194 lines
18 KiB
XML
194 lines
18 KiB
XML
<document ID-DOI="http://dx.doi.org/10.3897/zookeys.789.22781" ID-GBIF-Dataset="aa6fb882-7e3b-40a8-9458-2b9c746b7dc7" ID-PMC="PMC6193417" ID-Pensoft-Pub="1313-2970-789-51" ID-PubMed="30344435" ID-ZBK="496949FCA96A4489A0940182520DAB6C" ModsDocAuthor="" ModsDocDate="2018" ModsDocID="1313-2970-789-51" ModsDocOrigin="ZooKeys 789" ModsDocTitle="The phylogeny of pholcid spiders: a critical evaluation of relationships suggested by molecular data (Araneae, Pholcidae)" checkinTime="1539222998747" checkinUser="pensoft" docAuthor="Huber, Bernhard A., Eberle, Jonas & Dimitrov, Dimitar" docDate="2018" docId="285A0164DC475E0E155B4866F6DCC205" docLanguage="en" docName="ZooKeys 789: 51-101" docOrigin="ZooKeys 789" docSource="http://dx.doi.org/10.3897/zookeys.789.22781" docTitle="Ninetinae Simon 1890" docType="treatment" docVersion="6" lastPageNumber="54" masterDocId="FF94FFCCE316FFFCFFBAFFD4FFECFFF7" masterDocTitle="The phylogeny of pholcid spiders: a critical evaluation of relationships suggested by molecular data (Araneae, Pholcidae)" masterLastPageNumber="101" masterPageNumber="51" pageNumber="53" updateTime="1668166286259" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>The phylogeny of pholcid spiders: a critical evaluation of relationships suggested by molecular data (Araneae, Pholcidae)</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:role>
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<mods:namePart>Huber, Bernhard A.</mods:namePart>
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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:role>
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<mods:namePart>Eberle, Jonas</mods:namePart>
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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:role>
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<mods:namePart>Dimitrov, Dimitar</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>ZooKeys</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2018</mods:date>
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<mods:detail type="volume">
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<mods:number>789</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>51</mods:start>
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<mods:end>101</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:location>
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<mods:url>http://dx.doi.org/10.3897/zookeys.789.22781</mods:url>
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</mods:location>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.789.22781</mods:identifier>
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<mods:identifier type="Pensoft-Pub">1313-2970-789-51</mods:identifier>
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<mods:identifier type="ZBK">496949FCA96A4489A0940182520DAB6C</mods:identifier>
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<mods:identifier type="ZooBank">496949FCA96A4489A0940182520DAB6C</mods:identifier>
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</mods:mods>
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<treatment ID-GBIF-Taxon="148671576" LSID="urn:lsid:plazi:treatment:285A0164DC475E0E155B4866F6DCC205" httpUri="http://treatment.plazi.org/id/285A0164DC475E0E155B4866F6DCC205" lastPageId="3" lastPageNumber="54" pageId="2" pageNumber="53">
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<subSubSection pageId="2" pageNumber="53" type="nomenclature">
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<paragraph pageId="2" pageNumber="53">
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Subfamily
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<taxonomicName authority="Simon, 1890" authorityName="Simon" authorityYear="1890" family="Pholcidae" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="" order="Araneae" pageId="2" pageNumber="53" rank="subFamily" subFamily="Ninetinae">Ninetinae Simon, 1890</taxonomicName>
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Figure 2
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</paragraph>
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</subSubSection>
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<subSubSection pageId="2" pageNumber="53" type="reference_group">
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<paragraph pageId="2" pageNumber="53">
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<taxonomicName family="Pholcidae" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="" order="Araneae" pageId="2" pageNumber="53" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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Simon, 1890: 95. Type genus
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Ninetis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ninetis" order="Araneae" pageId="2" pageNumber="53" phylum="Arthropoda" rank="genus">Ninetis</taxonomicName>
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Simon, 1890, by monotypy.
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<bibRefCitation author="Huber, BA" journalOrPublisher="Journal of Arachnology" pageId="41" pageNumber="92" pagination="211 - 222" title="Phylogeny and classification of Pholcidae (Araneae): an update." url="https://doi.org/10.1636/CA10-57.1" volume="39" year="2011 b">Huber 2011b</bibRefCitation>
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: 212.
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="3" lastPageNumber="54" pageId="2" pageNumber="53" type="remarks">
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<paragraph pageId="2" pageNumber="53">Remarks.</paragraph>
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<paragraph pageId="2" pageNumber="53">
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<taxonomicName lsidName="" pageId="2" pageNumber="53" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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are small to tiny ground-dwelling spiders that are largely restricted to arid environments (
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<bibRefCitation author="Huber, BA" journalOrPublisher="Insect Systematics and Evolution" pageId="41" pageNumber="92" pagination="15 - 20" title="Ibotyporanga Mello-Leitao: tropical spiders in Brazilian semi-arid habitats (Araneae: Pholcidae)." url="https://doi.org/10.1163/187631203788964926" volume="34" year="2003">Huber and Brescovit 2003</bibRefCitation>
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; BA Huber, unpublished data). With only 31 described extant species, the subfamily is by far the smallest of the five currently recognized subfamilies in
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<taxonomicName family="Pholcidae" lsidName="" pageId="2" pageNumber="53" rank="family">Pholcidae</taxonomicName>
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.
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<taxonomicName lsidName="" pageId="2" pageNumber="53" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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seem to be diverse in the New World (ten named genera + about four unnamed genera; BA Huber, unpublished data) where they represent the most southern (Argentina) and most northern (Canada) autochthonous pholcid records worldwide. Only two genera (
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Ninetis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ninetis" order="Araneae" pageId="2" pageNumber="53" phylum="Arthropoda" rank="genus">Ninetis</taxonomicName>
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Simon, 1890 and one unnamed; BA Huber, unpublished data) are known from the Old World.
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</paragraph>
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<paragraph pageId="2" pageNumber="53">
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Their short legs make them superficially strikingly different from
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<normalizedToken originalValue="‘typical’">'typical'</normalizedToken>
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long-legged pholcids. This distinctness was recognized as early as 1893, when
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<normalizedToken originalValue="Eugène">Eugene</normalizedToken>
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Simon classified the only ninetine species available to him in a separate subfamily "
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<taxonomicName lsidName="" pageId="2" pageNumber="53" rank="subFamily" subFamily="Ninetidinae">Ninetidinae</taxonomicName>
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", as opposed to all other pholcids classified in
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<taxonomicName lsidName="" pageId="2" pageNumber="53" rank="subFamily" subFamily="Pholcinae">Pholcinae</taxonomicName>
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(
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<bibRefCitation pageId="2" pageNumber="53">Simon 1893</bibRefCitation>
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). Subsequent morphological and molecular phylogenies have partly supported this view (
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<bibRefCitation author="Huber, BA" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="40" pageNumber="91" pagination="1 - 348" title="New World pholcid spiders (Araneae: Pholcidae): a revision at generic level." url="https://doi.org/10.1206/0003-0090(2000)254<0001:NWPSAP>2.0.CO;2" volume="254" year="2000">Huber 2000</bibRefCitation>
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,
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<bibRefCitation author="Dimitrov, D" journalOrPublisher="Cladistics" pageId="39" pageNumber="90" pagination="132 - 146" title="Pholcid spider molecular systematics revisited, with new insights into the biogeography and the evolution of the group." url="https://doi.org/10.1111/j.1096-0031.2012.00419.x" volume="29" year="2013">Dimitrov et al. 2013</bibRefCitation>
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) but never convincingly with strong support.
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</paragraph>
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<paragraph pageId="2" pageNumber="53">
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Our present analyses include 15 species representing eight of the eleven described genera, originating from both the New World and the Old World (Figure 2). A sister-group relationship between
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<taxonomicName lsidName="" pageId="2" pageNumber="53" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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and all other pholcids is not supported by our analyses. Instead, all four analyses put
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<taxonomicName lsidName="" pageId="2" pageNumber="53" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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as sister to
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Artema" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Artema" order="Araneae" pageId="2" pageNumber="53" phylum="Arthropoda" rank="genus">Artema</taxonomicName>
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Walckenaer, 1837, and this clade is in turn sister to all other pholcids. For reasons discussed below (under
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<taxonomicName lsidName="" pageId="2" pageNumber="53" rank="subFamily" subFamily="Arteminae">Arteminae</taxonomicName>
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), we consider this relationship between
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Artema" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Artema" order="Araneae" pageId="2" pageNumber="53" phylum="Arthropoda" rank="genus">Artema</taxonomicName>
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and
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<taxonomicName lsidName="" pageId="2" pageNumber="53" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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dubious. The conclusion here is that
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<taxonomicName lsidName="" pageId="2" pageNumber="53" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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are
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<normalizedToken originalValue="‘basal’">'basal'</normalizedToken>
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, either with
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Artema" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Artema" order="Araneae" pageId="2" pageNumber="53" phylum="Arthropoda" rank="genus">Artema</taxonomicName>
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or without, but in any case the external relationships of
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<taxonomicName lsidName="" pageId="2" pageNumber="53" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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remain unsatisfactorily resolved and need further study.
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</paragraph>
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<caption pageId="2" pageNumber="53">
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<paragraph pageId="2" pageNumber="53">
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Figure 2.
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<taxonomicName lsidName="" pageId="2" pageNumber="53" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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and
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<taxonomicName lsidName="" pageId="2" pageNumber="53" rank="subFamily" subFamily="Arteminae">Arteminae</taxonomicName>
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a
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Artema" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Artema" order="Araneae" pageId="2" pageNumber="53" phylum="Arthropoda" rank="genus">Artema</taxonomicName>
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sp. n.
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<normalizedToken originalValue="“Om14”">"Om14"</normalizedToken>
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(Oman) b Gen. n. (Ninetinae) sp. n.
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<normalizedToken originalValue="“Om6”">"Om6"</normalizedToken>
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(Oman) c
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Chisosa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chisosa diluta" order="Araneae" pageId="2" pageNumber="53" phylum="Arthropoda" rank="species" species="diluta">Chisosa diluta</taxonomicName>
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(USA) d Gen. n. (Arteminae) sp. n.
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<normalizedToken originalValue="“Ind82”">"Ind82"</normalizedToken>
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(Sulawesi).
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</paragraph>
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</caption>
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<paragraph pageId="3" pageNumber="54">
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<pageBreakToken pageId="3" pageNumber="54" start="start">The</pageBreakToken>
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monophyly of the subfamily receives high to full support in all analyses but the composition is slightly different from previous concepts: the North American
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Chisosa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chisosa" order="Araneae" pageId="3" pageNumber="54" phylum="Arthropoda" rank="genus">Chisosa</taxonomicName>
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Huber, 2000, originally thought to be a representative of
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<taxonomicName lsidName="" pageId="3" pageNumber="54" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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(
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<bibRefCitation author="Huber, BA" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="40" pageNumber="91" pagination="1 - 348" title="New World pholcid spiders (Araneae: Pholcidae): a revision at generic level." url="https://doi.org/10.1206/0003-0090(2000)254<0001:NWPSAP>2.0.CO;2" volume="254" year="2000">Huber 2000</bibRefCitation>
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), is moved to
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<taxonomicName lsidName="" pageId="3" pageNumber="54" rank="subFamily" subFamily="Arteminae">Arteminae</taxonomicName>
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. This move is also supported by male genitalic characters (massive palpal femur; procursus with dorsal apophysis and ventral pocket) and by somatic characters (exposed tarsal organ; reduction of epiandrous spigots;
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<bibRefCitation author="Huber, BA" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="40" pageNumber="91" pagination="1 - 348" title="New World pholcid spiders (Araneae: Pholcidae): a revision at generic level." url="https://doi.org/10.1206/0003-0090(2000)254<0001:NWPSAP>2.0.CO;2" volume="254" year="2000">Huber 2000</bibRefCitation>
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). Another genus that was previously (
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<bibRefCitation author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="45 - 53" title="On Old World ninetine spiders (Araneae: Pholcidae), with a new genus and species and the first record for Madagascar." volume="1635" year="2007">Huber and El Hennawy 2007</bibRefCitation>
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,
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<bibRefCitation author="Huber, BA" journalOrPublisher="Journal of Arachnology" pageId="41" pageNumber="92" pagination="211 - 222" title="Phylogeny and classification of Pholcidae (Araneae): an update." url="https://doi.org/10.1636/CA10-57.1" volume="39" year="2011 b">Huber 2011b</bibRefCitation>
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) thought to be a member of
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<taxonomicName lsidName="" pageId="3" pageNumber="54" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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is
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Nita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nita" order="Araneae" pageId="3" pageNumber="54" phylum="Arthropoda" rank="genus">Nita</taxonomicName>
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Huber & El Hennawy, 2007. As already suggested in a previous analysis (
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<bibRefCitation author="Dimitrov, D" journalOrPublisher="Cladistics" pageId="39" pageNumber="90" pagination="132 - 146" title="Pholcid spider molecular systematics revisited, with new insights into the biogeography and the evolution of the group." url="https://doi.org/10.1111/j.1096-0031.2012.00419.x" volume="29" year="2013">Dimitrov et al. 2013</bibRefCitation>
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),
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Nita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nita" order="Araneae" pageId="3" pageNumber="54" phylum="Arthropoda" rank="genus">Nita</taxonomicName>
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is not a member of
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<taxonomicName lsidName="" pageId="3" pageNumber="54" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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but of
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<taxonomicName lsidName="" pageId="3" pageNumber="54" rank="subFamily" subFamily="Arteminae">Arteminae</taxonomicName>
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.
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</paragraph>
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<paragraph pageId="3" pageNumber="54">
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The internal relationships of
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<taxonomicName lsidName="" pageId="3" pageNumber="54" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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suggested by the molecular data are difficult to evaluate: they are mostly neither supported nor contradicted by morphological data. Two details are remarkable because they suggest that South America may not only be the most diverse region as far as
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<taxonomicName lsidName="" pageId="3" pageNumber="54" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
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are concerned but also the ancestral region of the subfamily. First, the analyses fully support a monophyletic North and Central American/Caribbean clade (
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcophora" order="Araneae" pageId="3" pageNumber="54" phylum="Arthropoda" rank="genus">Pholcophora</taxonomicName>
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Banks, 1896;
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Papiamenta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Papiamenta" order="Araneae" pageId="3" pageNumber="54" phylum="Arthropoda" rank="genus">Papiamenta</taxonomicName>
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Huber, 2000; and unidentified taxa from Cuba and Puerto Rico; "clade 2e" in
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<bibRefCitation author="Huber, BA" journalOrPublisher="Journal of Arachnology" pageId="41" pageNumber="92" pagination="211 - 222" title="Phylogeny and classification of Pholcidae (Araneae): an update." url="https://doi.org/10.1636/CA10-57.1" volume="39" year="2011 b">Huber 2011b</bibRefCitation>
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) that is either nested among South American ancestors or is sister to the South American
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Ibotyporanga" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ibotyporanga" order="Araneae" pageId="3" pageNumber="54" phylum="Arthropoda" rank="genus">Ibotyporanga</taxonomicName>
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<normalizedToken originalValue="Mello-Leitão">Mello-Leitao</normalizedToken>
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, 1944 (with reasonable support in the 4+ genes tree only). Based on its geographic distribution, we predict that the Mexican
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Tolteca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tolteca" order="Araneae" pageId="3" pageNumber="54" phylum="Arthropoda" rank="genus">Tolteca</taxonomicName>
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is also a member of this clade. Second, the two Old World genera (
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Ninetis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ninetis" order="Araneae" pageId="3" pageNumber="54" phylum="Arthropoda" rank="genus">Ninetis</taxonomicName>
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and an undescribed genus from Oman) are also sister taxa (with low to modest support) and in all analyses (except for the 4+ genes analysis where
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Ninetis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ninetis" order="Araneae" pageId="3" pageNumber="54" phylum="Arthropoda" rank="genus">Ninetis</taxonomicName>
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is missing) nested among South American taxa.
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</paragraph>
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</subSubSection>
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</treatment>
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</document> |