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214 lines
22 KiB
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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.769.25814" ID-GBIF-Dataset="24cfa1f5-171f-423b-8d0e-a66bf52b9fc2" ID-PMC="PMC6030207" ID-Pensoft-Pub="1313-2970-769-49" ID-Pensoft-UUID="FFADFF955D05FFC8857C7F46FFC4FF85" ID-PubMed="29988790" ID-Zenodo-Dep="1304507" ID-ZooBank="752922C5DD1842B698665716A341A94C" ModsDocID="1313-2970-769-49" checkinTime="1530090465466" checkinUser="pensoft" docAuthor="Hedin, Marshal, Derkarabetian, Shahan, Blair, Jennifer & Paquin, Pierre" docDate="2018" docId="33ADB0F519E06811BD0546508FA3F7B4" docLanguage="en" docName="ZooKeys 769: 49-76" docOrigin="ZooKeys 769" docPubDate="2018-06-26" docSource="http://dx.doi.org/10.3897/zookeys.769.25814" docTitle="Cicurina (Cicurella) vespera Gertsch 1992" docType="treatment" docVersion="4" id="FFADFF955D05FFC8857C7F46FFC4FF85" lastPageNumber="58" masterDocId="FFADFF955D05FFC8857C7F46FFC4FF85" masterDocTitle="Sequence capture phylogenomics of eyeless Cicurina spiders from Texas caves, with emphasis on US federally-endangered species from Bexar County (Araneae, Hahniidae)" masterLastPageNumber="76" masterPageNumber="49" pageNumber="58" updateTime="1668165938240" updateUser="ExternalLinkService">
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<mods:titleInfo>
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<mods:title>Sequence capture phylogenomics of eyeless Cicurina spiders from Texas caves, with emphasis on US federally-endangered species from Bexar County (Araneae, Hahniidae)</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Hedin, Marshal</mods:namePart>
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<mods:affiliation>Department of Biology, San Diego State University, 5500 Campanile Drive, San Diego CA, 92182, USA</mods:affiliation>
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<mods:nameIdentifier type="email">mhedin@mail.sdsu.edu</mods:nameIdentifier>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Derkarabetian, Shahan</mods:namePart>
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<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-9163-9277</mods:nameIdentifier>
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<mods:affiliation>Department of Biology, San Diego State University, 5500 Campanile Drive, San Diego CA, 92182, USA & Department of Organismic and Evolutionary Biology, Museum of Comparative Zoology, Harvard University, 26 Oxford St., Cambridge MA, 02138, USA</mods:affiliation>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Blair, Jennifer</mods:namePart>
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<mods:affiliation>Blair Wildlife Consulting, 3815 Dacy Lane, Kyle TX, 78640, USA</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Paquin, Pierre</mods:namePart>
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<mods:affiliation>Scienceinfuse Inc, 12 Saxby Sud, Shefford, QC, J 2 M 1 S 2, Canada</mods:affiliation>
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<mods:title>ZooKeys</mods:title>
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<mods:date>2018</mods:date>
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<mods:number>2018-06-26</mods:number>
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<mods:detail type="volume">
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<mods:number>769</mods:number>
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<mods:url>http://dx.doi.org/10.3897/zookeys.769.25814</mods:url>
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<subSection pageId="9" pageNumber="58" type="discussion">
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<treatment ID-GBIF-Taxon="182229730" LSID="urn:lsid:plazi:treatment:33ADB0F519E06811BD0546508FA3F7B4" httpUri="http://treatment.plazi.org/id/33ADB0F519E06811BD0546508FA3F7B4" lastPageNumber="58" pageId="9" pageNumber="58">
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<taxonomicName authority="Gertsch, 1992" authorityName="Gertsch" authorityYear="1992" class="Arachnida" family="Hahniidae" genus="Cicurina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cicurina (Cicurella) vespera" order="Araneae" pageId="9" pageNumber="58" phylum="Arthropoda" rank="species" species="vespera" subGenus="Cicurella">Cicurina (Cicurella) vespera Gertsch, 1992</taxonomicName>
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<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Phylogenetic tree from RAxML analysis of 399 UCE loci, 70 % (399 _ 70) occupancy matrix. Bootstrap values below 75 not shown. Geography and morphology highlighted. Abbreviations: ME (= mostly elongate) spermathecae clade, R (= rounded) spermathecae clade. Further details regarding specimen codes and locations found in Suppl. material 5. Inset: sampled Texas counties." figureDoi="10.3897/zookeys.769.25814.figure1" httpUri="https://binary.pensoft.net/fig/211928" pageId="9" pageNumber="58">Figs 1</figureCitation>
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<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Phylogenetic tree from BEAST analysis of 399 _ 70 UCE matrix. Posterior probability values above 0.95 shown. Inset: Boxplots of TI values (from Suppl. material 6) for ME and R clade members from Bexar County (for ME clade, two small juveniles and Stahl Cave specimens not included)." figureDoi="10.3897/zookeys.769.25814.figure2" httpUri="https://binary.pensoft.net/fig/211929" pageId="9" pageNumber="58">, 2</figureCitation>
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<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Phylogenetic tree from RAxML analysis of COI mitochondrial data. Previously published sequences with corresponding GenBank numbers (AY ), some with cave location codes as in Paquin and Hedin (2004). Bootstrap values above 70 shown only for major lineages. Bootstrap values within C. madla shown on Figure 6, those for C. loftini shown on Figure 10." figureDoi="10.3897/zookeys.769.25814.figure3" httpUri="https://binary.pensoft.net/fig/211930" pageId="9" pageNumber="58">, 3</figureCitation>
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<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Morphology of R clade with corresponding 399 _ 70 UCE RAxML phylogeny (not all bootstrap values shown). Holotype female spermathecal images from Paquin and Duperre (2009), used with permission. Images not to scale." figureDoi="10.3897/zookeys.769.25814.figure7" httpUri="https://binary.pensoft.net/fig/211934" pageId="9" pageNumber="58">, 7</figureCitation>
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<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Distribution of R clade in Bexar and Comal counties, with corresponding 399 _ 70 UCE RAxML phylogeny. Different species designated by different colors. Cave locations approximate." figureDoi="10.3897/zookeys.769.25814.figure8" httpUri="https://binary.pensoft.net/fig/211935" pageId="9" pageNumber="58">, 8</figureCitation>
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<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Morphology of adult C. loftini with corresponding 399 _ 70 UCE RAxML phylogeny. Imaged specimens highlighted on phylogeny; images of specimen TK _ 188520 from J. Ledford, specimen TK _ 188686 from J. Cokendolpher (via Symbiota Collections of Arthropods Network). Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.769.25814.figure9" httpUri="https://binary.pensoft.net/fig/211936" pageId="9" pageNumber="58">, 9</figureCitation>
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<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Distribution of C. loftini in Culebra Anticline KFR, with corresponding COI RAxML phylogeny. Different mtDNA microclades designated by different colors. Cave locations approximate." figureDoi="10.3897/zookeys.769.25814.figure10" httpUri="https://binary.pensoft.net/fig/211937" pageId="9" pageNumber="58">, 10</figureCitation>
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<figureCitation captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Anomalous geographic distribution of C. venii and C. vespera type specimens. All views ventral." figureDoi="10.3897/zookeys.769.25814.figure11" httpUri="https://binary.pensoft.net/fig/211938" pageId="9" pageNumber="58">, 11</figureCitation>
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<figureCitation captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Male palpal morphology of C. madla and C. vespera (left palp, ventral view). Specimens from Stevens Ranch Trash Hole Cave (CIC _ 1745) and Robbers Cave (TX _ 241) from PP personal collection. Image of specimen TK _ 188519 from Joel Ledford. Abbreviations: CB = cymbium, T = tegulum, E = embolus. Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.769.25814.figure12" httpUri="https://binary.pensoft.net/fig/211939" pageId="9" pageNumber="58">, 12</figureCitation>
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, Suppl. materials 1, 2, 3, 4, 5, 6.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="9" pageNumber="58" type="reference_group">
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<paragraph pageId="9" pageNumber="58">
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<taxonomicName authorityName="Gertsch" authorityYear="1992" class="Arachnida" family="Hahniidae" genus="Cicurina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cicurina vespera" order="Araneae" pageId="9" pageNumber="58" phylum="Arthropoda" rank="species" species="vespera">Cicurina vespera</taxonomicName>
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Gertsch, 1992: 111, figs 93-94.
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</paragraph>
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<paragraph pageId="9" pageNumber="58">
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<taxonomicName authorityName="Gertsch" authorityYear="1992" class="Arachnida" family="Hahniidae" genus="Cicurina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cicurina vespera" order="Araneae" pageId="9" pageNumber="58" phylum="Arthropoda" rank="species" species="vespera">Cicurina vespera</taxonomicName>
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Cokendolpher, 2004: 53, figs 65-66.
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</paragraph>
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<paragraph pageId="9" pageNumber="58">
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<taxonomicName authorityName="Gertsch" authorityYear="1992" class="Arachnida" family="Hahniidae" genus="Cicurina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cicurina vespera" order="Araneae" pageId="9" pageNumber="58" phylum="Arthropoda" rank="species" species="vespera">Cicurina vespera</taxonomicName>
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Paquin &
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<normalizedToken originalValue="Dupérré">Duperre</normalizedToken>
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, 2009: 53, figs 118-119, 134.
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</paragraph>
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<paragraph pageId="9" pageNumber="58">
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<taxonomicName authorityName="Cokendolpher" authorityYear="2004" class="Arachnida" family="Hahniidae" genus="Cicurina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cicurina loftini" order="Araneae" pageId="9" pageNumber="58" phylum="Arthropoda" rank="species" species="loftini">Cicurina loftini</taxonomicName>
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Cokendolpher, 2004: 41, figs 5, 10, 37-39;
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<emphasis bold="true" pageId="9" pageNumber="58">syn. n.</emphasis>
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</paragraph>
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<paragraph pageId="9" pageNumber="58">
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<taxonomicName authorityName="Cokendolpher" authorityYear="2004" class="Arachnida" family="Hahniidae" genus="Cicurina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cicurina loftini" order="Araneae" pageId="9" pageNumber="58" phylum="Arthropoda" rank="species" species="loftini">Cicurina loftini</taxonomicName>
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Paquin &
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<normalizedToken originalValue="Dupérré">Duperre</normalizedToken>
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, 2009: 27, figs 46-47, 134;
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<emphasis bold="true" pageId="9" pageNumber="58">syn. n.</emphasis>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="9" pageNumber="58" type="diagnosis">
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<paragraph pageId="9" pageNumber="58">Diagnosis.</paragraph>
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<paragraph pageId="9" pageNumber="58">
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Based on well-supported nuclear phylogenomic analyses (Figs
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<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Phylogenetic tree from RAxML analysis of 399 UCE loci, 70 % (399 _ 70) occupancy matrix. Bootstrap values below 75 not shown. Geography and morphology highlighted. Abbreviations: ME (= mostly elongate) spermathecae clade, R (= rounded) spermathecae clade. Further details regarding specimen codes and locations found in Suppl. material 5. Inset: sampled Texas counties." figureDoi="10.3897/zookeys.769.25814.figure1" httpUri="https://binary.pensoft.net/fig/211928" pageId="9" pageNumber="58">1</figureCitation>
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-
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<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Phylogenetic tree from BEAST analysis of 399 _ 70 UCE matrix. Posterior probability values above 0.95 shown. Inset: Boxplots of TI values (from Suppl. material 6) for ME and R clade members from Bexar County (for ME clade, two small juveniles and Stahl Cave specimens not included)." figureDoi="10.3897/zookeys.769.25814.figure2" httpUri="https://binary.pensoft.net/fig/211929" pageId="9" pageNumber="58">2</figureCitation>
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,
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<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Morphology of R clade with corresponding 399 _ 70 UCE RAxML phylogeny (not all bootstrap values shown). Holotype female spermathecal images from Paquin and Duperre (2009), used with permission. Images not to scale." figureDoi="10.3897/zookeys.769.25814.figure7" httpUri="https://binary.pensoft.net/fig/211934" pageId="9" pageNumber="58">7</figureCitation>
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), sister taxon to a clade including
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<taxonomicName lsidName="C. baronia" pageId="9" pageNumber="58" rank="species" species="baronia">
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<emphasis italics="true" pageId="9" pageNumber="58">C. baronia</emphasis>
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</taxonomicName>
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,
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<taxonomicName lsidName="C. neovespera" pageId="9" pageNumber="58" rank="species" species="neovespera">
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<emphasis italics="true" pageId="9" pageNumber="58">C. neovespera</emphasis>
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</taxonomicName>
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, and
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<taxonomicName lsidName="C. bullis" pageId="9" pageNumber="58" rank="species" species="bullis">
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<emphasis italics="true" pageId="9" pageNumber="58">C. bullis</emphasis>
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</taxonomicName>
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.
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<taxonomicName authorityName="Gertsch" authorityYear="1992" class="Arachnida" family="Hahniidae" genus="Cicurina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cicurina vespera" order="Araneae" pageId="9" pageNumber="58" phylum="Arthropoda" rank="species" species="vespera">
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<emphasis italics="true" pageId="9" pageNumber="58">Cicurina vespera</emphasis>
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</taxonomicName>
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is morphologically very similar to the above taxa (Figure
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<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Morphology of R clade with corresponding 399 _ 70 UCE RAxML phylogeny (not all bootstrap values shown). Holotype female spermathecal images from Paquin and Duperre (2009), used with permission. Images not to scale." figureDoi="10.3897/zookeys.769.25814.figure7" httpUri="https://binary.pensoft.net/fig/211934" pageId="9" pageNumber="58">7</figureCitation>
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,
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<bibRefCitation author="Cokendolpher, JC" journalOrPublisher="Texas Memorial Museum, Speleological Monographs" pageId="9" pageNumber="58" refId="B9" refString="Cokendolpher, JC, 2004. Cicurina spiders from caves in Bexar County, Texas. Texas Memorial Museum Speleological Monographs, 6. Studies on the cave and endogean fauna of North America IV: 13-58." title="Cicurina spiders from caves in Bexar County, Texas. Texas Memorial Museum Speleological Monographs, 6. Studies on the cave and endogean fauna of North America IV: 13 - 58." year="2004">Cokendolpher 2004</bibRefCitation>
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,
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<bibRefCitation author="Paquin, P" journalOrPublisher="Zootaxa" pageId="9" pageNumber="58" pagination="1 - 67" refId="B26" refString="Paquin, P, Duperre, N, 2009. A first step towards the revision of Cicurina: redescription of type specimens of 60 troglobitic species of the subgenus Cicurella (Araneae: Dictynidae), and a first visual assessment of their distribution. Zootaxa 2002: 1 - 67" title="A first step towards the revision of Cicurina: redescription of type specimens of 60 troglobitic species of the subgenus Cicurella (Araneae: Dictynidae), and a first visual assessment of their distribution." volume="2002" year="2009">
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Paquin and
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<normalizedToken originalValue="Dupérré">Duperre</normalizedToken>
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2009
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</bibRefCitation>
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), best separated by geographic allopatry (Figure
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<figureCitation captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Distribution of R clade in Bexar and Comal counties, with corresponding 399 _ 70 UCE RAxML phylogeny. Different species designated by different colors. Cave locations approximate." figureDoi="10.3897/zookeys.769.25814.figure8" httpUri="https://binary.pensoft.net/fig/211935" pageId="9" pageNumber="58">8</figureCitation>
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). This species can be distinguished from neighboring or sympatric ME clade members using genetic data, spermathecal morphology, male palpal morphology, or TI index.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="9" pageNumber="58" type="description">
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<paragraph pageId="9" pageNumber="58">Description.</paragraph>
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<paragraph pageId="9" pageNumber="58">
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Female spermathecal morphology as described in
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<bibRefCitation author="Paquin, P" journalOrPublisher="Zootaxa" pageId="9" pageNumber="58" pagination="1 - 67" refId="B26" refString="Paquin, P, Duperre, N, 2009. A first step towards the revision of Cicurina: redescription of type specimens of 60 troglobitic species of the subgenus Cicurella (Araneae: Dictynidae), and a first visual assessment of their distribution. Zootaxa 2002: 1 - 67" title="A first step towards the revision of Cicurina: redescription of type specimens of 60 troglobitic species of the subgenus Cicurella (Araneae: Dictynidae), and a first visual assessment of their distribution." volume="2002" year="2009">
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Paquin and
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<normalizedToken originalValue="Dupérré">Duperre</normalizedToken>
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(2009)
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</bibRefCitation>
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and
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<bibRefCitation author="Cokendolpher, JC" journalOrPublisher="Texas Memorial Museum, Speleological Monographs" pageId="9" pageNumber="58" refId="B9" refString="Cokendolpher, JC, 2004. Cicurina spiders from caves in Bexar County, Texas. Texas Memorial Museum Speleological Monographs, 6. Studies on the cave and endogean fauna of North America IV: 13-58." title="Cicurina spiders from caves in Bexar County, Texas. Texas Memorial Museum Speleological Monographs, 6. Studies on the cave and endogean fauna of North America IV: 13 - 58." year="2004">Cokendolpher (2004)</bibRefCitation>
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. Male palpus with relatively broad, truncate cymbium, compact tegulum, origin of embolus rotated slightly clockwise from 6
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<normalizedToken originalValue="o’clock">o'clock</normalizedToken>
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(Figs
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<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Morphology of adult C. loftini with corresponding 399 _ 70 UCE RAxML phylogeny. Imaged specimens highlighted on phylogeny; images of specimen TK _ 188520 from J. Ledford, specimen TK _ 188686 from J. Cokendolpher (via Symbiota Collections of Arthropods Network). Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.769.25814.figure9" httpUri="https://binary.pensoft.net/fig/211936" pageId="9" pageNumber="58">9</figureCitation>
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,
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<figureCitation captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Male palpal morphology of C. madla and C. vespera (left palp, ventral view). Specimens from Stevens Ranch Trash Hole Cave (CIC _ 1745) and Robbers Cave (TX _ 241) from PP personal collection. Image of specimen TK _ 188519 from Joel Ledford. Abbreviations: CB = cymbium, T = tegulum, E = embolus. Scale bar: 0.5 mm." figureDoi="10.3897/zookeys.769.25814.figure12" httpUri="https://binary.pensoft.net/fig/211939" pageId="9" pageNumber="58">12</figureCitation>
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).
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</paragraph>
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</subSubSection>
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<subSubSection pageId="9" pageNumber="58" type="distribution">
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<paragraph pageId="9" pageNumber="58">Distribution.</paragraph>
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<paragraph pageId="9" pageNumber="58">
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R clade member known from 16 cave or karst features in the Culebra Anticline KFR (Figure
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<figureCitation captionStart="Figure 10" captionStartId="F10" captionText="Figure 10. Distribution of C. loftini in Culebra Anticline KFR, with corresponding COI RAxML phylogeny. Different mtDNA microclades designated by different colors. Cave locations approximate." figureDoi="10.3897/zookeys.769.25814.figure10" httpUri="https://binary.pensoft.net/fig/211937" pageId="9" pageNumber="58">10</figureCitation>
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); possibly also from GCBC.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="9" pageNumber="58" type="discussion">
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<paragraph pageId="9" pageNumber="58">Discussion.</paragraph>
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<paragraph pageId="9" pageNumber="58">
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The low TI index, rounded spermathecae holotype
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<taxonomicName lsidName="C. vespera" pageId="9" pageNumber="58" rank="species" species="vespera">
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<emphasis italics="true" pageId="9" pageNumber="58">C. vespera</emphasis>
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</taxonomicName>
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type specimen is either 1) actually from Bracken Bat Cave, but was mislabeled or placed into an incorrect vial, or 2) is actually from GCBC, and represents a northern disjunct from most Culebra Anticline
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<taxonomicName lsidName="C. vespera" pageId="9" pageNumber="58" rank="species" species="vespera">
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<emphasis italics="true" pageId="9" pageNumber="58">C. vespera</emphasis>
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</taxonomicName>
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populations, although we note that direct evidence for such an extension does not exist in the Government Canyon KFR (Figure
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<figureCitation captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Anomalous geographic distribution of C. venii and C. vespera type specimens. All views ventral." figureDoi="10.3897/zookeys.769.25814.figure11" httpUri="https://binary.pensoft.net/fig/211938" pageId="9" pageNumber="58">11</figureCitation>
|
||
). If the latter, rare sympatry must exist in GCBC, as other known specimens from this location represent
|
||
<taxonomicName lsidName="C. madla" pageId="9" pageNumber="58" rank="species" species="madla">
|
||
<emphasis italics="true" pageId="9" pageNumber="58">C. madla</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<caption ID-Zenodo-Dep="1304621" doi="10.3897/zookeys.769.25814.figure12" httpUri="https://binary.pensoft.net/fig/211939" pageId="9" pageNumber="58" start="Figure 12" startId="F12">
|
||
<paragraph pageId="9" pageNumber="58">
|
||
<emphasis bold="true" pageId="9" pageNumber="58">Figure 12.</emphasis>
|
||
Male palpal morphology of
|
||
<taxonomicName lsidName="C. madla" pageId="9" pageNumber="58" rank="species" species="madla">
|
||
<emphasis italics="true" pageId="9" pageNumber="58">C. madla</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName lsidName="C. vespera" pageId="9" pageNumber="58" rank="species" species="vespera">
|
||
<emphasis italics="true" pageId="9" pageNumber="58">C. vespera</emphasis>
|
||
</taxonomicName>
|
||
(left palp, ventral view). Specimens from Stevens Ranch Trash Hole Cave (CIC_1745) and Robbers Cave (TX_241) from PP personal collection. Image of specimen TK_188519 from Joel Ledford. Abbreviations: CB = cymbium, T = tegulum, E = embolus. Scale bar: 0.5 mm.
|
||
</paragraph>
|
||
</caption>
|
||
</subSubSection>
|
||
</treatment>
|
||
</subSection>
|
||
</document> |