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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.1174.101251" ID-Pensoft-Pub="1313-2970-1174-219" ID-Pensoft-UUID="D0C94D1974975A4B9CD9EE6F18A21120" ID-ZooBank="608FAD80206A428E9743F8ED4F3139BB" ModsDocID="1313-2970-1174-219" checkinTime="1691771919581" checkinUser="pensoft" docAuthor="Ballarin, Francesco &amp; Eguchi, Katsuyuki" docDate="2023" docId="1F03361CB84A5821AA113CFAA248A60E" docLanguage="en" docName="ZooKeys 1174: 219-272" docOrigin="ZooKeys 1174" docPubDate="2023-08-11" docSource="http://dx.doi.org/10.3897/zookeys.1174.101251" docTitle="Howaia alba Ballarin &amp; Eguchi 2023, sp. nov." docType="treatment" docUuid="140385CD-7B9B-421C-9752-7182907C5369" docUuidSource="ZooBank" docVersion="1" id="D0C94D1974975A4B9CD9EE6F18A21120" lastPageNumber="219" masterDocId="D0C94D1974975A4B9CD9EE6F18A21120" masterDocTitle="Integrative taxonomic revision of the genera Nesticella and Howaia in Japan with the description of five new species (Araneae, Nesticidae, Nesticellini)" masterLastPageNumber="272" masterPageNumber="219" pageNumber="219" updateTime="1691771919581" updateUser="pensoft">
<mods:mods id="F47B1C6BB9AA65088DCFADA030C599D2" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="470D97CF5952E565B0D9B05E79478108">
<mods:title id="5CDCC0DB6CF07E4C267778C6AB8C7B5C">Integrative taxonomic revision of the genera Nesticella and Howaia in Japan with the description of five new species (Araneae, Nesticidae, Nesticellini)</mods:title>
</mods:titleInfo>
<mods:name id="F1EF72ADD059C9D370552CAFE1DB24A8" type="personal">
<mods:role id="74A31BE83CCE37F0035FC6F08696D241">
<mods:roleTerm id="56AB6168533261487E798A02683F4419">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="4C8621845887D437AF924C79ED39BC3B">Ballarin, Francesco</mods:namePart>
<mods:nameIdentifier id="25846EA2BA761EF53AE9AF42CFBC2DDF" type="ORCID">https://orcid.org/0000-0003-1417-2519</mods:nameIdentifier>
<mods:affiliation id="33F07BFAA0ED7840999BB5A9AD977758">Systematic Zoology Laboratory, Department of Biological Sciences, Tokyo Metropolitan University, 1 - 1 Minami-Osawa, Hachioji-shi, 192 - 0397, Tokyo, Japan &amp; Department of Zoology, Museo di Storia Naturale of Verona, Lungadige Porta Vittoria, 9, I- 37129 Verona, Italy</mods:affiliation>
<mods:nameIdentifier id="F20023D3A9298DE8EDE36776C2B8E499" type="email">ballarin.francesco@gmail.com</mods:nameIdentifier>
</mods:name>
<mods:name id="ADA0A28F65A5AF19DC901DC3589D8A20" type="personal">
<mods:role id="8A355E5165C3BE56349CDB4AD0176277">
<mods:roleTerm id="14E8172357361572AEF9994DAE067A9D">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="FD2BA7CF4A72CBFE79EBC25DBF0DE412">Eguchi, Katsuyuki</mods:namePart>
<mods:nameIdentifier id="407193DC42BB9EF9948B2CA039409545" type="ORCID">https://orcid.org/0000-0002-1054-1295</mods:nameIdentifier>
<mods:affiliation id="3D54FB6B01870EB9384FC556BE67DBAC">Systematic Zoology Laboratory, Department of Biological Sciences, Tokyo Metropolitan University, 1 - 1 Minami-Osawa, Hachioji-shi, 192 - 0397, Tokyo, Japan &amp; Department of International Health and Medical Anthropology, Institute of Tropical Medicine, Nagasaki University, 1 - 12 - 4 Sakamoto, Nagasaki-shi, 852 - 8523, Nagasaki, Japan</mods:affiliation>
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<mods:title id="1B580259090051868DEF20803350984D">ZooKeys</mods:title>
</mods:titleInfo>
<mods:part id="247F4F8E3AAE4EA71A1283A42DFCF15E">
<mods:date id="925BB3846DA144A7411BD07DDA4C64B5">2023</mods:date>
<mods:detail id="65124A72FA62E0BB6747F6ED3ADB31B9" type="pubDate">
<mods:number id="6257FC30AB72EB0BFA8F757C2343C715">2023-08-11</mods:number>
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<mods:detail id="D3E5893709C80D6BB7E6439A338C4357" type="volume">
<mods:number id="1869A62571500A2615B26D25E7050A32">1174</mods:number>
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<mods:start id="197EE9A2012FF3BFE5854AAD1A2AEA18">219</mods:start>
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<mods:location id="7552A605CDF9397B23F7CA408887BE15">
<mods:url id="4BC3E9B6D83C5085F912C17C82F70A36">http://dx.doi.org/10.3897/zookeys.1174.101251</mods:url>
</mods:location>
<mods:classification id="BD8A4EF18DA192861281BD359414531A">journal article</mods:classification>
<mods:identifier id="35532A76A026C984F96D516351F87351" type="DOI">http://dx.doi.org/10.3897/zookeys.1174.101251</mods:identifier>
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<mods:identifier id="53D788AAAF704379B11C337E3CAE0A2A" type="ZooBank">608FAD80206A428E9743F8ED4F3139BB</mods:identifier>
<mods:identifier id="B24720DE8E529FED70A80B08F2DB78F8" type="Pensoft-UUID">D0C94D1974975A4B9CD9EE6F18A21120</mods:identifier>
</mods:mods>
<treatment id="1F03361CB84A5821AA113CFAA248A60E" LSID="urn:lsid:zoobank.org:act:140385CD-7B9B-421C-9752-7182907C5369" httpUri="http://treatment.plazi.org/id/1F03361CB84A5821AA113CFAA248A60E" lastPageNumber="219" pageId="0" pageNumber="219">
<subSubSection id="ED9A3CD277CA12318C9F94CEA17A577C" pageId="0" pageNumber="219" type="nomenclature">
<paragraph id="8F5B4F3A2330CB41D0BB8B694854CB4E" pageId="0" pageNumber="219">
<taxonomicName id="CC082A1AE771553BC827054FD17675DD" LSID="https://zoobank.org/140385CD-7B9B-421C-9752-7182907C5369" authority="Ballarin &amp; Eguchi, 2023" authorityName="Ballarin &amp; Eguchi" authorityYear="2023" class="Arachnida" family="Nesticidae" genus="Howaia" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Howaia alba" order="Araneae" pageId="0" pageNumber="219" phylum="Arthropoda" rank="species" species="alba" status="sp. nov.">Howaia alba</taxonomicName>
<taxonomicNameLabel id="107B3A51A553DBDD13C4E6787282BBEC" pageId="0" pageNumber="219">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="9BAA84AB5F9EEE7ACB891DC61F8AB860" pageId="0" pageNumber="219" type="description">
<paragraph id="1176D6E73887C9D1D247502D854DCFAD" pageId="0" pageNumber="219">
<figureCitation id="DE8223001A668F83B3E05D10DF503CB3" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">Figs 2A-J</figureCitation>
<figureCitation id="1F8D7814766614A87EAF41A6DA364F50" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Genitalia of Howaia mogera and H. alba sp. nov. A male palp of H. mogera, ventral view B same, retrolateral view C detail or paracymbium, dorsal view E female epigyne, ventral view E male palp of H. alba sp. nov., ventral view F same, retrolateral view C detail or paracymbium, dorsal view H female epigyne, ventral view. Abbreviations: Cl - lobe of conductor; Cm - median process of conductor; Co - copulatory opening; Cp - prolateral process of conductor; Cr - retrolateral process of conductor; Di - distal process of paracymbium; E - embolus; P - paracymbium; Ra - radical apophysis; Rx - radix; S - spermatheca; Sc - scapus; Sd - sperm duct; St - subtegulum; Te - tegulum; Ve - ventral process of paracymbium. Scale bars: 0.2 mm." figureDoi="10.3897/zookeys.1174.101251.figure4" httpUri="https://binary.pensoft.net/fig/891288" pageId="0" pageNumber="219">, 4E-H</figureCitation>
<figureCitation id="09866DAC077C7858450B419A1476233C" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Nesticella and Howaia species in life and their natural habitats A female of H. mogera B female of N. terrestris C female of H. alba sp. nov. D female of N. occulta sp. nov. E female of N. okinawaensis F example of the epigean habitat of Nesticella species in the Ryukyus G habitat of H. alba sp nov. (dark zone of Tsuzupisuki-abu cave) H habitat of N. occulta sp. nov. (dark zone of Kabirano-ana cave) (all photos by F. Ballarin)." figureDoi="10.3897/zookeys.1174.101251.figure15" httpUri="https://binary.pensoft.net/fig/891299" pageId="0" pageNumber="219">, 15C</figureCitation>
<figureCitation id="01FC54CA510849F6889863E335F4BE31" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Distribution of Nesticella and Howaia species in mainland Japan and Ryukyu Archipelago A distribution of N. brevipes and N. terrestris in mainland Japan and neighboring countries B distribution of Nesticella and Howaia species endemic to the Ryukyu Archipelago C distribution of H. mogera in mainland Japan and neighboring countries D distribution of H. mogera in the Ryukyu Archipelago and neighboring countries. Colored squares refer to the type localities of the species, colored dots indicate the locations of specimens whose morphology has been checked during this study, dashed areas refer to the generally known distribution of the species, question marks refer to unclear distributions (see &quot; remarks on misidentifications &quot; of N. brevipes and N. terrestris)." figureDoi="10.3897/zookeys.1174.101251.figure16" httpUri="https://binary.pensoft.net/fig/891300" pageId="0" pageNumber="219">
, 16B (Japanese name: tsuzupisuki-horahimegumo
<normalizedToken id="E42D8D527C5D9CB16F785C7B35DEBF8E" originalValue="ツヅピスキホラヒメグモ">ツヅピスキホラヒメグモ</normalizedToken>
)
</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="0273451CBE86B3CF52DC5FB6FCB6E6E3" pageId="0" pageNumber="219" type="reference_group">
<paragraph id="5C33AAA74F1E192A023A713F915F61EB" pageId="0" pageNumber="219">
<taxonomicName id="F80CC10A160AF922D18F0631D4281226" authority="Ballarin &amp; Eguchi, 2023" authorityName="Ballarin &amp; Eguchi" authorityYear="2023" class="Arachnida" family="Nesticidae" genus="Nesticella" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Nesticella mogera" order="Araneae" pageId="0" pageNumber="219" phylum="Arthropoda" rank="species" species="mogera" status="sp. nov.">Nesticella mogera</taxonomicName>
<bibRefCitation id="A5FC7B49A4E29E62F0D16C1E3651B04E" DOI="https://doi.org/10.2476/asjaa.27.Specialnumber_337" author="Shimojana, M" journalOrPublisher="Zootaxa" pageId="0" pageNumber="219" refId="B45" refString="Shimojana, M, 1977. Preliminary report on the cave spider fauna of the Ryukyu Archipelago. Acta Arachnologica 27(Spec. No.): 337-365. https://doi.org/10.2476/asjaa.27.Specialnumber_337" title="Preliminary report on the cave spider fauna of the Ryukyu Archipelago. Acta Arachnologica 27 (Spec. No.): 337 - 365." url="https://doi.org/10.2476/asjaa.27.Specialnumber_337" year="1977">Shimojana 1977</bibRefCitation>
: 353, fig. 6 (♂, misidentification).
</paragraph>
</subSubSection>
<subSubSection id="A15C013BE24735F525383872828FC0DB" pageId="0" pageNumber="219" type="type material">
<paragraph id="5D60AC20DF772661508310A7E52F0ED3" pageId="0" pageNumber="219">Type material.</paragraph>
<paragraph id="80DBC3C3B3AB3AFB2A8FBD7344A34848" pageId="0" pageNumber="219">
♂
<emphasis id="D0A8C6159FD1D1E21794E661402EEC13" bold="true" italics="true" pageId="0" pageNumber="219">Holotype</emphasis>
(NMST-Ar. 25251):
<emphasis id="C5CD6CDEC8BA5B171859D1A81D632789" bold="true" pageId="0" pageNumber="219">Japan: Okinawa Pref.: Miyako-jima Is.</emphasis>
: Shimozato Hirara, Oharaminami Park, Tsuzupisuki-abu cave (ツヅピスキアブ), 32 m, long and humid cave, in the dark zone of the cave,
<geoCoordinate id="E9CF57F54643BE44F1F44A8C08526801" degrees="24.79468" direction="north" orientation="latitude" precision="1" value="24.79468">24.79468°N</geoCoordinate>
,
<geoCoordinate id="115145D656E6761C2A81E9B6C51411A9" degrees="125.28192" direction="east" orientation="longitude" precision="1" value="125.28192">125.28192°E</geoCoordinate>
, 12.Nov.2020, F. Ballarin leg.
</paragraph>
<paragraph id="0ACECD5857CB54B581785B18677A4209" pageId="0" pageNumber="219">
<emphasis id="EADB30F60390EF5DA47DC3FF1BF56BE2" bold="true" pageId="0" pageNumber="219">
<emphasis id="7A55D0DC9384AEE199C7B5E8D5C924DD" italics="true" pageId="0" pageNumber="219">Paratypes</emphasis>
: Japan: Miyako-jima Is.
</emphasis>
: 3♀, same data as the holotype (NSMT-Ar 25252); 1♀, same locality 14.Nov.2020, F. Ballarin leg. (RMUF); 5♀, same locality, 16.Sep.2022, F. Ballarin leg. (2♀ MNHAH, 3♀ FBPC); 4♀, Nobaru Ueno, Pinza-abu cave (ピンザアブ洞穴), 57 m, long and muddy cave, in the dark zone of the cave,
<geoCoordinate id="8FE191541C46EDCC0AB0E4BDD09D6C10" degrees="24.74853" direction="north" orientation="latitude" precision="1" value="24.74853">24.74853°N</geoCoordinate>
,
<geoCoordinate id="B08CB605E229092F40D6D4E8CD625A5C" degrees="125.33443" direction="east" orientation="longitude" precision="1" value="125.33443">125.33443°E</geoCoordinate>
, 13.Nov.2020, F. Ballarin leg. (RMUF); 3♀, same locality, 17.Sep.2022, F. Ballarin leg. (FBPC).
</paragraph>
</subSubSection>
<subSubSection id="AA2B17E429559F0B6F423DB51FA915F5" pageId="0" pageNumber="219" type="materials_examined">
<paragraph id="D5BDD17DA447F7CCE880FA089DE2CA11" pageId="0" pageNumber="219">Other material examined.</paragraph>
<paragraph id="607F14E020986BA8660954E5DFF06A47" pageId="0" pageNumber="219">
<materialsCitation id="2CDD29A381B0E5E97ECF567E5BAD4EDA" collectingDate="2021-12-29" collectorName="R. Miyata" country="Japan" latitude="24.73384" location="Nakabari Limestone Cave" longLatPrecision="1" longitude="125.3761" municipality="Nakabari" specimenCount="1" specimenCount-juvenile="1" stateProvince="Miyako-jima Is.">
<emphasis id="4AB55424080ADECD9507D4504C74151E" bold="true" pageId="0" pageNumber="219">
<collectingCountry id="E13F084E496C6B0E0D1658BF83A00511" name="Japan">Japan</collectingCountry>
:
<collectingRegion id="0D41355F7851CC0D5EDE81B49146A147">Miyako-jima Is.</collectingRegion>
</emphasis>
:
<specimenCount id="6A5A2A9959762B2184C048ADCB324467" type="generic">1 juv.</specimenCount>
,
<collectingMunicipality id="78991FF68FD4463C6FDB51C7FC27BEBA">Nakabari</collectingMunicipality>
,
<location id="B6EB206599786E793E16388AD89377D0" LSID="urn:lsid:plazi:treatment:1F03361CB84A5821AA113CFAA248A60E:B6EB206599786E793E16388AD89377D0" country="Japan" latitude="24.73384" longLatPrecision="1" longitude="125.3761" municipality="Nakabari" name="Nakabari Limestone Cave" stateProvince="Miyako-jima Is.">Nakabari Limestone Cave</location>
(仲原鍾乳洞),
<geoCoordinate id="E2E8A591DEC796ED60A3EB9BB496A879" degrees="24.73384" direction="north" orientation="latitude" precision="1" value="24.73384">24.73384°N</geoCoordinate>
,
<geoCoordinate id="C00049EF592DC6319F549EBC806A5E28" degrees="125.37610" direction="east" orientation="longitude" precision="1" value="125.3761">125.37610°E</geoCoordinate>
,
<collectingDate id="223707F10AFE4211E26F04EC58F33E3F" value="2021-12-29">29.Dec.2021</collectingDate>
,
<collectorName id="907FE09EF8E2985CDBAAA915DB558AD3">R. Miyata</collectorName>
leg. (FBPC)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="61029EF4875858BBC3EA73982AF438CD" pageId="0" pageNumber="219" type="etymology">
<paragraph id="C2E49F8FFF1F52C919833E88AF0504B9" pageId="0" pageNumber="219">Etymology.</paragraph>
<paragraph id="71316081470E9A3A593B0E8FD8A652E8" pageId="0" pageNumber="219">
The specific name is derived from the Latin word for the color white (
<emphasis id="F07F95B08AF6C35CD99103FCEB6CFF16" italics="true" pageId="0" pageNumber="219">albus</emphasis>
, adjective) referring to the whitish coloration of the species.
</paragraph>
</subSubSection>
<subSubSection id="02AA17F4AF1F5493C1965E6554DF88CD" pageId="0" pageNumber="219" type="diagnosis">
<paragraph id="520B3A424A111F34503AAF1E61DDECDE" pageId="0" pageNumber="219">Diagnosis.</paragraph>
<paragraph id="CA96227FD1B9B356CA63C6D02F51E7C9" pageId="0" pageNumber="219">
The new species is similar to
<taxonomicName id="690F4A922AE33B5749A5D821A40F591F" family="Nesticidae" kingdom="Animalia" lsidName="H. mogera" order="Araneae" pageId="0" pageNumber="219" rank="species" species="mogera">
<emphasis id="1BFAEC177D9BE002E2803376886D0675" italics="true" pageId="0" pageNumber="219">H. mogera</emphasis>
</taxonomicName>
and to the troglobitic species
<taxonomicName id="0C7B2EEBD2FA447F72B8202DAC716D26" family="Nesticidae" kingdom="Animalia" lsidName="H. rongtangensis" order="Araneae" pageId="0" pageNumber="219" rank="species" species="rongtangensis">
<emphasis id="4165FFB9A3BEDFC49B1EB25B1741A69E" italics="true" pageId="0" pageNumber="219">H. rongtangensis</emphasis>
</taxonomicName>
(Lin, Ballarin &amp; Li, 2016) from Hainan Island,
<taxonomicName id="95C5B2B961F801DFF8298A4B4F402BA0" family="Nesticidae" kingdom="Animalia" lsidName="H. subterranea" order="Araneae" pageId="0" pageNumber="219" rank="species" species="subterranea">
<emphasis id="32A107D501E24FEC87EA36B4981E1181" italics="true" pageId="0" pageNumber="219">H. subterranea</emphasis>
</taxonomicName>
sp. nov., and
<taxonomicName id="99E9F5E56AD8EB86D545B250A9C8C6A3" family="Nesticidae" kingdom="Animalia" lsidName="N. occulta" order="Araneae" pageId="0" pageNumber="219" rank="species" species="occulta">
<emphasis id="8BD41B1DDFAF57AE9C124C9CB0635390" italics="true" pageId="0" pageNumber="219">N. occulta</emphasis>
</taxonomicName>
sp. nov. Male of
<taxonomicName id="2AA43817AC2484A2058570D431398CA1" family="Nesticidae" kingdom="Animalia" lsidName="H. alba" order="Araneae" pageId="0" pageNumber="219" rank="species" species="alba">
<emphasis id="72C68505C7D8EACE54686EAB7F25CE81" italics="true" pageId="0" pageNumber="219">H. alba</emphasis>
</taxonomicName>
sp. nov. can be distinguished from male of
<taxonomicName id="BC0FF74D40209960E9CD10E77B69E26D" family="Nesticidae" kingdom="Animalia" lsidName="H. mogera" order="Araneae" pageId="0" pageNumber="219" rank="species" species="mogera">
<emphasis id="0B3D2C1AC508F53DD7633F6D7EB93D0E" italics="true" pageId="0" pageNumber="219">H. mogera</emphasis>
</taxonomicName>
and
<taxonomicName id="DF32BF6529F415D1D2A046C334A46637" family="Nesticidae" kingdom="Animalia" lsidName="H. rongtangensis" order="Araneae" pageId="0" pageNumber="219" rank="species" species="rongtangensis">
<emphasis id="2050894F5BCFFFC19A1063D2F33CBFBD" italics="true" pageId="0" pageNumber="219">H. rongtangensis</emphasis>
</taxonomicName>
by the different shape of the larger paracymbium (P) bearing a longer, slimmer, and sharper distal process (Di) (vs slimmer P with a shorter, larger, and blunter Di in
<taxonomicName id="51A82AB6BFD8F5C82F5B677BC271BB95" family="Nesticidae" kingdom="Animalia" lsidName="H. mogera" order="Araneae" pageId="0" pageNumber="219" rank="species" species="mogera">
<emphasis id="82CBB051C184CD910A157EA0661D7299" italics="true" pageId="0" pageNumber="219">H. mogera</emphasis>
</taxonomicName>
and
<taxonomicName id="E23393A0723ACDBFE27361AF1360807E" family="Nesticidae" kingdom="Animalia" lsidName="H. rongtangensis" order="Araneae" pageId="0" pageNumber="219" rank="species" species="rongtangensis">
<emphasis id="9C7EE8D9D03772FBBCA5CCE6AEA90BFD" italics="true" pageId="0" pageNumber="219">H. rongtangensis</emphasis>
</taxonomicName>
) (Figs
<figureCitation id="B8A1590037976503A27AE36C0795044F" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">2A-D</figureCitation>
,
<figureCitation id="AA54F71D6FEC6CBA3AAAF73F1BD3EB52" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Genitalia of Howaia mogera and H. alba sp. nov. A male palp of H. mogera, ventral view B same, retrolateral view C detail or paracymbium, dorsal view E female epigyne, ventral view E male palp of H. alba sp. nov., ventral view F same, retrolateral view C detail or paracymbium, dorsal view H female epigyne, ventral view. Abbreviations: Cl - lobe of conductor; Cm - median process of conductor; Co - copulatory opening; Cp - prolateral process of conductor; Cr - retrolateral process of conductor; Di - distal process of paracymbium; E - embolus; P - paracymbium; Ra - radical apophysis; Rx - radix; S - spermatheca; Sc - scapus; Sd - sperm duct; St - subtegulum; Te - tegulum; Ve - ventral process of paracymbium. Scale bars: 0.2 mm." figureDoi="10.3897/zookeys.1174.101251.figure4" httpUri="https://binary.pensoft.net/fig/891288" pageId="0" pageNumber="219">4E-G</figureCitation>
cf. Figs
<figureCitation id="7FF25FF390B42ADA969CF7E42641A3C2" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Howaia mogera A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne ventral view (specimen from Tokyo area) F same after dissection, shape variation G vulva, dorsal view H habitus of male (specimen from Tokyo area) I habitus of female J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure1" httpUri="https://binary.pensoft.net/fig/891285" pageId="0" pageNumber="219">1A-D</figureCitation>
,
<figureCitation id="10118A77FF847D54FA8A6E4E70E000EF" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Genitalia of Howaia mogera and H. alba sp. nov. A male palp of H. mogera, ventral view B same, retrolateral view C detail or paracymbium, dorsal view E female epigyne, ventral view E male palp of H. alba sp. nov., ventral view F same, retrolateral view C detail or paracymbium, dorsal view H female epigyne, ventral view. Abbreviations: Cl - lobe of conductor; Cm - median process of conductor; Co - copulatory opening; Cp - prolateral process of conductor; Cr - retrolateral process of conductor; Di - distal process of paracymbium; E - embolus; P - paracymbium; Ra - radical apophysis; Rx - radix; S - spermatheca; Sc - scapus; Sd - sperm duct; St - subtegulum; Te - tegulum; Ve - ventral process of paracymbium. Scale bars: 0.2 mm." figureDoi="10.3897/zookeys.1174.101251.figure4" httpUri="https://binary.pensoft.net/fig/891288" pageId="0" pageNumber="219">4A-C</figureCitation>
and
<bibRefCitation id="8128408B6ADC0208C0D40D28B185C6C7" DOI="https://doi.org/10.3897/zookeys.627.8629" author="Lin, YC" journalOrPublisher="ZooKeys" pageId="0" pageNumber="219" pagination="1 - 168" refId="B28" refString="Lin, YC, Ballarin, F, Li, SQ, 2016. A survey of the spider family Nesticidae (Arachnida, Araneae) in Asia and Madagascar, with the description of forty-three new species. ZooKeys 627: 1 - 168, DOI: https://doi.org/10.3897/zookeys.627.8629" title="A survey of the spider family Nesticidae (Arachnida, Araneae) in Asia and Madagascar, with the description of forty-three new species." url="https://doi.org/10.3897/zookeys.627.8629" volume="627" year="2016">Lin et al. 2016</bibRefCitation>
: fig. 44A, B, D). Female of the new species are distinguished from female of
<taxonomicName id="A7B32FEBBE2D51B19949CA96DB5140CE" family="Nesticidae" kingdom="Animalia" lsidName="H. mogera" order="Araneae" pageId="0" pageNumber="219" rank="species" species="mogera">
<emphasis id="1CEC8853A74B1723114EB7F97FC691D7" italics="true" pageId="0" pageNumber="219">H. mogera</emphasis>
</taxonomicName>
,
<taxonomicName id="DF8B4236891021C3D93F8DCF0A8C9967" family="Nesticidae" kingdom="Animalia" lsidName="H. subterranea" order="Araneae" pageId="0" pageNumber="219" rank="species" species="subterranea">
<emphasis id="69D07526A46ADAB58D6D3D52E99BD940" italics="true" pageId="0" pageNumber="219">H. subterranea</emphasis>
</taxonomicName>
sp. nov., and
<taxonomicName id="C5838CED059C93A578199CB6F6DE6B2B" family="Nesticidae" kingdom="Animalia" lsidName="N. occulta" order="Araneae" pageId="0" pageNumber="219" rank="species" species="occulta">
<emphasis id="B6655BCCCC3D7B550C16270F8DE48F2C" italics="true" pageId="0" pageNumber="219">N. occulta</emphasis>
</taxonomicName>
sp. nov. by the larger and stockier scapus (Sc) with a slightly rounded posterior margin (vs slimmer Sc with a flat posterior margin in
<taxonomicName id="E67ABD5234B0185A42C19F0CB0647012" family="Nesticidae" kingdom="Animalia" lsidName="H. mogera" order="Araneae" pageId="0" pageNumber="219" rank="species" species="mogera">
<emphasis id="AB3CF15D74C478FAAD8C59005C2C2BA5" italics="true" pageId="0" pageNumber="219">H. mogera</emphasis>
</taxonomicName>
, a longer Sc with a wider lobated tip in
<taxonomicName id="1A042FAE342F7279B0E8EC20A17C67AA" family="Nesticidae" kingdom="Animalia" lsidName="H. subterranea" order="Araneae" pageId="0" pageNumber="219" rank="species" species="subterranea">
<emphasis id="D6A759655A2EAE8E0785F9AC5CC03C29" italics="true" pageId="0" pageNumber="219">H. subterranea</emphasis>
</taxonomicName>
sp. nov., and a slimmer, tongue-like Sc ending with a strongly concave tip in
<taxonomicName id="D33F3818EC6EC84E29824EC068CBE95E" family="Nesticidae" kingdom="Animalia" lsidName="N. occulta" order="Araneae" pageId="0" pageNumber="219" rank="species" species="occulta">
<emphasis id="6459AE76F10E09CD7C63B203BB7E77A8" italics="true" pageId="0" pageNumber="219">N. occulta</emphasis>
</taxonomicName>
sp. nov.) (Figs
<figureCitation id="A9FC3BBD157BDC2BB76CAE14FF98FB85" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">2E-G</figureCitation>
,
<figureCitation id="E6B23C50BA9B6EFB0C3CAA2C80477FB5" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Genitalia of Howaia mogera and H. alba sp. nov. A male palp of H. mogera, ventral view B same, retrolateral view C detail or paracymbium, dorsal view E female epigyne, ventral view E male palp of H. alba sp. nov., ventral view F same, retrolateral view C detail or paracymbium, dorsal view H female epigyne, ventral view. Abbreviations: Cl - lobe of conductor; Cm - median process of conductor; Co - copulatory opening; Cp - prolateral process of conductor; Cr - retrolateral process of conductor; Di - distal process of paracymbium; E - embolus; P - paracymbium; Ra - radical apophysis; Rx - radix; S - spermatheca; Sc - scapus; Sd - sperm duct; St - subtegulum; Te - tegulum; Ve - ventral process of paracymbium. Scale bars: 0.2 mm." figureDoi="10.3897/zookeys.1174.101251.figure4" httpUri="https://binary.pensoft.net/fig/891288" pageId="0" pageNumber="219">4H</figureCitation>
cf. Figs
<figureCitation id="147D51E52504A1B94A30FBE738931E5E" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Howaia mogera A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne ventral view (specimen from Tokyo area) F same after dissection, shape variation G vulva, dorsal view H habitus of male (specimen from Tokyo area) I habitus of female J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure1" httpUri="https://binary.pensoft.net/fig/891285" pageId="0" pageNumber="219">1E-G</figureCitation>
,
<figureCitation id="D7B20E48DC91B3D52EE6E18B8A560ADF" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Howaia subterranea sp. nov. A habitus of female (holotype) B cephalic area of female, frontal view C habitus of female, lateral view D female epigyne, ventral view E vulva, dorsal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 1.0 mm (A, C); 0.2 mm (B, D, E)." figureDoi="10.3897/zookeys.1174.101251.figure3" httpUri="https://binary.pensoft.net/fig/891287" pageId="0" pageNumber="219">3D, E</figureCitation>
,
<figureCitation id="08CBE790974144B3CE05DEBEF063319B" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Genitalia of Howaia mogera and H. alba sp. nov. A male palp of H. mogera, ventral view B same, retrolateral view C detail or paracymbium, dorsal view E female epigyne, ventral view E male palp of H. alba sp. nov., ventral view F same, retrolateral view C detail or paracymbium, dorsal view H female epigyne, ventral view. Abbreviations: Cl - lobe of conductor; Cm - median process of conductor; Co - copulatory opening; Cp - prolateral process of conductor; Cr - retrolateral process of conductor; Di - distal process of paracymbium; E - embolus; P - paracymbium; Ra - radical apophysis; Rx - radix; S - spermatheca; Sc - scapus; Sd - sperm duct; St - subtegulum; Te - tegulum; Ve - ventral process of paracymbium. Scale bars: 0.2 mm." figureDoi="10.3897/zookeys.1174.101251.figure4" httpUri="https://binary.pensoft.net/fig/891288" pageId="0" pageNumber="219">4D</figureCitation>
,
<figureCitation id="45241E8B982BEE9BFFFC000731438C95" captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Nesticella occulta sp. nov. A habitus of female (holotype) B cephalic area of female, frontal view C female epigyne, ventral view D same, shape variation E vulva, dorsal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 1.0 mm (A, C); 0.2 mm (B, D, E)." figureDoi="10.3897/zookeys.1174.101251.figure11" httpUri="https://binary.pensoft.net/fig/891295" pageId="0" pageNumber="219">11C, D</figureCitation>
,
<figureCitation id="E7F1DCEDCCA0F73586D23E853503F481" captionStart="Figure 13" captionStartId="F13" captionText="Figure 13. Genitalia of Nesticella okinawaensis, Howaia subterranea sp. nov., and N. occulta sp. nov. A male palp of N. okinawaensis, ventral view B same, retrolateral view C detail or paracymbium, dorsal view D female epigyne, ventral view; E H. subterranea sp. nov., female epigyne, ventral view F N. occulta sp. nov. female epigyne, ventral view. Abbreviations: Ca - apophysis of the retrolateral process of conductor; Cl - lobe of conductor; Cm - median process of conductor; Co - copulatory opening; Cp - prolateral process of conductor; Cr - retrolateral process of conductor; Di I-II - distal processes I and II of paracymbium; E - embolus; P - paracymbium; Ra - radical apophysis; Rx - radix; S - spermatheca; Sc - scapus; Sd - sperm duct; St - subtegulum; Te - tegulum; Ve - ventral process of paracymbium. Scale bars: 0.2 mm." figureDoi="10.3897/zookeys.1174.101251.figure13" httpUri="https://binary.pensoft.net/fig/891297" pageId="0" pageNumber="219">13E, F</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="E083C342B31EC8EE55CF6B175A6CB5C8" pageId="0" pageNumber="219" type="description">
<paragraph id="A42E4EFF230632EB85EEFC5CA720DF3B" pageId="0" pageNumber="219">Description of male</paragraph>
<paragraph id="2B7479473F41BF630656574755787685" pageId="0" pageNumber="219">
<emphasis id="834CAF26392DCA8E0556AE36121B7C8D" bold="true" pageId="0" pageNumber="219">(holotype).</emphasis>
Habitus as in Fig.
<figureCitation id="FCA1CC6F0C92668563935EB8B29EF25B" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">2H</figureCitation>
. Total length 1.88. Prosoma 0.94 long, 0.83 wide. Carapace uniformly pale yellowish. Eyes completely degenerated and reduced to white maculae. Cervical groove and fovea indistinct. Chelicerae brownish. Labium, maxillae, and sternum of the same pale color as carapace. Legs uniformly pale yellowish. Leg formula: I, IV, II, III. Legs measurements as follows: I 6.17 (1.61, 0.47, 1.60, 1.72, 0.77), II 4.71 (1.32, 0.39, 1.20, 1.18, 0.62), III 3.89 (1.11, 0.31, 0.95, 1.02, 0.50), IV 5.17 (1.50, 0.37, 1.34, 1.31, 0.65). Opisthosoma uniformly greyish-yellow, covered with long, sparse hairs.
</paragraph>
<paragraph id="6D54126D087882B54079CA814DC4DD33" pageId="0" pageNumber="219">
Male palp as in Figs
<figureCitation id="47792C1DCC183FC5D409042E650F9EEE" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">2A-D</figureCitation>
,
<figureCitation id="848A319F5A3BD318579C4CD63EC5D5DB" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Genitalia of Howaia mogera and H. alba sp. nov. A male palp of H. mogera, ventral view B same, retrolateral view C detail or paracymbium, dorsal view E female epigyne, ventral view E male palp of H. alba sp. nov., ventral view F same, retrolateral view C detail or paracymbium, dorsal view H female epigyne, ventral view. Abbreviations: Cl - lobe of conductor; Cm - median process of conductor; Co - copulatory opening; Cp - prolateral process of conductor; Cr - retrolateral process of conductor; Di - distal process of paracymbium; E - embolus; P - paracymbium; Ra - radical apophysis; Rx - radix; S - spermatheca; Sc - scapus; Sd - sperm duct; St - subtegulum; Te - tegulum; Ve - ventral process of paracymbium. Scale bars: 0.2 mm." figureDoi="10.3897/zookeys.1174.101251.figure4" httpUri="https://binary.pensoft.net/fig/891288" pageId="0" pageNumber="219">4E-G</figureCitation>
. Cymbium relatively elongated, covered with thin sparse setae, bearing some thicker setae on the distal-prolateral margin (Fig.
<figureCitation id="83BB451508CCDF46B4434B9BA6A15D9D" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">2D</figureCitation>
). Paracymbium with a single distinctly sclerotized, stocky distal process (Di), slightly elongated near the tip and a single sharp, spine-like ventral process (Ve) (Figs
<figureCitation id="03CAD31B5D74B1300AC61CA20AC4F8CB" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">2A-D</figureCitation>
,
<figureCitation id="51D7B9945B6D6392EE76D13351AAA15F" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Genitalia of Howaia mogera and H. alba sp. nov. A male palp of H. mogera, ventral view B same, retrolateral view C detail or paracymbium, dorsal view E female epigyne, ventral view E male palp of H. alba sp. nov., ventral view F same, retrolateral view C detail or paracymbium, dorsal view H female epigyne, ventral view. Abbreviations: Cl - lobe of conductor; Cm - median process of conductor; Co - copulatory opening; Cp - prolateral process of conductor; Cr - retrolateral process of conductor; Di - distal process of paracymbium; E - embolus; P - paracymbium; Ra - radical apophysis; Rx - radix; S - spermatheca; Sc - scapus; Sd - sperm duct; St - subtegulum; Te - tegulum; Ve - ventral process of paracymbium. Scale bars: 0.2 mm." figureDoi="10.3897/zookeys.1174.101251.figure4" httpUri="https://binary.pensoft.net/fig/891288" pageId="0" pageNumber="219">4E-G</figureCitation>
). Embolus (E) long and filiform, origin of embolus positioned at ~ 6:00
<normalizedToken id="44ECE957A09745E0DB72B8F108320DD2" originalValue="o’clock">o'clock</normalizedToken>
on radix (Rx). Radical apophysis (Ra) broad, with a granulate surface. Conductor with 3 distinct processes (Cp, Cr, Cm) and a half-transparent distal lobe (Cl). Prolateral process of the conductor (Cp) flat, ribbon-like and headed counterclockwise, wrapped around embolus. Retrolateral process of conductor (Cr) wide and thick, curved internally, with a broadened, flat central part. Median process of conductor (Cm) stout, horn-like, strongly sclerotized bearing a smaller, stout ventral process. (Figs
<figureCitation id="A031977161FF85385950BEE0D5347C24" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">2A-C</figureCitation>
,
<figureCitation id="392150AF7E45802FF53007FFA609F1B6" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Genitalia of Howaia mogera and H. alba sp. nov. A male palp of H. mogera, ventral view B same, retrolateral view C detail or paracymbium, dorsal view E female epigyne, ventral view E male palp of H. alba sp. nov., ventral view F same, retrolateral view C detail or paracymbium, dorsal view H female epigyne, ventral view. Abbreviations: Cl - lobe of conductor; Cm - median process of conductor; Co - copulatory opening; Cp - prolateral process of conductor; Cr - retrolateral process of conductor; Di - distal process of paracymbium; E - embolus; P - paracymbium; Ra - radical apophysis; Rx - radix; S - spermatheca; Sc - scapus; Sd - sperm duct; St - subtegulum; Te - tegulum; Ve - ventral process of paracymbium. Scale bars: 0.2 mm." figureDoi="10.3897/zookeys.1174.101251.figure4" httpUri="https://binary.pensoft.net/fig/891288" pageId="0" pageNumber="219">4E, F</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="EB1961BDFE32AA9C52FC1BCB99DF7DCC" pageId="0" pageNumber="219" type="description">
<paragraph id="9CDBF5F125EC9C1B0E1D88F9AE6F5BCE" pageId="0" pageNumber="219">Description of female</paragraph>
<paragraph id="7171D4CF23FD1CC2A7A5619FFA2555FF" pageId="0" pageNumber="219">
<emphasis id="45142241C3F5C52D19E69BA9BAD1A2B9" bold="true" pageId="0" pageNumber="219">(one of the paratypes).</emphasis>
Habitus as in Figs
<figureCitation id="955D9E0BEA6E313BC2834D5211E7E97E" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">2I</figureCitation>
,
<figureCitation id="1854870B9896795446176143A5133FD9" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Nesticella and Howaia species in life and their natural habitats A female of H. mogera B female of N. terrestris C female of H. alba sp. nov. D female of N. occulta sp. nov. E female of N. okinawaensis F example of the epigean habitat of Nesticella species in the Ryukyus G habitat of H. alba sp nov. (dark zone of Tsuzupisuki-abu cave) H habitat of N. occulta sp. nov. (dark zone of Kabirano-ana cave) (all photos by F. Ballarin)." figureDoi="10.3897/zookeys.1174.101251.figure15" httpUri="https://binary.pensoft.net/fig/891299" pageId="0" pageNumber="219">15C</figureCitation>
. Total length 2.3. Prosoma 1.09 long, 0.92 wide. Cephalic area as in Fig.
<figureCitation id="5F39871B10B3805D7E07625EA9302C65" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">2J</figureCitation>
. Coloration and other details as in male. Legs measurements as follows: I 6.56 (1.83, 0.50, 1.74, 1.67, 0.82), II 5.09 (1.46, 0.42, 1.27, 1.22, 0.72), III 4.09 (1.31, 0.36, 0.90, 0.92, 0.60), IV 5.46 (1.65, 0.45, 1.42, 1.27, 0.67).
</paragraph>
<paragraph id="E32714AAB84086C356284B53D4E98462" pageId="0" pageNumber="219">
Epigyne and vulva as in Figs
<figureCitation id="C2E187557F915D35BD3FDA85578C46AA" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">2E-G</figureCitation>
,
<figureCitation id="4A6FB51BEA2D83F6708DC6E037E36D0A" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Genitalia of Howaia mogera and H. alba sp. nov. A male palp of H. mogera, ventral view B same, retrolateral view C detail or paracymbium, dorsal view E female epigyne, ventral view E male palp of H. alba sp. nov., ventral view F same, retrolateral view C detail or paracymbium, dorsal view H female epigyne, ventral view. Abbreviations: Cl - lobe of conductor; Cm - median process of conductor; Co - copulatory opening; Cp - prolateral process of conductor; Cr - retrolateral process of conductor; Di - distal process of paracymbium; E - embolus; P - paracymbium; Ra - radical apophysis; Rx - radix; S - spermatheca; Sc - scapus; Sd - sperm duct; St - subtegulum; Te - tegulum; Ve - ventral process of paracymbium. Scale bars: 0.2 mm." figureDoi="10.3897/zookeys.1174.101251.figure4" httpUri="https://binary.pensoft.net/fig/891288" pageId="0" pageNumber="219">4H</figureCitation>
. Scapus (Sc) short and stumpy, approximately as long as wide, ending with a slightly rounded posterior margin (Figs
<figureCitation id="16564DF54E437DE877DE569D6E100B33" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">2E, F</figureCitation>
,
<figureCitation id="459653F184E1A0A89CE5451FA2BCF8D7" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Genitalia of Howaia mogera and H. alba sp. nov. A male palp of H. mogera, ventral view B same, retrolateral view C detail or paracymbium, dorsal view E female epigyne, ventral view E male palp of H. alba sp. nov., ventral view F same, retrolateral view C detail or paracymbium, dorsal view H female epigyne, ventral view. Abbreviations: Cl - lobe of conductor; Cm - median process of conductor; Co - copulatory opening; Cp - prolateral process of conductor; Cr - retrolateral process of conductor; Di - distal process of paracymbium; E - embolus; P - paracymbium; Ra - radical apophysis; Rx - radix; S - spermatheca; Sc - scapus; Sd - sperm duct; St - subtegulum; Te - tegulum; Ve - ventral process of paracymbium. Scale bars: 0.2 mm." figureDoi="10.3897/zookeys.1174.101251.figure4" httpUri="https://binary.pensoft.net/fig/891288" pageId="0" pageNumber="219">4H</figureCitation>
). Copulatory opening (Co) at the inner-lateral sides of scapus. Internal ducts slightly visible through the transparent tegument, shaped as a narrow V. Copulatory ducts (Cd) short and thick, slightly divergent to each other, slightly twisted in the inner trait with 1 coil, curving outward and then inward before reaching the spermathecae. Insemination ducts (Id) thin, coiled around the Cd). Spermathecae (S) small and rounded, separated from each other by ~ 2
<normalizedToken id="DE7B078C70B15E4BCCE4D9F0E2BA3769" originalValue="×">x</normalizedToken>
their diameter (Fig.
<figureCitation id="B6B7FE110FDEB57405CA9961BD23B2A0" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Howaia alba sp. nov. A male palp (holotype), ventral view B same, retrolateral view C same, ventro-prolateral view D same, dorsal view E female epigyne (one of the paratypes), ventral view F same, shape variation G vulva, dorsal view H habitus of male (holotype) I habitus of female (one of the paratypes) J cephalic area of female, frontal view. Abbreviations: Cd - copulatory duct; Co - copulatory opening; Id - insemination duct; S - spermatheca; Sc - scapus. Scale bars: 0.2 mm (A-G, J); 1.0 mm (H, I)." figureDoi="10.3897/zookeys.1174.101251.figure2" httpUri="https://binary.pensoft.net/fig/891286" pageId="0" pageNumber="219">2G</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="BD8D0C8FAD9ED0A060340AA250CD1416" pageId="0" pageNumber="219" type="description">
<paragraph id="188815516E1ADFD5B745D5438AEB0AAB" pageId="0" pageNumber="219">Size variation.</paragraph>
<paragraph id="2DA197A09FD1916C24376E0C76BE686F" pageId="0" pageNumber="219">Female (based on 5 specimens): total length: 2.00-2.67, prosoma length: 1.05-1.12, prosoma width: 0.92-0.97.</paragraph>
</subSubSection>
<subSubSection id="C053B6725699618E06AB56434F778B35" pageId="0" pageNumber="219" type="distribution">
<paragraph id="632350DDE16BCBFA56C86F4E1D6BA323" pageId="0" pageNumber="219">Distribution.</paragraph>
<paragraph id="E09DD0843F9DDB94F0C090CC0B9E01F5" pageId="0" pageNumber="219">
Endemic to Miyako-jima Is., Ryukyus, Japan (Fig.
<figureCitation id="B8E56E0E893593186E5AD061B9F4026E" captionStart="Figure 16" captionStartId="F16" captionText="Figure 16. Distribution of Nesticella and Howaia species in mainland Japan and Ryukyu Archipelago A distribution of N. brevipes and N. terrestris in mainland Japan and neighboring countries B distribution of Nesticella and Howaia species endemic to the Ryukyu Archipelago C distribution of H. mogera in mainland Japan and neighboring countries D distribution of H. mogera in the Ryukyu Archipelago and neighboring countries. Colored squares refer to the type localities of the species, colored dots indicate the locations of specimens whose morphology has been checked during this study, dashed areas refer to the generally known distribution of the species, question marks refer to unclear distributions (see &quot; remarks on misidentifications &quot; of N. brevipes and N. terrestris)." figureDoi="10.3897/zookeys.1174.101251.figure16" httpUri="https://binary.pensoft.net/fig/891300" pageId="0" pageNumber="219">16B</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="9A27DBF64934F4A14094DE954F8BC5C7" pageId="0" pageNumber="219" type="habitat">
<paragraph id="6B8D4054FC14BFD64084188D380D3B07" pageId="0" pageNumber="219">Habitat and ecology.</paragraph>
<paragraph id="1B8E265B3FD458EBDBFB72814606A2A4" pageId="0" pageNumber="219">
<taxonomicName id="1D38E5FBC57451385D9A4C885968EF3D" authorityName="Ballarin &amp; Eguchi" authorityYear="2023" class="Arachnida" family="Nesticidae" genus="Howaia" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Howaia alba" order="Araneae" pageId="0" pageNumber="219" phylum="Arthropoda" rank="species" species="alba">
<emphasis id="E3B698B4C643ACD057A62B276B7B06CC" italics="true" pageId="0" pageNumber="219">Howaia alba</emphasis>
</taxonomicName>
sp. nov. is found in the natural caves in Miyako-jima Is. This species builds simple scaffold webs between rocks and in crevices at the base of the walls or on the floor of the caves. It dwells exclusively in the dark zone of the caves, in areas characterized by relatively high and uniform temperature and humidity (e.g., Tsuzupisuki-abi cave: temp: 25.2 °C, hum: 94.1%; Pinza-Abu cave: temp: 25.1 °C, hum: 92.6%) (Fig.
<figureCitation id="407BF98199313D77A98A22962D1918F4" captionStart="Figure 15" captionStartId="F15" captionText="Figure 15. Nesticella and Howaia species in life and their natural habitats A female of H. mogera B female of N. terrestris C female of H. alba sp. nov. D female of N. occulta sp. nov. E female of N. okinawaensis F example of the epigean habitat of Nesticella species in the Ryukyus G habitat of H. alba sp nov. (dark zone of Tsuzupisuki-abu cave) H habitat of N. occulta sp. nov. (dark zone of Kabirano-ana cave) (all photos by F. Ballarin)." figureDoi="10.3897/zookeys.1174.101251.figure15" httpUri="https://binary.pensoft.net/fig/891299" pageId="0" pageNumber="219">15G</figureCitation>
). Adults of
<taxonomicName id="E3064E1D4D6F3801BABAEC96716B44AC" authorityName="Ballarin &amp; Eguchi" authorityYear="2023" class="Arachnida" family="Nesticidae" genus="Howaia" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Howaia alba" order="Araneae" pageId="0" pageNumber="219" phylum="Arthropoda" rank="species" species="alba">
<emphasis id="C9BE1D4D9CA1262D697013BDB7E6550C" italics="true" pageId="0" pageNumber="219">Howaia alba</emphasis>
</taxonomicName>
sp. nov. were observed preying on
<taxonomicName id="A4BA52BD7BAC8488AE9F4F016579D736" authorityName="Lehtinen &amp; Saaristo" authorityYear="1980" class="Arachnida" kingdom="Animalia" lsidName="" order="Schizomida" pageId="0" pageNumber="219" rank="order">Schizomida</taxonomicName>
(
<taxonomicName id="86EF0E7434F18499632C7BE7BB1B151D" baseAuthorityName="Hansen" baseAuthorityYear="1905" class="Arachnida" family="Hubbardiidae" genus="Bamazomus" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Bamazomus siamensis" order="Schizomida" pageId="0" pageNumber="219" phylum="Arthropoda" rank="species" species="siamensis">
<emphasis id="2AC948FEBC4EF2BE6F6CA654723AF6A0" italics="true" pageId="0" pageNumber="219">Bamazomus siamensis</emphasis>
</taxonomicName>
(Hansen, 1905) which roam the floor of the caves in Miyako-jima Is. Females carrying the eggs cocoon attached to their spinnerets were also observed (but not collected) sitting on webs or wandering under rocks. Despite extensive surveying, no specimens were found in the numerous artificial tunnels or underground water reserves dug in the limestone rocks of the island. The complete absence of eyes and pigmentation, the lack of external records and the finding of the species only in the deepest areas of the caves identify
<taxonomicName id="DBEB6E9D005B2D3F7003222E1F1A9E0A" family="Nesticidae" kingdom="Animalia" lsidName="H. alba" order="Araneae" pageId="0" pageNumber="219" rank="species" species="alba">
<emphasis id="44A2D0F78668ADD30ED075161843F79E" italics="true" pageId="0" pageNumber="219">H. alba</emphasis>
</taxonomicName>
sp. nov. as a true troglobiont.
</paragraph>
</subSubSection>
<subSubSection id="AF2825E64F05FD292CFE2C10D3785ED1" pageId="0" pageNumber="219" type="remarks">
<paragraph id="1B2D2E9201AB956DF89FB5C6B679571C" pageId="0" pageNumber="219">Remarks on misidentifications.</paragraph>
<paragraph id="BFEF5E031FDB4AB4A0050BE757B01479" pageId="0" pageNumber="219">
This species was recorded and illustrated for the first time by
<bibRefCitation id="E606F01C55D4C635571C7270DD34D0D7" DOI="https://doi.org/10.2476/asjaa.27.Specialnumber_337" author="Shimojana, M" journalOrPublisher="Zootaxa" pageId="0" pageNumber="219" refId="B45" refString="Shimojana, M, 1977. Preliminary report on the cave spider fauna of the Ryukyu Archipelago. Acta Arachnologica 27(Spec. No.): 337-365. https://doi.org/10.2476/asjaa.27.Specialnumber_337" title="Preliminary report on the cave spider fauna of the Ryukyu Archipelago. Acta Arachnologica 27 (Spec. No.): 337 - 365." url="https://doi.org/10.2476/asjaa.27.Specialnumber_337" year="1977">Shimojana (1977</bibRefCitation>
: fig. 6A-C). Due to the general similarities in the shape of genitalia, it was identified as
<taxonomicName id="C2B88D55C9BF7D3733852E87534AE960" family="Nesticidae" kingdom="Animalia" lsidName="H. mogera" order="Araneae" pageId="0" pageNumber="219" rank="species" species="mogera">
<emphasis id="8B6BF8D025E549B6EBCB705C13CD35EE" italics="true" pageId="0" pageNumber="219">H. mogera</emphasis>
</taxonomicName>
although the author highlighted the lack of eyes in these specimens (
<bibRefCitation id="085CBEB102105E779147F84001389DA1" DOI="https://doi.org/10.2476/asjaa.27.Specialnumber_337" author="Shimojana, M" journalOrPublisher="Zootaxa" pageId="0" pageNumber="219" refId="B45" refString="Shimojana, M, 1977. Preliminary report on the cave spider fauna of the Ryukyu Archipelago. Acta Arachnologica 27(Spec. No.): 337-365. https://doi.org/10.2476/asjaa.27.Specialnumber_337" title="Preliminary report on the cave spider fauna of the Ryukyu Archipelago. Acta Arachnologica 27 (Spec. No.): 337 - 365." url="https://doi.org/10.2476/asjaa.27.Specialnumber_337" year="1977">Shimojana 1977</bibRefCitation>
: 353). Our analysis, based on both morphology and molecular data, supports
<taxonomicName id="6EB20CAF911BDDEF01073205FA84BDB5" family="Nesticidae" kingdom="Animalia" lsidName="H. alba" order="Araneae" pageId="0" pageNumber="219" rank="species" species="alba">
<emphasis id="8BBE5F4576F0293C9F32C12446810EE2" italics="true" pageId="0" pageNumber="219">H. alba</emphasis>
</taxonomicName>
sp. nov. as a closely related species but clearly distinct from
<taxonomicName id="9381304DAD0DB632C940224D26EC1261" family="Nesticidae" kingdom="Animalia" lsidName="H. mogera" order="Araneae" pageId="0" pageNumber="219" rank="species" species="mogera">
<emphasis id="94F8F27497FD3C45380C76A0477D2471" italics="true" pageId="0" pageNumber="219">H. mogera</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>
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