo.4648420" figureDoi-18="http://doi.org/10.5281/zenodo.4648422" figureDoi-19="http://doi.org/10.5281/zenodo.4648424" figureDoi-2="http://doi.org/10.5281/zenodo.4648387" figureDoi-20="http://doi.org/10.5281/zenodo.4648426" figureDoi-21="http://doi.org/10.5281/zenodo.4770191" figureDoi-22="http://doi.org/10.5281/zenodo.4648428" figureDoi-23="http://doi.org/10.5281/zenodo.4770195" figureDoi-24="http://doi.org/10.5281/zenodo.4648430" figureDoi-3="http://doi.org/10.5281/zenodo.4648389" figureDoi-4="http://doi.org/10.5281/zenodo.4648391" figureDoi-5="http://doi.org/10.5281/zenodo.4648393" figureDoi-6="http://doi.org/10.5281/zenodo.4648397" figureDoi-7="http://doi.org/10.5281/zenodo.4648399" figureDoi-8="http://doi.org/10.5281/zenodo.4648401" figureDoi-9="http://doi.org/10.5281/zenodo.4648403" httpUri-0="https://zenodo.org/record/4648383/files/figure.png" httpUri-1="https://zenodo.org/record/4648385/files/figure.png" httpUri-10="https://zenodo.org/record/4648406/files/figure.png" httpUri-11="https://zenodo.org/record/4648408/files/figure.png" httpUri-12="https://zenodo.org/record/4648410/files/figure.png" httpUri-13="https://zenodo.org/record/4648412/files/figure.png" httpUri-14="https://zenodo.org/record/4648414/files/figure.png" httpUri-15="https://zenodo.org/record/4648416/files/figure.png" httpUri-16="https://zenodo.org/record/4648418/files/figure.png" httpUri-17="https://zenodo.org/record/4648420/files/figure.png" httpUri-18="https://zenodo.org/record/4648422/files/figure.png" httpUri-19="https://zenodo.org/record/4648424/files/figure.png" httpUri-2="https://zenodo.org/record/4648387/files/figure.png" httpUri-20="https://zenodo.org/record/4648426/files/figure.png" httpUri-21="https://zenodo.org/record/4770191/files/figure.png" httpUri-22="https://zenodo.org/record/4648428/files/figure.png" httpUri-23="https://zenodo.org/record/4770195/files/figure.png" httpUri-24="https://zenodo.org/record/4648430/files/figure.png" httpUri-3="https://zenodo.org/record/4648389/files/figure.png" httpUri-4="https://zenodo.org/record/4648391/files/figure.png" httpUri-5="https://zenodo.org/record/4648393/files/figure.png" httpUri-6="https://zenodo.org/record/4648397/files/figure.png" httpUri-7="https://zenodo.org/record/4648399/files/figure.png" httpUri-8="https://zenodo.org/record/4648401/files/figure.png" httpUri-9="https://zenodo.org/record/4648403/files/figure.png" pageId="2" pageNumber="1">Figures 1211</figureCitation>
,
<tableCitation id="431A0377DC5F40196C83F966FA8956D4" box="[1170,1286,1691,1716]" captionStart-0="Table 1" captionStart-1="Table 2" captionStart-2="Table 3" captionStartId-0="12.[113,169,998,1019]" captionStartId-1="23.[115,171,942,963]" captionStartId-2="32.[115,171,447,468]" captionTargetId-0="graphics-407@12.[114,1468,242,970]" captionTargetId-1="graphics-574@23.[119,1476,234,918]" captionTargetId-2="graphics-722@32.[113,1474,244,422]" captionTargetPageId-0="12" captionTargetPageId-1="23" captionTargetPageId-2="32" captionText-0="Table 1. Comparative measurements of adults of Buthus berberensis and B. pococki sp. n. Abbreviations: length (L), width (W, in carapace it corresponds to posterior width), depth (D)." captionText-1="Table 2. Comparative measurements of adults of Buthus somalilandus sp. n. and B. zeylensis. Abbreviations: length (L), width (W, in carapace it corresponds to posterior width), depth (D)." captionText-2="Table 3. The diploid numbers, the postpachytene configuration (Ppt configuration), the percentage of the total chromosome length of the diploid set (% TCL) and the localities of Buthus species from the Horn of Africa." httpUri-0="http://table.plazi.org/id/5AE76644DC5140176860FC1BFD825479" httpUri-1="http://table.plazi.org/id/5AE76644DC4A400C6862FC53FDAD5380" httpUri-2="http://table.plazi.org/id/5AE76644DC7D403B6862FE42FCF25192" pageId="2" pageNumber="1" tableUuid-0="5AE76644DC5140176860FC1BFD825479" tableUuid-1="5AE76644DC4A400C6862FC53FDAD5380" tableUuid-2="5AE76644DC7D403B6862FE42FCF25192">Tables 13</tableCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="46826547DC5F40196B21F926FC125753" pageId="2" pageNumber="1" type="reference_group">
<paragraph id="0E2736CCDC5F40196B21F926FC125753" blockId="2.[816,1475,1755,1843]" pageId="2" pageNumber="1">
<treatmentCitationGroup id="2E8811E2DC5F40196B21F926FC125753" pageId="2" pageNumber="1">
<treatmentCitation id="8F3910DDDC5F40196B21F926FBB55694" author="LEACH, W. E." box="[816,1082,1755,1780]" page="391" pageId="2" pageNumber="1" year="1815">
<taxonomicName id="C9984D4FDC5F40196B21F926FBB55694" ID-CoL="62HV6" authority="Leach, 1815: 391" authorityName="Leach" authorityPageNumber="391" authorityYear="1815" box="[816,1082,1755,1780]" class="Arachnida" family="Buthidae" genus="Buthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="2" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="3CECEADEDC5F40196B21F926FCF55693" box="[816,890,1755,1779]" italics="true" pageId="2" pageNumber="1">Buthus</emphasis>
<bibRefCitation id="6A094B3DDC5F40196B91F926FBB55694" author="LEACH, W. E." box="[896,1082,1755,1780]" firstAuthor="Leach" page="391" pageId="2" pageNumber="1" pagination="306 - 400" refId="ref10154" refString="LEACH, W. E. 1815. A tabular view of the external characters of four classes of animals, which Linne arranged under Insecta; with the distribution of the genera composing three of these classes into orders, etc. and descriptions of several new genera and species. Transactions of the Linnean Society of London, 11: 306 - 400." type="journal article" year="1815">Leach, 1815: 391</bibRefCitation>
</taxonomicName>
</treatmentCitation>
;
<bibRefCitation id="6A094B3DDC5F40196C52F921FADD5694" author="FET, V. &amp; G. LOWE" bookContentInfo="689 pp." box="[1091,1362,1756,1780]" editor="Fet, V. &amp; W. D. Sissom &amp; G. Lowe &amp; M. E. Braunwalder" journalOrPublisher="New York: The New York Entomological Society" pageId="2" pageNumber="1" pagination="54 - 286" refId="ref9759" 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: 9197</bibRefCitation>
(complete references list until 1998);
<treatmentCitation id="8F3910DDDC5F40196C6CF901FA055773" author="SOUSA P., M. A. ARNEDO &amp; D. J. HARRIS. " box="[1149,1418,1787,1812]" httpUri="http://treatment.plazi.org/id/DA2A060B5DE6F5D024198E0FD0030BD3" page="15" pageId="2" pageNumber="1" year="2017">
<bibRefCitation id="6A094B3DDC5F40196C6CF901FA055773" author="SOUSA P., M. A. ARNEDO &amp; D. J. HARRIS. " box="[1149,1418,1787,1812]" etAl="et al." firstAuthor="Sousa" page="15 - 84" pageId="2" pageNumber="1" pagination="15 - 84" refId="ref10552" refString="SOUSA P., M. A. ARNEDO &amp; D. J. HARRIS. 2017. Updated catalogue and taxonomic notes on the Old-World scorpion genus Buthus Leach, 1815 (Scorpiones, Buthidae). ZooKeys, 686: 15 - 84." type="journal article" year="2017">Sousa et al., 2017: 1584</bibRefCitation>
</treatmentCitation>
, figs. 116.
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="46826547DC5F40196B21F8A1FABA5714" box="[816,1333,1884,1908]" pageId="2" pageNumber="1" type="type_taxon">
<paragraph id="0E2736CCDC5F40196B21F8A1FABA5714" blockId="2.[816,1333,1884,1908]" box="[816,1333,1884,1908]" pageId="2" pageNumber="1">
<typeStatus id="D123886EDC5F40196B21F8A1FCE95713" box="[816,870,1884,1907]" pageId="2" pageNumber="1">TYPE</typeStatus>
SPECIES.
<taxonomicName id="C9984D4FDC5F40196BD6F8A1FABA5714" authority="Amoreux, 1789" authorityName="Amoreux" authorityYear="1789" box="[967,1333,1884,1908]" class="Arachnida" family="Scorpionidae" genus="Scorpio" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="2" pageNumber="1" phylum="Arthropoda" rank="species" species="occitanus">
<emphasis id="3CECEADEDC5F40196BD6F8A1FB085713" box="[967,1159,1884,1907]" italics="true" pageId="2" pageNumber="1">Scorpio occitanus</emphasis>
Amoreux, 1789
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="46826547DC5F401F6B21F861FEC352CD" lastPageId="4" lastPageNumber="3" pageId="2" pageNumber="1" type="diagnosis">
<paragraph id="0E2736CCDC5F401F6B21F861FEC352CD" blockId="2.[816,1474,1947,2035]" lastBlockId="4.[113,771,245,685]" lastPageId="4" lastPageNumber="3" pageId="2" pageNumber="1">
DIAGNOSIS. Total length
<quantity id="C9609B29DC5F40196C51F866FB3457D3" box="[1088,1211,1947,1971]" metricMagnitude="-2" metricUnit="m" metricValue="5.75" metricValueMax="9.0" metricValueMin="2.5" pageId="2" pageNumber="1" unit="mm" value="57.5" valueMax="90.0" valueMin="25.0">2590 mm</quantity>
. Carapace with distinct carinae joined to form a lyre-shaped configuration, in lateral view with entire dorsal surface horizontal or nearly so. Five pairs of lateral eyes and eyespot present. Pectines with fulcra. Pectine teeth number ca 2040. Mesosoma tergites IVI with three carina which do not project beyond posterior margin as distinct spiniform processes. Telson without subaculear tubercle. Chelicera with typical buthid dentition, fixed finger with two ventral denticles. Orthobothriotaxic
<typeStatus id="D123886EDC59401F69AFFE4BFE6251AE" box="[446,493,438,462]" pageId="4" pageNumber="3">type</typeStatus>
B, dorsal trichobothria of pedipalp femur arranged in
<emphasis id="3CECEADEDC59401F69C1FE28FE70518D" box="[464,511,469,493]" italics="true" pageId="4" pageNumber="3">beta</emphasis>
-configuration. Patellar trichobothrium
<emphasis id="3CECEADEDC59401F690FFE08FEA4526D" box="[286,299,501,525]" italics="true" pageId="4" pageNumber="3">d</emphasis>
2 located externally to dorsomedian carina. Trichobothrium
<emphasis id="3CECEADEDC59401F6936FDE8FECE524D" box="[295,321,533,557]" italics="true" pageId="4" pageNumber="3">eb</emphasis>
located on fixed finger of chela. Dentate margin of pedipalp chela movable finger with distinct granules divided into 914 rows, 3 terminal granules and one basal terminal granule. Tibial spurs present on third and fourth pairs of legs.
</paragraph>
</subSubSection>
<caption id="5AE76644DC5E40186865F84AFD0F5789" ID-DOI="http://doi.org/10.5281/zenodo.4648383" ID-Zenodo-Dep="4648383" httpUri="https://zenodo.org/record/4648383/files/figure.png" pageId="3" pageNumber="2" startId="3.[116,193,1975,1997]" targetBox="[121,1459,257,1942]" targetPageId="3">
<paragraph id="0E2736CCDC5E40186865F84AFD0F5789" blockId="3.[116,1449,1975,2026]" pageId="3" pageNumber="2">
<emphasis id="3CECEADEDC5E40186865F84AFF6357AC" bold="true" box="[116,236,1975,1997]" pageId="3" pageNumber="2">Figures 14</emphasis>
:
<taxonomicName id="C9984D4FDC5E401868EBF84AFE3E57AC" authorityName="KovarIk" authorityYear="2011" box="[250,433,1975,1996]" class="Arachnida" family="Buthidae" genus="Buthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="3" pageNumber="2" phylum="Arthropoda" rank="species" species="awashensis">
<emphasis id="3CECEADEDC5E401868EBF84AFE3E57AC" box="[250,433,1975,1996]" italics="true" pageId="3" pageNumber="2">Buthus awashensis</emphasis>
</taxonomicName>
.
<emphasis id="3CECEADEDC5E401869AFF84AFDB957AC" bold="true" box="[446,566,1975,1997]" pageId="3" pageNumber="2">Figures 23</emphasis>
. Male paratopotype in dorsal (1) and ventral (2) views.
<emphasis id="3CECEADEDC5E40186C48F84AFB5D57AC" bold="true" box="[1113,1234,1975,1997]" pageId="3" pageNumber="2">Figures 34</emphasis>
. Female paratopotype in dorsal (3) and ventral (4) views. Scale bars: 10 mm.
</paragraph>
</caption>
<subSubSection id="46826547DC59401F6860FD28FD9957AD" pageId="4" pageNumber="3" type="discussion">
<paragraph id="0E2736CCDC59401F6860FD28FD9957AD" blockId="4.[113,772,725,1998]" pageId="4" pageNumber="3">
REMARKS ON KARYOTYPES (
<figureCitation id="96A32A49DC59401F69BCFD28FDDB528D" box="[429,596,725,749]" captionStart="Figures 195209" captionStartId="30.[113,190,1852,1874]" captionTargetBox="[181,1400,234,1839]" captionTargetId="figure-20@30.[159,1436,234,1839]" captionText="Figures 195209: Male mitotic metaphases (195, 198, 201, 204, 207), postpachytenes (196, 199, 202, 205, 208) and ideograms (197, 200, 203, 206, 206) (y axis - % of the diploid chromosome length, dark grey marks chromosomes in multivalent association) of Buthus species from the Horn of Africa. Figures 195197. B. awashensis, topotype from locality 12EX (2n=22, 6II+CX), Figures 198200. B. berberensis from locality 19SO (2n=21, 9II+CIII), Figures 201203. B. pococki sp. n., paratopotype (2n=22, 11II), Figures 204206. B. pococki sp. n., paratype from locality 19SN (2n=22, 11II), Figures 207209. B. zeylensis from locality 19SH (2n=22, 11II). Arrows show chromosomes in multivalent association during postpachytene. Asterisks show extra-large chromosome from multivalents. Arrowheads indicate large chromosome pair. Scale bar: 10 μm (195196, 198199, 201202, 204205, 207208)." figureDoi="http://doi.org/10.5281/zenodo.4770195" httpUri="https://zenodo.org/record/4770195/files/figure.png" pageId="4" pageNumber="3">Figs. 195209</figureCitation>
). We analyzed male karyotypes of four
<taxonomicName id="C9984D4FDC59401F698FFD08FE65536D" authorityName="Leach" authorityYear="1815" box="[414,490,757,781]" class="Arachnida" family="Buthidae" genus="Buthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="3CECEADEDC59401F698FFD08FE65536D" box="[414,490,757,781]" italics="true" pageId="4" pageNumber="3">Buthus</emphasis>
</taxonomicName>
species from the Horn of Africa (
<tableCitation id="431A0377DC59401F68FBFCE8FECC534D" box="[234,323,789,813]" captionStart="Table 3" captionStartId="32.[115,171,447,468]" captionTargetId="graphics-722@32.[113,1474,244,422]" captionTargetPageId="32" captionText="Table 3. The diploid numbers, the postpachytene configuration (Ppt configuration), the percentage of the total chromosome length of the diploid set (% TCL) and the localities of Buthus species from the Horn of Africa." httpUri="http://table.plazi.org/id/5AE76644DC7D403B6862FE42FCF25192" pageId="4" pageNumber="3" tableUuid="5AE76644DC7D403B6862FE42FCF25192">Table 3</tableCitation>
). The cytogenetic characteristics of all species correspond to the typical features of the family
<taxonomicName id="C9984D4FDC59401F6860FCA8FF5B530D" authorityName="C. L. Koch" authorityYear="1837" box="[113,212,853,877]" class="Arachnida" family="Buthidae" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="family">Buthidae</taxonomicName>
such as holocentric organization, achiasmatic meiosis in males, and lower number of chromosomes (e. g.
<bibRefCitation id="6A094B3DDC59401F689EFC6BFEE753CD" author="Mattos" box="[143,360,917,941]" etAl="et al." firstAuthor="Mattos" pageId="4" pageNumber="3" pagination="121 - 136" refId="ref10304" 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>
). All examined specimens of
<taxonomicName id="C9984D4FDC59401F6AA6FC68FF6153AD" authorityName="KovarIk" authorityYear="2011" class="Arachnida" family="Buthidae" genus="Buthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="awashensis">
<emphasis id="3CECEADEDC59401F6AA6FC68FF6153AD" italics="true" pageId="4" pageNumber="3">Buthus awashensis</emphasis>
</taxonomicName>
,
<taxonomicName id="C9984D4FDC59401F6914FC4BFE0F53AD" authority="Kovařík &amp; Šťáhlavský &amp; Elmi, 2020" authorityName="Kovařík &amp; Šťáhlavský &amp; Elmi" authorityYear="2020" box="[261,384,949,973]" class="Arachnida" family="Buthidae" genus="Buthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="pococki" status="sp. nov.">
<emphasis id="3CECEADEDC59401F6914FC4BFE0F53AD" box="[261,384,949,973]" italics="true" pageId="4" pageNumber="3">B. pococki</emphasis>
</taxonomicName>
<taxonomicNameLabel id="27DF57A5DC59401F699EFC48FE5A53AD" box="[399,469,949,973]" pageId="4" pageNumber="3" rank="species">
<emphasis id="3CECEADEDC59401F699EFC48FE2653AD" bold="true" box="[399,425,949,973]" pageId="4" pageNumber="3">sp</emphasis>
.
<emphasis id="3CECEADEDC59401F69AEFC48FE4153AD" bold="true" box="[447,462,949,973]" pageId="4" pageNumber="3">n</emphasis>
.
</taxonomicNameLabel>
and
<taxonomicName id="C9984D4FDC59401F6A08FC4BFD2E53AD" authorityName="Pocock" authorityYear="1900" box="[537,673,949,973]" class="Arachnida" family="Buthidae" genus="Buthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="zeylensis">
<emphasis id="3CECEADEDC59401F6A08FC4BFD2E53AD" box="[537,673,949,973]" italics="true" pageId="4" pageNumber="3">B. zeylensis</emphasis>
</taxonomicName>
possess 2n=22 (
<figureCitation id="96A32A49DC59401F68DBFC28FDAF538D" box="[202,544,981,1005]" captionStart="Figures 195209" captionStartId="30.[113,190,1852,1874]" captionTargetBox="[181,1400,234,1839]" captionTargetId="figure-20@30.[159,1436,234,1839]" captionText="Figures 195209: Male mitotic metaphases (195, 198, 201, 204, 207), postpachytenes (196, 199, 202, 205, 208) and ideograms (197, 200, 203, 206, 206) (y axis - % of the diploid chromosome length, dark grey marks chromosomes in multivalent association) of Buthus species from the Horn of Africa. Figures 195197. B. awashensis, topotype from locality 12EX (2n=22, 6II+CX), Figures 198200. B. berberensis from locality 19SO (2n=21, 9II+CIII), Figures 201203. B. pococki sp. n., paratopotype (2n=22, 11II), Figures 204206. B. pococki sp. n., paratype from locality 19SN (2n=22, 11II), Figures 207209. B. zeylensis from locality 19SH (2n=22, 11II). Arrows show chromosomes in multivalent association during postpachytene. Asterisks show extra-large chromosome from multivalents. Arrowheads indicate large chromosome pair. Scale bar: 10 μm (195196, 198199, 201202, 204205, 207208)." figureDoi="http://doi.org/10.5281/zenodo.4770195" httpUri="https://zenodo.org/record/4770195/files/figure.png" pageId="4" pageNumber="3">Figs. 195, 201, 204, 205, 208</figureCitation>
). We found only 21 chromosomes in karyotype of analysed
<taxonomicName id="C9984D4FDC59401F6A3EFC08FC8C546D" authorityName="Pocock" authorityYear="1900" box="[559,771,1013,1037]" class="Arachnida" family="Buthidae" genus="Buthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="berberensis">
<emphasis id="3CECEADEDC59401F6A3EFC08FC8C546D" box="[559,771,1013,1037]" italics="true" pageId="4" pageNumber="3">Buthus berberensis</emphasis>
</taxonomicName>
(
<figureCitation id="96A32A49DC59401F686BFBE8FF54544D" box="[122,219,1045,1069]" captionStart="Figures 195209" captionStartId="30.[113,190,1852,1874]" captionTargetBox="[181,1400,234,1839]" captionTargetId="figure-20@30.[159,1436,234,1839]" captionText="Figures 195209: Male mitotic metaphases (195, 198, 201, 204, 207), postpachytenes (196, 199, 202, 205, 208) and ideograms (197, 200, 203, 206, 206) (y axis - % of the diploid chromosome length, dark grey marks chromosomes in multivalent association) of Buthus species from the Horn of Africa. Figures 195197. B. awashensis, topotype from locality 12EX (2n=22, 6II+CX), Figures 198200. B. berberensis from locality 19SO (2n=21, 9II+CIII), Figures 201203. B. pococki sp. n., paratopotype (2n=22, 11II), Figures 204206. B. pococki sp. n., paratype from locality 19SN (2n=22, 11II), Figures 207209. B. zeylensis from locality 19SH (2n=22, 11II). Arrows show chromosomes in multivalent association during postpachytene. Asterisks show extra-large chromosome from multivalents. Arrowheads indicate large chromosome pair. Scale bar: 10 μm (195196, 198199, 201202, 204205, 207208)." figureDoi="http://doi.org/10.5281/zenodo.4770195" httpUri="https://zenodo.org/record/4770195/files/figure.png" pageId="4" pageNumber="3">Fig. 198</figureCitation>
). The reduction of the diploid number is probably consequence of heterozygous fusion of chromosomes. This
<typeStatus id="D123886EDC59401F6860FBABFF2F540E" box="[113,160,1110,1134]" pageId="4" pageNumber="3">type</typeStatus>
of chromosomal rearrangement forms conspicuous trivalent during postpachytene (
<figureCitation id="96A32A49DC59401F69C0FB88FDBD54ED" box="[465,562,1141,1165]" captionStart="Figures 195209" captionStartId="30.[113,190,1852,1874]" captionTargetBox="[181,1400,234,1839]" captionTargetId="figure-20@30.[159,1436,234,1839]" captionText="Figures 195209: Male mitotic metaphases (195, 198, 201, 204, 207), postpachytenes (196, 199, 202, 205, 208) and ideograms (197, 200, 203, 206, 206) (y axis - % of the diploid chromosome length, dark grey marks chromosomes in multivalent association) of Buthus species from the Horn of Africa. Figures 195197. B. awashensis, topotype from locality 12EX (2n=22, 6II+CX), Figures 198200. B. berberensis from locality 19SO (2n=21, 9II+CIII), Figures 201203. B. pococki sp. n., paratopotype (2n=22, 11II), Figures 204206. B. pococki sp. n., paratype from locality 19SN (2n=22, 11II), Figures 207209. B. zeylensis from locality 19SH (2n=22, 11II). Arrows show chromosomes in multivalent association during postpachytene. Asterisks show extra-large chromosome from multivalents. Arrowheads indicate large chromosome pair. Scale bar: 10 μm (195196, 198199, 201202, 204205, 207208)." figureDoi="http://doi.org/10.5281/zenodo.4770195" httpUri="https://zenodo.org/record/4770195/files/figure.png" pageId="4" pageNumber="3">Fig. 199</figureCitation>
). Moreover, fusion of two chromosomes form one extra large chromosome in the karyotype of this species (
<figureCitation id="96A32A49DC59401F69F6FB48FDC354AD" box="[487,588,1205,1229]" captionStart="Figures 195209" captionStartId="30.[113,190,1852,1874]" captionTargetBox="[181,1400,234,1839]" captionTargetId="figure-20@30.[159,1436,234,1839]" captionText="Figures 195209: Male mitotic metaphases (195, 198, 201, 204, 207), postpachytenes (196, 199, 202, 205, 208) and ideograms (197, 200, 203, 206, 206) (y axis - % of the diploid chromosome length, dark grey marks chromosomes in multivalent association) of Buthus species from the Horn of Africa. Figures 195197. B. awashensis, topotype from locality 12EX (2n=22, 6II+CX), Figures 198200. B. berberensis from locality 19SO (2n=21, 9II+CIII), Figures 201203. B. pococki sp. n., paratopotype (2n=22, 11II), Figures 204206. B. pococki sp. n., paratype from locality 19SN (2n=22, 11II), Figures 207209. B. zeylensis from locality 19SH (2n=22, 11II). Arrows show chromosomes in multivalent association during postpachytene. Asterisks show extra-large chromosome from multivalents. Arrowheads indicate large chromosome pair. Scale bar: 10 μm (195196, 198199, 201202, 204205, 207208)." figureDoi="http://doi.org/10.5281/zenodo.4770195" httpUri="https://zenodo.org/record/4770195/files/figure.png" pageId="4" pageNumber="3">Fig. 200</figureCitation>
). Except
<taxonomicName id="C9984D4FDC59401F6AA6FB48FF61548D" authorityName="KovarIk" authorityYear="2011" class="Arachnida" family="Buthidae" genus="Buthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="awashensis">
<emphasis id="3CECEADEDC59401F6AA6FB48FF61548D" italics="true" pageId="4" pageNumber="3">Buthus awashensis</emphasis>
</taxonomicName>
, the karyotypes of the all analysed species are typical with one pair of chromosomes with distinctive length whereas the subsequent chromosomes are shorter and gradually decrease in length (
<figureCitation id="96A32A49DC59401F6A1FFAC8FF10550D" captionStart="Figures 195209" captionStartId="30.[113,190,1852,1874]" captionTargetBox="[181,1400,234,1839]" captionTargetId="figure-20@30.[159,1436,234,1839]" captionText="Figures 195209: Male mitotic metaphases (195, 198, 201, 204, 207), postpachytenes (196, 199, 202, 205, 208) and ideograms (197, 200, 203, 206, 206) (y axis - % of the diploid chromosome length, dark grey marks chromosomes in multivalent association) of Buthus species from the Horn of Africa. Figures 195197. B. awashensis, topotype from locality 12EX (2n=22, 6II+CX), Figures 198200. B. berberensis from locality 19SO (2n=21, 9II+CIII), Figures 201203. B. pococki sp. n., paratopotype (2n=22, 11II), Figures 204206. B. pococki sp. n., paratype from locality 19SN (2n=22, 11II), Figures 207209. B. zeylensis from locality 19SH (2n=22, 11II). Arrows show chromosomes in multivalent association during postpachytene. Asterisks show extra-large chromosome from multivalents. Arrowheads indicate large chromosome pair. Scale bar: 10 μm (195196, 198199, 201202, 204205, 207208)." figureDoi="http://doi.org/10.5281/zenodo.4770195" httpUri="https://zenodo.org/record/4770195/files/figure.png" pageId="4" pageNumber="3">Figs. 200, 203, 206, 209</figureCitation>
). Similar one longer pair of chromosomes is known also in karyotypes of all karyotyped
<taxonomicName id="C9984D4FDC59401F6A0CFA88FD2655ED" authorityName="Ehrenberg" authorityYear="1828" box="[541,681,1397,1421]" class="Arachnida" family="Buthidae" genus="Androctonus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="3CECEADEDC59401F6A0CFA88FD2655ED" box="[541,681,1397,1421]" italics="true" pageId="4" pageNumber="3">Androctonus</emphasis>
</taxonomicName>
species (
<bibRefCitation id="6A094B3DDC59401F6868FA6BFEEC55CD" author="SadIlek" box="[121,355,1429,1456]" etAl="et al." firstAuthor="SadIlek" pageId="4" pageNumber="3" pagination="69 - 76" refId="ref10433" 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>
). This similarity of karyotypes may reflect phylogenetic relationships that was already proposed in previous phylogenetic analysis (
<bibRefCitation id="6A094B3DDC59401F6A1BFA28FD47558D" author="Fet" box="[522,712,1493,1517]" etAl="et al." firstAuthor="Fet" pageId="4" pageNumber="3" pagination="1 - 10" refId="ref9714" refString="FET, V., B. GANTENBEIN, A. V. GROMOV, G. LOWE &amp; W. R. LOURENCO. 2003. The first molecular phylogeny of Buthidae (Scorpiones). Euscorpius, 4: 1 - 10." type="journal article" year="2003">Fet et al., 2003</bibRefCitation>
). We observed multivalent association (ten chromosomes of different length) in
<taxonomicName id="C9984D4FDC59401F6952F9E8FD9D564D" authorityName="KovarIk" authorityYear="2011" box="[323,530,1557,1581]" class="Arachnida" family="Buthidae" genus="Buthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="awashensis">
<emphasis id="3CECEADEDC59401F6952F9E8FD9D564D" box="[323,530,1557,1581]" italics="true" pageId="4" pageNumber="3">Buthus awashensis</emphasis>
</taxonomicName>
during postpachytene (
<figureCitation id="96A32A49DC59401F686BF9C8FE94562D" box="[122,283,1589,1613]" captionStart="Figures 195209" captionStartId="30.[113,190,1852,1874]" captionTargetBox="[181,1400,234,1839]" captionTargetId="figure-20@30.[159,1436,234,1839]" captionText="Figures 195209: Male mitotic metaphases (195, 198, 201, 204, 207), postpachytenes (196, 199, 202, 205, 208) and ideograms (197, 200, 203, 206, 206) (y axis - % of the diploid chromosome length, dark grey marks chromosomes in multivalent association) of Buthus species from the Horn of Africa. Figures 195197. B. awashensis, topotype from locality 12EX (2n=22, 6II+CX), Figures 198200. B. berberensis from locality 19SO (2n=21, 9II+CIII), Figures 201203. B. pococki sp. n., paratopotype (2n=22, 11II), Figures 204206. B. pococki sp. n., paratype from locality 19SN (2n=22, 11II), Figures 207209. B. zeylensis from locality 19SH (2n=22, 11II). Arrows show chromosomes in multivalent association during postpachytene. Asterisks show extra-large chromosome from multivalents. Arrowheads indicate large chromosome pair. Scale bar: 10 μm (195196, 198199, 201202, 204205, 207208)." figureDoi="http://doi.org/10.5281/zenodo.4770195" httpUri="https://zenodo.org/record/4770195/files/figure.png" pageId="4" pageNumber="3">Figs. 196197</figureCitation>
). This chain of chromosomes is consequence of multiple reciprocal translocations. Although this
<typeStatus id="D123886EDC59401F6AA4F9ABFD6B560E" box="[693,740,1622,1646]" pageId="4" pageNumber="3">type</typeStatus>
of chromosome rearrangement does not affect chromosome numbers, it may however considerably change the size of the chromosomes (e. g.
<bibRefCitation id="6A094B3DDC59401F696BF94BFDD256AD" author="KovarIk" box="[378,605,1717,1744]" etAl="et al." firstAuthor="KovarIk" pageId="4" pageNumber="3" pagination="1 - 31" refId="ref9978" refString="KOVARIK, F., G. LOWE, M. SEITER, J. PLISKOVA &amp; F. STAHLAVSKY. 2015. Scorpions of Ethiopia (Arachnida: Scorpiones). Part II. Genus Babycurus Karsch, 1886 (Buthidae), with description of two new species. Euscorpius, 196: 1 - 31." type="journal article" year="2015">KovařÍk et al., 2015</bibRefCitation>
). It is probably the reason why this species has not one extra large pair of chromosomes, the typical feature of the remaining species with only bivalents (
<figureCitation id="96A32A49DC59401F6970F8E8FDC7574D" box="[353,584,1813,1837]" captionStart="Figures 195209" captionStartId="30.[113,190,1852,1874]" captionTargetBox="[181,1400,234,1839]" captionTargetId="figure-20@30.[159,1436,234,1839]" captionText="Figures 195209: Male mitotic metaphases (195, 198, 201, 204, 207), postpachytenes (196, 199, 202, 205, 208) and ideograms (197, 200, 203, 206, 206) (y axis - % of the diploid chromosome length, dark grey marks chromosomes in multivalent association) of Buthus species from the Horn of Africa. Figures 195197. B. awashensis, topotype from locality 12EX (2n=22, 6II+CX), Figures 198200. B. berberensis from locality 19SO (2n=21, 9II+CIII), Figures 201203. B. pococki sp. n., paratopotype (2n=22, 11II), Figures 204206. B. pococki sp. n., paratype from locality 19SN (2n=22, 11II), Figures 207209. B. zeylensis from locality 19SH (2n=22, 11II). Arrows show chromosomes in multivalent association during postpachytene. Asterisks show extra-large chromosome from multivalents. Arrowheads indicate large chromosome pair. Scale bar: 10 μm (195196, 198199, 201202, 204205, 207208)." figureDoi="http://doi.org/10.5281/zenodo.4770195" httpUri="https://zenodo.org/record/4770195/files/figure.png" pageId="4" pageNumber="3">Figs. 202, 205, 208</figureCitation>
). Despite of the mentioned small differences among analysed species, the karyotypes of species from the Horn of Africa fully correspond to those of
<taxonomicName id="C9984D4FDC59401F6989F888FDEF57ED" baseAuthorityName="Amoreux" baseAuthorityYear="1789" box="[408,608,1909,1933]" class="Arachnida" family="Buthidae" genus="Buthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="species" species="occitanus">
<emphasis id="3CECEADEDC59401F6989F888FDEF57ED" box="[408,608,1909,1933]" italics="true" pageId="4" pageNumber="3">Buthus occitanus</emphasis>
</taxonomicName>
from
<collectingCountry id="768F765CDC59401F6AA9F88BFC8C57ED" box="[696,771,1910,1933]" name="France" pageId="4" pageNumber="3">France</collectingCountry>
(
<bibRefCitation id="6A094B3DDC59401F6868F868FE9057CD" author="Guenin" box="[121,287,1941,1965]" firstAuthor="Guenin" pageId="4" pageNumber="3" pagination="89 - 96" refId="ref9832" refString="GUENIN, H. A. 1961. Contribution a la connaissance cytologique des scorpions: les chromosomes de Buthus occitanus Amor. (I). Vie et Milieu, 12: 89 - 96." type="journal article" year="1961">Guénin, 1961</bibRefCitation>
), and the karyotypes seem to be very conservative within the genus
<taxonomicName id="C9984D4FDC59401F69D2F848FD8057AD" authorityName="Leach" authorityYear="1815" box="[451,527,1973,1997]" class="Arachnida" family="Buthidae" genus="Buthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Scorpiones" pageId="4" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="3CECEADEDC59401F69D2F848FD8057AD" box="[451,527,1973,1997]" italics="true" pageId="4" pageNumber="3">Buthus</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>