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<mods:title id="463ACE463954675C6586C55556919496">The phylogeny of pholcid spiders: a critical evaluation of relationships suggested by molecular data (Araneae, Pholcidae)</mods:title>
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<treatment id="2C6A08DB4D12751A58C844D35E7C10F5" ID-GBIF-Taxon="148671575" LSID="urn:lsid:plazi:treatment:2C6A08DB4D12751A58C844D35E7C10F5" httpUri="http://treatment.plazi.org/id/2C6A08DB4D12751A58C844D35E7C10F5" lastPageId="12" lastPageNumber="63" pageId="9" pageNumber="60" scope_family="Pholcidae" scope_order="Araneae">
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<paragraph id="E3CB491F65D6CA2B6C944B03CF77636D" pageId="9" pageNumber="60">
Subfamily
<taxonomicName id="C66548ABA39B23A654DDF5FC615DD4D1" authority="Simon, 1893" authorityName="Simon" authorityYear="1893" class="Arachnida" family="Pholcidae" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="" order="Araneae" pageId="9" pageNumber="60" phylum="Arthropoda" rank="subFamily" subFamily="Smeringopinae">Smeringopinae Simon, 1893</taxonomicName>
Figure 6
</paragraph>
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<paragraph id="16F37E53AA31D704A474D24B5F215ABC" pageId="10" pageNumber="61">
<taxonomicName id="EF82E63D93E9BC2FE05C82A64448E4BD" class="Arachnida" family="Smeringopodeae" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Modisimeae" order="Araneae" pageId="10" pageNumber="61" phylum="Arthropoda" rank="family">
<pageBreakToken id="D83DFDEE41EB8FC5ECA83BF74046CD51" pageId="10" pageNumber="61" start="start">Smeringopodeae</pageBreakToken>
</taxonomicName>
Simon, 1893: 474. Type genus
<taxonomicName id="974D2E71F6A576A26EC35C88CD76F9F9" class="Arachnida" family="Pholcidae" genus="Smeringopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopus" order="Araneae" pageId="10" pageNumber="61" phylum="Arthropoda" rank="genus">Smeringopus</taxonomicName>
Simon, 1890, by subsequent designation (
<bibRefCitation id="8EF831E545BC4E47B69CF389A6F294FC" 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>
).
</paragraph>
<paragraph id="1F0FE0ACFDF14D6E2F4B8A78D1D4306A" pageId="10" pageNumber="61">
<taxonomicName id="7B372341292BFCC2D42548433281D91F" class="Arachnida" family="Pholcidae" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="" order="Araneae" pageId="10" pageNumber="61" phylum="Arthropoda" rank="subFamily" subFamily="Smeringopinae">Smeringopinae</taxonomicName>
Simon;
<bibRefCitation id="69823C4780431BF5CC84E4620BA89A00" 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>
: 217.
</paragraph>
</subSubSection>
<subSubSection id="44D4C455DA911464A732D622F0E7D9CD" lastPageId="12" lastPageNumber="63" pageId="10" pageNumber="61" type="remarks">
<paragraph id="7ACCECDF47E6B1948B0698A32E53B869" pageId="10" pageNumber="61">Remarks.</paragraph>
<paragraph id="BDE91078EDF9A5667D25885B9AEEE538" pageId="10" pageNumber="61">
<taxonomicName id="F70842AD1CB436611BE37DB1D44116C3" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Smeringopinae">Smeringopinae</taxonomicName>
is a relatively homogeneous subfamily (with respect to body shapes, colour, webs, and microhabitats), and in this sense similar to
<taxonomicName id="EDB725AD1A38C25F140B69FC4F685294" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
and
<taxonomicName id="E372E9CE970DDE14A5F05C966165C2F7" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Arteminae">Arteminae</taxonomicName>
but very unlike
<taxonomicName id="0C72B58B3E0B78BCE34E5D0A8FE53C17" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Modisiminae">Modisiminae</taxonomicName>
and
<taxonomicName id="941C3D840470838BABF85039F60A17AA" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Pholcinae">Pholcinae</taxonomicName>
. Most of the 125 known species of
<taxonomicName id="7187551B7D09B1DDC9A0423112DD0267" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Smeringopinae">Smeringopinae</taxonomicName>
are medium-size to large, have long legs, elongated to cylindrical abdomens, and all have eight eyes. Another similarity to
<taxonomicName id="490DD40B0860DE78B8A1A50A14AF0E87" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
and
<taxonomicName id="DACF510261E195C59DCA04BF218C8B37" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Arteminae">Arteminae</taxonomicName>
is that
<taxonomicName id="7E40313E0A064505E928B0061C9F2118" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Smeringopinae">Smeringopinae</taxonomicName>
are often found in rather arid regions. The most obvious exception is the largely humid tropical genus
<taxonomicName id="2AB18D230D838E8F25CDD62D689CB3B6" class="Arachnida" family="Pholcidae" genus="Smeringopina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopina" order="Araneae" pageId="10" pageNumber="61" phylum="Arthropoda" rank="genus">Smeringopina</taxonomicName>
Kraus, 1957, which is also the genus with the smallest and largest representatives in the subfamily (with body lengths ranging from 2.5-10 mm) and with the widest range of microhabitats used (leaf litter to large sheltered spaces) (
<bibRefCitation id="8CDA69DAA96D430D76A099F0272F8803" author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="1 - 160" title="Revision and cladistic analysis of the Guineo-Congolian spider genus Smeringopina Kraus (Araneae, Pholcidae)." url="https://doi.org/10.11646/zootaxa.3713.1.1" volume="3713" year="2013">Huber 2013</bibRefCitation>
). The original distribution of the subfamily is Africa, the Mediterranean, and the Middle East. Three species have attained much wider distributions, resulting from human-mediated dispersal (
<bibRefCitation id="1F339847B1ADD5E2E012DD12E5CA0D11" 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>
).
</paragraph>
<paragraph id="A964C7E64C657FAC4021995A75602FD1" pageId="10" pageNumber="61">
As in previous molecular analyses (
<bibRefCitation id="54ADD98449B96B2F5BFB8D69D563ED93" author="Bruvo-Mađaric, B" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="39" pageNumber="90" pagination="661 - 673" title="Phylogeny of pholcid spiders (Araneae: Pholcidae): combined analysis using morphology and molecules." url="https://doi.org/10.1016/j.ympev.2005.08.016" volume="37" year="2005">
<normalizedToken id="7F347596D21C071E35977F97A8ED1600" originalValue="Bruvo-Mađarić">Bruvo-Mađaric</normalizedToken>
et al. 2005
</bibRefCitation>
,
<bibRefCitation id="8C7306BF74C371637F366A962501BA4C" author="Astrin, JJ" journalOrPublisher="Zootaxa" pageId="39" pageNumber="90" pagination="17 - 30" title="The pitfalls of exaggeration: molecular and morphological evidence suggests Kaliana is a synonym of Mesabolivar (Araneae: Pholcidae)." volume="1646" year="2007">Astrin et al. 2007</bibRefCitation>
,
<bibRefCitation id="BD7544C6B196B396F0FE9F3A9CC030DD" author="Dimitrov, D" journalOrPublisher="Cladistics" pageId="39" pageNumber="90" pagination="132 - 146" title="Pholcid spider molecular systematics revisited, with new insights into the biogeography and the evolution of the group." url="https://doi.org/10.1111/j.1096-0031.2012.00419.x" volume="29" year="2013">Dimitrov et al. 2013</bibRefCitation>
),
<taxonomicName id="86301D70EBBE3D214CF48DD17316A973" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Smeringopinae">Smeringopinae</taxonomicName>
is sister to
<taxonomicName id="7451282CDD9A8D0F0F6B540E3B412C93" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Pholcinae">Pholcinae</taxonomicName>
(Figure 1) with reasonable to high support. This relationship is also supported by morphology: the two taxa share tarsus IV comb-hairs spread over the entire length of the tarsus (
<bibRefCitation id="98A9CAEB4ECB42C7D76C3B04282A1FC2" author="Huber, BA" journalOrPublisher="Journal of Arachnology" pageId="41" pageNumber="92" pagination="232 - 240" title="Comb-hairs on the fourth tarsi in pholcid spiders (Araneae, Pholcidae)." url="https://doi.org/10.1636/CSh07-71.1" volume="36" year="2008">Huber and Fleckenstein 2008</bibRefCitation>
).
</paragraph>
<paragraph id="DC03B30A36C7767C8AC0DED63F22F3C4" pageId="10" pageNumber="61">
The monophyly of
<taxonomicName id="7CAAE9036E26CE8E6B593DF760E85057" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Smeringopinae">Smeringopinae</taxonomicName>
receives reasonable to high support in all our analyses. Previous molecular analyses have partly supported
<taxonomicName id="61F431C14409AB8E7EE89B9133708664" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Smeringopinae">Smeringopinae</taxonomicName>
, but also suggested rather obscure relationships [e.g., the position of
<taxonomicName id="BFE102063E81811197F190BC3BF8D332" class="Arachnida" family="Pholcidae" genus="Holocnemus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Holocnemus pluchei" order="Araneae" pageId="10" pageNumber="61" phylum="Arthropoda" rank="species" species="pluchei">Holocnemus pluchei</taxonomicName>
(Scopoli, 1763) among
<taxonomicName id="4C580DC4ECCC0C96818D4B9E4A54A1C5" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Ninetinae">Ninetinae</taxonomicName>
in
<bibRefCitation id="2B216E65EF2FE38A1A10A720004D0E6D" author="Astrin, JJ" journalOrPublisher="Zootaxa" pageId="39" pageNumber="90" pagination="17 - 30" title="The pitfalls of exaggeration: molecular and morphological evidence suggests Kaliana is a synonym of Mesabolivar (Araneae: Pholcidae)." volume="1646" year="2007">Astrin et al. 2007</bibRefCitation>
].
<taxonomicName id="34E339EBF6E3B9273EEF647731813524" class="Arachnida" family="Pholcidae" genus="Holocnemus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Holocnemus pluchei" order="Araneae" pageId="10" pageNumber="61" phylum="Arthropoda" rank="species" species="pluchei">Holocnemus pluchei</taxonomicName>
was included in preliminary analyses of the present data but its position was drastically unstable, so we decided to exclude it from the final analyses.
<taxonomicName id="94BFCF4183A1C1C480AB46DE10D96CF2" lsidName="" pageId="10" pageNumber="61" rank="subFamily" subFamily="Smeringopinae">Smeringopinae</taxonomicName>
monophyly is rather weakly supported by morphology, i.e., by the presence of a large thoracic pit on the carapace (rather than a narrow furrow or an evenly domed carapace; cf.
<bibRefCitation id="54775CE24A3B9972E0229B371A53AEE5" 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>
).
</paragraph>
<paragraph id="6DB13A210DE6AF8DC1FD4A52F1A8145B" pageId="11" pageNumber="62">
<pageBreakToken id="D9ADE29C3865CE92FB52A429E77AF559" pageId="11" pageNumber="62" start="start">Within</pageBreakToken>
<taxonomicName id="36D551D341021E937BC7E82023D5D021" lsidName="" pageId="11" pageNumber="62" rank="subFamily" subFamily="Smeringopinae">Smeringopinae</taxonomicName>
, our data strongly support a basal split between a northern clade (Mediterranean, northern Africa, Middle East, Central Asia) and a southern clade (Sub-Sahara) (Figure 6). This basal split was also recovered in a morphological cladistic analysis (
<bibRefCitation id="5BC192C5EBEA3A0237382E170B1D33AF" 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>
). Within the northern clade,
<taxonomicName id="160962F756293188DF39C3C5C7F49BDC" class="Arachnida" family="Pholcidae" genus="Hoplopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hoplopholcus" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="genus">Hoplopholcus</taxonomicName>
Kulczynski, 1908 is sister to all other genera and not close to
<taxonomicName id="9D2220A952B8D3590AB4B0A8EA2A9347" class="Arachnida" family="Pholcidae" genus="Stygopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Stygopholcus" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="genus">Stygopholcus</taxonomicName>
Kratochvil, 1932 as repeatedly claimed by
<bibRefCitation id="28B363C60D128104A4FF384DA0A85EE1" author="Brignoli, PM" journalOrPublisher="Mitteilungen des Zoologischen Museums Berlin" pageId="39" pageNumber="90" pagination="255 - 267" title="Beitrag zur Kenntnis der mediterranen Pholcidae (Arachnida, Araneae)." url="https://doi.org/10.1002/mmnz.19710470203" volume="47" year="1971">Brignoli (1971</bibRefCitation>
,
<bibRefCitation id="8896A977635055C9B79F28BB97AE899E" author="Brignoli, PM" journalOrPublisher="Revue suisse de Zoologie" pageId="39" pageNumber="90" pagination="539 - 578" title="Ragni di Grecia IX. Specie nuove o interessanti delle famiglie Leptonetidae, Dysderidae, Pholcidae ed Agelenidae (Araneae)." volume="83" year="1976">1976</bibRefCitation>
,
<bibRefCitation id="EC995F8CBDC307937A5BC2A8E14EAE8D" author="Brignoli, PM" journalOrPublisher="Bulletin of the British arachnological Society" pageId="39" pageNumber="90" pagination="350 - 352" title="Spiders from Lebanon, V. On Hoplopholcuscecconii Kulczynski, 1908 (Pholcidae)." volume="4" year="1979">1979</bibRefCitation>
) but contested by
<bibRefCitation id="7BB1261862A500B95B1DE85C033104A8" pageId="11" pageNumber="62">Senglet (1971</bibRefCitation>
,
<bibRefCitation id="A488C4AD39F62966C1E7BD7637D0563C" author="Senglet, A" journalOrPublisher="Mitteilungen der schweizerischen entomologischen Gesellschaft" pageId="43" pageNumber="94" pagination="43 - 67" title="Copulatory mechanisms in Hoplopholcus, Stygopholcus (revalidated), Pholcus, Spermophora and Spermophorides (Araneae, Pholcidae), with additional faunistic and taxonomic data." volume="74" year="2001">2001</bibRefCitation>
). The genera
<taxonomicName id="9B2FD9428E4FEC2311039DE01F5D841C" class="Arachnida" family="Pholcidae" genus="Hoplopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hoplopholcus" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="genus">Hoplopholcus</taxonomicName>
,
<taxonomicName id="3C0015FCD155B9D323D801256AA83A3F" class="Arachnida" family="Pholcidae" genus="Stygopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Stygopholcus" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="genus">Stygopholcus</taxonomicName>
, and
<taxonomicName id="5BE6ECE44AE88217301A930F98FF93B0" class="Arachnida" family="Pholcidae" genus="Crossopriza" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Crossopriza" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="genus">Crossopriza</taxonomicName>
Simon, 1893 all receive full support, but the small Mediterranean genus
<taxonomicName id="EEF75DCCAA894500D909A8608B36D5E5" class="Arachnida" family="Pholcidae" genus="Holocnemus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Holocnemus" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="genus">Holocnemus</taxonomicName>
Simon, 1873 (only three described species) continues to be problematic even after the exclusion of
<taxonomicName id="BA2B5E51E9388095AA72E997436E82A5" lsidName="H. pluchei" pageId="11" pageNumber="62" rank="species" species="pluchei">H. pluchei</taxonomicName>
. The two species of
<taxonomicName id="A1E94F8071A3764958252A828E6E2246" class="Arachnida" family="Pholcidae" genus="Holocnemus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Holocnemus" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="genus">Holocnemus</taxonomicName>
included in our analyses never group together, and no morphological synapomorphy is known to suggest their sister-group relationship (in fact,
<taxonomicName id="3D4B803280A9E59C080C1F9AF637FED1" class="Arachnida" family="Pholcidae" genus="Holocnemus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Holocnemus" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="genus">Holocnemus</taxonomicName>
has never been revised).
</paragraph>
<caption id="8AC6A2CB8C949436D712B6F9EA9CEB8E" pageId="11" pageNumber="62">
<paragraph id="1CA11D671F79380F63F39D8213F82811" pageId="11" pageNumber="62">
Figure 6. Smeringopinae a
<taxonomicName id="7F4076C1890911210728981ED2D151C0" class="Arachnida" family="Pholcidae" genus="Hoplopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hoplopholcus" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="genus">Hoplopholcus</taxonomicName>
sp. n.
<normalizedToken id="846B189F0410A83C0E8F47474CAE55DE" originalValue="“Mar66”">&quot;Mar66&quot;</normalizedToken>
(Turkey) b
<taxonomicName id="C2D9ED0603F050FDF90320D339FF7377" class="Arachnida" family="Pholcidae" genus="Stygopholcus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Stygopholcus absoloni" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="species" species="absoloni">Stygopholcus absoloni</taxonomicName>
? (Bosnia and Herzegovina) c
<taxonomicName id="B73E060FD67EAE6C593C7F333E1F837F" class="Arachnida" family="Pholcidae" genus="Crossopriza" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Crossopriza" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="genus">Crossopriza</taxonomicName>
sp. n.
<normalizedToken id="6F7CA0A04560A142B61DE6C85FB514A9" originalValue="“Om11”">&quot;Om11&quot;</normalizedToken>
(Oman) d
<taxonomicName id="CDB3BFC6EBB194325ECBD7D9D8B0BD0C" class="Arachnida" family="Pholcidae" genus="Smeringopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopus pallidus" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="species" species="pallidus">Smeringopus pallidus</taxonomicName>
(Philippines) e
<taxonomicName id="C1C2CD6B7687F2DEE11AAEC1F57F05EC" class="Arachnida" family="Pholcidae" genus="Smeringopina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopina pulchra" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="species" species="pulchra">Smeringopina pulchra</taxonomicName>
(Ghana) f
<taxonomicName id="F37D43AAD6672FF9D447543A18488596" class="Arachnida" family="Pholcidae" genus="Smeringopina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopina ankasa" order="Araneae" pageId="11" pageNumber="62" phylum="Arthropoda" rank="species" species="ankasa">Smeringopina ankasa</taxonomicName>
(Ghana).
</paragraph>
</caption>
<paragraph id="EBC31628DEDC8795EE2D7859350AFCC6" pageId="12" pageNumber="63">
<pageBreakToken id="4766D9B613C35BB7588B4CA9FCD3B253" pageId="12" pageNumber="63" start="start">The</pageBreakToken>
southern (Sub-Saharan) clade includes
<taxonomicName id="C7E82FDE349F36B0243E0A5C64AC2C5F" class="Arachnida" family="Pholcidae" genus="Smeringopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopus" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopus</taxonomicName>
Simon, 1890 and
<taxonomicName id="6090F4BBAC5FD6082A54AEF492C0989A" class="Arachnida" family="Pholcidae" genus="Smeringopina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopina" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopina</taxonomicName>
, and is also supported by a unique number of epiandrous spigots (two) (
<bibRefCitation id="EA323C12432C6D263FA3F71F654E3F51" 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>
). The paraphyly of
<taxonomicName id="80F943C7B1875391AAE071867B081B6D" class="Arachnida" family="Pholcidae" genus="Smeringopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopus" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopus</taxonomicName>
has been suggested before (
<bibRefCitation id="C0C46BBE00097303A5CB8F9275A5119E" author="Dimitrov, D" journalOrPublisher="Cladistics" pageId="39" pageNumber="90" pagination="132 - 146" title="Pholcid spider molecular systematics revisited, with new insights into the biogeography and the evolution of the group." url="https://doi.org/10.1111/j.1096-0031.2012.00419.x" volume="29" year="2013">Dimitrov et al. 2013</bibRefCitation>
), and our larger data set supports this view, but with low support values. Two of the species groups of
<taxonomicName id="F27FEA9F68A26B88D4B84F4C1CF86461" class="Arachnida" family="Pholcidae" genus="Smeringopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopus" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopus</taxonomicName>
proposed in
<bibRefCitation id="017AA2E502E57AB66FC0DD7967E0351A" 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>
appear closer to
<taxonomicName id="5C784C0AEA3530BBCB15C362897D2462" class="Arachnida" family="Pholcidae" genus="Smeringopina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopina" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopina</taxonomicName>
than to other
<taxonomicName id="4A36A050996784AFC3EB5E2EFD9A7149" class="Arachnida" family="Pholcidae" genus="Smeringopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopus" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopus</taxonomicName>
: the
<taxonomicName id="86134A9D132261370D2E67B6C5D270F9" genus="Modisimeae" lsidName="Modisimeae chogoria" pageId="12" pageNumber="63" rank="species" species="chogoria">chogoria</taxonomicName>
group and the rubrotinctus group. Morphological data do not support this view but they neither strongly contradict it: the two species groups lack the distinctive arrangement of pores on the pore plates (in groups or
<normalizedToken id="C0B4DDFD3B6880ADD7ED4E4343F35952" originalValue="islands">'islands'</normalizedToken>
) and the retrolateral furrow on the male palpal femur present in all other species of
<taxonomicName id="8ABDEE4BE794171B1B491CF595CDABA6" class="Arachnida" family="Pholcidae" genus="Smeringopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopus" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopus</taxonomicName>
(
<bibRefCitation id="4A2A6054D72BD8511856AF32FC0739C9" 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>
). Remarkably,
<taxonomicName id="913F52556FDA480991EB902E7C063E69" class="Arachnida" family="Pholcidae" genus="Smeringopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopus" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopus</taxonomicName>
and
<taxonomicName id="051B622A0F4AFB0004B5E7E35F699D33" class="Arachnida" family="Pholcidae" genus="Smeringopina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopina" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopina</taxonomicName>
are largely separated geographically, with
<taxonomicName id="DD9E257488189A4BAF18EFA467384575" class="Arachnida" family="Pholcidae" genus="Smeringopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopus" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopus</taxonomicName>
being most diverse in southern and eastern Africa, and
<taxonomicName id="6C85C0B2A014A5E2E885F67A08ED1CCD" class="Arachnida" family="Pholcidae" genus="Smeringopina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopina" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopina</taxonomicName>
in western and central Africa (
<bibRefCitation id="5CA029195EF1DA4D2C87CC735D0F1C24" 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>
,
<bibRefCitation id="0E9F964BD43CD475EF6E69C2FA265A31" author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="1 - 160" title="Revision and cladistic analysis of the Guineo-Congolian spider genus Smeringopina Kraus (Araneae, Pholcidae)." url="https://doi.org/10.11646/zootaxa.3713.1.1" volume="3713" year="2013">2013</bibRefCitation>
). The
<taxonomicName id="AEDE51B291E7D564EF14B8FCAE2311A9" genus="Modisimeae" lsidName="Modisimeae chogoria" pageId="12" pageNumber="63" rank="species" species="chogoria">chogoria</taxonomicName>
and rubrotinctus groups are geographically restricted to an area where Central Africa (the Guineo-Congolian center of endemism) meets East Africa (
<bibRefCitation id="0456ED6E6C63A0075DD5A63DC494E7CA" 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>
). Other than that, our sampling in
<taxonomicName id="1B5089D026DDEE56C32CE27D36FF3B19" class="Arachnida" family="Pholcidae" genus="Smeringopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopus" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopus</taxonomicName>
is not dense enough to test the species groups proposed in
<bibRefCitation id="FC863E7F5463BBCCC32975DAC2FBE80E" 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>
. Remarkably, though, the isolated
<normalizedToken id="DE4F10F78C6D6CA608ACDAF45CD4D335" originalValue="basal">'basal'</normalizedToken>
position of
<taxonomicName id="3550A94C73A15BFB99CAD03FCB2EB9AB" lsidName="S. ngangao" pageId="12" pageNumber="63" rank="species" species="ngangao">S. ngangao</taxonomicName>
Huber, 2012 is supported by the present analyses.
</paragraph>
<paragraph id="70C4F90C99BE7B1B036BD69E88797857" pageId="12" pageNumber="63">
Our analyses include 30 of the 44 described species of
<taxonomicName id="6BDF6D81084D7E6DD394CC8C51DE5E36" class="Arachnida" family="Pholcidae" genus="Smeringopina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopina" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopina</taxonomicName>
(68%), and all species groups proposed in
<bibRefCitation id="4BFC6B61594ADCF32FD3A5A9FFBD0EBD" author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="1 - 160" title="Revision and cladistic analysis of the Guineo-Congolian spider genus Smeringopina Kraus (Araneae, Pholcidae)." url="https://doi.org/10.11646/zootaxa.3713.1.1" volume="3713" year="2013">Huber (2013)</bibRefCitation>
except two monotypic
<normalizedToken id="5B4B2D0E5F3E07C116DA7619FE818900" originalValue="groups">'groups'</normalizedToken>
(
<taxonomicName id="2BDF1E5C4AEE3C091AF185D8EF1D9C97" lsidName="S. fon" pageId="12" pageNumber="63" rank="species" species="fon">S. fon</taxonomicName>
Huber, 2013;
<taxonomicName id="7B2304B50D18ADF9C9D810992D6F8E53" lsidName="S. ngungu" pageId="12" pageNumber="63" rank="species" species="ngungu">S. ngungu</taxonomicName>
Huber, 2013). Even though for some species only one gene (CO1) was sequenced, our analyses support several species groups and deeper relationships proposed previously (
<bibRefCitation id="5DB5F0D0442B792D41A5253301B028EC" author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="1 - 160" title="Revision and cladistic analysis of the Guineo-Congolian spider genus Smeringopina Kraus (Araneae, Pholcidae)." url="https://doi.org/10.11646/zootaxa.3713.1.1" volume="3713" year="2013">Huber 2013</bibRefCitation>
), based on cladistic analysis of morphological characters. Morphology placed the West African guineensis group as sister to all other
<taxonomicName id="0EBF39B0B55B163B5F1DD6530FF41A25" class="Arachnida" family="Pholcidae" genus="Smeringopina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Smeringopina" order="Araneae" pageId="12" pageNumber="63" phylum="Arthropoda" rank="genus">Smeringopina</taxonomicName>
; all our analyses support both the monophyly of the guineensis group and its sister-group relationship with all other congeners. The next two branches are composed of representatives of the lekoni group, which is thus here considered paraphyletic rather than monophyletic. The
<taxonomicName id="36948D800054E431B00AFF02ACD33018" genus="Modisimeae" lsidName="Modisimeae ankasa" pageId="12" pageNumber="63" rank="species" species="ankasa">ankasa</taxonomicName>
and cornigera groups are both supported, as is their sister group relationship to each other. The attuleh group is supported, but not as sister to the
<taxonomicName id="E4147584E4A34649A911577B754659D9" genus="Modisimeae" lsidName="Modisimeae ankasa" pageId="12" pageNumber="63" rank="species" species="ankasa">ankasa</taxonomicName>
+ cornigera groups but as sister to the following group. The last clade is composed of representatives of the simplex and beninensis groups, but the clear dichotomy in the molecular trees is not equivalent to these groups. Instead, the simplex group includes all
<normalizedToken id="D774FABB53574554674588B4F01655B6" originalValue="basal">'basal'</normalizedToken>
representatives originally assigned to the beninensis group; the beninensis group includes only those species that have a light transversal element ventrally on the abdomen (character 9 in
<bibRefCitation id="C541239F0FE65C718CC57C0575C610F3" author="Huber, BA" journalOrPublisher="Zootaxa" pageId="41" pageNumber="92" pagination="1 - 160" title="Revision and cladistic analysis of the Guineo-Congolian spider genus Smeringopina Kraus (Araneae, Pholcidae)." url="https://doi.org/10.11646/zootaxa.3713.1.1" volume="3713" year="2013">Huber 2013</bibRefCitation>
, which is thus less homoplastic than previously thought).
</paragraph>
</subSubSection>
</treatment>
</document>