910 lines
108 KiB
XML
910 lines
108 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="740EFDBBFB32C5739C37A21FAEE475D3" docLanguage="en" docName="ZooKeys 789: 51-101" docOrigin="ZooKeys 789" docSource="http://dx.doi.org/10.3897/zookeys.789.22781" docTitle="Pholcinae group subsp. 3 '" docType="treatment" docVersion="6" lastPageNumber="79" masterDocId="FF94FFCCE316FFFCFFBAFFD4FFECFFF7" masterDocTitle="The phylogeny of pholcid spiders: a critical evaluation of relationships suggested by molecular data (Araneae, Pholcidae)" masterLastPageNumber="101" masterPageNumber="51" pageNumber="69" updateTime="1668166286259" updateUser="ExternalLinkService">
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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:role>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Huber, Bernhard A.</mods:namePart>
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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: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: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="148671572" LSID="urn:lsid:plazi:treatment:740EFDBBFB32C5739C37A21FAEE475D3" httpUri="http://treatment.plazi.org/id/740EFDBBFB32C5739C37A21FAEE475D3" lastPageId="28" lastPageNumber="79" pageId="18" pageNumber="69">
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<subSubSection pageId="18" pageNumber="69" type="nomenclature">
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<paragraph pageId="18" pageNumber="69">
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<taxonomicName family="Pholcidae" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Pholcinae group subsp. 3" order="Araneae" pageId="18" pageNumber="69" rank="subFamily" subFamily="Pholcinae 'group 3'">Pholcinae 'group 3'</taxonomicName>
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Figs 9, 10, 11, 12
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="28" lastPageNumber="79" pageId="19" pageNumber="70" type="remarks">
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<paragraph pageId="19" pageNumber="70">
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<pageBreakToken pageId="19" pageNumber="70" start="start">Remarks</pageBreakToken>
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.
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</paragraph>
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<paragraph pageId="19" pageNumber="70">
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A sister-group relationship between the African genus
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Quamtana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Quamtana" order="Araneae" pageId="19" pageNumber="70" phylum="Arthropoda" rank="genus">Quamtana</taxonomicName>
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and the
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="19" pageNumber="70" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
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group of genera (Figure 9) is recovered in all our analyses. Support values are low, but a morphological cladistic analysis has partly suggested the same relationship (based on a distinct sclerite connecting the genital bulb to the palpal tarsus;
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<bibRefCitation author="Huber, BA" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="40" pageNumber="91" pagination="477 - 527" title="Southern African pholcid spiders: revision and cladistic analysis of Quamtana gen. nov. and Spermophora Hentz (Araneae: Pholcidae), with notes on male-female covariation." url="https://doi.org/10.1046/j.0024-4082.2003.00082.x" volume="139" year="2003 c">Huber 2003c</bibRefCitation>
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). The monophyly of
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Quamtana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Quamtana" order="Araneae" pageId="19" pageNumber="70" phylum="Arthropoda" rank="genus">Quamtana</taxonomicName>
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is highly supported in all analyses (except for the 4+ genes analysis). It was also supported by morphological data when using character weighting (but not in the equal weights analysis;
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<bibRefCitation author="Huber, BA" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="40" pageNumber="91" pagination="477 - 527" title="Southern African pholcid spiders: revision and cladistic analysis of Quamtana gen. nov. and Spermophora Hentz (Araneae: Pholcidae), with notes on male-female covariation." url="https://doi.org/10.1046/j.0024-4082.2003.00082.x" volume="139" year="2003 c">Huber 2003c</bibRefCitation>
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).
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</paragraph>
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<paragraph pageId="19" pageNumber="70">
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Within
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Quamtana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Quamtana" order="Araneae" pageId="19" pageNumber="70" phylum="Arthropoda" rank="genus">Quamtana</taxonomicName>
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(Figure 9), our data suggest that there is no simple geographic pattern with respect to South African species (the large majority) versus species from other parts of Africa (marked in Figure 9). By contrast, three species groups with reasonable to full support include species from both South Africa and other regions: the South African
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<taxonomicName lsidName="Q. filmeri" pageId="19" pageNumber="70" rank="species" species="filmeri">Q. filmeri</taxonomicName>
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Huber, 2003 is sister to the Madagascan undescribed species
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<normalizedToken originalValue="“CAS5”">"CAS5"</normalizedToken>
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; the South African
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<taxonomicName lsidName="Q. vidal" pageId="19" pageNumber="70" rank="species" species="vidal">Q. vidal</taxonomicName>
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Huber, 2003 and
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<taxonomicName lsidName="Q. umzinto" pageId="19" pageNumber="70" rank="species" species="umzinto">Q. umzinto</taxonomicName>
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Huber, 2003 are placed in a group with species from East and Central Africa (
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<taxonomicName lsidName="Q. kabale" pageId="19" pageNumber="70" rank="species" species="kabale">Q. kabale</taxonomicName>
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Huber, 2003,
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<normalizedToken originalValue="“Cam117”">"Cam117"</normalizedToken>
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); and the group including the South African
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<taxonomicName lsidName="Q. embuleni" pageId="19" pageNumber="70" rank="species" species="embuleni">Q. embuleni</taxonomicName>
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Huber, 2003 and
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<taxonomicName lsidName="Q. bonamanzi" pageId="19" pageNumber="70" rank="species" species="bonamanzi">Q. bonamanzi</taxonomicName>
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Huber, 2003 also includes species from East and Central Africa (
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<taxonomicName lsidName="Q. kitahurira" pageId="19" pageNumber="70" rank="species" species="kitahurira">Q. kitahurira</taxonomicName>
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Huber, 2003,
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<taxonomicName lsidName="Q. oku" pageId="19" pageNumber="70" rank="species" species="oku">Q. oku</taxonomicName>
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Huber, 2003). We suspect that
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Quamtana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Quamtana" order="Araneae" pageId="19" pageNumber="70" phylum="Arthropoda" rank="genus">Quamtana</taxonomicName>
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was once widely distributed throughout Africa but largely replaced by more modern taxa in humid regions and extinguished in northern Africa. The Paris amber fossil
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Quamtana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Quamtana huberi" order="Araneae" pageId="19" pageNumber="70" phylum="Arthropoda" rank="species" species="huberi">Quamtana huberi</taxonomicName>
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Penney, 2007 supports this view, but its generic assignment is uncertain (
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<bibRefCitation author="Penney, D" journalOrPublisher="Journal of Arachnology" pageId="43" pageNumber="94" pagination="592 - 598" title="The oldest pholcid and selenopid spiders (Araneae) in lowermost Eocene amber from the Paris Basin, France." url="https://doi.org/10.1636/H05-61.1" volume="34" year="2007">Penney 2007</bibRefCitation>
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).
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</paragraph>
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<paragraph pageId="19" pageNumber="70">
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All remaining clades together (Figs 10-12) represent the
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="19" pageNumber="70" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
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group of genera sensu
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<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber (2011a)</bibRefCitation>
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. This clade was first proposed in Huber & Fleckenstein (2009) based on the distinctive simplified shape of the tarsus IV comb-hairs, and later supported in a cladistic morphological analysis by an additional character (female epigynal
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<normalizedToken originalValue="‘knob’">'knob'</normalizedToken>
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) (
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<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
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). All our analyses fully support this clade. The previous morphological analysis (
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<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
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) identified two major problems within this clade: (1) relationships among genera were basically unresolved, resulting in large polytomies; and (2) several species groups assigned to
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="19" pageNumber="70" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
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appeared more closely related to other genera. The present analysis strongly supports the polyphyly of
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="19" pageNumber="70" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
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in its previous composition, and it provides for the first time a reasonable framework to redefine generic limits in this large group (currently 501 species).
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</paragraph>
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<paragraph pageId="20" pageNumber="71">
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<pageBreakToken pageId="20" pageNumber="71" start="start">The</pageBreakToken>
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first major clade within the
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
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group of genera (Figure 10) is composed of three Southeast Asian genera (
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Calapnita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Calapnita" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Calapnita</taxonomicName>
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,
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
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,
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Uthina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Uthina" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Uthina</taxonomicName>
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Simon, 1893) as well as several Southeast Asian and Sri Lankan species groups that were originally tentatively assigned to
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
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(
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<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
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;
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<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 55" title="The Southeast Asian Pholcushalabala species group (Araneae, Pholcidae), new data from field observations and ultrastructure." url="https://doi.org/10.5852/ejt.2016.190" volume="190" year="2016 a">Huber et al. 2016a</bibRefCitation>
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,
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<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 45" title="New leaf- and litter-dwelling species of the genus Pholcus from Southeast Asia (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2016.200" volume="200" year="2016 b">2016b</bibRefCitation>
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,
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<bibRefCitation author="Huber, BA" journalOrPublisher="Zoologischer Anzeiger" pageId="41" pageNumber="92" pagination="394 - 403" title="Slow genital and genetic but rapid non-genital and ecological differentiation in a pair of spider species (Araneae, Pholcidae)." url="https://doi.org/10.1016/j.jcz.2014.04.001" volume="253" year="2014">Huber and Dimitrov 2014</bibRefCitation>
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). We informally call it the '
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Calapnita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Calapnita" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Calapnita</taxonomicName>
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-
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
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<normalizedToken originalValue="clade’">clade'</normalizedToken>
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because many species in this group are leaf-dwellers, and representatives of
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Calapnita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Calapnita" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Calapnita</taxonomicName>
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and
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
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are particularly strongly adapted to life on green leaves. Remarkably, even some of the species collected in the leaf litter (under large dead leaves on the ground) look like leaf-dwellers rather than litter dwellers (i.e., they have long abdomens, long legs, light colouration; e.g.,
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Kintaqa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kintaqa satun" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="species" species="satun">Kintaqa satun</taxonomicName>
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(Huber, 2011) and
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<taxonomicName lsidName="K. schwendingeri" pageId="20" pageNumber="71" rank="species" species="schwendingeri">K. schwendingeri</taxonomicName>
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(Huber, 2011); and Malaysian representatives of
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Tissahamia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tissahamia" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Tissahamia</taxonomicName>
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, previously the
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus ethagala" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="species" species="ethagala">Pholcus ethagala</taxonomicName>
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group). Ancestral character state reconstruction suggests that the ancestor of the entire clade was leaf-dwelling (
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<bibRefCitation author="Eberle, J" journalOrPublisher="Cladistics" pageId="39" pageNumber="90" url="https://doi.org/10.1186/s12862-018-1244-8" year="2018">Eberle et al. 2018</bibRefCitation>
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).
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</paragraph>
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<paragraph pageId="20" pageNumber="71">
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The present analyses reject the monophyly of
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Calapnita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Calapnita" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Calapnita</taxonomicName>
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(Figure 10). A recent cladistic analysis of morphological data (
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<bibRefCitation author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="1 - 63" title="Revision and cladistic analysis of the Southeast Asian leaf-dwelling spider genus Calapnita (Araneae, Pholcidae)." url="https://doi.org/10.11646/zootaxa.4219.1.1" volume="4219" year="2017">Huber 2017</bibRefCitation>
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) resolved
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Calapnita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Calapnita" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Calapnita</taxonomicName>
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as monophyletic but with low support (<50 using Jackknifing). On the other hand, support for the two subgroups, previously called phyllicola group and
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<taxonomicName genus="Pholcinae" lsidName="Pholcinae vermiformis" pageId="20" pageNumber="71" rank="species" species="vermiformis">vermiformis</taxonomicName>
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group, is full in all analyses. The two species groups have been identified long ago (
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<bibRefCitation author="Deeleman-Reinhold, CL" journalOrPublisher="Memoirs of the Queensland Museum" pageId="39" pageNumber="90" pagination="205 - 224" title="Studies on tropical Pholcidae II. Redescription of Micromerysgracilis Bradley and Calapnitavermiformis Simon (Araneae, Pholcidae) and description of some related new species." volume="22" year="1986 b">Deeleman-Reinhold 1986b</bibRefCitation>
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), and have been supported by cladistic analysis (
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<bibRefCitation author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="1 - 63" title="Revision and cladistic analysis of the Southeast Asian leaf-dwelling spider genus Calapnita (Araneae, Pholcidae)." url="https://doi.org/10.11646/zootaxa.4219.1.1" volume="4219" year="2017">Huber 2017</bibRefCitation>
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). Our analyses strongly suggest that the
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<taxonomicName genus="Pholcinae" lsidName="Pholcinae vermiformis" pageId="20" pageNumber="71" rank="species" species="vermiformis">vermiformis</taxonomicName>
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group is closer to species previously in
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
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than to the phyllicola group (see below). The phyllicola group is thus elevated to genus rank (
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Nipisa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nipisa" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Nipisa</taxonomicName>
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; see Taxonomy section below).
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</paragraph>
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<paragraph pageId="20" pageNumber="71">
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Within
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<taxonomicName class="Arachnida" family="Pholcidae" genus="Nipisa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nipisa" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Nipisa</taxonomicName>
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(Figure 10), the internal relationships proposed previously (
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<bibRefCitation author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="1 - 63" title="Revision and cladistic analysis of the Southeast Asian leaf-dwelling spider genus Calapnita (Araneae, Pholcidae)." url="https://doi.org/10.11646/zootaxa.4219.1.1" volume="4219" year="2017">Huber 2017</bibRefCitation>
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) are mostly supported even though data gaps are severe in this genus (several species with only two genes): (1)
|
||
<taxonomicName lsidName="N. lehi" pageId="20" pageNumber="71" rank="species" species="lehi">N. lehi</taxonomicName>
|
||
(Huber, 2017) [but not
|
||
<taxonomicName lsidName="N. kubah" pageId="20" pageNumber="71" rank="species" species="kubah">N. kubah</taxonomicName>
|
||
(Huber, 2017)] is a
|
||
<normalizedToken originalValue="‘basal’">'basal'</normalizedToken>
|
||
species, i.e., sister to all other species (reasonable to high support); (2) a clade including the species with egg-sacs that have all eggs aligned in a single row (weak support, possibly because
|
||
<taxonomicName lsidName="N. kubah" pageId="20" pageNumber="71" rank="species" species="kubah">N. kubah</taxonomicName>
|
||
is included, which is contradicted by morphology and egg-sac shape); (3) a clade including
|
||
<taxonomicName lsidName="N. semengoh" pageId="20" pageNumber="71" rank="species" species="semengoh">N. semengoh</taxonomicName>
|
||
(Huber, 2011) and its sister group, characterized by the position of the tarsal organ on a turret, a serrate embolus, and the shape of the pore-plates (full support).
|
||
</paragraph>
|
||
<paragraph pageId="20" pageNumber="71">
|
||
The relationships within
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Calapnita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Calapnita" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Calapnita</taxonomicName>
|
||
(previously
|
||
<taxonomicName genus="Pholcinae" lsidName="Pholcinae vermiformis" pageId="20" pageNumber="71" rank="species" species="vermiformis">vermiformis</taxonomicName>
|
||
group) proposed in
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="1 - 63" title="Revision and cladistic analysis of the Southeast Asian leaf-dwelling spider genus Calapnita (Araneae, Pholcidae)." url="https://doi.org/10.11646/zootaxa.4219.1.1" volume="4219" year="2017">Huber (2017)</bibRefCitation>
|
||
are only partly supported: (1) a clade with a continuous connection between epigynal plate and
|
||
<normalizedToken originalValue="‘knob’">'knob'</normalizedToken>
|
||
(all species in the present analysis except
|
||
<taxonomicName lsidName="C. bario" pageId="20" pageNumber="71" rank="species" species="bario">C. bario</taxonomicName>
|
||
Huber, 2017 and
|
||
<taxonomicName lsidName="C. saluang" pageId="20" pageNumber="71" rank="species" species="saluang">C. saluang</taxonomicName>
|
||
Huber, 2011; high support); (2) within the previous clade, a clade characterized by a prolateral process at the tip of the procursus (in the present analysis:
|
||
<taxonomicName lsidName="C. nunezae" pageId="20" pageNumber="71" rank="species" species="nunezae">C. nunezae</taxonomicName>
|
||
Huber, 2017 and
|
||
<taxonomicName lsidName="C. dinagat" pageId="20" pageNumber="71" rank="species" species="dinagat">C. dinagat</taxonomicName>
|
||
Huber, 2017; full support).
|
||
</paragraph>
|
||
<paragraph lastPageId="21" lastPageNumber="72" pageId="20" pageNumber="71">
|
||
The present analyses also reject the monophyly of
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
|
||
(Figure 10). They split the genus into two unrelated lineages, one of which is equivalent to what was previously called the nigrifrons group; the other is equivalent to the previous
|
||
<taxonomicName genus="Pholcinae" lsidName="Pholcinae vermiformis" pageId="20" pageNumber="71" rank="species" species="vermiformis">vermiformis</taxonomicName>
|
||
+ cavicola groups (
|
||
<bibRefCitation author="Deeleman-Reinhold, CL" journalOrPublisher="Journal of the New York Entomological Society" pageId="39" pageNumber="90" pagination="559 - 561" title="A new Panjange from northern Borneo (Araneae, Pholcidae)." volume="94" year="1986">Deeleman-Reinhold and Platnick 1986</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 46" title="Evolution of genital asymmetry, exaggerated eye stalks, and extreme palpal elongation in Panjange spiders (Araneae: Pholcidae)." url="https://doi.org/10.5852/ejt.2015.169" volume="169" year="2015">
|
||
Huber and
|
||
<normalizedToken originalValue="Nuñeza">Nuneza</normalizedToken>
|
||
2015
|
||
</bibRefCitation>
|
||
). Our analyses place each group with reasonable to full support in clades together with species previously assigned to
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="20" pageNumber="71" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
. A morphological cladistic analysis has recently
|
||
<pageBreakToken pageId="21" pageNumber="72" start="start">supported</pageBreakToken>
|
||
the monophyly of
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
|
||
based on the presence of parallel ridges ventrally on the procursus and on the reduction of the bulbal uncus (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 46" title="Evolution of genital asymmetry, exaggerated eye stalks, and extreme palpal elongation in Panjange spiders (Araneae: Pholcidae)." url="https://doi.org/10.5852/ejt.2015.169" volume="169" year="2015">
|
||
Huber and
|
||
<normalizedToken originalValue="Nuñeza">Nuneza</normalizedToken>
|
||
2015
|
||
</bibRefCitation>
|
||
). However, the monophyly was lost when using specific weighting parameters (implied weighting with K = 1 and K = 2), and some morphological characters do in fact support the split of
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
|
||
: (1) the loss of distal cheliceral apophyses in
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
|
||
and its closest relatives according to the present analyses; (2) the loss of an uncus in
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
|
||
and its closest relatives according to the present analyses. The nigrifrons group is thus elevated to genus rank (
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Apokayana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Apokayana" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="genus">Apokayana</taxonomicName>
|
||
; see Taxonomy section below).
|
||
</paragraph>
|
||
<paragraph pageId="21" pageNumber="72">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Apokayana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Apokayana" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="genus">Apokayana</taxonomicName>
|
||
is recovered with full support. This is remarkable considering the fact that in the morphological analysis its equivalent (the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange nigrifrons" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="species" species="nigrifrons">Panjange nigrifrons</taxonomicName>
|
||
group) was supported by a single homoplastic character only (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 46" title="Evolution of genital asymmetry, exaggerated eye stalks, and extreme palpal elongation in Panjange spiders (Araneae: Pholcidae)." url="https://doi.org/10.5852/ejt.2015.169" volume="169" year="2015">
|
||
Huber and
|
||
<normalizedToken originalValue="Nuñeza">Nuneza</normalizedToken>
|
||
2015
|
||
</bibRefCitation>
|
||
). Within the genus, our analyses identify two subgroups with full support each. These groups do not correspond to the relationships suggested in
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 46" title="Evolution of genital asymmetry, exaggerated eye stalks, and extreme palpal elongation in Panjange spiders (Araneae: Pholcidae)." url="https://doi.org/10.5852/ejt.2015.169" volume="169" year="2015">
|
||
Huber and
|
||
<normalizedToken originalValue="Nuñeza">Nuneza</normalizedToken>
|
||
(2015)
|
||
</bibRefCitation>
|
||
. In that analysis, each node was based on a single character, some of them not particularly convincing. We thus tend to prefer the present grouping even though our matrix is particularly incomplete in this genus (we did not manage to get 28S and CO1 sequences for any of the six species included).
|
||
</paragraph>
|
||
<paragraph pageId="21" pageNumber="72">
|
||
The monophyly of
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
|
||
(
|
||
<taxonomicName genus="Pholcinae" lsidName="Pholcinae vermiformis" pageId="21" pageNumber="72" rank="species" species="vermiformis">vermiformis</taxonomicName>
|
||
+ cavicola groups) is supported by several morphological characters (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 46" title="Evolution of genital asymmetry, exaggerated eye stalks, and extreme palpal elongation in Panjange spiders (Araneae: Pholcidae)." url="https://doi.org/10.5852/ejt.2015.169" volume="169" year="2015">
|
||
Huber and
|
||
<normalizedToken originalValue="Nuñeza">Nuneza</normalizedToken>
|
||
2015
|
||
</bibRefCitation>
|
||
), and receives high support in our present analyses (except for the 4+ genes analysis). The cavicola group (including also the two undescribed species
|
||
<normalizedToken originalValue="“Ind103”">"Ind103"</normalizedToken>
|
||
and
|
||
<normalizedToken originalValue="“Ind109”">"Ind109"</normalizedToken>
|
||
) was recovered as paraphyletic in
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 46" title="Evolution of genital asymmetry, exaggerated eye stalks, and extreme palpal elongation in Panjange spiders (Araneae: Pholcidae)." url="https://doi.org/10.5852/ejt.2015.169" volume="169" year="2015">
|
||
Huber and
|
||
<normalizedToken originalValue="Nuñeza">Nuneza</normalizedToken>
|
||
(2015)
|
||
</bibRefCitation>
|
||
but is here resolved as monophyletic. By contrast, the lanthana group which was supported by two morphological characters, one of them considered particularly strong (the unique direction of the embolus, pointing in the opposite direction of the appendix) is resolved as monophyletic only in the RogueNaRok tree; in the IQ-TREE and RAxML trees it is paraphyletic with respect to the cavicola group (actually, these trees suggest a basal trichotomy). Within the lanthana group, three species (
|
||
<taxonomicName lsidName="P. malagos" pageId="21" pageNumber="72" rank="species" species="malagos">P. malagos</taxonomicName>
|
||
Huber, 2015;
|
||
<taxonomicName lsidName="P. casaroro" pageId="21" pageNumber="72" rank="species" species="casaroro">P. casaroro</taxonomicName>
|
||
Huber, 2015;
|
||
<taxonomicName lsidName="P. camiguin" pageId="21" pageNumber="72" rank="species" species="camiguin">P. camiguin</taxonomicName>
|
||
Huber, 2015) share asymmetric male pedipalps, a character that is extremely rare in spiders (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Biological Reviews" pageId="42" pageNumber="93" pagination="647 - 698" title="The evolution of asymmetric genitalia in spiders and insects." url="https://doi.org/10.1111/j.1469-185X.2007.00029.x" volume="82" year="2007">Huber et al. 2007</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 46" title="Evolution of genital asymmetry, exaggerated eye stalks, and extreme palpal elongation in Panjange spiders (Araneae: Pholcidae)." url="https://doi.org/10.5852/ejt.2015.169" volume="169" year="2015">
|
||
Huber and
|
||
<normalizedToken originalValue="Nuñeza">Nuneza</normalizedToken>
|
||
2015
|
||
</bibRefCitation>
|
||
). This group is not recovered in any of the present analyses, where it consistently includes the symmetric
|
||
<taxonomicName lsidName="P. lanthana" pageId="21" pageNumber="72" rank="species" species="lanthana">P. lanthana</taxonomicName>
|
||
Deeleman-Reinhold & Deeleman, 1983 (requiring a regain of symmetry or two origins of asymmetry). Only the sister group relationship between
|
||
<taxonomicName lsidName="P. dinagat" pageId="21" pageNumber="72" rank="species" species="dinagat">P. dinagat</taxonomicName>
|
||
Huber, 2015 and
|
||
<taxonomicName lsidName="P. marilog" pageId="21" pageNumber="72" rank="species" species="marilog">P. marilog</taxonomicName>
|
||
Huber, 2015 is strongly supported by both morphology and molecules. In conclusion, alternative topologies within the lanthana group are supported by seemingly strong molecular and morphological data, respectively.
|
||
</paragraph>
|
||
<paragraph pageId="21" pageNumber="72">
|
||
Ten species groups previously assigned to
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
(in
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 55" title="The Southeast Asian Pholcushalabala species group (Araneae, Pholcidae), new data from field observations and ultrastructure." url="https://doi.org/10.5852/ejt.2016.190" volume="190" year="2016 a">Huber et al. 2016a</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 45" title="New leaf- and litter-dwelling species of the genus Pholcus from Southeast Asia (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2016.200" volume="200" year="2016 b">2016b</bibRefCitation>
|
||
) are representatives of the '
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Calapnita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Calapnita" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="genus">Calapnita</taxonomicName>
|
||
-
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
|
||
<normalizedToken originalValue="clade’">clade'</normalizedToken>
|
||
(Figure 10). Of these, nine are entirely Southeast Asian; only the ethagala group (now
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Tissahamia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tissahamia" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="genus">Tissahamia</taxonomicName>
|
||
) has representatives in Southeast Asia and Sri Lanka. For some of these species groups, our data provide strong evidence about the sister-group or close relatives. All of these groups are here transferred from
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="21" pageNumber="72" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
to new genera (see Taxonomy section below).
|
||
</paragraph>
|
||
<paragraph pageId="22" pageNumber="73">
|
||
<pageBreakToken pageId="22" pageNumber="73" start="start">A</pageBreakToken>
|
||
sister-group relationship between
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Kelabita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kelabita" order="Araneae" pageId="22" pageNumber="73" phylum="Arthropoda" rank="genus">Kelabita</taxonomicName>
|
||
(previously the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus andulau" order="Araneae" pageId="22" pageNumber="73" phylum="Arthropoda" rank="species" species="andulau">Pholcus andulau</taxonomicName>
|
||
group) and
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Apokayana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Apokayana" order="Araneae" pageId="22" pageNumber="73" phylum="Arthropoda" rank="genus">Apokayana</taxonomicName>
|
||
(previously the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange nigrifrons" order="Araneae" pageId="22" pageNumber="73" phylum="Arthropoda" rank="species" species="nigrifrons">Panjange nigrifrons</taxonomicName>
|
||
group) is fully supported in all our analyses (except for the 4+ genes tree where
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Kelabita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kelabita" order="Araneae" pageId="22" pageNumber="73" phylum="Arthropoda" rank="genus">Kelabita</taxonomicName>
|
||
is not represented). Both genera are restricted to Borneo and share habitus, colouration, web structure, and microhabitat (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="41" pageNumber="92" pagination="1 - 32" title="The Panjangenigrifrons group in Borneo (Araneae: Pholcidae): high diversity in Sarawak, apparent absence in Sabah." url="https://doi.org/10.5852/ejt.2016.184" volume="184" year="2016">Huber and Leh Moi Ung 2016</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 55" title="The Southeast Asian Pholcushalabala species group (Araneae, Pholcidae), new data from field observations and ultrastructure." url="https://doi.org/10.5852/ejt.2016.190" volume="190" year="2016 a">Huber et al. 2016a</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph lastPageId="23" lastPageNumber="74" pageId="22" pageNumber="73">
|
||
The western Indonesian '
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus kerinci" order="Araneae" pageId="22" pageNumber="73" phylum="Arthropoda" rank="species" species="kerinci">Pholcus kerinci</taxonomicName>
|
||
<normalizedToken originalValue="group’">group'</normalizedToken>
|
||
(
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
) and the Philippine '
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus domingo" order="Araneae" pageId="22" pageNumber="73" phylum="Arthropoda" rank="species" species="domingo">Pholcus domingo</taxonomicName>
|
||
<normalizedToken originalValue="group’">group'</normalizedToken>
|
||
(
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 45" title="New leaf- and litter-dwelling species of the genus Pholcus from Southeast Asia (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2016.200" volume="200" year="2016 b">Huber et al. 2016b</bibRefCitation>
|
||
) are both fully supported, as is their sister-group relationship (Figure 10). They are joined in the new genus
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Teranga" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Teranga" order="Araneae" pageId="22" pageNumber="73" phylum="Arthropoda" rank="genus">Teranga</taxonomicName>
|
||
(see Taxonomy section below). Together with
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Tissahamia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tissahamia" order="Araneae" pageId="22" pageNumber="73" phylum="Arthropoda" rank="genus">Tissahamia</taxonomicName>
|
||
(previously the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus ethagala" order="Araneae" pageId="22" pageNumber="73" phylum="Arthropoda" rank="species" species="ethagala">Pholcus ethagala</taxonomicName>
|
||
group) and with
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="22" pageNumber="73" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
|
||
they form a clade that is recovered in all analyses with reasonable support. This clade was supported in almost exactly the same composition by morphological cladistic analysis (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
) except that the '
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="22" pageNumber="73" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
|
||
'
|
||
<taxonomicName genus="Pholcinae" lsidName="Pholcinae nigrifrons" pageId="22" pageNumber="73" rank="species" species="nigrifrons">nigrifrons</taxonomicName>
|
||
<pageBreakToken pageId="23" pageNumber="74" start="start">group</pageBreakToken>
|
||
was also included (the '
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus domingo" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="species" species="domingo">Pholcus domingo</taxonomicName>
|
||
<normalizedToken originalValue="group’">group'</normalizedToken>
|
||
was not yet known in 2011). The clade is supported by the loss of the bulbal uncus and by the loss of distal male cheliceral apophyses [in
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
, the latter character supports a more inclusive taxon (including
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
Simon, 1893) that is strongly rejected by the present molecular data].
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Tissahamia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tissahamia" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Tissahamia</taxonomicName>
|
||
consists of two subgroups, a Sri Lankan subgroup and a Southeast Asian (Malaysian Peninsula, Sumatra) subgroup. The subgroups are consistently recovered in all our analyses with modest to reasonable support, but the monophyly of the entire group is only recovered in the IQ-TREE analysis (low support). Morphological analysis recovered the group, but with varying support depending on weighting regime (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph pageId="23" pageNumber="74">
|
||
Three genera composed of species that were previously assigned to
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
are consistently placed in a highly supported clade together with
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Calapnita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Calapnita" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Calapnita</taxonomicName>
|
||
(Figure 10):
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Paiwana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paiwana" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Paiwana</taxonomicName>
|
||
(previously not assigned to a group),
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Muruta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Muruta" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Muruta</taxonomicName>
|
||
(previously the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus tambunan" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="species" species="tambunan">Pholcus tambunan</taxonomicName>
|
||
group;
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 45" title="New leaf- and litter-dwelling species of the genus Pholcus from Southeast Asia (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2016.200" volume="200" year="2016 b">Huber et al. 2016b</bibRefCitation>
|
||
), and
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Meraha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Meraha" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Meraha</taxonomicName>
|
||
(previously the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus krabi" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="species" species="krabi">Pholcus krabi</taxonomicName>
|
||
group;
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 45" title="New leaf- and litter-dwelling species of the genus Pholcus from Southeast Asia (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2016.200" volume="200" year="2016 b">Huber et al. 2016b</bibRefCitation>
|
||
). We know of no convincing morphological synapomorphy for this group but note two interesting similarities: representatives of
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Calapnita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Calapnita" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Calapnita</taxonomicName>
|
||
and of
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Meraha" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Meraha" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Meraha</taxonomicName>
|
||
share the loss of piriform gland spigots on the anterior lateral spinnerets (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="1 - 63" title="Revision and cladistic analysis of the Southeast Asian leaf-dwelling spider genus Calapnita (Araneae, Pholcidae)." url="https://doi.org/10.11646/zootaxa.4219.1.1" volume="4219" year="2017">2017</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 45" title="New leaf- and litter-dwelling species of the genus Pholcus from Southeast Asia (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2016.200" volume="200" year="2016 b">Huber et al. 2016b</bibRefCitation>
|
||
); representatives of
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Calapnita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Calapnita" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Calapnita</taxonomicName>
|
||
and of
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Muruta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Muruta" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Muruta</taxonomicName>
|
||
and
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Paiwana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paiwana" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Paiwana</taxonomicName>
|
||
share the distinctive shape of the epigynum (roughly triangular, with
|
||
<normalizedToken originalValue="‘knob’">'knob'</normalizedToken>
|
||
directed towards anterior;
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 45" title="New leaf- and litter-dwelling species of the genus Pholcus from Southeast Asia (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2016.200" volume="200" year="2016 b">Huber et al. 2016b</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="1 - 63" title="Revision and cladistic analysis of the Southeast Asian leaf-dwelling spider genus Calapnita (Araneae, Pholcidae)." url="https://doi.org/10.11646/zootaxa.4219.1.1" volume="4219" year="2017">Huber 2017</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Zoologischer Anzeiger" pageId="41" pageNumber="92" pagination="394 - 403" title="Slow genital and genetic but rapid non-genital and ecological differentiation in a pair of spider species (Araneae, Pholcidae)." url="https://doi.org/10.1016/j.jcz.2014.04.001" volume="253" year="2014">Huber and Dimitrov 2014</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph pageId="23" pageNumber="74">
|
||
For two further genera composed of species previously assigned to
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
the present analysis supports the monophyly but gives not clear indication about their closest relatives within the '
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Calapnita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Calapnita" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Calapnita</taxonomicName>
|
||
-
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
|
||
<normalizedToken originalValue="clade’">clade'</normalizedToken>
|
||
(Figure 10):
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pribumia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pribumia" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Pribumia</taxonomicName>
|
||
(previously the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus minang" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="species" species="minang">Pholcus minang</taxonomicName>
|
||
group;
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
) and
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Kintaqa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kintaqa" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Kintaqa</taxonomicName>
|
||
(previously the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus buatong" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="species" species="buatong">Pholcus buatong</taxonomicName>
|
||
group;
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 55" title="The Southeast Asian Pholcushalabala species group (Araneae, Pholcidae), new data from field observations and ultrastructure." url="https://doi.org/10.5852/ejt.2016.190" volume="190" year="2016 a">Huber et al. 2016a</bibRefCitation>
|
||
). All analyses except the IQ-TREE analysis place
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Kintaqa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kintaqa" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Kintaqa</taxonomicName>
|
||
as sister to
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Uthina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Uthina" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Uthina</taxonomicName>
|
||
, but with low support. We know of no potential morphological synapomorphy that links these two groups.
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pribumia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pribumia" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Pribumia</taxonomicName>
|
||
is in our analysis represented by four species. Of these,
|
||
<taxonomicName lsidName="P. diopsis" pageId="23" pageNumber="74" rank="species" species="diopsis">P. diopsis</taxonomicName>
|
||
(Simon, 1901) is never placed within the group; together with
|
||
<taxonomicName lsidName="P. atrigularis" pageId="23" pageNumber="74" rank="species" species="atrigularis">P. atrigularis</taxonomicName>
|
||
(Simon, 1901) it is detected as a rogue taxon and excluded in the RogueNaRok tree. External relationships of
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pribumia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pribumia" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Pribumia</taxonomicName>
|
||
remain dubious. The hypothesis that the genus might be close to
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Tissahamia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tissahamia" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Tissahamia</taxonomicName>
|
||
(previously the '
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus ethagala" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="species" species="ethagala">Pholcus ethagala</taxonomicName>
|
||
<normalizedToken originalValue="group’">group'</normalizedToken>
|
||
;
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
) is supported by numerous distinctive morphological similarities but it is not supported by the present data. However, note that in our analysis
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pribumia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pribumia" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Pribumia</taxonomicName>
|
||
suffers seriously from missing data (we were not able to sequence 28S for any of the four species).
|
||
</paragraph>
|
||
<paragraph lastPageId="24" lastPageNumber="75" pageId="23" pageNumber="74">
|
||
The second major clade within the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
group of genera (Figure 11) is composed of four
|
||
<normalizedToken originalValue="‘old’">'old'</normalizedToken>
|
||
genera (
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
;
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
;
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micromerys" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micromerys" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Micromerys</taxonomicName>
|
||
Bradley, 1877;
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pehrforsskalia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pehrforsskalia" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Pehrforsskalia</taxonomicName>
|
||
Deeleman-Reinhold & van Harten, 2001) and
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Cantikus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cantikus" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Cantikus</taxonomicName>
|
||
(previously the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus halabala" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="species" species="halabala">Pholcus halabala</taxonomicName>
|
||
group;
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 55" title="The Southeast Asian Pholcushalabala species group (Araneae, Pholcidae), new data from field observations and ultrastructure." url="https://doi.org/10.5852/ejt.2016.190" volume="190" year="2016 a">Huber et al. 2016a</bibRefCitation>
|
||
). Except for one clade of Neotropical
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
, all representatives are Old World taxa. We informally call it the '
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="23" pageNumber="74" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
<normalizedToken originalValue="clade’">clade'</normalizedToken>
|
||
. This clade receives full support in all our analyses, and major internal relationships are also well resolved. Three subclades are
|
||
<pageBreakToken pageId="24" pageNumber="75" start="start">fully</pageBreakToken>
|
||
supported each:
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
;
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Cantikus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cantikus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Cantikus</taxonomicName>
|
||
; and a subclade including
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
,
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micromerys" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micromerys" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Micromerys</taxonomicName>
|
||
, and
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pehrforsskalia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pehrforsskalia" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Pehrforsskalia</taxonomicName>
|
||
. All analyses put
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
as sister to
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Cantikus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cantikus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Cantikus</taxonomicName>
|
||
, but with modest support.
|
||
</paragraph>
|
||
<paragraph pageId="24" pageNumber="75">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
is ecologically diverse, including ground-dwelling as well as rock- and leaf-dwelling species, and together with
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
it is also the only genus with autochthonous species in both the New and Old World. Our analysis rejects the previous idea that
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
is
|
||
<normalizedToken originalValue="‘basal’">'basal'</normalizedToken>
|
||
in the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
group of genera (i.e., in a basal trichotomy, with
|
||
<taxonomicName family="Pholcidae" genus="Sihala" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sihala" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Sihala</taxonomicName>
|
||
occupying the second branch and all other taxa the third branch;
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
). Within
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
, our analyses all support a monophyletic New World clade, but with low support values (reasonable support in the 4+ genes analysis). Within the New World clade, a Caribbean clade is fully supported. A remarkable sister-group relationship that is highly supported by the present data is between the Moroccan '
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
'
|
||
<taxonomicName genus="Pholcinae" lsidName="Pholcinae agadir" pageId="24" pageNumber="75" rank="species" species="agadir">agadir</taxonomicName>
|
||
(now transferred to
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
; see Taxonomy section below) and the undescribed Philippine species
|
||
<normalizedToken originalValue="“Phi114”">"Phi114"</normalizedToken>
|
||
. Both have very limited distributions; only one further species of
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
(other than the pantropical
|
||
<taxonomicName lsidName="M. fauroti" pageId="24" pageNumber="75" rank="species" species="fauroti">M. fauroti</taxonomicName>
|
||
) is known from between Morocco and the Philippines:
|
||
<taxonomicName lsidName="M. jacominae" pageId="24" pageNumber="75" rank="species" species="jacominae">M. jacominae</taxonomicName>
|
||
Deeleman-Reinhold & van Harten, 2001 from Yemen. We suspect that
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
in the Old World has a relict distribution, just as it has been hypothesized for South American
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
(
|
||
<bibRefCitation pageId="24" pageNumber="75">Huber et al. 2005a</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Invertebrate Systematics" pageId="42" pageNumber="93" pagination="432 - 450" title="On the New World spiders previously misplaced in Leptopholcus: molecular and morphological analyses and descriptions of four new species (Araneae, Pholcidae)." url="https://doi.org/10.1071/IS13050" volume="28" year="2014">2014</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph lastPageId="25" lastPageNumber="76" pageId="24" pageNumber="75">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Cantikus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cantikus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Cantikus</taxonomicName>
|
||
was recently revised (as '
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="24" pageNumber="75" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
'
|
||
<taxonomicName genus="Pholcinae" lsidName="Pholcinae halabala" pageId="24" pageNumber="75" rank="species" species="halabala">halabala</taxonomicName>
|
||
group;
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 55" title="The Southeast Asian Pholcushalabala species group (Araneae, Pholcidae), new data from field observations and ultrastructure." url="https://doi.org/10.5852/ejt.2016.190" volume="190" year="2016 a">Huber et al. 2016a</bibRefCitation>
|
||
) and divided into a 'core
|
||
<normalizedToken originalValue="group’">group'</normalizedToken>
|
||
that was supported by numerous morphological and behavioral similarities, and a group of species that were assigned to the group tentatively.
|
||
<pageBreakToken pageId="25" pageNumber="76" start="start">This</pageBreakToken>
|
||
tentative assignment was based mainly on preliminary results from the present molecular analysis; a putative morphological synapomorphy for the entire genus
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Cantikus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cantikus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Cantikus</taxonomicName>
|
||
was and is not known. The present analyses fully support both the entire genus and the core group; the genus includes
|
||
<taxonomicName lsidName="C. quinquenotatus" pageId="25" pageNumber="76" rank="species" species="quinquenotatus">C. quinquenotatus</taxonomicName>
|
||
(Thorell, 1878), making the quinquenotatus group proposed in
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber (2011a)</bibRefCitation>
|
||
obsolete; and it highly to fully supports the sister group relationship between the two rock-dwelling species
|
||
<taxonomicName lsidName="C. kuhapimuk" pageId="25" pageNumber="76" rank="species" species="kuhapimuk">C. kuhapimuk</taxonomicName>
|
||
Huber, 2016 and
|
||
<taxonomicName lsidName="C. khaolek" pageId="25" pageNumber="76" rank="species" species="khaolek">C. khaolek</taxonomicName>
|
||
Huber, 2016.
|
||
</paragraph>
|
||
<paragraph pageId="25" pageNumber="76">
|
||
The clade including
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
,
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micromerys" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micromerys" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Micromerys</taxonomicName>
|
||
, and
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pehrforsskalia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pehrforsskalia" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Pehrforsskalia</taxonomicName>
|
||
(Figure 11) was only partly supported in a previous cladistic analysis of morphological data (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
): while
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
and
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micromerys" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micromerys" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Micromerys</taxonomicName>
|
||
were consistently seen as sister taxa (with a mono- or paraphyletic
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
), the position of
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pehrforsskalia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pehrforsskalia" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Pehrforsskalia</taxonomicName>
|
||
varied widely. The characters supporting a close relationship among the three genera are the distal position of the lateral apophyses on the male chelicerae, and the absence of frontal cheliceral apophyses (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
). The present analyses fully support this clade. Within the clade,
|
||
<normalizedToken originalValue="‘basal’">'basal'</normalizedToken>
|
||
relationships are unresolved, essentially resulting in a tetrachotomy: (1) '
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
'
|
||
<taxonomicName genus="Pholcinae" lsidName="Pholcinae podophthalmus" pageId="25" pageNumber="76" rank="species" species="podophthalmus">podophthalmus</taxonomicName>
|
||
(Simon, 1893) is not clearly included in
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
. (2) The Australasian
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micromerys" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micromerys" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Micromerys</taxonomicName>
|
||
receives full support in all analyses. (3) The African
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pehrforsskalia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pehrforsskalia" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Pehrforsskalia</taxonomicName>
|
||
is only represented by its type species. (4)
|
||
<normalizedToken originalValue="‘True’">'True'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
receives reasonable to full support and includes both African and Asian representatives but not the Asian
|
||
<taxonomicName lsidName="L. podophthalmus" pageId="25" pageNumber="76" rank="species" species="podophthalmus">L. podophthalmus</taxonomicName>
|
||
(and its putative close relative
|
||
<taxonomicName lsidName="L. tanikawai" pageId="25" pageNumber="76" rank="species" species="tanikawai">L. tanikawai</taxonomicName>
|
||
Irie, 1999 that is not included in our analyses). Within
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
, our data provide little resolution, but an Asian clade (represented by
|
||
<taxonomicName lsidName="L. borneensis" pageId="25" pageNumber="76" rank="species" species="borneensis">L. borneensis</taxonomicName>
|
||
Deeleman-Reinhold, 1986 and
|
||
<taxonomicName lsidName="L. kandy" pageId="25" pageNumber="76" rank="species" species="kandy">L. kandy</taxonomicName>
|
||
Huber, 2011) receives reasonable support. Among these four clades,
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pehrforsskalia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pehrforsskalia" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Pehrforsskalia</taxonomicName>
|
||
is the only one that does not share the distinctively serrated tip of the male palpal trochanter apophysis (
|
||
<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>
|
||
), suggesting that it may be sister to the other three clades.
|
||
</paragraph>
|
||
<paragraph pageId="25" pageNumber="76">
|
||
The third and last major clade within the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
group of genera is
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
(Figure 12). Support for this group is very low in the IQ-TREE analysis, which reflects the fact that one of the two basal subclades (including the phungiformes and bidentatus groups and
|
||
<taxonomicName lsidName="P. mentawir" pageId="25" pageNumber="76" rank="species" species="mentawir">P. mentawir</taxonomicName>
|
||
Huber, 2011) is closer to the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
-
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
clade than to
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
in some analyses (RAxML, RogueNaRok). By contrast, the 4+ genes analysis recovers the monophyly of
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
with reasonable support, suggesting that the poor support or non-monophyly of
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
in some analyses may result from the many missing data in our full matrix.
|
||
</paragraph>
|
||
<paragraph pageId="25" pageNumber="76">
|
||
Even after removing the eleven species groups that are here placed in the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Calapnita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Calapnita" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Calapnita</taxonomicName>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Panjange" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Panjange" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Panjange</taxonomicName>
|
||
clade and in the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Micropholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Micropholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Micropholcus</taxonomicName>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Leptopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Leptopholcus</taxonomicName>
|
||
clade,
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
continues to be the most species rich genus in
|
||
<taxonomicName family="Pholcidae" lsidName="" pageId="25" pageNumber="76" rank="family">Pholcidae</taxonomicName>
|
||
. It now contains 321 species, most of which are distributed in tropical and subtropical Old World regions. The only exception is the kingi group with ten species in the southeastern USA (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
). Most species of
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="25" pageNumber="76" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
resemble the synanthropic type species
|
||
<taxonomicName lsidName="P. phalangioides" pageId="25" pageNumber="76" rank="species" species="phalangioides">P. phalangioides</taxonomicName>
|
||
in being relatively large, long-legged, brown, and in having a cylindrical abdomen; most or all of these species build their webs in large sheltered spaces. However, the genus is ecologically diverse and includes small litter dwellers with relatively short legs, rock- and ground dwellers with oval abdomens, and pale leaf-dwellers with worm-shaped abdomens.
|
||
</paragraph>
|
||
<paragraph pageId="26" pageNumber="77">
|
||
<pageBreakToken pageId="26" pageNumber="77" start="start">In</pageBreakToken>
|
||
a first effort to structure the known diversity of
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="26" pageNumber="77" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
, the genus was divided into 29 operational species groups (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
), including 25 species groups in the 'core
|
||
<normalizedToken originalValue="group’">group'</normalizedToken>
|
||
, i.e., in
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="26" pageNumber="77" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
. Even though the aim was to identify monophyla, some groups were explicitly proposed as 'waste baskets
|
||
<normalizedToken originalValue="groups’">groups'</normalizedToken>
|
||
(e.g., the bamboutos group) or as "probably not monophyletic" (e.g., the circularis group). The present analysis clarifies a number of relationships, it supports several of the species groups and rejects others, and it confirms the non-monophyly of some groups as suspected. However, we acknowledge that internal relationships in
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="26" pageNumber="77" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
remain highly uncertain and need considerably more work. Our data seem to suffer from two main problems that result in variable topologies among different types of analyses: (1) Even though
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="26" pageNumber="77" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
is in our analyses represented by more species than any other genus (59), our sample is still highly incomplete, including only 18% of the described species and entirely missing seven of the previously suggested species groups (alticeps, nenjukovi, ponticus, zham, yichengicus, taishan, and nagasakiensis groups). (2) The percentage of missing sequences is high in
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="26" pageNumber="77" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
, partly due to the fact that we identified paralogs for 28S and 18S that we excluded, partly due to other unidentified problems.
|
||
</paragraph>
|
||
<paragraph pageId="27" pageNumber="78">
|
||
<pageBreakToken pageId="27" pageNumber="78" start="start">Of</pageBreakToken>
|
||
the 25 operational species groups within
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="27" pageNumber="78" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
proposed previously (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
), ten are supported by the present data: phungiformes group, bidentatus group, calligaster group, Macaronesian group, gracillimus group (excl.
|
||
<taxonomicName lsidName="P. mentawir" pageId="27" pageNumber="78" rank="species" species="mentawir">P. mentawir</taxonomicName>
|
||
),
|
||
<taxonomicName genus="Pholcinae" lsidName="Pholcinae bicornutus" pageId="27" pageNumber="78" rank="species" species="bicornutus">bicornutus</taxonomicName>
|
||
group, chappuisi group, lamperti group, debilis group (incl.
|
||
<taxonomicName lsidName="P. nkoetye" pageId="27" pageNumber="78" rank="species" species="nkoetye">P. nkoetye</taxonomicName>
|
||
Huber, 2011 and
|
||
<taxonomicName lsidName="P. kribi" pageId="27" pageNumber="78" rank="species" species="kribi">P. kribi</taxonomicName>
|
||
Huber, 2011), and guineensis group. Four groups are represented by single species (taarab group, ancoralis group,
|
||
<taxonomicName genus="Pholcinae" lsidName="Pholcinae phalangioides" pageId="27" pageNumber="78" rank="species" species="phalangioides">phalangioides</taxonomicName>
|
||
group, kingi group). Seven groups are missing in the analyses (see above). For the remaining four groups, the present analyses reject the monophyly: (1) The bamboutos group is polyphyletic as expected and the six species in our analyses split into four parts; of these
|
||
<taxonomicName lsidName="P. kribi" pageId="27" pageNumber="78" rank="species" species="kribi">P. kribi</taxonomicName>
|
||
is moved to the debilis group;
|
||
<taxonomicName lsidName="P. bamboutos" pageId="27" pageNumber="78" rank="species" species="bamboutos">P. bamboutos</taxonomicName>
|
||
Huber, 2011 is close to the guineensis group; the affinities of the other four species are unclear. (2) The circularis group is represented by three species; of these,
|
||
<taxonomicName lsidName="P. nkoetye" pageId="27" pageNumber="78" rank="species" species="nkoetye">P. nkoetye</taxonomicName>
|
||
is moved to the debilis group;
|
||
<taxonomicName lsidName="P. leruthi" pageId="27" pageNumber="78" rank="species" species="leruthi">P. leruthi</taxonomicName>
|
||
Lessert, 1935 and
|
||
<taxonomicName lsidName="P. rawiriae" pageId="27" pageNumber="78" rank="species" species="rawiriae">P. rawiriae</taxonomicName>
|
||
Huber, 2014 are sister species and close to the guineensis group. (3/4) The
|
||
<taxonomicName genus="Pholcinae" lsidName="Pholcinae opilionoides" pageId="27" pageNumber="78" rank="species" species="opilionoides">opilionoides</taxonomicName>
|
||
and crypticolens groups are both rejected, but together with the North American kingi group they form a monophylum with reasonable to high support (except for the 4+ genes analysis) but with unknown affinities with other groups.
|
||
</paragraph>
|
||
<paragraph pageId="27" pageNumber="78">
|
||
The present analysis identifies two major clades within
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="27" pageNumber="78" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
that are remarkable even though support values are low to modest. (1) A clade combining the ancoralis, gracillimus, and
|
||
<taxonomicName genus="Pholcinae" lsidName="Pholcinae bicornutus" pageId="27" pageNumber="78" rank="species" species="bicornutus">bicornutus</taxonomicName>
|
||
groups is composed of large dark Southeast Asian and Australasian species; a close relationship between the ancoralis group and the
|
||
<taxonomicName genus="Pholcinae" lsidName="Pholcinae bicornutus" pageId="27" pageNumber="78" rank="species" species="bicornutus">bicornutus</taxonomicName>
|
||
group has been suspected before, based on male ocular area modifications (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
: 314). (2) A large clade including all Subsaharan African taxa. This clade has low bootstrap support but SH values range from 81 to 96, so we consider this a first tentative indication that tropical African
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="27" pageNumber="78" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
might form a large monophylum. The two species that disrupt this picture were both identified as rogue taxa:
|
||
<taxonomicName lsidName="P. taarab" pageId="27" pageNumber="78" rank="species" species="taarab">P. taarab</taxonomicName>
|
||
Huber, 2011 (which is not included in the clade but is African), and
|
||
<taxonomicName lsidName="P. phalangioides" pageId="27" pageNumber="78" rank="species" species="phalangioides">P. phalangioides</taxonomicName>
|
||
(which is included but is most probably not originally African). On the other hand, the inclusion of the Sri Lankan genus
|
||
<taxonomicName family="Pholcidae" genus="Sihala" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sihala" order="Araneae" pageId="27" pageNumber="78" phylum="Arthropoda" rank="genus">Sihala</taxonomicName>
|
||
Huber, 2011 in this clade is plausible, even though weakly supported. Our data highly support the inclusion of
|
||
<taxonomicName family="Pholcidae" genus="Sihala" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sihala" order="Araneae" pageId="27" pageNumber="78" phylum="Arthropoda" rank="genus">Sihala</taxonomicName>
|
||
in
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="27" pageNumber="78" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
, but neither morphology nor molecules seem to give an indication about its sister taxon.
|
||
</paragraph>
|
||
<subSection lastPageId="28" lastPageNumber="79" pageId="27" pageNumber="78" type="notes on genera not included in the present analyses">
|
||
<paragraph pageId="27" pageNumber="78">Notes on genera not included in the present analyses</paragraph>
|
||
<paragraph pageId="27" pageNumber="78">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Aucana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Aucana" order="Araneae" pageId="27" pageNumber="78" phylum="Arthropoda" rank="genus">Aucana</taxonomicName>
|
||
Huber, 2000. This Chilean genus (four species; formally including the mysterious New Caledonian
|
||
<taxonomicName lsidName="A. kaala" pageId="27" pageNumber="78" rank="species" species="kaala">A. kaala</taxonomicName>
|
||
Huber, 2000) was previously thought to be a member of
|
||
<taxonomicName lsidName="" pageId="27" pageNumber="78" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
|
||
(
|
||
<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>
|
||
,
|
||
<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">2011b</bibRefCitation>
|
||
). However, the procursus (dorsal apophysis and corresponding ventral pocket) suggests a placement in
|
||
<taxonomicName lsidName="" pageId="27" pageNumber="78" rank="subFamily" subFamily="Arteminae">Arteminae</taxonomicName>
|
||
. Within
|
||
<taxonomicName lsidName="" pageId="27" pageNumber="78" rank="subFamily" subFamily="Arteminae">Arteminae</taxonomicName>
|
||
, it shares an exposed tarsal organ with
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Chisosa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chisosa" order="Araneae" pageId="27" pageNumber="78" phylum="Arthropoda" rank="genus">Chisosa</taxonomicName>
|
||
and
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Nita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nita" order="Araneae" pageId="27" pageNumber="78" phylum="Arthropoda" rank="genus">Nita</taxonomicName>
|
||
(
|
||
<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>
|
||
,
|
||
<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">2011b</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph pageId="27" pageNumber="78">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Blancoa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Blancoa" order="Araneae" pageId="27" pageNumber="78" phylum="Arthropoda" rank="genus">Blancoa</taxonomicName>
|
||
Huber, 2000. A small Venezuelan genus (two species), probably member of
|
||
<taxonomicName lsidName="" pageId="27" pageNumber="78" rank="subFamily" subFamily="Modisiminae">Modisiminae</taxonomicName>
|
||
(
|
||
<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>
|
||
), but the sister group remains entirely obscure.
|
||
</paragraph>
|
||
<paragraph lastPageId="28" lastPageNumber="79" pageId="27" pageNumber="78">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Canaima" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Canaima" order="Araneae" pageId="27" pageNumber="78" phylum="Arthropoda" rank="genus">Canaima</taxonomicName>
|
||
Huber, 2000. Also probably member of
|
||
<taxonomicName lsidName="" pageId="27" pageNumber="78" rank="subFamily" subFamily="Modisiminae">Modisiminae</taxonomicName>
|
||
, with only two species restricted to Trinidad and Venezuela (
|
||
<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>
|
||
). The shape of the ventral apo
|
||
<pageBreakToken pageId="28" pageNumber="79" start="start">physis</pageBreakToken>
|
||
on the male palpal femur is reminiscent of the Venezuelan clade including
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Mecolaesthus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Mecolaesthus" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Mecolaesthus</taxonomicName>
|
||
,
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Stenosfemuraia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Stenosfemuraia" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Stenosfemuraia</taxonomicName>
|
||
,
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Systenita" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Systenita" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Systenita</taxonomicName>
|
||
, and
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Coryssocnemis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Coryssocnemis" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Coryssocnemis</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph pageId="28" pageNumber="79">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Cenemus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cenemus" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Cenemus</taxonomicName>
|
||
Saaristo, 2001. A small Seychellois genus (three species), member of
|
||
<taxonomicName lsidName="" pageId="28" pageNumber="79" rank="subFamily" subFamily="Smeringopinae">Smeringopinae</taxonomicName>
|
||
; a morphological cladistic analysis (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="1 - 138" title="Revision and cladistic analysis of the Afrotropical endemic genus Smeringopus Simon, 1890 (Araneae: Pholcidae)." volume="3461" year="2012">Huber 2012</bibRefCitation>
|
||
) suggested a placement in the 'northern
|
||
<normalizedToken originalValue="clade’">clade'</normalizedToken>
|
||
of
|
||
<taxonomicName lsidName="" pageId="28" pageNumber="79" rank="subFamily" subFamily="Smeringopinae">Smeringopinae</taxonomicName>
|
||
even though the Seychelles are geographically much closer to the 'southern
|
||
<normalizedToken originalValue="clade’">clade'</normalizedToken>
|
||
.
|
||
</paragraph>
|
||
<paragraph pageId="28" pageNumber="79">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Enetea" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Enetea" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Enetea</taxonomicName>
|
||
Huber, 2000. A monotypic Bolivian genus, member of
|
||
<taxonomicName lsidName="" pageId="28" pageNumber="79" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
|
||
(
|
||
<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>
|
||
); the sister group remains entirely obscure.
|
||
</paragraph>
|
||
<paragraph pageId="28" pageNumber="79">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Galapa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Galapa" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Galapa</taxonomicName>
|
||
Huber, 2000. A small genus (two species) restricted to the Galapagos Islands, member of
|
||
<taxonomicName lsidName="" pageId="28" pageNumber="79" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
|
||
(
|
||
<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>
|
||
); the sister group remains entirely obscure.
|
||
</paragraph>
|
||
<paragraph pageId="28" pageNumber="79">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Ossinissa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ossinissa" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Ossinissa</taxonomicName>
|
||
Dimitrov & Ribera, 2005. A monotypic genus from the Canary Islands, member of the
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
group of genera (
|
||
<bibRefCitation author="Huber, BA" journalOrPublisher="Bonner zoologische Monographien" pageId="40" pageNumber="91" pagination="1 - 509" title="Revision and cladistic analysis of Pholcus and closely related taxa (Araneae, Pholcidae)." volume="58" year="2011 a">Huber 2011a</bibRefCitation>
|
||
); the sister group is dubious, but we suspect a close relationship with other Canary Island cavernicole species in
|
||
<normalizedToken originalValue="‘true’">'true'</normalizedToken>
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcus" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Pholcus</taxonomicName>
|
||
(
|
||
<taxonomicName lsidName="P. baldiosensis" pageId="28" pageNumber="79" rank="species" species="baldiosensis">P. baldiosensis</taxonomicName>
|
||
Wunderlich, 1992;
|
||
<taxonomicName lsidName="P. corniger" pageId="28" pageNumber="79" rank="species" species="corniger">P. corniger</taxonomicName>
|
||
Dimitrov & Ribera, 2006).
|
||
</paragraph>
|
||
<paragraph pageId="28" pageNumber="79">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pomboa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pomboa" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Pomboa</taxonomicName>
|
||
Huber, 2000. Member of
|
||
<taxonomicName lsidName="" pageId="28" pageNumber="79" rank="subFamily" subFamily="Modisiminae">Modisiminae</taxonomicName>
|
||
, with currently four species restricted to Colombia. The vertical hairs in high density on the leg tibiae suggest an affinity to
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pisaboa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pisaboa" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Pisaboa</taxonomicName>
|
||
and
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Waunana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Waunana" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Waunana</taxonomicName>
|
||
(
|
||
<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>
|
||
).
|
||
</paragraph>
|
||
<paragraph pageId="28" pageNumber="79">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Queliceria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Queliceria" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Queliceria</taxonomicName>
|
||
<normalizedToken originalValue="González-Sponga">Gonzalez-Sponga</normalizedToken>
|
||
, 2003. A monotypic Venezuelan genus, probably member of
|
||
<taxonomicName lsidName="" pageId="28" pageNumber="79" rank="subFamily" subFamily="Modisiminae">Modisiminae</taxonomicName>
|
||
; the sister group remains entirely obscure.
|
||
</paragraph>
|
||
<paragraph pageId="28" pageNumber="79">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Tibetia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tibetia" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Tibetia</taxonomicName>
|
||
Zhang, Zhu & Song, 2006. A monotypic Chinese (Tibetan) genus, probably member of
|
||
<taxonomicName lsidName="" pageId="28" pageNumber="79" rank="subFamily" subFamily="Arteminae">Arteminae</taxonomicName>
|
||
; the sister group remains entirely obscure.
|
||
</paragraph>
|
||
<paragraph pageId="28" pageNumber="79">
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Tolteca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tolteca" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Tolteca</taxonomicName>
|
||
Huber, 2000. A small Mexican genus (two species), member of
|
||
<taxonomicName lsidName="" pageId="28" pageNumber="79" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
|
||
. We predict that
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Tolteca" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tolteca" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Tolteca</taxonomicName>
|
||
is member of the North and Central American & Caribbean clade (Figure 2), together with
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcophora" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Pholcophora</taxonomicName>
|
||
and
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Papiamenta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Papiamenta" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Papiamenta</taxonomicName>
|
||
. The frontal humps on the male sternum and the shape of the procursus are reminiscent of
|
||
<taxonomicName class="Arachnida" family="Pholcidae" genus="Pholcophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pholcophora" order="Araneae" pageId="28" pageNumber="79" phylum="Arthropoda" rank="genus">Pholcophora</taxonomicName>
|
||
(
|
||
<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>
|
||
).
|
||
</paragraph>
|
||
</subSection>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |