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<mods:title id="6AF05B3322A11737A4CBA420EF0940C0">A stable phylogenomic classification of Travunioidea (Arachnida, Opiliones, Laniatores) based on sequence capture of ultraconserved elements</mods:title>
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<mods:namePart id="0BA9D750AA304E9DA1FBD526975C55B0">Derkarabetian, Shahan</mods:namePart>
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<mods:namePart id="A2C93D66DF56972F7F09049A441B87A2">Starrett, James</mods:namePart>
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<mods:namePart id="9C5F5DE3AD9CA8DB32EE5100AAC13E7F">Tsurusaki, Nobuo</mods:namePart>
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<mods:namePart id="712856E073121FC158DA15026CADBABD">Ubick, Darrell</mods:namePart>
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<mods:namePart id="F03FE0E42E4CDF172300F6E4E3468BD7">Castillo, Stephanie</mods:namePart>
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<mods:namePart id="49D1C8B4D567E8F81E71F4E6A51ECBC8">Hedin, Marshal</mods:namePart>
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<mods:date id="6C33B7160A19B6A67998FF1B85014F5D">2018</mods:date>
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<treatment id="43981D382E814BEE609448197A2BDD66" ID-GBIF-Taxon="144148279" LSID="urn:lsid:plazi:treatment:43981D382E814BEE609448197A2BDD66" httpUri="http://treatment.plazi.org/id/43981D382E814BEE609448197A2BDD66" lastPageNumber="1" pageId="0" pageNumber="1" scope_class="Arachnida" scope_order="Opiliones" scope_subOrder="Laniatores">
<subSubSection id="41BE6B07F96EBF2FED004AC61EB1FEC6" pageId="0" pageNumber="1" type="nomenclature">
<paragraph id="56657EA62915A08E84306DA733DD36B7" pageId="0" pageNumber="1">
Family
<taxonomicName id="534B3C2CC959794C58EC5FCB91E6D341" ID-CoL="DZF" authority="Briggs, 1971" authorityName="Briggs" authorityYear="1971" class="Arachnida" family="Paranonychidae" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">PARANONYCHIDAE Briggs, 1971</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="D34AD6D54CAB434D0E474B7E04146D03" pageId="0" pageNumber="1" type="type genus">
<paragraph id="8EFF026C319061968CB1706A8CCB7CCA" pageId="0" pageNumber="1">Type genus.</paragraph>
<paragraph id="FD171DC127A866C2C2D41AD4E2717099" pageId="0" pageNumber="1">
<taxonomicName id="8AFA79DB986D45B108FA43E5265D76D2" class="Arachnida" family="Paranonychidae" genus="Paranonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paranonychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Paranonychus</taxonomicName>
Briggs, 1971
</paragraph>
</subSubSection>
<subSubSection id="D35F2E1AC04DC3C6D8703EBF1F5181A0" pageId="0" pageNumber="1" type="type species">
<paragraph id="5698802EC6DDA3D930607DEFE13E3B6D" pageId="0" pageNumber="1">Type species.</paragraph>
<paragraph id="86D89A23D7F851BB0BCC24DD38D58B90" pageId="0" pageNumber="1">
<taxonomicName id="14EC10A42E7BC41305B144967C74D743" class="Arachnida" family="Paranonychidae" genus="Paranonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paranonychus brunneus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="brunneus">Paranonychus brunneus</taxonomicName>
(Banks, 1893).
</paragraph>
</subSubSection>
<subSubSection id="B74D1F3D2EE56570C996A8EC0D632EE8" pageId="0" pageNumber="1" type="diagnosis">
<paragraph id="ADCD8AC7407DFD347FA92AB5AE5E494C" pageId="0" pageNumber="1">Diagnosis.</paragraph>
<paragraph id="8B065844AAF36523DC7DD60BA6CBF7FB" pageId="0" pageNumber="1">
The
<taxonomicName id="748CF776D49607794E06E6A600298819" class="Arachnida" family="Paranonychidae" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Paranonychidae</taxonomicName>
can be diagnosed by their relatively complex glans (except
<taxonomicName id="126317F5C80F15E3BC5BA9A41AF6D877" class="Arachnida" family="Paranonychidae" genus="Paranonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paranonychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Paranonychus</taxonomicName>
) (Figure 7 and Suppl. material 2: Figure 4), and by their intestinal complex (Suppl. material 2: Figure 2). For all taxa that have been examined, the paranonychids possess a small D1 that is circular to subtriangular, and a simple and shorter OD3. The paranonychids are restricted to western North America and East Asia. In southern Japan the paranonychids are sympatric with
<taxonomicName id="3DAFF93B4D58035139E667DEB0978C13" class="Arachnida" genus="Yuria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Yuria" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Yuria</taxonomicName>
and can be diagnosed by several characteristics:
<taxonomicName id="7E7BDEA57CEAC08CF3A816FA1E0629DF" class="Arachnida" genus="Yuria" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Yuria" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Yuria</taxonomicName>
possesses a free ninth tergite, and the penis has a dorsal plate with fused stylus; the paranonychids do not have a free ninth tergite and the penis glans lacks a dorsal plate. In western North America, the paranonychids are sympatric and syntopic in surface habitats with the
<taxonomicName id="863ED48C27213E00A26840E4D7861BFE" class="Arachnida" family="Cladonychiidae" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Cladonychiidae</taxonomicName>
(
<taxonomicName id="90A0F3D53F29166781B867A08DC43EB8" class="Arachnida" family="Cladonychiidae" genus="Briggsus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Briggsus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Briggsus</taxonomicName>
,
<taxonomicName id="4D3CD6835ED67A7EEFF1BA1F8BF9D06D" class="Arachnida" family="Cladonychiidae" genus="Isolachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Isolachus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Isolachus</taxonomicName>
) and
<taxonomicName id="D25EBD4BE38BDDB7373A9A0DCF0D4286" class="Arachnida" family="Cryptomastridae" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Cryptomastridae</taxonomicName>
(
<taxonomicName id="18CBE622241DE3ECFF8972A755BA6B69" class="Arachnida" family="Cryptomastridae" genus="Cryptomaster" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Cryptomaster" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Cryptomaster</taxonomicName>
). The paranonychids can be differentiated from these families by the structure of D1: paranonychids possess a small circular to subtriangular unbranched D1, while
<taxonomicName id="807C93D53FB1100DE9A3608929AB9965" class="Arachnida" family="Cladonychiidae" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Cladonychiidae</taxonomicName>
and
<taxonomicName id="DE69469405A1B3D7D3202A84A1A006AD" class="Arachnida" family="Cryptomastridae" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Cryptomastridae</taxonomicName>
possess an elongate, triangular, and branched D1.
</paragraph>
</subSubSection>
<subSubSection id="9341A58A765CA084436D0B6C5C912E42" pageId="0" pageNumber="1" type="included genera and species">
<paragraph id="ACA4FF779D2103833C463AB6242D2B0D" pageId="0" pageNumber="1">Included genera and species.</paragraph>
<paragraph id="2B34996B5D4C187C81CCCCEE75CC81EB" pageId="0" pageNumber="1">
<taxonomicName id="BC94C6C0D1FA735B62D6C4ED98CFB306" class="Arachnida" family="Paranonychidae" genus="Paranonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paranonychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Paranonychus</taxonomicName>
(Figure 1K). This trans-Beringian genus includes three known species:
<taxonomicName id="0C90581D9B59057D849AB4949AAAF59E" lsidName="P. brunneus" pageId="0" pageNumber="1" rank="species" species="brunneus">P. brunneus</taxonomicName>
(Banks, 1893) distributed in the Coast and Cascade Ranges of Oregon and Washington with records extending north to Alaska;
<taxonomicName id="62D16ABA0979ED8861AFA39AD456F558" lsidName="P. concolor" pageId="0" pageNumber="1" rank="species" species="concolor">P. concolor</taxonomicName>
Briggs, 1971, recorded from a single location in the southern Cascade Range of Oregon; and
<taxonomicName id="07CEAF71F0744E2DFA4E232847D24B5E" lsidName="P. fuscus" pageId="0" pageNumber="1" rank="species" species="fuscus">P. fuscus</taxonomicName>
found throughout northern Honshu in Japan.
</paragraph>
<paragraph id="EF56476B7692CA5F89A6C2C625D094F1" pageId="0" pageNumber="1">
<taxonomicName id="A1D27BC49DD6683998029D3BEBC24BCB" class="Arachnida" family="Paranonychidae" genus="Metanonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Metanonychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Metanonychus</taxonomicName>
Briggs, 1971. This genus and all species were described by Briggs (1971) and are restricted to the moist forests of the Pacific Northwest of North America.
<taxonomicName id="BD8355273DACB5CB23DC656046D2E0A5" class="Arachnida" family="Paranonychidae" genus="Metanonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Metanonychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Metanonychus</taxonomicName>
includes three species:
<taxonomicName id="5296A14B3574A2B30EA20F31DCE520D8" lsidName="M. nigricans" pageId="0" pageNumber="1" rank="species" species="nigricans">M. nigricans</taxonomicName>
Briggs, 1971 with two subspecies,
<taxonomicName id="A6ABEEA9A6CED31F71563AB38EF98328" lsidName="M. n." pageId="0" pageNumber="1" rank="subspecies" species="n." subspecies="nigricans">M. n. nigricans</taxonomicName>
and
<taxonomicName id="AC13F33CE1BB5812072A261D552E28A7" lsidName="M. n." pageId="0" pageNumber="1" rank="subspecies" species="n." subspecies="oregonus">M. n. oregonus</taxonomicName>
, found in Oregon;
<taxonomicName id="5EC8E5A04880852FBEDB00E637AA1FAA" lsidName="M. setulus" pageId="0" pageNumber="1" rank="species" species="setulus">M. setulus</taxonomicName>
Briggs, 1971 with five subspecies,
<taxonomicName id="7D1B6352A08A845EFE8E557CCCC6C431" lsidName="M. s." pageId="0" pageNumber="1" rank="subspecies" species="s." subspecies="setulus">M. s. setulus</taxonomicName>
,
<taxonomicName id="FB881679CEED64A4FEFD45B534C8CF02" lsidName="M. s." pageId="0" pageNumber="1" rank="subspecies" species="s." subspecies="cascadus">M. s. cascadus</taxonomicName>
,
<taxonomicName id="C9BAEFD1C23965F4557BA7370FB77213" lsidName="M. s." pageId="0" pageNumber="1" rank="subspecies" species="s." subspecies="mazamus">M. s. mazamus</taxonomicName>
,
<taxonomicName id="AF3CE068388EC4FFA044475A69FCF28F" lsidName="M. s." pageId="0" pageNumber="1" rank="subspecies" species="s." subspecies="navarrus">M. s. navarrus</taxonomicName>
, and
<taxonomicName id="AE2A6C249D3ABE92A3AA7117193B6507" lsidName="M. s." pageId="0" pageNumber="1" rank="subspecies" species="s." subspecies="obrieni">M. s. obrieni</taxonomicName>
, found in Oregon, Washington, and northern California; and
<taxonomicName id="DB6EDC022500A5D4EA53274A208F4BB0" lsidName="M. idahoensis" pageId="0" pageNumber="1" rank="species" species="idahoensis">M. idahoensis</taxonomicName>
Briggs, 1971 found in northern Idaho.
</paragraph>
<paragraph id="346BDD95368422F4140B605B29C3148E" pageId="0" pageNumber="1">
<taxonomicName id="76FFB05A2A553A8BA57E9EEA54D174A3" class="Arachnida" family="Paranonychidae" genus="Sclerobunus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sclerobunus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Sclerobunus</taxonomicName>
Banks, 1893 (Figure 1L). Recently revised by
<bibRefCitation id="2BCFF8FE7F4CF43D12D49F0CF7031F42" author="Derkarabetian, S" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="1" url="https://doi.org/10.1371/journal.pone.0104982" year="2014">Derkarabetian and Hedin (2014)</bibRefCitation>
,
<taxonomicName id="325EEE64D62EBE413BB1F6D6202AE3E1" class="Arachnida" family="Paranonychidae" genus="Sclerobunus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sclerobunus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Sclerobunus</taxonomicName>
is distributed throughout western North America and currently includes 12 species divided into three species groups. The nondimorphicus group includes
<taxonomicName id="69BCEAE33BD87E0E2A1DE106A1B0D2E6" lsidName="S. nondimorphicus" pageId="0" pageNumber="1" rank="species" species="nondimorphicus">S. nondimorphicus</taxonomicName>
Briggs, 1971 from Oregon, Washington, and British Columbia, and
<taxonomicName id="21AD0A07BD794FFA80BF3E7F499F36BF" lsidName="S. idahoensis" pageId="0" pageNumber="1" rank="species" species="idahoensis">S. idahoensis</taxonomicName>
Briggs, 1971 from northern Idaho. The cave-obligate cavicolens group includes:
<taxonomicName id="388FA510506D61C5E440DF2075C752C3" class="Arachnida" family="Paranonychidae" genus="Sclerobunus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sclerobunus cavicolens" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="cavicolens">Sclerobunus cavicolens</taxonomicName>
(Banks, 1905) restricted to Lewis and Clark Caverns, Montana;
<taxonomicName id="D554386430B306BE4323278F18352EF9" class="Arachnida" family="Paranonychidae" genus="Sclerobunus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sclerobunus ungulatus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="ungulatus">Sclerobunus ungulatus</taxonomicName>
(Briggs, 1971) from caves in Great Basin National Park, Nevada;
<taxonomicName id="170F2F30F8AF7A81F93932970039B4C3" class="Arachnida" family="Paranonychidae" genus="Sclerobunus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sclerobunus madhousensis" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="madhousensis">Sclerobunus madhousensis</taxonomicName>
(Briggs, 1971) from caves near Provo, Utah. The robustus group includes the widespread
<taxonomicName id="EB952298348A2A2148F7E8FDC5E7F507" lsidName="S. robustus" pageId="0" pageNumber="1" rank="species" species="robustus">S. robustus</taxonomicName>
(Packard, 1877),
<taxonomicName id="41A0C9FF05571CAC9CCC4AF579CA9641" lsidName="S. glorietus" pageId="0" pageNumber="1" rank="species" species="glorietus">S. glorietus</taxonomicName>
Briggs, 1971, and
<taxonomicName id="11740F7DFEFE1E1B6B82BE36A3EC6D76" lsidName="S. skywalkeri" pageId="0" pageNumber="1" rank="species" species="skywalkeri">S. skywalkeri</taxonomicName>
Derkarabetian &amp; Hedin, 2014, all distributed throughout the high elevation forests of the southwestern United States, and
<taxonomicName id="F70BFB77ABEB8FA6DA9B73BA5D15850D" lsidName="S. jemez" pageId="0" pageNumber="1" rank="species" species="jemez">S. jemez</taxonomicName>
Derkarabetian &amp; Hedin, 2014,
<taxonomicName id="58C66A58617F007579BD38218D551487" lsidName="S. klomax" pageId="0" pageNumber="1" rank="species" species="klomax">S. klomax</taxonomicName>
Derkarabetian &amp; Hedin, 2014,
<taxonomicName id="559EBF6D5047F1BEB21341B09AD32C23" lsidName="S. speoventus" pageId="0" pageNumber="1" rank="species" species="speoventus">S. speoventus</taxonomicName>
Derkarabetian &amp; Hedin, 2014, and
<taxonomicName id="F5DB07EDFAF66941C4BFCEFE13D6175E" lsidName="S. steinmanni" pageId="0" pageNumber="1" rank="species" species="steinmanni">S. steinmanni</taxonomicName>
Derkarabetian &amp; Hedin, 2014, which are all troglomorphic species restricted to cave and talus habitats along the eastern edge of the southern Rocky Mountains in New Mexico and Colorado.
</paragraph>
<paragraph id="5952411C7F223528734BA9401F22B95D" pageId="0" pageNumber="1">
<taxonomicName id="C65A3DEDE80828A7B179A9BF6691CB2B" class="Arachnida" family="Paranonychidae" genus="Kaolinonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kaolinonychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Kaolinonychus</taxonomicName>
Suzuki, 1975. This monotypic genus endemic to South Korea is recorded mostly from caves.
<taxonomicName id="DE23B4975AC90450AD1DDD39BD88663A" class="Arachnida" family="Paranonychidae" genus="Kaolinonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kaolinonychus coreanus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="coreanus">Kaolinonychus coreanus</taxonomicName>
(Suzuki, 1966) includes two subspecies
<taxonomicName id="7AF89DC49995B5F1DCF13DCC27C9FC2E" lsidName="K. c." pageId="0" pageNumber="1" rank="subspecies" species="c." subspecies="coreanus">K. c. coreanus</taxonomicName>
and
<taxonomicName id="DAB93D1C85B5A4FF41C2231B09DCF1B6" lsidName="K. c." pageId="0" pageNumber="1" rank="subspecies" species="c." subspecies="longipes">K. c. longipes</taxonomicName>
.
</paragraph>
<paragraph id="3811D1B1F4F3ACB8C24BB33F0AD5EAEE" pageId="0" pageNumber="1">
<taxonomicName id="0A909613C6C26AFF7D2EF448428BB4CE" class="Arachnida" family="Paranonychidae" genus="Metanippononychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Metanippononychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Metanippononychus</taxonomicName>
Suzuki, 1975. (Figure 1M). Endemic to Japan,
<taxonomicName id="7D63398BD6B3BC6CA2F1B5652055CADD" class="Arachnida" family="Paranonychidae" genus="Metanippononychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Metanippononychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Metanippononychus</taxonomicName>
is restricted to southern Honshu, Shikoku, and Kyushu and includes four species:
<taxonomicName id="538B1D625C9681EC1D0B012641678712" lsidName="M. daisenensis" pageId="0" pageNumber="1" rank="species" species="daisenensis">M. daisenensis</taxonomicName>
Suzuki, 1975;
<taxonomicName id="C7D29B749AFAD63336D031531E1779A7" lsidName="M. iriei" pageId="0" pageNumber="1" rank="species" species="iriei">M. iriei</taxonomicName>
Suzuki, 1975, with two subspecies
<taxonomicName id="4E34033E31612D13B76A45EA1F467B43" lsidName="M. i." pageId="0" pageNumber="1" rank="subspecies" species="i." subspecies="iriei">M. i. iriei</taxonomicName>
and
<taxonomicName id="F7C53361D1C88967A7B87D96D31CFC2F" lsidName="M. i." pageId="0" pageNumber="1" rank="subspecies" species="i." subspecies="yakuensis">M. i. yakuensis</taxonomicName>
;
<taxonomicName id="D4556ACC26066B884B41EE7BA0F0CFA9" lsidName="M. iyanus" pageId="0" pageNumber="1" rank="species" species="iyanus">M. iyanus</taxonomicName>
Suzuki, 1975;
<taxonomicName id="F0F8959EE2B8AAED22C3330BC358595F" lsidName="M. tomishimai" pageId="0" pageNumber="1" rank="species" species="tomishimai">M. tomishimai</taxonomicName>
Suzuki, 1975, with two subspecies
<taxonomicName id="72BB294E041DC239C8707E65B7D35C7A" lsidName="M. t." pageId="0" pageNumber="1" rank="subspecies" species="t." subspecies="tomishimai">M. t. tomishimai</taxonomicName>
and
<taxonomicName id="BF18AD7955FE85AE92FC3B1DB5E38E89" lsidName="M. t." pageId="0" pageNumber="1" rank="subspecies" species="t." subspecies="awanus">M. t. awanus</taxonomicName>
.
</paragraph>
<paragraph id="C01F19BF069D31EF29D7DA9B8A73639C" pageId="0" pageNumber="1">
<taxonomicName id="0856DE8B933AD8F3034443D72F07739E" class="Arachnida" family="Paranonychidae" genus="Nippononychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nippononychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Nippononychus</taxonomicName>
Suzuki, 1975. A monotypic genus endemic to Japan,
<taxonomicName id="8AE35E3D5F5F5AEE21ED93AF78584689" class="Arachnida" family="Paranonychidae" genus="Nippononychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nippononychus japonicus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="japonicus">Nippononychus japonicus</taxonomicName>
(Miyosi, 1957) is restricted to southern Honshu and Shikoku.
</paragraph>
<paragraph id="87F07A8696F1A541A484CDA66ADEC24F" pageId="0" pageNumber="1">
<taxonomicName id="D87D4E13F24D7BDEF84BFA0E204F3C4E" class="Arachnida" family="Paranonychidae" genus="Zuma" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuma" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Zuma</taxonomicName>
(Figure 1N).
<taxonomicName id="26498075DD630C586D1910E166FD9E00" class="Arachnida" family="Paranonychidae" genus="Zuma" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuma" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Zuma</taxonomicName>
includes two species restricted to forests of central and northern California:
<taxonomicName id="AB2607AF48BF1C95A6C9ECCA855BEA17" class="Arachnida" family="Paranonychidae" genus="Zuma" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuma acuta" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="acuta">Zuma acuta</taxonomicName>
Goodnight &amp; Goodnight, 1942 restricted to the coastal forests south of San Francisco;
<taxonomicName id="62466AA19E749FA5E4BF9ADA9C0DD62E" class="Arachnida" family="Paranonychidae" genus="Zuma" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zuma tioga" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="tioga">Zuma tioga</taxonomicName>
Briggs, 1971 found in the central and northern Sierra Nevada range.
</paragraph>
<paragraph id="3D4C74691C75A43F59F143BC5C05AC9C" pageId="0" pageNumber="1">
<taxonomicName id="C04469436DFE11747D075F9178C98A4B" class="Arachnida" family="Paranonychidae" genus="Izunonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Izunonychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Izunonychus</taxonomicName>
Suzuki, 1975. A monotypic genus endemic to Japan,
<taxonomicName id="FF015E5257DB781A938629E6D1A940B9" class="Arachnida" family="Paranonychidae" genus="Izunonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Izunonychus ohruii" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="ohruii">Izunonychus ohruii</taxonomicName>
Suzuki, 1975 is restricted to the Izu peninsula and Hakone area in central Honshu.
</paragraph>
<paragraph id="387F08D1C170C0E8443DD3D5CBD0E821" pageId="0" pageNumber="1">
<taxonomicName id="85E2F0A0DAE5A63BB6198988A188E580" class="Arachnida" family="Paranonychidae" genus="Kainonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kainonychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Kainonychus</taxonomicName>
Suzuki, 1975 (Figure 1O). A monotypic genus endemic to Japan,
<taxonomicName id="FCFA801CC0BCF96B17EDAE449BE289CE" class="Arachnida" family="Paranonychidae" genus="Kainonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Kainonychus akamai" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="akamai">Kainonychus akamai</taxonomicName>
(Suzuki, 1972) includes two subspecies,
<taxonomicName id="3F092FDC0CCBEBE25D049E0E883D7309" lsidName="K. a." pageId="0" pageNumber="1" rank="subspecies" species="a." subspecies="akamai">K. a. akamai</taxonomicName>
distributed throughout northern Honshu and
<taxonomicName id="B33EB913F84DF9AD716BB7372BDDEE5B" lsidName="K. a." pageId="0" pageNumber="1" rank="subspecies" species="a." subspecies="esoensis">K. a. esoensis</taxonomicName>
restricted to Hokkaido.
</paragraph>
</subSubSection>
<subSubSection id="E764CEBD76FCF0A15726EBAB2C6725EE" pageId="0" pageNumber="1" type="remarks">
<paragraph id="450A303AE37D9C22C7FA00A9BEB1A90E" pageId="0" pageNumber="1">Remarks.</paragraph>
<paragraph id="62D780CD8772BB977A4C6F59EF101677" pageId="0" pageNumber="1">
In this study all genera in the
<taxonomicName id="05A8786DEA804C4CBF0992922F63EF2B" class="Arachnida" family="Paranonychidae" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Paranonychidae</taxonomicName>
have been sampled and the generic relationships are consistent and highly supported across all analyses (Figs 4, 5). Although the study of
<bibRefCitation id="0FC4E1BF20A7CAB2A6BBB5045D31B064" author="Derkarabetian, S" journalOrPublisher="Journal of Theoretical Biology" pageId="0" pageNumber="1" url="https://doi.org/10.1371/journal.pone.0010388" year="2010">Derkarabetian et al. (2010)</bibRefCitation>
only included North American taxa, the relationships of paranonychids recovered here are the same, notably
<taxonomicName id="5400CA6275BFFDF87CCE2EA376FA21F1" class="Arachnida" family="Paranonychidae" genus="Paranonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paranonychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Paranonychus</taxonomicName>
as the earliest diverging genus, and a sister relationship between
<taxonomicName id="9211C9C5C1327A2550548BF0C423C4D8" class="Arachnida" family="Paranonychidae" genus="Sclerobunus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sclerobunus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Sclerobunus</taxonomicName>
and
<taxonomicName id="26FFFB4BFC611414D66FC18872E2DB24" class="Arachnida" family="Paranonychidae" genus="Metanonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Metanonychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Metanonychus</taxonomicName>
. The familial name
<taxonomicName id="A68E3AF9EE247517AB25151A8A24A135" family="Sclerobunidae" lsidName="" pageId="0" pageNumber="1" rank="family">Sclerobunidae</taxonomicName>
has been used previously (
<bibRefCitation id="1551E1EE051E6050DE826CAE78DD6907" author="Giribet, G" journalOrPublisher="Cladistics" pageId="0" pageNumber="1" pagination="408 - 437" title="A multilocus approach to harvestman (Arachnida: Opiliones) phylogeny with emphasis on biogeography and the systematics of Laniatores." url="https://doi.org/10.1111/j.1096-0031.2009.00296.x" volume="26" year="2010">Giribet et al. 2010</bibRefCitation>
) for the &quot;northern triaenonychids&quot;. However,
<taxonomicName id="7183F8B31FEA9D1FBA19E5BC1711ABDE" class="Arachnida" family="Paranonychidae" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Paranonychidae</taxonomicName>
and the subfamily
<taxonomicName id="476594CA899E3672868648C0B32C405C" lsidName="" pageId="0" pageNumber="1" rank="subfamily" subfamily="Paranonychinae">Paranonychinae</taxonomicName>
Briggs, 1971 have priority over the names
<taxonomicName id="E13D6D8FD68BAA5A9C427784C37A8D43" family="Sclerobunidae" lsidName="" pageId="0" pageNumber="1" rank="family">Sclerobunidae</taxonomicName>
and
<taxonomicName id="87A1B0F61D52B099858C09144B3E8FCF" lsidName="" pageId="0" pageNumber="1" rank="subfamily" subfamily="Sclerobuninae">Sclerobuninae</taxonomicName>
<bibRefCitation id="54C58F2534AE72679EC2D04EA8EBDEB6" author="Dumitrescu, DO" journalOrPublisher="Travaux du Museum d'Histoire naturelle (Grigore Antipa)" pageId="0" pageNumber="1" pagination="17 - 30" title="Recherches morphologiques sur l'appareil digestif (intestin moyen) des Gonyleptomorphi (Arachnida, Opilionida)." volume="17" year="1976">Dumitrescu 1976</bibRefCitation>
.
</paragraph>
<paragraph id="041A6CDA145B4551C3221BF457FB1608" pageId="0" pageNumber="1">
The Japanese genera
<taxonomicName id="5A6A16C57059568D55A230860CC18CDA" class="Arachnida" family="Paranonychidae" genus="Metanippononychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Metanippononychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Metanippononychus</taxonomicName>
and
<taxonomicName id="AA8EF87799CB2B50A906EE8ECF1A3D6B" class="Arachnida" family="Paranonychidae" genus="Nippononychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nippononychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Nippononychus</taxonomicName>
show levels of UCE divergence consistent with congeners (Figs 4, 5). Intermediate morphological forms between
<taxonomicName id="207FF64E9132B778404FA3D529460591" class="Arachnida" family="Paranonychidae" genus="Nippononychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nippononychus japonicus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="japonicus">Nippononychus japonicus</taxonomicName>
and
<taxonomicName id="6DA8EC51F4C609AA324EDAB5F1D4A1F8" class="Arachnida" family="Paranonychidae" genus="Metanippononychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Metanippononychus daisenensis" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="daisenensis">Metanippononychus daisenensis</taxonomicName>
can be found where the two species come into contact (Tsurusaki pers. obs.). These genera are differentiated only by tarsal claw structure:
<taxonomicName id="B203F28CAEB9F929DAC1778656BF3327" class="Arachnida" family="Paranonychidae" genus="Metanippononychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Metanippononychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Metanippononychus</taxonomicName>
possessing a ventral tooth on the median prong of the hind claws. The original drawings of male genitalia show that
<taxonomicName id="44295B077A78EB176A8E05BBDBE99363" lsidName="M. daisenensis" pageId="0" pageNumber="1" rank="species" species="daisenensis">M. daisenensis</taxonomicName>
and
<taxonomicName id="1BF4051BC9034A06C665006B2358327D" lsidName="N. japonicus" pageId="0" pageNumber="1" rank="species" species="japonicus">N. japonicus</taxonomicName>
differ in the width of the stylus (
<bibRefCitation id="0FB360F8C01FA7F67DD7E21B47EE438A" pageId="0" pageNumber="1">Suzuki 1975b</bibRefCitation>
). However, the penis of
<taxonomicName id="F967EBEB876860E101D4305B64DEC5A9" lsidName="N. japonicus" pageId="0" pageNumber="1" rank="species" species="japonicus">N. japonicus</taxonomicName>
is highly similar to that of the geographically proximate
<taxonomicName id="42A4016A549862016B6A897ECFED1DD1" lsidName="M. tomishimai" pageId="0" pageNumber="1" rank="subspecies" species="tomishimai" subspecies="tomishimai">M. tomishimai tomishimai</taxonomicName>
.
</paragraph>
<paragraph id="1CD07F019E58E3B3B02C0D2DA898F3B2" pageId="0" pageNumber="1">
<bibRefCitation id="7858ECC304301B20CA53CAB638BD9145" author="Kury, AB" journalOrPublisher="Munis Entomology &amp; Zoology" pageId="0" pageNumber="1" url="https://doi.org/10.3897/BDJ.2.e4094" year="2014">Kury et al. (2014)</bibRefCitation>
includes
<taxonomicName id="01DF0FAB3B532FF9E541417502752A1D" class="Arachnida" family="Paranonychidae" genus="Paranonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paranonychus fuscus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="fuscus">Paranonychus fuscus</taxonomicName>
(formerly
<taxonomicName id="6956D53D9AFFEE173E73D37F198CC2D4" genus="Mutsunonychus" lsidName="Mutsunonychus fuscus" pageId="0" pageNumber="1" rank="species" species="fuscus">Mutsunonychus fuscus</taxonomicName>
) as a synonym of
<taxonomicName id="516D1B72C2DF201E2706D7A4A84E44FC" class="Arachnida" family="Paranonychidae" genus="Paranonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paranonychus brunneus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="brunneus">Paranonychus brunneus</taxonomicName>
(Banks, 1893) based on
<bibRefCitation id="92375C2C97397969B213A524091843B0" author="Shear, WA" journalOrPublisher="American Museum Novitates" pageId="0" pageNumber="1" pagination="1 - 29" title="A cladistic analysis of the opilionid superfamily Ischyropsalidoidea, with descriptions of the new family Ceratolasmatidae, the new genus Acuclavella, and four new species." url="http://hdl.handle.net/2246/3579" volume="2844" year="1986">
<normalizedToken id="4360BB6E09CC659A132715D1434AB54B" originalValue="Shear’s">Shear's</normalizedToken>
(1986)
</bibRefCitation>
statement &quot;
<taxonomicName id="677C8BBF3A2E90991E298B07158782B0" class="Arachnida" family="Paranonychidae" genus="Paranonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paranonychus brunneus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="brunneus">Paranonychus brunneus</taxonomicName>
(=
<taxonomicName id="9FB1DA77C3E2CFDD500DFA775835E9B8" genus="Mutsunonychus" lsidName="Mutsunonychus fuscus" pageId="0" pageNumber="1" rank="species" species="fuscus">Mutsunonychus fuscus</taxonomicName>
Suzuki;
<taxonomicName id="159F402FCB7ECFB4831215701A31E873" class="Arachnida" family="Paranonychidae" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Paranonychidae</taxonomicName>
)&quot;. Later in
<bibRefCitation id="91C4E167BEA3843E668B71BF4ECBB648" author="Shear, WA" journalOrPublisher="Zootaxa" pageId="0" pageNumber="1" pagination="67 - 68" title="Nomenclatorial changes in Triaenonychidae: Sclerobunusparvus Roewer is a junior synonym of Paranonychusbrunneus (Banks), Mutsunonychus Suzuki is a junior synonym of Paranonychus Briggs, and Kaolinonychinae Suzuki is a junior synonym of Paranonychinae Briggs (Opiliones: Triaenonychidae)." volume="1809" year="2008">Shear and Derkarabetian (2008)</bibRefCitation>
, the genus
<taxonomicName id="7849A798F3DADFC32967DCECE55426CF" genus="Mutsunonychus" lsidName="Mutsunonychus" pageId="0" pageNumber="1" rank="genus">Mutsunonychus</taxonomicName>
was formally synonymized under
<taxonomicName id="C79AE420A95BC04E41179120988E3FDA" class="Arachnida" family="Paranonychidae" genus="Paranonychus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paranonychus" order="Opiliones" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Paranonychus</taxonomicName>
, and although a potential species level synonymy was noted, it was not formally established. The levels of UCE divergence between
<taxonomicName id="5CE3D38F26B5D10522676B89275D241D" lsidName="P. brunneus" pageId="0" pageNumber="1" rank="species" species="brunneus">P. brunneus</taxonomicName>
and
<taxonomicName id="9B667853E99DE46B83792DAE50B89E9E" lsidName="P. fuscus" pageId="0" pageNumber="1" rank="species" species="fuscus">P. fuscus</taxonomicName>
are consistent with species level divergences compared to other pairs of congeneric taxa included (Figs 4 and 5), and as such,
<taxonomicName id="90DB770683BF873024E76FF565180CBF" lsidName="P. fuscus" pageId="0" pageNumber="1" rank="species" species="fuscus">P. fuscus</taxonomicName>
is again treated as a distinct species here.
</paragraph>
</subSubSection>
</treatment>
</document>
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