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301 lines
40 KiB
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<mods:title id="6489506F62BB40B78E7D52A636740131">Tracking the trends of karyotype differentiation in the phylogenetic context of Gint, a scorpion genus endemic to the Horn of Africa (Scorpiones: Buthidae)</mods:title>
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<mods:namePart id="76BA8063E3D2271D2C4E690B39639620">Just, Pavel</mods:namePart>
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<mods:namePart id="F7B17C07E404F875CF89FCBD648C02BD">Kovařík, František</mods:namePart>
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<mods:namePart id="EBA7AB3C7F1D8EEE5A3E7199317F76DC">Štundlová, Jana</mods:namePart>
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<treatment id="03AACF58FFA71F22FEF41709FECF94C9" ID-DOI="http://doi.org/10.5281/zenodo.7194672" ID-GBIF-Taxon="202609290" ID-Zenodo-Dep="7194672" LSID="urn:lsid:plazi:treatment:03AACF58FFA71F22FEF41709FECF94C9" httpUri="http://treatment.plazi.org/id/03AACF58FFA71F22FEF41709FECF94C9" lastPageId="10" lastPageNumber="893" pageId="8" pageNumber="893">
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<subSubSection id="C3192DC5FFA71F20FEF41709FDAC9437" box="[348,557,1046,1070]" pageId="8" pageNumber="893" type="nomenclature">
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<paragraph id="8BBC7E4EFFA71F20FEF41709FDAC9437" blockId="8.[348,557,1046,1070]" box="[348,557,1046,1070]" pageId="8" pageNumber="893">
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<heading id="D0F4C922FFA71F20FEF41709FDAC9437" box="[348,557,1046,1070]" centered="true" fontSize="9" level="2" pageId="8" pageNumber="893" reason="2">
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<taxonomicName id="4C0305CDFFA71F20FEF41709FDAC9437" ID-CoL="3G3J5" authorityName="Kovarik, Lowe, Just, Awale, Elmi & Stahlavsky" authorityYear="2018" box="[348,557,1046,1070]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="gubanensis">
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<emphasis id="B977A25CFFA71F20FEF41709FDAC9437" box="[348,557,1046,1070]" italics="true" pageId="8" pageNumber="893">
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<emphasis id="B977A25CFFA71F20FEF41709FE169435" box="[348,407,1046,1069]" italics="true" pageId="8" pageNumber="893">GINT</emphasis>
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GUBANENSIS
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</emphasis>
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</taxonomicName>
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</heading>
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</paragraph>
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<subSubSection id="C3192DC5FFA71F20FF391720FE8395DD" pageId="8" pageNumber="893" type="discussion">
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<paragraph id="8BBC7E4EFFA71F20FF391720FE8395DD" blockId="8.[145,762,1086,1476]" pageId="8" pageNumber="893">
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Karyotype of
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<taxonomicName id="4C0305CDFFA71F20FEEE1720FE7A944A" box="[326,507,1086,1108]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="gubanensis">
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<emphasis id="B977A25CFFA71F20FEEE1720FE7A944A" box="[326,507,1086,1108]" italics="true" pageId="8" pageNumber="893">G. gubanensis</emphasis>
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</taxonomicName>
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comprises 2
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<emphasis id="B977A25CFFA71F20FD0B1720FD33944D" box="[675,690,1087,1108]" italics="true" pageId="8" pageNumber="893">n</emphasis>
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= 45 chromosomes, which decrease in length from 2.97% to 1.02% of DSL (Supporting Information,
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<figureCitation id="133862CBFFA71F20FD611763FF4794A9" captionStart="Figure 2" captionStartId="5.[163,241,1512,1534]" captionTargetBox="[171,1436,200,1467]" captionTargetId="figure-117@5.[166,1440,195,1472]" captionTargetPageId="5" captionText="Figure 2. Post-pachytene cells of Gint species after Giemsa staining (A, C, E, G, I, K, M, O) and FISH with 18S rDNA (red signals) (B, D, F, H, J,L, N, P).A, B,G. banfasae cytotype I(2n=18 –7II + IV).C, D, G. banfasae cytotype II (2n= 18 – 6II +VI). E, F, G. banfasae cytotype III (2n = 19 – 5II + III + VI). G, H, G. dabakalo cytotype I (2n = 23 – 9II + V). I, J,G. dabakalo cytotype II (2n = 24 – 8II + III + V). K, L, G. dabakalo cytotype III (2n = 27 – 8II + 2III + V). M, N, G. gaitako (2n = 30 – 13II + IV). O, P, G. maidensis (2n = 34). Abbreviations: II, bivalent; III, trivalent; IV, quadrivalent; V, pentavalent; VI, hexavalent. Arrowheads indicate the position of 18S rDNA. Scale bar = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.7184498" httpUri="https://zenodo.org/record/7184498/files/figure.png" pageId="8" pageNumber="893">Fig. S2G</figureCitation>
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; Table S2). Overall, 21 bivalents and a single trivalent, composed of chromosomes 7, 33 and 45, are detected in post-pachytene nuclei (
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<figureCitation id="133862CBFFA71F20FD8817C7FDEB94F4" box="[544,618,1240,1262]" captionStart="Figure 3" captionStartId="6.[145,223,1192,1214]" captionTargetBox="[153,1417,198,1149]" captionTargetId="figure-348@6.[150,1420,195,1152]" captionTargetPageId="6" captionText="Figure 3. Post-pachytene cells of Gint amoudensis and G. gubanensis after Giemsa staining (A, C, E, G, I) and FISH with 18S rDNA (red signals) (B, D, F, H, J). A, B, G. amoudensis cytotype I (2n = 36 – 16II + IV). C, D, G. amoudensis cytotype II (2n = 35 – 14II + III + IV). E, F, G. amoudensis cytotype III (2n = 36 – 15II + VI). G, H, G. amoudensis cytotype IV (2n = 35 – 13II + III + VI). I, J, G.gubanensis (2n = 45 – 21II + III). Abbreviations: II, bivalent; III, trivalent; IV, quadrivalent; VI, hexavalent. Arrowheads indicate the position of 18S rDNA. Scale bar = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.7184502" httpUri="https://zenodo.org/record/7184502/files/figure.png" pageId="8" pageNumber="893">Fig. 3I</figureCitation>
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; Supporting Information,
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<figureCitation id="133862CBFFA71F20FE9A17E9FE1D9515" box="[306,412,1270,1292]" captionStart="Figure 2" captionStartId="5.[163,241,1512,1534]" captionTargetBox="[171,1436,200,1467]" captionTargetId="figure-117@5.[166,1440,195,1472]" captionTargetPageId="5" captionText="Figure 2. Post-pachytene cells of Gint species after Giemsa staining (A, C, E, G, I, K, M, O) and FISH with 18S rDNA (red signals) (B, D, F, H, J,L, N, P).A, B,G. banfasae cytotype I(2n=18 –7II + IV).C, D, G. banfasae cytotype II (2n= 18 – 6II +VI). E, F, G. banfasae cytotype III (2n = 19 – 5II + III + VI). G, H, G. dabakalo cytotype I (2n = 23 – 9II + V). I, J,G. dabakalo cytotype II (2n = 24 – 8II + III + V). K, L, G. dabakalo cytotype III (2n = 27 – 8II + 2III + V). M, N, G. gaitako (2n = 30 – 13II + IV). O, P, G. maidensis (2n = 34). Abbreviations: II, bivalent; III, trivalent; IV, quadrivalent; V, pentavalent; VI, hexavalent. Arrowheads indicate the position of 18S rDNA. Scale bar = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.7184498" httpUri="https://zenodo.org/record/7184498/files/figure.png" pageId="8" pageNumber="893">Fig. S2G</figureCitation>
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). One pair of 18S rDNA loci is found on a terminal part of the heteromorphic chromosome pair (chromosomes 26 and 44). A single large 18S rDNA cluster is located on the overhanging part of chromosome 26 (the larger chromosome of the heteromorphic pair) (
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<figureCitation id="133862CBFFA71F20FE2E168FFE5295BC" box="[390,467,1424,1446]" captionStart="Figure 3" captionStartId="6.[145,223,1192,1214]" captionTargetBox="[153,1417,198,1149]" captionTargetId="figure-348@6.[150,1420,195,1152]" captionTargetPageId="6" captionText="Figure 3. Post-pachytene cells of Gint amoudensis and G. gubanensis after Giemsa staining (A, C, E, G, I) and FISH with 18S rDNA (red signals) (B, D, F, H, J). A, B, G. amoudensis cytotype I (2n = 36 – 16II + IV). C, D, G. amoudensis cytotype II (2n = 35 – 14II + III + IV). E, F, G. amoudensis cytotype III (2n = 36 – 15II + VI). G, H, G. amoudensis cytotype IV (2n = 35 – 13II + III + VI). I, J, G.gubanensis (2n = 45 – 21II + III). Abbreviations: II, bivalent; III, trivalent; IV, quadrivalent; VI, hexavalent. Arrowheads indicate the position of 18S rDNA. Scale bar = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.7184502" httpUri="https://zenodo.org/record/7184502/files/figure.png" pageId="8" pageNumber="893">Fig. 3J</figureCitation>
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; Supporting Information,
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<figureCitation id="133862CBFFA71F20FF3916B1FF7295DD" box="[145,243,1454,1476]" captionStart="Figure 2" captionStartId="5.[163,241,1512,1534]" captionTargetBox="[171,1436,200,1467]" captionTargetId="figure-117@5.[166,1440,195,1472]" captionTargetPageId="5" captionText="Figure 2. Post-pachytene cells of Gint species after Giemsa staining (A, C, E, G, I, K, M, O) and FISH with 18S rDNA (red signals) (B, D, F, H, J,L, N, P).A, B,G. banfasae cytotype I(2n=18 –7II + IV).C, D, G. banfasae cytotype II (2n= 18 – 6II +VI). E, F, G. banfasae cytotype III (2n = 19 – 5II + III + VI). G, H, G. dabakalo cytotype I (2n = 23 – 9II + V). I, J,G. dabakalo cytotype II (2n = 24 – 8II + III + V). K, L, G. dabakalo cytotype III (2n = 27 – 8II + 2III + V). M, N, G. gaitako (2n = 30 – 13II + IV). O, P, G. maidensis (2n = 34). Abbreviations: II, bivalent; III, trivalent; IV, quadrivalent; V, pentavalent; VI, hexavalent. Arrowheads indicate the position of 18S rDNA. Scale bar = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.7184498" httpUri="https://zenodo.org/record/7184498/files/figure.png" pageId="8" pageNumber="893">Fig. S2G</figureCitation>
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).
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</paragraph>
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</subSubSection>
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<subSubSection id="C3192DC5FFA71F22FF5F151BFECF94C9" lastPageId="10" lastPageNumber="895" pageId="8" pageNumber="893" type="description">
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<paragraph id="8BBC7E4EFFA71F20FF5F151BFD139601" blockId="8.[247,658,1537,1562]" box="[247,658,1537,1562]" pageId="8" pageNumber="893">MTDNA DIVERSITY AND PHYLOGENY</paragraph>
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<paragraph id="8BBC7E4EFFA71F20FF391536FD5B96C1" blockId="8.[145,761,1577,1906]" pageId="8" pageNumber="893">
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The sequences for both 16S and
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<emphasis id="B977A25CFFA71F20FDB11535FDC89626" box="[537,585,1578,1599]" italics="true" pageId="8" pageNumber="893">COI</emphasis>
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were obtained from 22
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<taxonomicName id="4C0305CDFFA71F20FF581557FEA29644" box="[240,291,1608,1629]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="genus">
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<emphasis id="B977A25CFFA71F20FF581557FEA29644" box="[240,291,1608,1629]" italics="true" pageId="8" pageNumber="893">Gint</emphasis>
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</taxonomicName>
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individuals. In
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<taxonomicName id="4C0305CDFFA71F20FE701557FDDC9644" baseAuthorityName="Pocock" baseAuthorityYear="1900" box="[472,605,1608,1629]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="calviceps">
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<emphasis id="B977A25CFFA71F20FE701557FDDC9644" box="[472,605,1608,1629]" italics="true" pageId="8" pageNumber="893">G. calviceps</emphasis>
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</taxonomicName>
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, only the 16S gene fragment was successfully sequenced. The final alignment with a total length of 963 bp (16S: 358 bp;
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<emphasis id="B977A25CFFA71F20FF3915BBFF4396A0" box="[145,194,1700,1721]" italics="true" pageId="8" pageNumber="893">COI</emphasis>
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: 605 bp) consisted of 765 conserved sites, 194 variable sites and 172 parsimony-informative sites.
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</paragraph>
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<paragraph id="8BBC7E4EFFA71F20FF0115FEFB7A944D" blockId="8.[145,761,1577,1906]" lastBlockId="8.[808,1426,197,1660]" pageId="8" pageNumber="893">
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Based on
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<emphasis id="B977A25CFFA71F20FEB315FDFECB96EE" box="[283,330,1762,1783]" italics="true" pageId="8" pageNumber="893">COI</emphasis>
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data,
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<taxonomicName id="4C0305CDFFA71F20FE3D15FDFE4896EE" box="[405,457,1762,1783]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="genus">
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<emphasis id="B977A25CFFA71F20FE3D15FDFE4896EE" box="[405,457,1762,1783]" italics="true" pageId="8" pageNumber="893">Gint</emphasis>
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</taxonomicName>
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exhibits genetic variation at both inter- and intraspecific levels, with the pairwise genetic distances ranging from 3.28% to 11.52% between species, and from 0.00% to 2.64% within species (Supporting Information, Tables S3, S4). The haplotype network reconstruction showed a presence of 15
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<emphasis id="B977A25CFFA71F20FCC413FBFC1D90E0" box="[876,924,228,249]" italics="true" pageId="8" pageNumber="893">COI</emphasis>
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sequence haplotypes in
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<taxonomicName id="4C0305CDFFA71F20FB6D13FBFB7B90E0" box="[1221,1274,228,249]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="genus">
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<emphasis id="B977A25CFFA71F20FB6D13FBFB7B90E0" box="[1221,1274,228,249]" italics="true" pageId="8" pageNumber="893">Gint</emphasis>
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</taxonomicName>
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(Supporting Information, (
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<figureCitation id="133862CBFFA71F20FC78121DFBA59101" box="[976,1060,258,280]" captionStart="Figure 3" captionStartId="6.[145,223,1192,1214]" captionTargetBox="[153,1417,198,1149]" captionTargetId="figure-348@6.[150,1420,195,1152]" captionTargetPageId="6" captionText="Figure 3. Post-pachytene cells of Gint amoudensis and G. gubanensis after Giemsa staining (A, C, E, G, I) and FISH with 18S rDNA (red signals) (B, D, F, H, J). A, B, G. amoudensis cytotype I (2n = 36 – 16II + IV). C, D, G. amoudensis cytotype II (2n = 35 – 14II + III + IV). E, F, G. amoudensis cytotype III (2n = 36 – 15II + VI). G, H, G. amoudensis cytotype IV (2n = 35 – 13II + III + VI). I, J, G.gubanensis (2n = 45 – 21II + III). Abbreviations: II, bivalent; III, trivalent; IV, quadrivalent; VI, hexavalent. Arrowheads indicate the position of 18S rDNA. Scale bar = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.7184502" httpUri="https://zenodo.org/record/7184502/files/figure.png" pageId="8" pageNumber="893">Fig. S3</figureCitation>
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; Table S3). Within the species studied,
|
||
<taxonomicName id="4C0305CDFFA71F20FC21123DFB8C912F" authorityName="Kovarik & Lowe" authorityYear="2019" box="[905,1037,289,311]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="banfasae">
|
||
<emphasis id="B977A25CFFA71F20FC21123DFB8C912F" box="[905,1037,289,311]" italics="true" pageId="8" pageNumber="893">G. banfasae</emphasis>
|
||
</taxonomicName>
|
||
shows the highest within-location haplotype diversity (four haplotypes per locality), but with minor pairwise genetic differences (
|
||
<emphasis id="B977A25CFFA71F20FAB91240FA9E916D" box="[1297,1311,351,372]" italics="true" pageId="8" pageNumber="893">p</emphasis>
|
||
-distance: ≤0.66%). In this species, all individuals of the same cytotype have distinct haplotypes. The haplotypes of cytotype I individuals (S1531 and S1534) are separated from one another by four mutational steps. Cytotype II (S1532) shares its haplotype with the cytotype III individual (S1530), which is one mutational step distant from the haplotype of the cytotype III specimen (S1533) (Supporting Information,
|
||
<figureCitation id="133862CBFFA71F20FA98114BFA039270" box="[1328,1410,596,618]" captionStart="Figure 3" captionStartId="6.[145,223,1192,1214]" captionTargetBox="[153,1417,198,1149]" captionTargetId="figure-348@6.[150,1420,195,1152]" captionTargetPageId="6" captionText="Figure 3. Post-pachytene cells of Gint amoudensis and G. gubanensis after Giemsa staining (A, C, E, G, I) and FISH with 18S rDNA (red signals) (B, D, F, H, J). A, B, G. amoudensis cytotype I (2n = 36 – 16II + IV). C, D, G. amoudensis cytotype II (2n = 35 – 14II + III + IV). E, F, G. amoudensis cytotype III (2n = 36 – 15II + VI). G, H, G. amoudensis cytotype IV (2n = 35 – 13II + III + VI). I, J, G.gubanensis (2n = 45 – 21II + III). Abbreviations: II, bivalent; III, trivalent; IV, quadrivalent; VI, hexavalent. Arrowheads indicate the position of 18S rDNA. Scale bar = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.7184502" httpUri="https://zenodo.org/record/7184502/files/figure.png" pageId="8" pageNumber="893">Fig. S3</figureCitation>
|
||
).
|
||
<taxonomicName id="4C0305CDFFA71F20FC81116CFC4B929E" authorityName="Kovarik & Mazuch" authorityYear="2015" box="[809,970,626,648]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="dabakalo">
|
||
<emphasis id="B977A25CFFA71F20FC81116CFC4B929E" box="[809,970,626,648]" italics="true" pageId="8" pageNumber="893">Gint dabakalo</emphasis>
|
||
</taxonomicName>
|
||
, comprising three haplotypes, shows the highest intraspecific genetic variation with a
|
||
<emphasis id="B977A25CFFA71F20FAB5118DFAAA92BE" box="[1309,1323,658,679]" italics="true" pageId="8" pageNumber="893">p</emphasis>
|
||
-distance of 0.17–2.64%. In this species, the genetic distance between individuals of cytotype I and cytotype II from the same locality is higher than between each of them and a specimen of cytotype III from a distinct locality (Supporting Information,
|
||
<figureCitation id="133862CBFFA71F20FBC31035FB459359" box="[1131,1220,810,832]" captionStart="Figure 3" captionStartId="6.[145,223,1192,1214]" captionTargetBox="[153,1417,198,1149]" captionTargetId="figure-348@6.[150,1420,195,1152]" captionTargetPageId="6" captionText="Figure 3. Post-pachytene cells of Gint amoudensis and G. gubanensis after Giemsa staining (A, C, E, G, I) and FISH with 18S rDNA (red signals) (B, D, F, H, J). A, B, G. amoudensis cytotype I (2n = 36 – 16II + IV). C, D, G. amoudensis cytotype II (2n = 35 – 14II + III + IV). E, F, G. amoudensis cytotype III (2n = 36 – 15II + VI). G, H, G. amoudensis cytotype IV (2n = 35 – 13II + III + VI). I, J, G.gubanensis (2n = 45 – 21II + III). Abbreviations: II, bivalent; III, trivalent; IV, quadrivalent; VI, hexavalent. Arrowheads indicate the position of 18S rDNA. Scale bar = 10 µm." figureDoi="http://doi.org/10.5281/zenodo.7184502" httpUri="https://zenodo.org/record/7184502/files/figure.png" pageId="8" pageNumber="893">Fig. S3</figureCitation>
|
||
; Table S1).
|
||
<taxonomicName id="4C0305CDFFA71F20FAF21034FC1F9347" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="maidensis">
|
||
<emphasis id="B977A25CFFA71F20FAF21034FC1F9347" italics="true" pageId="8" pageNumber="893">Gint maidensis</emphasis>
|
||
</taxonomicName>
|
||
, a species with a stable karyotype 2
|
||
<emphasis id="B977A25CFFA71F20FAE91055FAD19346" box="[1345,1360,842,863]" italics="true" pageId="8" pageNumber="893">n</emphasis>
|
||
= 34, is represented by three haplotypes with a
|
||
<emphasis id="B977A25CFFA71F20FAAA1076FA919367" box="[1282,1296,873,894]" italics="true" pageId="8" pageNumber="893">p</emphasis>
|
||
-distance of 0.33–0.50%.
|
||
<emphasis id="B977A25CFFA71F20FC191099FBB89382" box="[945,1081,902,923]" italics="true" pageId="8" pageNumber="893">Gintgaitako</emphasis>
|
||
presentstwohaplotypes, where one haplogroup, formed by
|
||
<specimenCount id="9D05B5C7FFA71F20FBC110BAFA9493A2" box="[1129,1301,933,955]" pageId="8" pageNumber="893" type="generic">five specimens</specimenCount>
|
||
, is distant from the other by one mutational step (
|
||
<emphasis id="B977A25CFFA71F20FB7E10DBFB6593C0" box="[1238,1252,964,985]" italics="true" pageId="8" pageNumber="893">p</emphasis>
|
||
-distance: 0.17). In
|
||
<taxonomicName id="4C0305CDFFA71F20FCEF10FCFC6693EE" box="[839,999,994,1016]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="amoudensis">
|
||
<emphasis id="B977A25CFFA71F20FCEF10FCFC6693EE" box="[839,999,994,1016]" italics="true" pageId="8" pageNumber="893">G. amoudensis</emphasis>
|
||
</taxonomicName>
|
||
, individuals of cytotype II, cytotype III and cytotype IV share the same haplotype, whereas the specimen of cytotype I differs from this haplogroup by eight point mutations (
|
||
<emphasis id="B977A25CFFA71F20FB821720FBB9944D" box="[1066,1080,1087,1108]" italics="true" pageId="8" pageNumber="893">p</emphasis>
|
||
-distance: 1.49%).
|
||
</paragraph>
|
||
<paragraph id="8BBC7E4EFFA71F20FCE91742FAF99665" blockId="8.[808,1426,197,1660]" pageId="8" pageNumber="893">
|
||
Both BI and ML mtDNA analyses provide similar tree topologies and nodal supports for clades. Phylogenetic reconstruction depicts
|
||
<taxonomicName id="4C0305CDFFA71F20FABB1784FACD94A9" box="[1299,1356,1179,1200]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977A25CFFA71F20FABB1784FACD94A9" box="[1299,1356,1179,1200]" italics="true" pageId="8" pageNumber="893">Gint</emphasis>
|
||
</taxonomicName>
|
||
as a monophyletic group (
|
||
<figureCitation id="133862CBFFA71F20FB8817A6FBE094D6" box="[1056,1121,1209,1231]" captionStart="Figure 5" captionStartId="9.[163,244,1583,1605]" captionTargetBox="[170,1434,197,1542]" captionTargetId="figure-99@9.[169,1436,195,1543]" captionTargetPageId="9" captionText="Figure 5. Bayesian tree of the Gint species based on the concatenated data set (16S and COI), complemented with summarized cytogenetic data of the species studied. Numbers above branches correspond to values for highly supported nodes as follows: Bayesian posterior probabilities (PP)> 0.95/maximum likelihood bootstrap> 70%. Specimen IDs depicted in bold indicate individuals based on chromosome counts determined for the corresponding Gint species in previous studies (see: Kovařík et al., 2013; Kovařík & Mazuch, 2015; Kovařík et al., 2018). Abbreviations: cyt, cytotype; 2n, diploid number of chromosomes; II, bivalent; III, trivalent; IV, quadrivalent; V, pentavalent; VI, hexavalent." figureDoi="http://doi.org/10.5281/zenodo.7184506" httpUri="https://zenodo.org/record/7184506/files/figure.png" pageId="8" pageNumber="893">Fig. 5</figureCitation>
|
||
). Most of the
|
||
<taxonomicName id="4C0305CDFFA71F20FAAB17A6FA1094D6" box="[1283,1425,1209,1231]" pageId="8" pageNumber="893">
|
||
<emphasis id="B977A25CFFA71F20FAAB17A6FAB694D7" box="[1283,1335,1209,1230]" italics="true" pageId="8" pageNumber="893">Gint</emphasis>
|
||
species
|
||
</taxonomicName>
|
||
form well-supported clades [bootstrap support (bs) ≥ 83, posterior probability (PP) ≥ 0.98]. The monophyly of the
|
||
<taxonomicName id="4C0305CDFFA71F20FBA31609FB1A9533" authorityName="Kovarik & Lowe" authorityYear="2019" box="[1035,1179,1301,1323]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="banfasae">
|
||
<emphasis id="B977A25CFFA71F20FBA31609FB1A9533" box="[1035,1179,1301,1323]" italics="true" pageId="8" pageNumber="893">G. banfasae</emphasis>
|
||
</taxonomicName>
|
||
specimens remains unresolved due to low support values.
|
||
<taxonomicName id="4C0305CDFFA71F20FB4A162BFA119550" box="[1250,1424,1332,1353]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="maidensis">
|
||
<emphasis id="B977A25CFFA71F20FB4A162BFA119550" box="[1250,1424,1332,1353]" italics="true" pageId="8" pageNumber="893">Gint maidensis</emphasis>
|
||
</taxonomicName>
|
||
forms an early-branching clade, which is sister to the clade comprising all of the remaining
|
||
<taxonomicName id="4C0305CDFFA71F20FB50166EFA10959E" box="[1272,1425,1393,1415]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="undetermined">
|
||
<emphasis id="B977A25CFFA71F20FB50166EFAAC959F" box="[1272,1325,1393,1414]" italics="true" pageId="8" pageNumber="893">Gint</emphasis>
|
||
species.
|
||
</taxonomicName>
|
||
This clade is further subdivided into two groups: (1) a clade in which
|
||
<taxonomicName id="4C0305CDFFA71F20FC5D16B0FB1B95DA" box="[1013,1178,1454,1476]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="gubanensis">
|
||
<emphasis id="B977A25CFFA71F20FC5D16B0FB1B95DA" box="[1013,1178,1454,1476]" italics="true" pageId="8" pageNumber="893">G. gubanensis</emphasis>
|
||
</taxonomicName>
|
||
is a sister-species to
|
||
<emphasis id="B977A25CFFA71F20FC8216D1FC5295FB" box="[810,979,1485,1507]" italics="true" pageId="8" pageNumber="893">
|
||
<taxonomicName id="4C0305CDFFA71F20FC8216D1FC4E95FB" box="[810,975,1485,1507]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="amoudensis">G. amoudensis</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and (2) a clade consisting of
|
||
<taxonomicName id="4C0305CDFFA71F20FAB016D1FA0A95FB" box="[1304,1419,1485,1507]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="gaitako">
|
||
<emphasis id="B977A25CFFA71F20FAB016D1FA0A95FB" box="[1304,1419,1485,1507]" italics="true" pageId="8" pageNumber="893">G. gaitako</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C0305CDFFA71F20FC8116F3FC3B9618" baseAuthorityName="Pocock" baseAuthorityYear="1900" box="[809,954,1516,1537]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="calviceps">
|
||
<emphasis id="B977A25CFFA71F20FC8116F3FC3B9618" box="[809,954,1516,1537]" italics="true" pageId="8" pageNumber="893">G. calviceps</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C0305CDFFA71F20FC6416F3FBE19618" authorityName="Kovarik & Mazuch" authorityYear="2015" box="[972,1120,1516,1537]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="dabakalo">
|
||
<emphasis id="B977A25CFFA71F20FC6416F3FBE19618" box="[972,1120,1516,1537]" italics="true" pageId="8" pageNumber="893">G. dabakalo</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C0305CDFFA71F20FB0B16F3FAB29618" authorityName="Kovarik & Lowe" authorityYear="2019" box="[1187,1331,1516,1537]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="banfasae">
|
||
<emphasis id="B977A25CFFA71F20FB0B16F3FAB29618" box="[1187,1331,1516,1537]" italics="true" pageId="8" pageNumber="893">G. banfasae</emphasis>
|
||
</taxonomicName>
|
||
. In the latter,
|
||
<taxonomicName id="4C0305CDFFA71F20FCD01514FC739606" box="[888,1010,1546,1568]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="gaitako">
|
||
<emphasis id="B977A25CFFA71F20FCD01514FC739606" box="[888,1010,1546,1568]" italics="true" pageId="8" pageNumber="893">G. gaitako</emphasis>
|
||
</taxonomicName>
|
||
forms a sister-lineage to the clade comprising
|
||
<taxonomicName id="4C0305CDFFA71F20FC071535FBB59627" baseAuthorityName="Pocock" baseAuthorityYear="1900" box="[943,1076,1577,1599]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="calviceps">
|
||
<emphasis id="B977A25CFFA71F20FC071535FBB59627" box="[943,1076,1577,1599]" italics="true" pageId="8" pageNumber="893">G. calviceps</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C0305CDFFA71F20FBEA1535FB4A9627" authorityName="Kovarik & Mazuch" authorityYear="2015" box="[1090,1227,1577,1599]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="dabakalo">
|
||
<emphasis id="B977A25CFFA71F20FBEA1535FB4A9627" box="[1090,1227,1577,1599]" italics="true" pageId="8" pageNumber="893">G. dabakalo</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C0305CDFFA71F20FAAE1535FA0A9627" authorityName="Kovarik & Lowe" authorityYear="2019" box="[1286,1419,1577,1599]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="species" species="banfasae">
|
||
<emphasis id="B977A25CFFA71F20FAAE1535FA0A9627" box="[1286,1419,1577,1599]" italics="true" pageId="8" pageNumber="893">G. banfasae</emphasis>
|
||
</taxonomicName>
|
||
. The mutual relationships between the species in this group remain unresolved due to low support values.
|
||
</paragraph>
|
||
<paragraph id="8BBC7E4EFFA71F20FC9A15A6FB149717" blockId="8.[818,1415,1721,1806]" pageId="8" pageNumber="893">THE RECONSTRUCTION OF ANCESTRAL CHROMOSOME NUMBER AND POSITION OF 18S RDNA GENE CLUSTERS</paragraph>
|
||
<paragraph id="8BBC7E4EFFA71F22FC811401FECF94C9" blockId="8.[809,1426,1822,1906]" lastBlockId="10.[145,762,1088,1232]" lastPageId="10" lastPageNumber="895" pageId="8" pageNumber="893">
|
||
Maximum parsimony analysis inferred 2
|
||
<emphasis id="B977A25CFFA71F20FAB01400FAA6972D" box="[1304,1319,1823,1844]" italics="true" pageId="8" pageNumber="893">n</emphasis>
|
||
= 32–34 as the ancestral chromosome number for the genus
|
||
<taxonomicName id="4C0305CDFFA71F20FC811443FCE19768" box="[809,864,1884,1905]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="8" pageNumber="893" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977A25CFFA71F20FC811443FCE19768" box="[809,864,1884,1905]" italics="true" pageId="8" pageNumber="893">Gint</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="133862CBFFA71F20FCDD1443FC51976B" box="[885,976,1884,1906]" captionStart="Figure 6" captionStartId="10.[147,227,939,961]" captionTargetBox="[184,1385,198,897]" captionTargetId="figure-433@10.[181,1388,195,900]" captionTargetPageId="10" captionText="Figure 6. Ancestral state reconstruction analyses based on the Bayesian tree from this study, depicting: A, number of chromosomes; B, position of 18S rDNA. Results of maximum parsimony analysis reconstructing ancestral states for both cytogenetic characters are visualized by colours of the circles. Arrowheads indicate hypothetical inversion on 18S rDNAbearing chromosomes." figureDoi="http://doi.org/10.5281/zenodo.7184508" httpUri="https://zenodo.org/record/7184508/files/figure.png" pageId="8" pageNumber="893">Fig. 6A</figureCitation>
|
||
). Two independent increases of 2
|
||
<emphasis id="B977A25CFFA71F20FA2A1442FA10976B" box="[1410,1425,1885,1906]" italics="true" pageId="8" pageNumber="893">n</emphasis>
|
||
were detected within
|
||
<taxonomicName id="4C0305CDFFA61F21FE0115E1FE5F970A" box="[425,478,1790,1811]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="9" pageNumber="894" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977A25CFFA61F21FE0115E1FE5F970A" box="[425,478,1790,1811]" italics="true" pageId="9" pageNumber="894">Gint</emphasis>
|
||
</taxonomicName>
|
||
: (1) the clade containing all cytotypes of
|
||
<taxonomicName id="4C0305CDFFA61F21FECF1402FD989728" box="[359,537,1820,1842]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="9" pageNumber="894" phylum="Arthropoda" rank="species" species="amoudensis">
|
||
<emphasis id="B977A25CFFA61F21FECF1402FD989728" box="[359,537,1820,1842]" italics="true" pageId="9" pageNumber="894">G. amoudensis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis id="B977A25CFFA61F21FDF21402FC8B9728" box="[602,778,1820,1842]" italics="true" pageId="9" pageNumber="894">
|
||
<taxonomicName id="4C0305CDFFA61F21FDF21402FC879728" box="[602,774,1820,1842]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="9" pageNumber="894" phylum="Arthropoda" rank="species" species="gubanensis">G. gubanensis</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and (2) cytotype III within the
|
||
<taxonomicName id="4C0305CDFFA61F21FD811424FD3D9749" authorityName="Kovarik & Mazuch" authorityYear="2015" box="[553,700,1851,1872]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="9" pageNumber="894" phylum="Arthropoda" rank="species" species="dabakalo">
|
||
<emphasis id="B977A25CFFA61F21FD811424FD3D9749" box="[553,700,1851,1872]" italics="true" pageId="9" pageNumber="894">G. dabakalo</emphasis>
|
||
</taxonomicName>
|
||
clade. When compared with the ancestral 2
|
||
<emphasis id="B977A25CFFA61F21FDFA1445FDE09776" box="[594,609,1882,1903]" italics="true" pageId="9" pageNumber="894">n</emphasis>
|
||
, a tendency of decreasing of 2
|
||
<emphasis id="B977A25CFFA61F21FC4715E1FC7F970A" box="[1007,1022,1790,1811]" italics="true" pageId="9" pageNumber="894">n</emphasis>
|
||
from 30 to 18 chromosomes can be observed in
|
||
<taxonomicName id="4C0305CDFFA61F21FC611402FBBE9728" box="[969,1087,1820,1842]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="9" pageNumber="894" phylum="Arthropoda" rank="species" species="gaitako">
|
||
<emphasis id="B977A25CFFA61F21FC611402FBBE9728" box="[969,1087,1820,1842]" italics="true" pageId="9" pageNumber="894">G. gaitako</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C0305CDFFA61F21FBE51402FB589728" authorityName="Kovarik & Mazuch" authorityYear="2015" box="[1101,1241,1820,1842]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="9" pageNumber="894" phylum="Arthropoda" rank="species" species="dabakalo">
|
||
<emphasis id="B977A25CFFA61F21FBE51402FB589728" box="[1101,1241,1820,1842]" italics="true" pageId="9" pageNumber="894">G. dabakalo</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C0305CDFFA61F21FABD1402FA1C9728" authorityName="Kovarik & Lowe" authorityYear="2019" box="[1301,1437,1820,1842]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="9" pageNumber="894" phylum="Arthropoda" rank="species" species="banfasae">
|
||
<emphasis id="B977A25CFFA61F21FABD1402FA1C9728" box="[1301,1437,1820,1842]" italics="true" pageId="9" pageNumber="894">G. banfasae</emphasis>
|
||
</taxonomicName>
|
||
. Concerning the position of 18S rRNA gene clusters, maximum parsimony inferred their terminal position as the ancestral state for the genus
|
||
<taxonomicName id="4C0305CDFFA51F22FDE3175FFD01944C" box="[587,640,1088,1109]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="10" pageNumber="895" phylum="Arthropoda" rank="genus">
|
||
<emphasis id="B977A25CFFA51F22FDE3175FFD01944C" box="[587,640,1088,1109]" italics="true" pageId="10" pageNumber="895">Gint</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="133862CBFFA51F22FD3B175FFD68944F" box="[659,745,1088,1110]" captionStart="Figure 6" captionStartId="10.[147,227,939,961]" captionTargetBox="[184,1385,198,897]" captionTargetId="figure-433@10.[181,1388,195,900]" captionTargetPageId="10" captionText="Figure 6. Ancestral state reconstruction analyses based on the Bayesian tree from this study, depicting: A, number of chromosomes; B, position of 18S rDNA. Results of maximum parsimony analysis reconstructing ancestral states for both cytogenetic characters are visualized by colours of the circles. Arrowheads indicate hypothetical inversion on 18S rDNAbearing chromosomes." figureDoi="http://doi.org/10.5281/zenodo.7184508" httpUri="https://zenodo.org/record/7184508/files/figure.png" pageId="10" pageNumber="895">Fig. 6B</figureCitation>
|
||
). Two independent changes of the ancestral terminal position to a subterminal position were identified in
|
||
<taxonomicName id="4C0305CDFFA51F22FF3A1783FEB494A8" box="[146,309,1180,1201]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="10" pageNumber="895" phylum="Arthropoda" rank="species" species="amoudensis">
|
||
<emphasis id="B977A25CFFA51F22FF3A1783FEB494A8" box="[146,309,1180,1201]" italics="true" pageId="10" pageNumber="895">G. amoudensis</emphasis>
|
||
</taxonomicName>
|
||
and in the clade containing
|
||
<taxonomicName id="4C0305CDFFA51F22FDD81783FD7894A8" authorityName="Kovarik & Mazuch" authorityYear="2015" box="[624,761,1180,1201]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="10" pageNumber="895" phylum="Arthropoda" rank="species" species="dabakalo">
|
||
<emphasis id="B977A25CFFA51F22FDD81783FD7894A8" box="[624,761,1180,1201]" italics="true" pageId="10" pageNumber="895">G. dabakalo</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C0305CDFFA51F22FF6B17A4FEC794D6" authorityName="Kovarik & Lowe" authorityYear="2019" box="[195,326,1210,1232]" class="Arachnida" family="Buthidae" genus="Gint" kingdom="Animalia" order="Scorpiones" pageId="10" pageNumber="895" phylum="Arthropoda" rank="species" species="banfasae">
|
||
<emphasis id="B977A25CFFA51F22FF6B17A4FEC794D6" box="[195,326,1210,1232]" italics="true" pageId="10" pageNumber="895">G. banfasae</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |