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<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 &amp; Dimitrov, Dimitar" docDate="2018" docId="0F52150CBF7AE849A8B1F3E398408FE1" docLanguage="en" docName="ZooKeys 789: 51-101" docOrigin="ZooKeys 789" docSource="http://dx.doi.org/10.3897/zookeys.789.22781" docTitle="Pholcinae group subsp. 1 '" docType="treatment" docVersion="6" lastPageNumber="67" masterDocId="FF94FFCCE316FFFCFFBAFFD4FFECFFF7" masterDocTitle="The phylogeny of pholcid spiders: a critical evaluation of relationships suggested by molecular data (Araneae, Pholcidae)" masterLastPageNumber="101" masterPageNumber="51" pageNumber="64" updateTime="1668166286259" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>The phylogeny of pholcid spiders: a critical evaluation of relationships suggested by molecular data (Araneae, Pholcidae)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Huber, Bernhard A.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Eberle, Jonas</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Dimitrov, Dimitar</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2018</mods:date>
<mods:detail type="volume">
<mods:number>789</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>51</mods:start>
<mods:end>101</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.789.22781</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.789.22781</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-789-51</mods:identifier>
<mods:identifier type="ZBK">496949FCA96A4489A0940182520DAB6C</mods:identifier>
<mods:identifier type="ZooBank">496949FCA96A4489A0940182520DAB6C</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="148671578" LSID="urn:lsid:plazi:treatment:0F52150CBF7AE849A8B1F3E398408FE1" httpUri="http://treatment.plazi.org/id/0F52150CBF7AE849A8B1F3E398408FE1" lastPageId="16" lastPageNumber="67" pageId="13" pageNumber="64">
<subSubSection pageId="13" pageNumber="64" type="nomenclature">
<paragraph pageId="13" pageNumber="64">
<taxonomicName family="Pholcidae" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Pholcinae group subsp. 1" order="Araneae" pageId="13" pageNumber="64" rank="subFamily" subFamily="Pholcinae 'group 1'">
<pageBreakToken pageId="13" pageNumber="64" start="start">Pholcinae</pageBreakToken>
'group 1'
</taxonomicName>
Figs 7, 8
</paragraph>
</subSubSection>
<subSubSection lastPageId="16" lastPageNumber="67" pageId="14" pageNumber="65" type="remarks">
<paragraph pageId="14" pageNumber="65">
<pageBreakToken pageId="14" pageNumber="65" start="start">Remarks</pageBreakToken>
.
</paragraph>
<paragraph pageId="14" pageNumber="65">
This group includes some genera named long ago, like
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
,
<taxonomicName class="Arachnida" family="Pholcidae" genus="Belisana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Belisana" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Belisana</taxonomicName>
, and
<taxonomicName class="Arachnida" family="Pholcidae" genus="Paramicromerys" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paramicromerys" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Paramicromerys</taxonomicName>
Millot, 1946. Most other genera were described relatively recently and resulted either from splitting of
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
(e.g.,
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophorides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophorides" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Spermophorides</taxonomicName>
Wunderlich, 1992;
<taxonomicName class="Arachnida" family="Pholcidae" genus="Buitinga" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Buitinga" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Buitinga</taxonomicName>
Huber, 2003;
<taxonomicName class="Arachnida" family="Pholcidae" genus="Savarna" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Savarna" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Savarna</taxonomicName>
Huber, 2005;
<taxonomicName class="Arachnida" family="Pholcidae" genus="Khorata" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Khorata" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Khorata</taxonomicName>
Huber, 2005) or from the discovery and description of new species (
<taxonomicName class="Arachnida" family="Pholcidae" genus="Aetana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Aetana" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Aetana</taxonomicName>
Huber, 2005;
<taxonomicName class="Arachnida" family="Pholcidae" genus="Wanniyala" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Wanniyala" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Wanniyala</taxonomicName>
Huber &amp; Benjamin, 2005;
<taxonomicName class="Arachnida" family="Pholcidae" genus="Hantu" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hantu" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Hantu</taxonomicName>
Huber, 2016).
</paragraph>
<paragraph pageId="14" pageNumber="65">
A Southeast Asian clade that is consistently resolved with high to full support but variably placed either inside 'group 1' (IQ-TREE, RogueNaRok) or outside of the three operational groups as an isolated fourth group (4+ genes, RAxML) is composed of
<taxonomicName class="Arachnida" family="Pholcidae" genus="Aetana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Aetana" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Aetana</taxonomicName>
, Southeast Asian '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
', and an undescribed new genus from Indonesia (
<normalizedToken originalValue="“Ind206”">&quot;Ind206&quot;</normalizedToken>
). Morphological data have suggested a close relationship of
<taxonomicName class="Arachnida" family="Pholcidae" genus="Aetana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Aetana" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Aetana</taxonomicName>
with
<taxonomicName class="Arachnida" family="Pholcidae" genus="Savarna" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Savarna" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Savarna</taxonomicName>
,
<taxonomicName class="Arachnida" family="Pholcidae" genus="Khorata" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Khorata" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Khorata</taxonomicName>
, and
<taxonomicName class="Arachnida" family="Pholcidae" genus="Hantu" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hantu" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Hantu</taxonomicName>
(
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 78" title="Revision, phylogeny, and microhabitat shifts in the Southeast Asian spider genus Aetana (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2015.162" volume="162" year="2015">Huber et al. 2015</bibRefCitation>
). The positions of those three genera in our molecular trees are all unstable and problematic (see below). Thus, we consider it premature to conclude that the morphological data were misleading, and suggest that the positions of
<taxonomicName class="Arachnida" family="Pholcidae" genus="Savarna" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Savarna" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Savarna</taxonomicName>
,
<taxonomicName class="Arachnida" family="Pholcidae" genus="Khorata" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Khorata" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Khorata</taxonomicName>
, and
<taxonomicName class="Arachnida" family="Pholcidae" genus="Hantu" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hantu" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Hantu</taxonomicName>
need further analysis. A similar problem occurs with Southeast Asian '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
'. The monophyly of the five species included receives reasonable to high support, but this group does not seem to be close to the type species
<taxonomicName lsidName="S. senoculata." pageId="14" pageNumber="65" rank="species" species="senoculata.">S. senoculata.</taxonomicName>
However, the position of
<taxonomicName lsidName="S. senoculata" pageId="14" pageNumber="65" rank="species" species="senoculata">S. senoculata</taxonomicName>
varies strongly among analyses, and the idea that Southeast Asian taxa are in fact congeneric with
<taxonomicName lsidName="S. senoculata" pageId="14" pageNumber="65" rank="species" species="senoculata">S. senoculata</taxonomicName>
(
<bibRefCitation pageId="14" pageNumber="65">Huber 2005a</bibRefCitation>
) should not yet be discarded based on the present molecular data.
</paragraph>
<paragraph pageId="14" pageNumber="65">
In
<taxonomicName class="Arachnida" family="Pholcidae" genus="Aetana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Aetana" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Aetana</taxonomicName>
, our analyses include 16 of 18 (89%) described species plus two undescribed species. The monophyly of the genus is highly to fully supported even though morphological support appeared weak (
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 78" title="Revision, phylogeny, and microhabitat shifts in the Southeast Asian spider genus Aetana (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2015.162" volume="162" year="2015">Huber et al. 2015</bibRefCitation>
). All four species groups proposed after cladistic analysis of morphological characters (
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 78" title="Revision, phylogeny, and microhabitat shifts in the Southeast Asian spider genus Aetana (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2015.162" volume="162" year="2015">Huber et al. 2015</bibRefCitation>
) are supported, but with different relationships among each other. Most of these relationships among species groups receive low support, but the kiukoki group is resolved as sister of the omayan group (with modest support) and this is in conflict with the results from morphology (
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 78" title="Revision, phylogeny, and microhabitat shifts in the Southeast Asian spider genus Aetana (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2015.162" volume="162" year="2015">Huber et al. 2015</bibRefCitation>
). The two unnamed subgroups within the
<taxonomicName genus="Pholcinae" lsidName="Pholcinae kinabalu" pageId="14" pageNumber="65" rank="species" species="kinabalu">kinabalu</taxonomicName>
group and within the omayan group, respectively, proposed in
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 78" title="Revision, phylogeny, and microhabitat shifts in the Southeast Asian spider genus Aetana (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2015.162" volume="162" year="2015">Huber et al. (2015)</bibRefCitation>
are all recovered (with modest to full support).
</paragraph>
<paragraph lastPageId="15" lastPageNumber="66" pageId="14" pageNumber="65">
The next clade within 'group 1' (Figure 7) includes three taxa whose position varies strongly among different analyses (see above): the type species of
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
,
<taxonomicName lsidName="S. senoculata" pageId="14" pageNumber="65" rank="species" species="senoculata">S. senoculata</taxonomicName>
, and the Southeast Asian genera
<taxonomicName class="Arachnida" family="Pholcidae" genus="Khorata" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Khorata" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Khorata</taxonomicName>
and
<taxonomicName class="Arachnida" family="Pholcidae" genus="Savarna" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Savarna" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Savarna</taxonomicName>
.
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora senoculata" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="species" species="senoculata">Spermophora senoculata</taxonomicName>
is alternatively resolved as sister to the African '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
'
<taxonomicName genus="Pholcinae" lsidName="Pholcinae akwamu" pageId="14" pageNumber="65" rank="species" species="akwamu">akwamu</taxonomicName>
group (RAxML) or to the African '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
'
<taxonomicName genus="Pholcinae" lsidName="Pholcinae kyambura" pageId="14" pageNumber="65" rank="species" species="kyambura">kyambura</taxonomicName>
Huber &amp; Warui, 2012 (4+ genes, RogueNaRok). Its sister group is essentially unknown. As indicated above, a close relationship with Southeast Asian '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
', even though never recovered by our analyses, should not be definitely discarded.
<taxonomicName class="Arachnida" family="Pholcidae" genus="Khorata" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Khorata" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Khorata</taxonomicName>
and
<taxonomicName class="Arachnida" family="Pholcidae" genus="Savarna" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Savarna" order="Araneae" pageId="14" pageNumber="65" phylum="Arthropoda" rank="genus">Savarna</taxonomicName>
are sister taxa in some analyses (low support; IQ-TREE, RogueNaRok), but wide apart in
<pageBreakToken pageId="15" pageNumber="66" start="start">others</pageBreakToken>
. The former result is considered more plausible for two reasons: (1) morphology supports a close relationship between
<taxonomicName class="Arachnida" family="Pholcidae" genus="Khorata" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Khorata" order="Araneae" pageId="15" pageNumber="66" phylum="Arthropoda" rank="genus">Khorata</taxonomicName>
and
<taxonomicName class="Arachnida" family="Pholcidae" genus="Savarna" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Savarna" order="Araneae" pageId="15" pageNumber="66" phylum="Arthropoda" rank="genus">Savarna</taxonomicName>
(
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 78" title="Revision, phylogeny, and microhabitat shifts in the Southeast Asian spider genus Aetana (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2015.162" volume="162" year="2015">Huber et al. 2015</bibRefCitation>
); (2) the alternative topology (4+ genes, RAxML) places the Southeast Asian
<taxonomicName class="Arachnida" family="Pholcidae" genus="Savarna" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Savarna" order="Araneae" pageId="15" pageNumber="66" phylum="Arthropoda" rank="genus">Savarna</taxonomicName>
as sister to an East African clade.
</paragraph>
<paragraph lastPageId="16" lastPageNumber="67" pageId="15" pageNumber="66">
The large Asian genus
<taxonomicName class="Arachnida" family="Pholcidae" genus="Belisana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Belisana" order="Araneae" pageId="15" pageNumber="66" phylum="Arthropoda" rank="genus">Belisana</taxonomicName>
(Figure 8) is well represented in our analyses (30 species) but seems to suffer from rogue taxa, paralogs, and/or other unidentified problems. Only the RogueNaRok tree resolves a monophyletic
<taxonomicName class="Arachnida" family="Pholcidae" genus="Belisana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Belisana" order="Araneae" pageId="15" pageNumber="66" phylum="Arthropoda" rank="genus">Belisana</taxonomicName>
. In other analysis, either
<taxonomicName class="Arachnida" family="Pholcidae" genus="Hantu" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hantu" order="Araneae" pageId="15" pageNumber="66" phylum="Arthropoda" rank="genus">Hantu</taxonomicName>
(RAxML) or
<taxonomicName class="Arachnida" family="Pholcidae" genus="Hantu" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hantu" order="Araneae" pageId="15" pageNumber="66" phylum="Arthropoda" rank="genus">Hantu</taxonomicName>
and '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="15" pageNumber="66" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
'
<taxonomicName genus="Pholcinae" lsidName="Pholcinae kyambura" pageId="15" pageNumber="66" rank="species" species="kyambura">kyambura</taxonomicName>
are nested within
<taxonomicName class="Arachnida" family="Pholcidae" genus="Belisana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Belisana" order="Araneae" pageId="15" pageNumber="66" phylum="Arthropoda" rank="genus">Belisana</taxonomicName>
(IQ-TREE). A close relationship between
<taxonomicName class="Arachnida" family="Pholcidae" genus="Belisana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Belisana" order="Araneae" pageId="15" pageNumber="66" phylum="Arthropoda" rank="genus">Belisana</taxonomicName>
and
<taxonomicName class="Arachnida" family="Pholcidae" genus="Hantu" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hantu" order="Araneae" pageId="15" pageNumber="66" phylum="Arthropoda" rank="genus">Hantu</taxonomicName>
(that is also suggested in the RogueNaRok tree) is strongly contradicted by morphology: several
<pageBreakToken pageId="16" pageNumber="67" start="start">characters</pageBreakToken>
support a close relationship between
<taxonomicName class="Arachnida" family="Pholcidae" genus="Hantu" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hantu" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Hantu</taxonomicName>
,
<taxonomicName class="Arachnida" family="Pholcidae" genus="Khorata" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Khorata" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Khorata</taxonomicName>
, and
<taxonomicName class="Arachnida" family="Pholcidae" genus="Savarna" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Savarna" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Savarna</taxonomicName>
(
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="42" pageNumber="93" pagination="1 - 78" title="Revision, phylogeny, and microhabitat shifts in the Southeast Asian spider genus Aetana (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2015.162" volume="162" year="2015">Huber et al. 2015</bibRefCitation>
). We have no explanation for the position of
<taxonomicName class="Arachnida" family="Pholcidae" genus="Hantu" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hantu" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Hantu</taxonomicName>
in our trees. Intriguingly,
<taxonomicName lsidName="H. niah" pageId="16" pageNumber="67" rank="species" species="niah">H. niah</taxonomicName>
Huber, 2016 (but not
<taxonomicName lsidName="H. kapit" pageId="16" pageNumber="67" rank="species" species="kapit">H. kapit</taxonomicName>
Huber, 2016) was placed in a clade together with
<taxonomicName class="Arachnida" family="Pholcidae" genus="Khorata" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Khorata" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Khorata</taxonomicName>
and
<taxonomicName class="Arachnida" family="Pholcidae" genus="Savarna" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Savarna" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Savarna</taxonomicName>
in preliminary analyses of the present data. On the other hand, the African '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
'
<taxonomicName genus="Pholcinae" lsidName="Pholcinae kyambura" pageId="16" pageNumber="67" rank="species" species="kyambura">kyambura</taxonomicName>
might indeed be close to
<taxonomicName class="Arachnida" family="Pholcidae" genus="Belisana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Belisana" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Belisana</taxonomicName>
. In fact, had it been collected in Southeast Asia, it would probably have been assigned to
<taxonomicName class="Arachnida" family="Pholcidae" genus="Belisana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Belisana" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Belisana</taxonomicName>
. It was tentatively assigned to
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
because African '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
' were polyphyletic anyway and because the closest known record of
<taxonomicName class="Arachnida" family="Pholcidae" genus="Belisana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Belisana" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Belisana</taxonomicName>
was from India, more than 5000 km east. However, the position of '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
'
<taxonomicName genus="Pholcinae" lsidName="Pholcinae kyambura" pageId="16" pageNumber="67" rank="species" species="kyambura">kyambura</taxonomicName>
varies among analyses and should be considered unresolved.
</paragraph>
<paragraph pageId="16" pageNumber="67">
Our sample of
<taxonomicName class="Arachnida" family="Pholcidae" genus="Belisana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Belisana" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Belisana</taxonomicName>
includes numerous representatives from different microhabitats (litter and leaves) and with different types of webs (
<normalizedToken originalValue="‘usual’">'usual'</normalizedToken>
pholcid domed sheets and highly regular
<normalizedToken originalValue="‘curtain’">'curtain'</normalizedToken>
webs; Figs 8
<normalizedToken originalValue="e–f">e-f</normalizedToken>
; see also
<bibRefCitation pageId="16" pageNumber="67">Deeleman-Reinhold 1986a</bibRefCitation>
,
<bibRefCitation author="Huber, BA" journalOrPublisher="Zoologica" pageId="40" pageNumber="91" pagination="1 - 126" title="High species diversity, male-female coevolution, and metaphyly in Southeast Asian pholcid spiders: the case of Belisana Thorell 1898 (Araneae, Pholcidae)." volume="155" year="2005 b">Huber 2005b</bibRefCitation>
). The present data suggest multiple microhabitat shifts within
<taxonomicName class="Arachnida" family="Pholcidae" genus="Belisana" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Belisana" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Belisana</taxonomicName>
, but note that many nodes within the genus have very low support values. These low values also impede a proper interpretation of the fact that the two species with a
<normalizedToken originalValue="‘usual’">'usual'</normalizedToken>
domed web (marked with D in Figure 8) included in the analyses (
<taxonomicName genus="B." lsidName="B." pageId="16" pageNumber="67" rank="genus">B.</taxonomicName>
<normalizedToken originalValue="“Mal77”">&quot;Mal77&quot;</normalizedToken>
,
<taxonomicName lsidName="B. tambligan" pageId="16" pageNumber="67" rank="species" species="tambligan">B. tambligan</taxonomicName>
Huber, 2005) are not
<normalizedToken originalValue="‘basal’">'basal'</normalizedToken>
but nested among species with highly regular
<normalizedToken originalValue="‘curtain’">'curtain'</normalizedToken>
webs (marked with R in Figure 8) [confirmed for
<taxonomicName lsidName="B. bohorok" pageId="16" pageNumber="67" rank="species" species="bohorok">B. bohorok</taxonomicName>
Huber, 2005;
<taxonomicName lsidName="B. leuser" pageId="16" pageNumber="67" rank="species" species="leuser">B. leuser</taxonomicName>
Huber, 2005;
<taxonomicName genus="B." lsidName="B." pageId="16" pageNumber="67" rank="genus">B.</taxonomicName>
<normalizedToken originalValue="“Bor85”">&quot;Bor85&quot;</normalizedToken>
;
<taxonomicName lsidName="B. junkoae" pageId="16" pageNumber="67" rank="species" species="junkoae">B. junkoae</taxonomicName>
(Irie, 1997);
<taxonomicName lsidName="B. sabah" pageId="16" pageNumber="67" rank="species" species="sabah">B. sabah</taxonomicName>
Huber, 2005; BA Huber, unpubl. data].
</paragraph>
<paragraph pageId="16" pageNumber="67">
Except for the Sri Lankan genus
<taxonomicName class="Arachnida" family="Pholcidae" genus="Wanniyala" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Wanniyala" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Wanniyala</taxonomicName>
, all remaining taxa of
<taxonomicName lsidName="" pageId="16" pageNumber="67" rank="subFamily" subFamily="Pholcinae">Pholcinae</taxonomicName>
'group 1' (Figure 7) are African, Madagascan, and Mediterranean. They are grouped together but with very low support. South African and Madagascan '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
' were not available for sequencing and are thus not included in our analyses; we predict they are members of this clade. As mentioned above, some analyses (RAxML, 4+ genes) placed the East Asian genus
<taxonomicName class="Arachnida" family="Pholcidae" genus="Savarna" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Savarna" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Savarna</taxonomicName>
within this clade; we consider this topology dubious.
</paragraph>
<paragraph pageId="16" pageNumber="67">
A close relationship between the West African '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
'
<taxonomicName genus="Pholcinae" lsidName="Pholcinae tonkoui" pageId="16" pageNumber="67" rank="species" species="tonkoui">tonkoui</taxonomicName>
group and
<taxonomicName class="Arachnida" family="Pholcidae" genus="Wanniyala" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Wanniyala" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Wanniyala</taxonomicName>
is suggested in all our analyses, even though with low support (only SH values are consistently at 96-97). This relationship is also supported by morphology: the two taxa share a hinged procursus with a membranous process arising from the proximal part (see
<bibRefCitation author="Huber, BA" journalOrPublisher="Journal of Natural History" pageId="41" pageNumber="92" pagination="3305 - 3319" title="The pholcid spiders from Sri Lanka: redescription of Pholcusceylonicus and description of Wanniyala new genus (Araneae: Pholcidae)." url="https://doi.org/10.1080/00222930500145123" volume="39" year="2005">Huber and Benjamin 2005</bibRefCitation>
: fig. 7,
<bibRefCitation author="Huber, BA" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="40" pageNumber="91" pagination="555 - 619" title="High species diversity in one of the dominant groups of spiders in East African montane forests (Araneae: Pholcidae: Buitinga n. gen., Spermophora Hentz)." url="https://doi.org/10.1046/j.1096-3642.2003.00053.x" volume="137" year="2003 b">Huber 2003b</bibRefCitation>
: fig. 293,
<bibRefCitation author="Huber, BA" journalOrPublisher="European Journal of Taxonomy" pageId="41" pageNumber="92" pagination="1 - 44" title="West African pholcid spiders: an overview, with descriptions of five new species (Araneae, Pholcidae)." url="https://doi.org/10.5852/ejt.2013.59" volume="59" year="2013">Huber and Kwapong 2013</bibRefCitation>
: fig. 101).
</paragraph>
<paragraph pageId="16" pageNumber="67">
The following clade (Figure 7) places the Central African '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
'
<taxonomicName genus="Pholcinae" lsidName="Pholcinae awalai" pageId="16" pageNumber="67" rank="species" species="awalai">awalai</taxonomicName>
group as sister to the Macaronesian and Mediterranean genus
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophorides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophorides" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Spermophorides</taxonomicName>
, both together sister to the Madagascan genus
<taxonomicName class="Arachnida" family="Pholcidae" genus="Paramicromerys" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paramicromerys" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Paramicromerys</taxonomicName>
, and all together sister to an undescribed Madagascan genus (
<normalizedToken originalValue="“CAS13”">&quot;CAS13&quot;</normalizedToken>
). Support for these relationships is modest, and the clade is different in composition in the 4+ genes tree (
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophorides" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophorides" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Spermophorides</taxonomicName>
is missing from this analysis).
</paragraph>
<paragraph pageId="16" pageNumber="67">
The last clade in
<taxonomicName lsidName="" pageId="16" pageNumber="67" rank="subFamily" subFamily="Pholcinae">Pholcinae</taxonomicName>
'group 1' is highly to fully supported in all analyses and includes the East African genus
<taxonomicName class="Arachnida" family="Pholcidae" genus="Buitinga" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Buitinga" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Buitinga</taxonomicName>
and East African '
<taxonomicName class="Arachnida" family="Pholcidae" genus="Spermophora" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Spermophora" order="Araneae" pageId="16" pageNumber="67" phylum="Arthropoda" rank="genus">Spermophora</taxonomicName>
', each with full support in all analyses. The sister group relationship between these two taxa makes sense geographically but is not evident from morphology.
</paragraph>
</subSubSection>
</treatment>
</document>
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