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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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(complete reference list until 1998); 
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<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>
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, figs. 777–941; 
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<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; 
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<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; 
<bibRefCitation id="DD0B7C9CFFA4107697A0A06D5DFFFBF9" author="KovarIk" box="[193,430,1038,1064]" 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>
), 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>