treatments-xml/data/03/FF/E7/03FFE764FFDBFFADFF6A7982FD95FE0B.xml

<document id="A62EFC10E23CAD09AAED2709FA672D59" ID-DOI="10.11646/zootaxa.3866.1.3" ID-GBIF-Dataset="14e55954-4068-455f-add4-8ed6b83f67ae" ID-ISSN="1175-5326" ID-Zenodo-Dep="287184" ID-ZooBank="6334A96F-7716-4A4D-81AF-A81EAA06B9B7" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1460915749551" checkinUser="plazi" docAuthor="Yoshikawa, Natsuhiko &amp; Matsui, Masafumi" docDate="2014" docId="03FFE764FFDBFFADFF6A7982FD95FE0B" docLanguage="en" docName="zt03866p078.pdf" docOrigin="Zootaxa 3866 (1)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Onychodactylus fuscus Yoshikawa &amp; Matsui, 2014, sp. nov." docType="treatment" docVersion="5" lastPageNumber="74" masterDocId="FFC69F1CFFD4FFB8FFFD7A03FFD0FF8C" masterDocTitle="Two new Salamanders of the genus Onychodactylus from Eastern Honshu, Japan (Amphibia, Caudata, Hynobiidae)" masterLastPageNumber="78" masterPageNumber="53" pageNumber="68" updateTime="1698361655219" updateUser="plazi">
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<mods:title id="6168037EDEB7943F401A6EDF0283EF01">Two new Salamanders of the genus Onychodactylus from Eastern Honshu, Japan (Amphibia, Caudata, Hynobiidae)</mods:title>
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<mods:namePart id="4AFCB990C9A136260D7CBF3BE05BECB7">Yoshikawa, Natsuhiko</mods:namePart>
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<mods:namePart id="946BF4D4C43E26403EAD34DAC131383D">Matsui, Masafumi</mods:namePart>
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<heading id="D0A1E11EFFDBFFB7FF6A7982FDC3FC17" bold="true" box="[151,531,897,923]" fontSize="11" level="1" pageId="15" pageNumber="68" reason="1">
<emphasis id="B9228A60FFDBFFB7FF6A7982FDC3FC17" bold="true" box="[151,531,897,923]" pageId="15" pageNumber="68">
<taxonomicName id="4C562DF1FFDBFFB7FF6A7982FE7EFC17" ID-CoL="49R6B" box="[151,430,897,923]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="15" pageNumber="68" phylum="Chordata" rank="species" species="fuscus" status="sp. nov.">
<emphasis id="B9228A60FFDBFFB7FF6A7982FE7EFC17" bold="true" box="[151,430,897,923]" italics="true" pageId="15" pageNumber="68">Onychodactylus fuscus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A211371BFFDBFFB7FE487982FDC3FC17" box="[437,531,897,923]" pageId="15" pageNumber="68" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
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<paragraph id="8BE95672FFDBFFB7FF6A79A7FE92FB88" blockId="15.[151,682,897,1029]" pageId="15" pageNumber="68">
(Japanese name: Tadami-hakone-sanshou-uwo) (English name: Tadami clawed salamander) (
<figureCitation id="136D4AF7FFDBFFB7FF6279EFFF2AFB88" box="[159,250,1004,1029]" captionStart="FIGURE 10" captionStartId="16.[151,250,1959,1981]" captionTargetBox="[150,1435,720,1922]" captionTargetId="figure@16.[150,1436,720,1955]" captionTargetPageId="16" captionText="FIGURE 10. Dorsal and ventral views of male holotype (A and B; KUHE 48284, SVL = 69.0 mm) and a female paratype of O. fuscus sp. nov. (C and D; KUHE 48129, SVL = 74.8 mm). Scale bar indicates 10 mm." httpUri="https://zenodo.org/record/287194/files/figure.png" pageId="15" pageNumber="68">Figs. 10</figureCitation>
&amp; 
<figureCitation id="136D4AF7FFDBFFB7FEE379EFFEE9FB88" box="[286,313,1004,1028]" captionStart="FIGURE 11. A" captionStartId="17.[151,250,1258,1280]" captionTargetBox="[151,1436,589,1236]" captionTargetId="figure@17.[151,1436,589,1237]" captionTargetPageId="17" captionText="FIGURE 11. A male paratype of O. fuscus sp. nov. (KUHE 48283) in life." httpUri="https://zenodo.org/record/287195/files/figure.png" pageId="15" pageNumber="68">11</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C34C05F9FFDBFFB7FF6A7E2DFEB4FB3F" pageId="15" pageNumber="68" type="etymology">
<paragraph id="8BE95672FFDBFFB7FF6A7E2DFEB4FB3F" blockId="15.[151,1437,1070,2034]" pageId="15" pageNumber="68">
<emphasis id="B9228A60FFDBFFB7FF6A7E2DFECEFBCB" bold="true" box="[151,286,1070,1095]" pageId="15" pageNumber="68">Etymology.</emphasis>
The specific epithet “ 
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<emphasis id="B9228A60FFDBFFB7FDCD7E2CFDA6FBCB" box="[560,630,1071,1095]" italics="true" pageId="15" pageNumber="68">fuscus</emphasis>
</taxonomicName>
”, a Latin word meaning dark or black, is derived from unstriped, uniformly dark dorsal coloration of the new species, which is quite unique among species in the 
<taxonomicName id="4C562DF1FFDBFFB7FAF47E50FA4CFBE6" box="[1289,1436,1107,1130]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="15" pageNumber="68" phylum="Chordata" rank="species" species="japonicus">
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<collectingCountry id="F34116E2FFDBFFB7FAF47E50FAF0FBE6" box="[1289,1312,1107,1130]" name="Iceland" pageId="15" pageNumber="68">O</collectingCountry>
. japonicus
</emphasis>
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species complex. Suggested Japanese and English names are after Tadami-machi (Tadami Town), where the 
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locality is located.
</paragraph>
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<subSubSection id="C34C05F9FFDBFFB7FF3A7EC2FD46FA76" pageId="15" pageNumber="68" type="materials_examined">
<paragraph id="8BE95672FFDBFFB7FF3A7EC2FA88FB72" blockId="15.[151,1437,1070,2034]" pageId="15" pageNumber="68">
<emphasis id="B9228A60FFDBFFB7FF3A7EC2FEEBFB56" bold="true" box="[199,315,1217,1242]" pageId="15" pageNumber="68">
<typeStatus id="54EDE8D0FFDBFFB7FF3A7EC2FEE6FB56" box="[199,310,1217,1242]" pageId="15" pageNumber="68" type="holotype">Holotype</typeStatus>
.
</emphasis>
<collectionCode id="ED47CEB7FFDBFFB7FEBB7EC1FE44FB55" box="[326,404,1218,1241]" httpUri="http://grbio.org/cool/jxgt-7i97" name="Kyoto University, Graduate School of Human and Environmental Studies" pageId="15" pageNumber="68">KUHE</collectionCode>
48284, an adult male from environs of Mt. Yogai (
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, 
<geoCoordinate id="EE6230B5FFDBFFB7FB847EC1FB04FB56" box="[1145,1236,1214,1242]" direction="north" orientation="latitude" pageId="15" pageNumber="68" precision="925" value="37.35">37o21'N</geoCoordinate>
, 
<quantity id="4CAEFB97FFDBFFB7FB187EC1FAE1FB55" box="[1253,1329,1218,1242]" metricMagnitude="2" metricUnit="m" metricValue="4.3" pageId="15" pageNumber="68" unit="m" value="430.0">430 m</quantity>
a.s.l.) in Tadami-machi, Fukushima Prefecture, 
<collectingCountry id="F34116E2FFDBFFB7FDB67EE5FD5EFB71" box="[587,654,1254,1277]" name="Japan" pageId="15" pageNumber="68">Japan</collectingCountry>
, collected by N. Yoshikawa and 
<collectingCountry id="F34116E2FFDBFFB7FC067EE5FBC2FB72" box="[1019,1042,1254,1278]" name="Iceland" pageId="15" pageNumber="68">O</collectingCountry>
. Inaba on 
<date id="FFE870B2FFDBFFB7FB7B7EE5FA84FB72" box="[1158,1364,1253,1278]" pageId="15" pageNumber="68" value="2013-11-09">9 November 2013</date>
.
</paragraph>
<paragraph id="8BE95672FFDBFFB7FF3A7F0AFE4AFA5A" blockId="15.[151,1437,1070,2034]" pageId="15" pageNumber="68">
<emphasis id="B9228A60FFDBFFB7FF3A7F0AFE96FAAE" bold="true" box="[199,326,1289,1314]" pageId="15" pageNumber="68">
<typeStatus id="54EDE8D0FFDBFFB7FF3A7F0AFE91FAAE" box="[199,321,1289,1314]" pageId="15" pageNumber="68" type="paratype">Paratypes</typeStatus>
.
</emphasis>
A total of 
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: four males (
<collectionCode id="ED47CEB7FFDBFFB7FD017F09FC9CFAAE" box="[764,844,1290,1314]" httpUri="http://grbio.org/cool/jxgt-7i97" name="Kyoto University, Graduate School of Human and Environmental Studies" pageId="15" pageNumber="68">KUHE</collectionCode>
48282–48283, 48285 and 20131109-1[stored at the Tadami Beech Center]) and one female (
<collectionCode id="ED47CEB7FFDBFFB7FD987F2DFD65FACA" box="[613,693,1326,1350]" httpUri="http://grbio.org/cool/jxgt-7i97" name="Kyoto University, Graduate School of Human and Environmental Studies" pageId="15" pageNumber="68">KUHE</collectionCode>
48129), collected on 
<date id="FFE870B2FFDBFFB7FC537F2DFBABFACA" box="[942,1147,1325,1350]" pageId="15" pageNumber="68" value="2013-11-09">9 November 2013</date>
by N. Yoshikawa and 
<collectingCountry id="F34116E2FFDBFFB7FA7C7F2DFA48FACA" box="[1409,1432,1326,1350]" name="Iceland" pageId="15" pageNumber="68">O</collectingCountry>
. Inaba) from the 
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locality; one female (
<collectionCode id="ED47CEB7FFDBFFB7FD847F51FD19FAE6" box="[633,713,1362,1386]" httpUri="http://grbio.org/cool/jxgt-7i97" name="Kyoto University, Graduate School of Human and Environmental Studies" pageId="15" pageNumber="68">KUHE</collectionCode>
48286 collected on 
<date id="FFE870B2FFDBFFB7FC4D7F51FBADFAE6" box="[944,1149,1361,1386]" pageId="15" pageNumber="68" value="2013-11-09">9 November 2013</date>
by N. Yoshikawa and 
<collectingCountry id="F34116E2FFDBFFB7FA7C7F51FA48FAE6" box="[1409,1432,1362,1386]" name="Iceland" pageId="15" pageNumber="68">O</collectingCountry>
. Inaba) from environs of Mt. Hiraishi, Tadami-machi, Fukushima Prefecture; one male and three females (
<collectionCode id="ED47CEB7FFDBFFB7FAB07F75FA4DFA02" box="[1357,1437,1398,1422]" httpUri="http://grbio.org/cool/jxgt-7i97" name="Kyoto University, Graduate School of Human and Environmental Studies" pageId="15" pageNumber="68">KUHE</collectionCode>
44847 and 
<collectionCode id="ED47CEB7FFDBFFB7FEEB7F99FEB4FA3D" box="[278,356,1434,1457]" httpUri="http://grbio.org/cool/jxgt-7i97" name="Kyoto University, Graduate School of Human and Environmental Studies" pageId="15" pageNumber="68">KUHE</collectionCode>
44848–44850, respectively, collected on 
<date id="FFE870B2FFDBFFB7FCC17F99FC15FA3E" box="[828,965,1434,1458]" pageId="15" pageNumber="68" value="2011-06-02">2 June 2011</date>
by 
<collectingCountry id="F34116E2FFDBFFB7FC0F7F99FBD6FA3D" box="[1010,1030,1434,1457]" name="Italy" pageId="15" pageNumber="68">Y</collectingCountry>
. Misawa) from Shionofuchi, Sanjoshi, Niigata Prefecture.
</paragraph>
<paragraph id="8BE95672FFDBFFB7FF3A7FE2FD46FA76" blockId="15.[151,1437,1070,2034]" box="[199,662,1505,1530]" pageId="15" pageNumber="68">
<emphasis id="B9228A60FFDBFFB7FF3A7FE2FE69FA76" bold="true" box="[199,441,1505,1530]" pageId="15" pageNumber="68">Referred specimens.</emphasis>
See APPENDIX 1.
</paragraph>
</subSubSection>
<subSubSection id="C34C05F9FFDBFFB7FF3A7C06FA59F8B2" pageId="15" pageNumber="68" type="diagnosis">
<paragraph id="8BE95672FFDBFFB7FF3A7C06FA59F8B2" blockId="15.[151,1437,1070,2034]" pageId="15" pageNumber="68">
<emphasis id="B9228A60FFDBFFB7FF3A7C06FE91F992" bold="true" box="[199,321,1541,1566]" pageId="15" pageNumber="68">Diagnosis.</emphasis>
The new species is a member of the genus 
<taxonomicName id="4C562DF1FFDBFFB7FCCE7C06FC38F991" box="[819,1000,1541,1565]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="15" pageNumber="68" phylum="Chordata" rank="genus">
<emphasis id="B9228A60FFDBFFB7FCCE7C06FC38F991" box="[819,1000,1541,1565]" italics="true" pageId="15" pageNumber="68">Onychodactylus</emphasis>
</taxonomicName>
diagnosed by the characteristics listed for 
<taxonomicName id="4C562DF1FFDBFFB7FF437C29FEB4F9CD" box="[190,356,1577,1601]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="15" pageNumber="68" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="B9228A60FFDBFFB7FF437C29FEB4F9CD" box="[190,356,1577,1601]" italics="true" pageId="15" pageNumber="68">
<collectingCountry id="F34116E2FFDBFFB7FF437C29FF05F9CD" box="[190,213,1578,1601]" name="Iceland" pageId="15" pageNumber="68">O</collectingCountry>
. intermedius
</emphasis>
</taxonomicName>
. It is further assigned to a member of the 
<taxonomicName id="4C562DF1FFDBFFB7FCC47C29FC18F9CD" box="[825,968,1578,1601]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="15" pageNumber="68" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFDBFFB7FCC47C29FC18F9CD" box="[825,968,1578,1601]" italics="true" pageId="15" pageNumber="68">
<collectingCountry id="F34116E2FFDBFFB7FCC47C29FC80F9CD" box="[825,848,1578,1601]" name="Iceland" pageId="15" pageNumber="68">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
species complex by the genetic evidence, although this species completely lacks dorsal stripe which is a diagnostic characteristic of the complex. Dorsum uniformly dark-brown without stripe, sometimes with slight indistinct ochre small flecks; slightly large body size with mean SVL (±1SD) of 69.5±3.0 mm in males and 69.0± 
<quantity id="4CAEFB97FFDBFFB7FCC37C96FC44F921" box="[830,916,1685,1710]" metricMagnitude="-3" metricUnit="m" metricValue="4.8" pageId="15" pageNumber="68" unit="mm" value="4.8">4.8 mm</quantity>
in females; tail longer than SVL in males, but nearly equal in females; hindlimb comparatively short; head and internarial space comparatively wide; usually lacking wedge-shaped dark marking on chest; ventrum purplish brown or purplish gray, with silvery white dots; presacral vertebrae including atlas 18 or 19; costal grooves 12 or 13; right and left vomerine tooth series interrupted by gap, series tending to curve shallowly, with comparatively small number of teeth, 12–15 on one side of series.
</paragraph>
</subSubSection>
<subSubSection id="C34C05F9FFDBFFADFF3A7D4AFD95FE0B" lastPageId="21" lastPageNumber="74" pageId="15" pageNumber="68" type="description">
<paragraph id="8BE95672FFDBFFA8FF3A7D4AFB78FDB1" blockId="15.[151,1437,1070,2034]" lastBlockId="16.[151,1437,151,680]" lastPageId="16" lastPageNumber="69" pageId="15" pageNumber="68">
<emphasis id="B9228A60FFDBFFB7FF3A7D4AFE3FF8EE" bold="true" box="[199,495,1865,1890]" pageId="15" pageNumber="68">
Description of 
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.
</emphasis>
An adult male with SVL 69.0 mm (
<figureCitation id="136D4AF7FFDBFFB7FC507D4AFBD2F8EE" box="[941,1026,1865,1890]" captionStart="FIGURE 10" captionStartId="16.[151,250,1959,1981]" captionTargetBox="[150,1435,720,1922]" captionTargetId="figure@16.[150,1436,720,1955]" captionTargetPageId="16" captionText="FIGURE 10. Dorsal and ventral views of male holotype (A and B; KUHE 48284, SVL = 69.0 mm) and a female paratype of O. fuscus sp. nov. (C and D; KUHE 48129, SVL = 74.8 mm). Scale bar indicates 10 mm." httpUri="https://zenodo.org/record/287194/files/figure.png" pageId="15" pageNumber="68">Fig. 10</figureCitation>
A–B); body moderately stout; skin smooth; head oval and depressed, longer than wide; neck slightly narrower than head; snout rounded, projecting beyond lower jaw; nostril close to eye; eye large, shorter than snout, prominently protruded; gular fold posteroventral to head; parotoid gland well developed, oval, ca. 1.6 times longer than wide, projecting more posteriorly than gular fold; postorbital groove obvious; vomerine teeth in two transverse, shallowly curved archshaped series with a gap in between (
<figureCitation id="136D4AF7FFC4FFA8FDBE7A94FD58FF3C" box="[579,648,151,176]" captionStart="FIGURE 6" captionStartId="9.[151,250,1391,1413]" captionTargetBox="[155,1428,948,1363]" captionTargetId="figure@9.[151,1436,941,1369]" captionTargetPageId="9" captionText="FIGURE 6. Open mouth of holotype of O. intermedius sp. nov. (A: KUHE 47455) and O. fuscus sp. nov. (B: KUHE 48284) showing the shape of vomerine tooth series. Scale bars indicate 1 mm." httpUri="https://zenodo.org/record/287190/files/figure.png" pageId="16" pageNumber="69">Fig. 6</figureCitation>
B); 15 teeth on right and 14 on left side of series; few dark pigments scattered around vomerine tooth series; forelimb thin; relative length of fingers I&lt;IV&lt;II&lt;III; hindlimb slightly longer and distinctly more robust than forelimb; relative length of toes I&lt;V&lt;II&lt;III=IV; adpressed limbs touching each other; secondary sexual characters obvious; dermal flap on posterior edge of hindlimb extremely developed; black horny claws on tips of fingers and toes; black tubercles and asperities on palm and sole; trunk elongated and cylindrical but slightly depressed; middorsal groove indistinct; well-developed 13 costal grooves on both sides of trunk; presacral vertebrae including atlas numbering 18; base of tail including cloacal region well swollen; cloaca longitudinal slit, length 11.3% SVL, with anterior two-fifth of its edge slightly swollen, forming inverse shallow Vshaped precloacal skin fold; precloacal skin fold as wide as length of cloaca; a series of small black spines lining along front edge of precloacal skin fold; tail long, 127.8% length of SVL, cylindrical at base, increasingly compressed posteriorly; tail slightly lowered toward tip, laterally compressed posterior two-thirds and extremely at tip, forming finny shape; tip of tail rounded in lateral view; tail highest (11.3% SVL) at base.
</paragraph>
<paragraph id="8BE95672FFC4FFA8FF3A7844FB65FD24" blockId="16.[151,1437,151,680]" pageId="16" pageNumber="69">
<emphasis id="B9228A60FFC4FFA8FF3A7844FD4DFDEC" bold="true" box="[199,669,583,608]" pageId="16" pageNumber="69">
Measurements (in mm) of the 
<typeStatus id="54EDE8D0FFC4FFA8FDCD7844FD49FDEC" box="[560,665,583,608]" pageId="16" pageNumber="69" type="holotype">holotype</typeStatus>
.
</emphasis>
SVL 69.0, HL 15.8, HW 10.1, TAL 88.2, AGD 34.7, FLL (L) 18.8, FLL (R) 18.7, HLL (L) 20.1, HLL (R) 20.0, UEL (L) 4.1, UEL (R) 4.2, IOD 3.4, 
<collectingCountry id="F34116E2FFC4FFA8FBD2786EFBB9FD08" box="[1071,1129,621,644]" name="India" pageId="16" pageNumber="69">END</collectingCountry>
(L) 2.3, 
<collectingCountry id="F34116E2FFC4FFA8FB30786EFAD7FD08" box="[1229,1287,621,644]" name="India" pageId="16" pageNumber="69">END</collectingCountry>
(R) 2.2, ICD 6.1, 
<collectingCountry id="F34116E2FFC4FFA8FF357893FF2AFD2B" box="[200,250,656,679]" name="India" pageId="16" pageNumber="69">IND</collectingCountry>
5.1, SL (L) 5.5, SL (R) 5.4, CW 9.8, BTAH 7.8, BTAW 8.2, MTAH 6.6, MTAW 4.4.
</paragraph>
<caption id="DF2906FAFFC4FFA8FF6A7DA4FC25F857" httpUri="https://zenodo.org/record/287194/files/figure.png" pageId="16" pageNumber="69" targetBox="[150,1435,720,1922]" targetPageId="16">
<paragraph id="8BE95672FFC4FFA8FF6A7DA4FC25F857" blockId="16.[151,1436,1959,2011]" pageId="16" pageNumber="69">
<emphasis id="B9228A60FFC4FFA8FF6A7DA4FECFF831" bold="true" box="[151,287,1959,1981]" pageId="16" pageNumber="69">FIGURE 10.</emphasis>
Dorsal and ventral views of male holotype (A and B; KUHE 48284, SVL=69.0 mm) and a female paratype of 
<taxonomicName id="4C562DF1FFC4FFA8FA7E7DABFF06F857" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="16" pageNumber="69" phylum="Chordata" rank="species" species="fuscus" status="sp. nov.">
<emphasis id="B9228A60FFC4FFA8FA7E7DABFF06F857" italics="true" pageId="16" pageNumber="69">O. fuscus</emphasis>
</taxonomicName>
<emphasis id="B9228A60FFC4FFA8FF217DC6FEFDF857" bold="true" box="[220,301,1989,2011]" pageId="16" pageNumber="69">
<taxonomicNameLabel id="A211371BFFC4FFA8FF217DC6FEFDF857" box="[220,301,1989,2011]" pageId="16" pageNumber="69" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(C and D; KUHE 48129, SVL=74.8 mm). Scale bar indicates 10 mm.
</paragraph>
</caption>
<paragraph id="8BE95672FFC5FFA9FF3A7A94FEA2FECC" blockId="17.[151,1437,151,536]" pageId="17" pageNumber="70">
<emphasis id="B9228A60FFC5FFA9FF3A7A94FEC4FF3C" bold="true" box="[199,276,151,176]" pageId="17" pageNumber="70">Color:</emphasis>
In life, dorsum uniformly dark-brown with a few indistinct ochre flecks; dorsal stripe completely absent; lateral and ventral sides of body dark-brown with silvery white dots, but background color sometimes gradually fading toward grayish ventrum; iris mottled with dark-brown and gold in upper half, uniformly darkbrown in lower half. In alcohol, color and pattern fading generally, but no obvious change except for bleach of dorsal ochre flecks.
</paragraph>
<paragraph id="8BE95672FFC5FFA9FF3A7B4FFAB6FE04" blockId="17.[151,1437,151,536]" pageId="17" pageNumber="70">
<emphasis id="B9228A60FFC5FFA9FF3A7B4FFEEBFEE9" bold="true" box="[199,315,332,357]" pageId="17" pageNumber="70">Variation.</emphasis>
The following description of variation is based on the maximum number of six adult males and five adult females. Morphometric data are summarized in Table 1 together with those of seven congeneric species.
</paragraph>
<paragraph id="8BE95672FFC5FFA9FF3A7B96FBB9FD94" blockId="17.[151,1437,151,536]" pageId="17" pageNumber="70">
In 
<taxonomicName id="4C562DF1FFC5FFA9FF177B97FE88FE20" box="[234,344,404,428]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="70" phylum="Chordata" rank="species" species="fuscus" status="sp. nov.">
<emphasis id="B9228A60FFC5FFA9FF177B97FE88FE20" box="[234,344,404,428]" italics="true" pageId="17" pageNumber="70">
<collectingCountry id="F34116E2FFC5FFA9FF177B97FED1FE27" box="[234,257,404,427]" name="Iceland" pageId="17" pageNumber="70">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
<emphasis id="B9228A60FFC5FFA9FE997B97FE12FE21" bold="true" box="[356,450,404,429]" pageId="17" pageNumber="70">
<taxonomicNameLabel id="A211371BFFC5FFA9FE997B97FE12FE21" box="[356,450,404,429]" pageId="17" pageNumber="70" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, there was no significant difference in SVL (t = 0.183, 
<emphasis id="B9228A60FFC5FFA9FB9A7B96FBA5FE20" box="[1127,1141,405,428]" italics="true" pageId="17" pageNumber="70">p</emphasis>
= 0.859) between males (69.5±3.0 mm, n = 6) and females (69.0± 
<quantity id="4CAEFB97FFC5FFA9FD8E7BB4FD1EFE43" box="[627,718,439,464]" metricMagnitude="-3" metricUnit="m" metricValue="4.8" pageId="17" pageNumber="70" unit="mm" value="4.8">4.8 mm</quantity>
, n = 5: Table 1). In values relative to SVL, sexes significantly differed in longer tail and shorter trunk in males than females (Table 1). 
<collectingCountry id="F34116E2FFC5FFA9FC1C7BDFFBD0FE78" box="[993,1024,476,500]" name="American Samoa" pageId="17" pageNumber="70">As</collectingCountry>
seen in other congeners, males had more robust hindlimb and posteriorly more projecting distal end of fibula than females.
</paragraph>
<caption id="DF2906FAFFC5FFA9FF6A7EE9FC5FFA8C" box="[151,911,1258,1280]" httpUri="https://zenodo.org/record/287195/files/figure.png" pageId="17" pageNumber="70" targetBox="[151,1436,589,1236]" targetPageId="17">
<paragraph id="8BE95672FFC5FFA9FF6A7EE9FC5FFA8C" blockId="17.[151,911,1258,1280]" box="[151,911,1258,1280]" pageId="17" pageNumber="70">
<emphasis id="B9228A60FFC5FFA9FF6A7EE9FEC8FA8C" bold="true" box="[151,280,1258,1280]" pageId="17" pageNumber="70">FIGURE 11</emphasis>
. A male paratype of 
<taxonomicName id="4C562DF1FFC5FFA9FE107EE8FD9BFA8C" box="[493,587,1258,1280]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="70" phylum="Chordata" rank="species" species="fuscus" status="sp. nov.">
<emphasis id="B9228A60FFC5FFA9FE107EE8FD9BFA8C" box="[493,587,1258,1280]" italics="true" pageId="17" pageNumber="70">O. fuscus</emphasis>
</taxonomicName>
<emphasis id="B9228A60FFC5FFA9FDAF7EE9FD72FA8C" bold="true" box="[594,674,1258,1280]" pageId="17" pageNumber="70">
<taxonomicNameLabel id="A211371BFFC5FFA9FDAF7EE9FD72FA8C" box="[594,674,1258,1280]" pageId="17" pageNumber="70" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(KUHE 48283) in life.
</paragraph>
</caption>
<paragraph id="8BE95672FFC5FFA9FF3A7F2FFD3EFA5A" blockId="17.[151,1437,1324,2034]" pageId="17" pageNumber="70">Adult specimens showed the following characteristics in the breeding season: well-developed dermal skin fold on posterior edge of hindlimb, black tubercles and asperities on palm and sole, cloaca well swollen, and precloacal skin fold forming inverse V-shaped in males; precloacal skin fold forming inverse narrow U-shaped in females; black claws on fingers and toes, parotoid gland well swollen, black precloacal spine series, and tail laterally compressed in both sexes, more remarkable in males.</paragraph>
<paragraph id="8BE95672FFC5FFA9FF3A7FE2FDC6F979" blockId="17.[151,1437,1324,2034]" pageId="17" pageNumber="70">
The number of costal grooves was significantly largerer in males (mode = 
<quantity id="4CAEFB97FFC5FFA9FBE17FE2FB87FA76" box="[1052,1111,1505,1530]" metricMagnitude="-1" metricUnit="m" metricValue="3.302" pageId="17" pageNumber="70" unit="in" value="13.0">13 in</quantity>
males vs. 
<quantity id="4CAEFB97FFC5FFA9FB2F7FE2FADCFA76" box="[1234,1292,1505,1530]" metricMagnitude="-1" metricUnit="m" metricValue="3.048" pageId="17" pageNumber="70" unit="in" value="12.0">12 in</quantity>
females: t = 2.455, 
<emphasis id="B9228A60FFC5FFA9FF187C05FF23F991" box="[229,243,1542,1565]" italics="true" pageId="17" pageNumber="70">p</emphasis>
= 0.036 for each side: Table 2). The number of presacral vertebrae including atlas varied from 18 to 19 (Table 2) and was not sexually different (t = 0.980, 
<emphasis id="B9228A60FFC5FFA9FD2C7C29FD0FF9CD" box="[721,735,1578,1601]" italics="true" pageId="17" pageNumber="70">p</emphasis>
= 0.353). The number of vomerine teeth, tended to be larger in females (9 to 15, mean = 12.4± 
<quantity id="4CAEFB97FFC5FFA9FDE87C4EFD89F9EA" box="[533,601,1613,1638]" metricMagnitude="-2" metricUnit="m" metricValue="4.318" pageId="17" pageNumber="70" unit="in" value="1.7">1.7 in</quantity>
males and 11 to 19, 15.0±3.0 in females), but the difference was not significant (t = 1.755, 
<emphasis id="B9228A60FFC5FFA9FE707C71FE4BF905" box="[397,411,1650,1673]" italics="true" pageId="17" pageNumber="70">p</emphasis>
= 0.113). Gap between left and right vomerine tooth series was present in both sexes except for a single female (present in 90.9%). Short anteriorly directed sub-branch of vomerine tooth series at inner meeting occurred in 27.3% of specimens that were all females (Table 3), and the frequency between sexes was significantly different (
<emphasis id="B9228A60FFC5FFA9FE677CDDFE78F979" box="[410,424,1758,1781]" italics="true" pageId="17" pageNumber="70">p</emphasis>
= 0.033).
</paragraph>
<paragraph id="8BE95672FFC5FFA9FF3A7D02FDC8F80A" blockId="17.[151,1437,1324,2034]" pageId="17" pageNumber="70">Color pattern in life was stable among individuals, only amount of ochre flecks on dorsal and ventral sides of the body varied, but neither stripe nor distinct markings was present. Density of silvery white dots on venter also varied from moderate to dense, frosted appearance. On lateral to ventral sides, the ground color was same as dorsum or fading to purplish-gray.</paragraph>
<paragraph id="8BE95672FFC5FFAAFF3A7D92FC2BFEE9" blockId="17.[151,1437,1324,2034]" lastBlockId="18.[151,1437,151,2032]" lastPageId="18" lastPageNumber="71" pageId="17" pageNumber="70">
<emphasis id="B9228A60FFC5FFA9FF3A7D92FEC7F826" bold="true" box="[199,279,1937,1962]" pageId="17" pageNumber="70">Larva.</emphasis>
We could only find five larvae of 
<taxonomicName id="4C562DF1FFC5FFA9FD5C7D92FCDAF825" box="[673,778,1937,1961]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="17" pageNumber="70" phylum="Chordata" rank="species" species="fuscus" status="sp. nov.">
<emphasis id="B9228A60FFC5FFA9FD5C7D92FCDAF825" box="[673,778,1937,1961]" italics="true" pageId="17" pageNumber="70">
<collectingCountry id="F34116E2FFC5FFA9FD5C7D92FD68F824" box="[673,696,1937,1960]" name="Iceland" pageId="17" pageNumber="70">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
<emphasis id="B9228A60FFC5FFA9FCEF7D92FCBDF826" bold="true" box="[786,877,1937,1962]" pageId="17" pageNumber="70">
<taxonomicNameLabel id="A211371BFFC5FFA9FCEF7D92FCBDF826" box="[786,877,1937,1962]" pageId="17" pageNumber="70" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
All the specimens were at premetamorphic stage (stage 69–70), and 
<quantity id="4CAEFB97FFC5FFA9FE8A7DB6FDCDF841" box="[375,541,1973,1998]" metricMagnitude="-2" metricUnit="m" metricValue="3.96" metricValueMax="4.31" metricValueMin="3.61" pageId="17" pageNumber="70" unit="mm" value="39.6" valueMax="43.1" valueMin="36.1">36.1–43.1 mm</quantity>
in SVL and 33.0– 
<quantity id="4CAEFB97FFC5FFA9FD057DB6FC8FF841" box="[760,863,1973,1997]" metricMagnitude="-2" metricUnit="m" metricValue="4.18" pageId="17" pageNumber="70" unit="mm" value="41.8">41.8 mm</quantity>
in TAL, with the resultant TL of 
<quantity id="4CAEFB97FFC5FFA9FB127DB6FA49F841" box="[1263,1433,1973,1998]" metricMagnitude="-2" metricUnit="m" metricValue="7.7" metricValueMax="8.49" metricValueMin="6.909999999999999" pageId="17" pageNumber="70" unit="mm" value="77.0" valueMax="84.9" valueMin="69.1">69.1–84.9 mm</quantity>
. General morphological characteristics are as follows (
<figureCitation id="136D4AF7FFC5FFA9FCF07DDAFCB1F87D" box="[781,865,2009,2034]" captionStart="FIGURE 12" captionStartId="20.[151,250,975,997]" captionTargetBox="[180,1435,193,953]" captionTargetId="figure@20.[151,1435,193,954]" captionTargetPageId="20" captionText="FIGURE 12. Larvae of O. fuscus sp. nov. (A and B) and a sympatric O. japonicus larva (C) from Sanjo-shi, Niigata Prefecture (In preservation). Larvae of O. japonicus in life (D – F) collected from Tadami-machi, Fukushima Prefecture, near the type locality of O. fuscus sp. nov. Scale bar indicates 10 mm." httpUri="https://zenodo.org/record/287196/files/figure.png" pageId="17" pageNumber="70">Fig. 12</figureCitation>
A–B): head rectangular and blunt at snout; body relatively stout; eye slightly prominent; three pairs of short external gills; labial fold well developed at posterior half of upper jaw; caudal fin relatively well-developed dorsally and ventrally; dorsal fin higher than ventral fin; origin of dorsal fin at level of posterior hindlimb to cloaca; ventral fin originating from around posterior one-sixth to two-thirds of tail; tail tip moderately rounded; skin fold on posterior edge of limb; surfaces of palm and sole with dark asperities; digits with acute and curved black claws; dorsum uniformly dark-brown, sometimes with indistinct ochre flecks and/or pale spots; ventrally whitish or grayish without markings.
</paragraph>
<paragraph id="8BE95672FFC6FFAAFF3A7B6CFCFBFE79" blockId="18.[151,1437,151,2032]" pageId="18" pageNumber="71">
<emphasis id="B9228A60FFC6FFAAFF3A7B6CFEBDFE04" bold="true" box="[199,365,367,392]" pageId="18" pageNumber="71">Comparisons.</emphasis>
Morphological data are summarized in Tables 1–4, and some data are from 
<bibRefCitation id="EFC72B83FFC6FFAAFB167B6CFF31FE20" pageId="18" pageNumber="71" refString="Poyarkov, N. A., Che, J., Min, M. S., Kuro-o, M., Yan, F., Li, C., Iizuka, K. &amp; Vieites, D. R. (2012) Review of the systematics, morphology and distribution of Asian clawed salamanders, genus Onychodactylus (Amphibia, Caudata: Hynobiidae), with the description of four new species. Zootaxa, 3465, 1 - 106." type="journal article">
Poyarkov 
<emphasis id="B9228A60FFC6FFAAFA9C7B72FA4CFE04" box="[1377,1436,368,392]" italics="true" pageId="18" pageNumber="71">et al.</emphasis>
(2012)
</bibRefCitation>
and Yoshikawa 
<emphasis id="B9228A60FFC6FFAAFE657B96FE18FE20" box="[408,456,404,428]" italics="true" pageId="18" pageNumber="71">et al</emphasis>
. (2013). 
<taxonomicName id="4C562DF1FFC6FFAAFDD27B97FCFFFE20" box="[559,815,404,428]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus" status="sp. nov.">
<emphasis id="B9228A60FFC6FFAAFDD27B97FCFFFE20" box="[559,815,404,428]" italics="true" pageId="18" pageNumber="71">Onychodactylus fuscus</emphasis>
</taxonomicName>
<emphasis id="B9228A60FFC6FFAAFCCB7B97FC5FFE21" bold="true" box="[822,911,404,429]" pageId="18" pageNumber="71">
<taxonomicNameLabel id="A211371BFFC6FFAAFCCB7B97FC5FFE21" box="[822,911,404,429]" pageId="18" pageNumber="71" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is different from the continental 
<taxonomicName id="4C562DF1FFC6FFAAFB007B97FAA3FE20" box="[1277,1395,404,428]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fischeri">
<emphasis id="B9228A60FFC6FFAAFB007B97FAA3FE20" box="[1277,1395,404,428]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFB007B97FAC4FE27" box="[1277,1300,404,427]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fischeri
</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C562DF1FFC6FFAAFA7C7B97FF2DFE5C" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="koreanus">
<emphasis id="B9228A60FFC6FFAAFA7C7B97FF2DFE5C" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFA7C7B97FA48FE27" box="[1409,1432,404,427]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. koreanus
</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C562DF1FFC6FFAAFEF67BBBFE14FE5C" box="[267,452,440,464]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="zhangyapingi">
<emphasis id="B9228A60FFC6FFAAFEF67BBBFE14FE5C" box="[267,452,440,464]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFEF67BBBFEF2FE43" box="[267,290,440,463]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. zhangyapingi
</emphasis>
</taxonomicName>
, and 
<taxonomicName id="4C562DF1FFC6FFAAFDFF7BBBFD45FE5C" box="[514,661,440,464]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="zhaoermii">
<emphasis id="B9228A60FFC6FFAAFDFF7BBBFD45FE5C" box="[514,661,440,464]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFDFF7BBBFDC9FE43" box="[514,537,440,463]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. zhaoermii
</emphasis>
</taxonomicName>
in complete lack of distinct dorsal markings (vs. presence of distinct spots, mottlings, or indistinct stripe in continental species).
</paragraph>
<paragraph id="8BE95672FFC6FFAAFF3A7803FED3FD08" blockId="18.[151,1437,151,2032]" pageId="18" pageNumber="71">
<taxonomicName id="4C562DF1FFC6FFAAFF3A7803FE1BFD94" box="[199,459,512,536]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFF3A7803FE1BFD94" box="[199,459,512,536]" italics="true" pageId="18" pageNumber="71">Onychodactylus fuscus</emphasis>
</taxonomicName>
has smaller SVL, relatively shorter trunk, longer forelimb and hindlimb, longer and wider head, and larger 
<collectingCountry id="F34116E2FFC6FFAAFE607826FE02FDB0" box="[413,466,549,572]" name="India" pageId="18" pageNumber="71">IND</collectingCountry>
, 
<collectingCountry id="F34116E2FFC6FFAAFE237826FDCAFDB0" box="[478,538,549,572]" name="India" pageId="18" pageNumber="71">END</collectingCountry>
, and CW (Table 1) in both sexes, and relatively shorter tail in females, than 
<taxonomicName id="4C562DF1FFC6FFAAFA7C7827FF3BFDEC" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fischeri">
<emphasis id="B9228A60FFC6FFAAFA7C7827FF3BFDEC" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFA7C7827FA48FDB7" box="[1409,1432,548,571]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fischeri
</emphasis>
</taxonomicName>
. 
<taxonomicName id="4C562DF1FFC6FFAAFF05784BFE2AFDEC" box="[248,506,584,608]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFF05784BFE2AFDEC" box="[248,506,584,608]" italics="true" pageId="18" pageNumber="71">Onychodactylus fuscus</emphasis>
</taxonomicName>
also has smaller numbers of presacral vertebrae and costal grooves than 
<taxonomicName id="4C562DF1FFC6FFAAFADB784BFA4CFDEC" box="[1318,1436,584,608]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fischeri">
<emphasis id="B9228A60FFC6FFAAFADB784BFA4CFDEC" box="[1318,1436,584,608]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFADB784BFAEDFDD3" box="[1318,1341,584,607]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fischeri
</emphasis>
</taxonomicName>
(Table 2).
</paragraph>
<paragraph id="8BE95672FFC6FFAAFF3A7893FBB7FD7C" blockId="18.[151,1437,151,2032]" pageId="18" pageNumber="71">
<emphasis id="B9228A60FFC6FFAAFF3A7893FE05FD24" box="[199,469,656,680]" italics="true" pageId="18" pageNumber="71">
Oncychodactylus 
<taxonomicName id="4C562DF1FFC6FFAAFE727893FE05FD24" box="[399,469,656,680]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">fuscus</taxonomicName>
</emphasis>
differs from 
<taxonomicName id="4C562DF1FFC6FFAAFD977893FD22FD24" box="[618,754,656,680]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="koreanus">
<emphasis id="B9228A60FFC6FFAAFD977893FD22FD24" box="[618,754,656,680]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFD977893FD51FD2B" box="[618,641,656,679]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. koreanus
</emphasis>
</taxonomicName>
in having shorter tail, smaller number of presacral vertebrae in both sexes, relatively shorter hindlimb, wider internarial space and chest (Table 1), and a smaller number of vomerine teeth in males (Table 3), as well as fewer costal grooves in females (Table 2).
</paragraph>
<paragraph id="8BE95672FFC6FFAAFF3A78FFFCBAFC0C" blockId="18.[151,1437,151,2032]" pageId="18" pageNumber="71">
<taxonomicName id="4C562DF1FFC6FFAAFF3A78FFFE1EFC98" box="[199,462,764,788]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFF3A78FFFE1EFC98" box="[199,462,764,788]" italics="true" pageId="18" pageNumber="71">Onychodactylus fuscus</emphasis>
</taxonomicName>
is different from the other species of the 
<taxonomicName id="4C562DF1FFC6FFAAFC3278FFFBB4FC98" box="[975,1124,764,788]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC6FFAAFC3278FFFBB4FC98" box="[975,1124,764,788]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFC3278FFFC36FC9F" box="[975,998,764,787]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
complex in possession of uniformly dark-brown dorsum with slight, indistinct ochre flecks and completely lacking markings (vs. possessing distinct dorsal stripe or markings). In addition, the gap between vomerine tooth series is much more frequently present (90.9% of specimens) than in other species (6.3–41.7%).
</paragraph>
<paragraph id="8BE95672FFC6FFAAFF3A798FFD4DFBF5" blockId="18.[151,1437,151,2032]" pageId="18" pageNumber="71">
<taxonomicName id="4C562DF1FFC6FFAAFF3A798FFE1EFC28" box="[199,462,908,932]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFF3A798FFE1EFC28" box="[199,462,908,932]" italics="true" pageId="18" pageNumber="71">Onychodactylus fuscus</emphasis>
</taxonomicName>
differs from 
<taxonomicName id="4C562DF1FFC6FFAAFD8E798FFCD7FC28" box="[627,775,908,932]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC6FFAAFD8E798FFCD7FC28" box="[627,775,908,932]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFD8E798FFD5AFC2F" box="[627,650,908,931]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
in: larger SVL in males (
<quantity id="4CAEFB97FFC6FFAAFBB8798FFB60FC28" box="[1093,1200,908,933]" metricMagnitude="-2" metricUnit="m" metricValue="6.95" pageId="18" pageNumber="71" unit="mm" value="69.5">69.5 mm</quantity>
vs. 
<quantity id="4CAEFB97FFC6FFAAFB14798FFA82FC28" box="[1257,1362,908,933]" metricMagnitude="-2" metricUnit="m" metricValue="6.57" pageId="18" pageNumber="71" unit="mm" value="65.7">65.7 mm</quantity>
in 
<taxonomicName id="4C562DF1FFC6FFAAFA7C798FFED4FC4B" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC6FFAAFA7C798FFED4FC4B" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFA7C798FFA48FC2F" box="[1409,1432,908,931]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
); relatively shorter tail (121.9% of SVL in males and 99.7% in females vs. 125.0% and 106.7%) in both sexes; wider head (15.5% of SVL vs. 14.8%) and narrower intercanthal space (57.1% of HW vs. 59.6%) in females; and larger number of costal grooves in males (mode = 13 vs. 12). In coloration, 
<taxonomicName id="4C562DF1FFC6FFAAFB8079F4FB35FB82" box="[1149,1253,1014,1038]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFB8079F4FB35FB82" box="[1149,1253,1014,1038]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFB8079F4FB44FB82" box="[1149,1172,1015,1038]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
has a uniformly dark-brown dorsum lacking markings which differs from 
<taxonomicName id="4C562DF1FFC6FFAAFCDF7E19FC61FBBD" box="[802,945,1050,1073]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC6FFAAFCDF7E19FC61FBBD" box="[802,945,1050,1073]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFCDF7E19FCE9FBBD" box="[802,825,1050,1073]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
with dorsal markings (99.3% of specimens had dorsal markings). A pair of dark markings on the chest tends to be absent in 
<taxonomicName id="4C562DF1FFC6FFAAFBC37E3DFB78FBD9" box="[1086,1192,1085,1109]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFBC37E3DFB78FBD9" box="[1086,1192,1085,1109]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFBC37E3DFB85FBD9" box="[1086,1109,1086,1109]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
(present in 18.2% of specimens vs. 68.6% present in 
<taxonomicName id="4C562DF1FFC6FFAAFE007E62FD5CFBF4" box="[509,652,1121,1144]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC6FFAAFE007E62FD5CFBF4" box="[509,652,1121,1144]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFE007E62FDC4FBF4" box="[509,532,1121,1144]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="8BE95672FFC6FFAAFF3A7E87FC04FA19" blockId="18.[151,1437,151,2032]" pageId="18" pageNumber="71">
<emphasis id="B9228A60FFC6FFAAFF3A7E87FE06FB10" box="[199,470,1156,1180]" italics="true" pageId="18" pageNumber="71">
Oncychodactylus 
<taxonomicName id="4C562DF1FFC6FFAAFE6D7E87FE06FB10" box="[400,470,1156,1180]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">fuscus</taxonomicName>
</emphasis>
significantly differs from 
<taxonomicName id="4C562DF1FFC6FFAAFCF97E86FC43FB10" box="[772,915,1156,1180]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="kinneburi">
<emphasis id="B9228A60FFC6FFAAFCF97E86FC43FB10" box="[772,915,1156,1180]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFCF97E86FCCBFB10" box="[772,795,1157,1180]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. kinneburi
</emphasis>
</taxonomicName>
in having: smaller SVL in females (69.0 mm vs. 
<quantity id="4CAEFB97FFC6FFAAFF407EABFEF4FB4C" box="[189,292,1192,1216]" metricMagnitude="-2" metricUnit="m" metricValue="7.84" pageId="18" pageNumber="71" unit="mm" value="78.4">78.4 mm</quantity>
); wider internarial space (51.3% of SVL vs. 49.1%) in males, wider head (15.5% of SVL vs. 14.5%) in females, and wider chest (13.5% of SVL in males and 13.8% in females vs. 12.3% and 12.2%) in both sexes. 
<taxonomicName id="4C562DF1FFC6FFAAFF6A7EEDFE70FA8B" box="[151,416,1262,1287]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFF6A7EEDFE70FA8B" box="[151,416,1262,1287]" italics="true" pageId="18" pageNumber="71">Onychodactylus fuscus</emphasis>
</taxonomicName>
also differs from 
<taxonomicName id="4C562DF1FFC6FFAAFD787EECFCCAFA8A" box="[645,794,1262,1286]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="kinneburi">
<emphasis id="B9228A60FFC6FFAAFD787EECFCCAFA8A" box="[645,794,1262,1286]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFD787EECFD4CFA8A" box="[645,668,1263,1286]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. kinneburi
</emphasis>
</taxonomicName>
in having significantly smaller numbers of presacral vertebrae in both sexes (mode = 18 or 
<quantity id="4CAEFB97FFC6FFAAFDBC7F11FDAAFAA7" box="[577,634,1298,1323]" metricMagnitude="-1" metricUnit="m" metricValue="4.826" pageId="18" pageNumber="71" unit="in" value="19.0">19 in</quantity>
males and 
<quantity id="4CAEFB97FFC6FFAAFD0A7F11FCFFFAA7" box="[759,815,1298,1323]" metricMagnitude="-1" metricUnit="m" metricValue="4.572" pageId="18" pageNumber="71" unit="in" value="18.0">18 in</quantity>
females vs. 
<quantity id="4CAEFB97FFC6FFAAFC447F11FC21FAA7" box="[953,1009,1298,1323]" metricMagnitude="-1" metricUnit="m" metricValue="4.826" pageId="18" pageNumber="71" unit="in" value="19.0">19 in</quantity>
both sexes in 
<taxonomicName id="4C562DF1FFC6FFAAFB727F11FACCFAA6" box="[1167,1308,1298,1322]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="kinneburi">
<emphasis id="B9228A60FFC6FFAAFB727F11FACCFAA6" box="[1167,1308,1298,1322]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFB727F11FB76FAA5" box="[1167,1190,1298,1321]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. kinneburi
</emphasis>
</taxonomicName>
) and costal grooves in females (mode = 12 vs. 
<quantity id="4CAEFB97FFC6FFAAFDD97F35FD8DFAC2" box="[548,605,1333,1358]" metricMagnitude="-1" metricUnit="m" metricValue="3.302" pageId="18" pageNumber="71" unit="in" value="13.0">13 in</quantity>
<taxonomicName id="4C562DF1FFC6FFAAFD997F35FD22FAC1" box="[612,754,1333,1357]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="kinneburi">
<emphasis id="B9228A60FFC6FFAAFD997F35FD22FAC1" box="[612,754,1333,1357]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFD997F35FDABFAC1" box="[612,635,1334,1357]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. kinneburi
</emphasis>
</taxonomicName>
). In coloration, 
<taxonomicName id="4C562DF1FFC6FFAAFC547F35FBC2FAC2" box="[937,1042,1334,1358]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFC547F35FBC2FAC2" box="[937,1042,1334,1358]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFC547F35FC10FAC1" box="[937,960,1334,1357]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
has uniformly dark-brown dorsum and silvery-dotted ventrolateral sides, and differs from 
<taxonomicName id="4C562DF1FFC6FFAAFCF27F5AFC4FFAFD" box="[783,927,1369,1393]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="kinneburi">
<emphasis id="B9228A60FFC6FFAAFCF27F5AFC4FFAFD" box="[783,927,1369,1393]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFCF27F5AFCF6FAFC" box="[783,806,1369,1392]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. kinneburi
</emphasis>
</taxonomicName>
which has yellowish-orange dorsal stripe or blotches (100% of specimens examined) and no ventrolateral silvery dots.
</paragraph>
<paragraph id="8BE95672FFC6FFAAFF3A7FA3FC51F9CA" blockId="18.[151,1437,151,2032]" pageId="18" pageNumber="71">
<emphasis id="B9228A60FFC6FFAAFF3A7FA3FE08FA34" box="[199,472,1440,1464]" italics="true" pageId="18" pageNumber="71">
Oncychodactylus 
<taxonomicName id="4C562DF1FFC6FFAAFE6F7FA3FE08FA34" box="[402,472,1440,1464]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">fuscus</taxonomicName>
</emphasis>
differs from 
<taxonomicName id="4C562DF1FFC6FFAAFD887FA3FC81FA34" box="[629,849,1440,1464]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="nipponoborealis">
<emphasis id="B9228A60FFC6FFAAFD887FA3FC81FA34" box="[629,849,1440,1464]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFD887FA3FD5CFA3B" box="[629,652,1440,1463]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. nipponoborealis
</emphasis>
</taxonomicName>
in having larger SVL in males (
<quantity id="4CAEFB97FFC6FFAAFB327F9CFAE7FA34" box="[1231,1335,1439,1464]" metricMagnitude="-2" metricUnit="m" metricValue="6.95" pageId="18" pageNumber="71" unit="mm" value="69.5">69.5 mm</quantity>
vs. 
<quantity id="4CAEFB97FFC6FFAAFA967F9CFF17FA50" metricMagnitude="-2" metricUnit="m" metricValue="6.42" pageId="18" pageNumber="71" unit="mm" value="64.2">64.2 mm</quantity>
), and shorter tail (99.7% of SVL vs. 106.7%) in females, and wider internarial space (51.3% of HW vs. 49.7%) in males. In coloration, 
<taxonomicName id="4C562DF1FFC6FFAAFE4D7FE4FDCCFA73" box="[432,540,1511,1535]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFE4D7FE4FDCCFA73" box="[432,540,1511,1535]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFE4D7FE4FE17FA72" box="[432,455,1511,1534]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
is different from 
<taxonomicName id="4C562DF1FFC6FFAAFD0D7FE4FC1CFA73" box="[752,972,1511,1535]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="nipponoborealis">
<emphasis id="B9228A60FFC6FFAAFD0D7FE4FC1CFA73" box="[752,972,1511,1535]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFD0D7FE4FCD7FA72" box="[752,775,1511,1534]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. nipponoborealis
</emphasis>
</taxonomicName>
in lacking dorsal markings (vs. usually having dorsal markings [97.8%]). Dark marking on chest is also less frequently observed in 
<taxonomicName id="4C562DF1FFC6FFAAFB4F7C08FACBF9AE" box="[1202,1307,1546,1570]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFB4F7C08FACBF9AE" box="[1202,1307,1546,1570]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFB4F7C08FB19F9AE" box="[1202,1225,1547,1570]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
(present in 18.2% of specimens) than in 
<taxonomicName id="4C562DF1FFC6FFAAFE207C2DFD66F9CA" box="[477,694,1582,1606]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="nipponoborealis">
<emphasis id="B9228A60FFC6FFAAFE207C2DFD66F9CA" box="[477,694,1582,1606]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFE207C2DFE24F9C9" box="[477,500,1582,1605]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. nipponoborealis
</emphasis>
</taxonomicName>
(65.2%: Table 4).
</paragraph>
<paragraph id="8BE95672FFC6FFAAFF3A7C51FED2F8B3" blockId="18.[151,1437,151,2032]" pageId="18" pageNumber="71">
In comparison with 
<taxonomicName id="4C562DF1FFC6FFAAFE517C51FD8FF9E5" box="[428,607,1617,1641]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="tsukubaensis">
<emphasis id="B9228A60FFC6FFAAFE517C51FD8FF9E5" box="[428,607,1617,1641]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFE517C51FE13F9E5" box="[428,451,1618,1641]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. tsukubaensis
</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C562DF1FFC6FFAAFD927C51FD09F9E5" box="[623,729,1617,1641]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFD927C51FD09F9E5" box="[623,729,1617,1641]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFD927C51FD56F9E5" box="[623,646,1618,1641]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
differs in having longer tail in both sexes (121.9% of SVL in males and 99.7% in females vs. 102.3% and 90.4%) and wider head in females (54.7% of SVL vs. 52.4%), and wider internarial space in males (51.3% of HW vs. 48.3%) and wider interorbital space in both sexes (32.2% of HW in males and 30.8% in females vs. 28.3% and 27.8%). 
<taxonomicName id="4C562DF1FFC6FFAAFCC47CBFFBEBF958" box="[825,1083,1724,1748]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFCC47CBFFBEBF958" box="[825,1083,1724,1748]" italics="true" pageId="18" pageNumber="71">Onychodactylus fuscus</emphasis>
</taxonomicName>
had the gap between vomerine tooth series more frequently than 
<taxonomicName id="4C562DF1FFC6FFAAFDE27CE3FD03F97B" box="[543,723,1759,1783]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="tsukubaensis">
<emphasis id="B9228A60FFC6FFAAFDE27CE3FD03F97B" box="[543,723,1759,1783]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFDE27CE3FDE6F97B" box="[543,566,1760,1783]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. tsukubaensis
</emphasis>
</taxonomicName>
(90.9% present in 
<taxonomicName id="4C562DF1FFC6FFAAFC4E7CE3FBCEF97B" box="[947,1054,1759,1783]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFC4E7CE3FBCEF97B" box="[947,1054,1759,1783]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFC4E7CE3FC1AF97B" box="[947,970,1760,1783]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
vs. 6.3% in 
<taxonomicName id="4C562DF1FFC6FFAAFB4B7CE3FABAF97B" box="[1206,1386,1759,1783]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="tsukubaensis">
<emphasis id="B9228A60FFC6FFAAFB4B7CE3FABAF97B" box="[1206,1386,1759,1783]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFB4B7CE3FB1DF97B" box="[1206,1229,1760,1783]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. tsukubaensis
</emphasis>
</taxonomicName>
). In coloration, 
<taxonomicName id="4C562DF1FFC6FFAAFEE47D00FE53F897" box="[281,387,1795,1819]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFEE47D00FE53F897" box="[281,387,1795,1819]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFEE47D00FEE0F896" box="[281,304,1795,1818]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
differs from 
<taxonomicName id="4C562DF1FFC6FFAAFDE37D00FD02F897" box="[542,722,1795,1819]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="tsukubaensis">
<emphasis id="B9228A60FFC6FFAAFDE37D00FD02F897" box="[542,722,1795,1819]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFDE37D00FDE5F896" box="[542,565,1795,1818]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. tsukubaensis
</emphasis>
</taxonomicName>
by lacking dorsal markings (vs. typically having dorsal stripe [93.8%]).
</paragraph>
<paragraph id="8BE95672FFC6FFABFF3A7D49FBD6FF7B" blockId="18.[151,1437,151,2032]" lastBlockId="19.[151,1437,151,1934]" lastPageId="19" lastPageNumber="72" pageId="18" pageNumber="71">
In comparison with 
<taxonomicName id="4C562DF1FFC6FFAAFE5B7D49FD9BF8ED" box="[422,587,1865,1889]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="B9228A60FFC6FFAAFE5B7D49FD9BF8ED" box="[422,587,1865,1889]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFE5B7D49FE6DF8ED" box="[422,445,1866,1889]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. intermedius
</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C562DF1FFC6FFAAFDA47D49FD11F8ED" box="[601,705,1865,1889]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFDA47D49FD11F8ED" box="[601,705,1865,1889]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFDA47D49FDA0F8ED" box="[601,624,1866,1889]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
differs in having significantly larger SVL in males (69.5±3.0 mm vs. 63.0±4.9) and tendency of larger SVL in females (69.0± 
<quantity id="4CAEFB97FFC6FFAAFCB87D6EFC4DF809" box="[837,925,1901,1926]" metricMagnitude="-3" metricUnit="m" metricValue="4.8" pageId="18" pageNumber="71" unit="mm" value="4.8">4.8 mm</quantity>
vs. 65.5±4.1), significantly wider internarial space relative to HW in males (51.3% vs. 49.4%) and narrower intercanthal space relative to HW in females (57.1% vs. 59.3%). Relative HLL also tended to be shorter in both sexes (30.0% in males and 30.9% in females vs. 31.5% and 32.1%).Number of vomerine teeth was significantly larger in 
<taxonomicName id="4C562DF1FFC6FFAAFC207DDBFB97F87C" box="[989,1095,2008,2032]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC6FFAAFC207DDBFB97F87C" box="[989,1095,2008,2032]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFC207DDBFC24F863" box="[989,1012,2008,2031]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
than 
<taxonomicName id="4C562DF1FFC6FFAAFB777DDBFAE2F863" box="[1162,1330,2007,2031]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="18" pageNumber="71" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="B9228A60FFC6FFAAFB777DDBFAE2F863" box="[1162,1330,2007,2031]" italics="true" pageId="18" pageNumber="71">
<collectingCountry id="F34116E2FFC6FFAAFB777DDBFB71F863" box="[1162,1185,2008,2031]" name="Iceland" pageId="18" pageNumber="71">O</collectingCountry>
. intermedius
</emphasis>
</taxonomicName>
in males (12.4±1.7 vs. 15.9±2.4). Presence of the gap between vomerine tooth series is more frequent in 
<taxonomicName id="4C562DF1FFC7FFABFB427A9BFAF7FF3C" box="[1215,1319,152,176]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC7FFABFB427A9BFAF7FF3C" box="[1215,1319,152,176]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFB427A9BFB06FF23" box="[1215,1238,152,175]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
than in 
<taxonomicName id="4C562DF1FFC7FFABFA7C7A9BFECCFF5F" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="B9228A60FFC7FFABFA7C7A9BFECCFF5F" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFA7C7A9BFA48FF23" box="[1409,1432,152,175]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. intermedius
</emphasis>
</taxonomicName>
(90.9% present in 
<taxonomicName id="4C562DF1FFC7FFABFE017ABFFDB8FF5F" box="[508,616,187,211]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC7FFABFE017ABFFDB8FF5F" box="[508,616,187,211]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFE017ABFFDC3FF5F" box="[508,531,188,211]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
vs. 29.6% in 
<taxonomicName id="4C562DF1FFC7FFABFCF27ABFFC68FF5F" box="[783,952,187,211]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="B9228A60FFC7FFABFCF27ABFFC68FF5F" box="[783,952,187,211]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFCF27ABFFCF6FF5F" box="[783,806,188,211]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. intermedius
</emphasis>
</taxonomicName>
). In coloration, 
<taxonomicName id="4C562DF1FFC7FFABFB8A7ABFFB33FF5F" box="[1143,1251,187,211]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC7FFABFB8A7ABFFB33FF5F" box="[1143,1251,187,211]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFB8A7ABFFB5EFF5F" box="[1143,1166,188,211]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
differs from 
<taxonomicName id="4C562DF1FFC7FFABFA7C7ABFFECCFF7B" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="B9228A60FFC7FFABFA7C7ABFFECCFF7B" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFA7C7ABFFA48FF5F" box="[1409,1432,188,211]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. intermedius
</emphasis>
</taxonomicName>
by lacking dorsal markings (vs. having dorsal stripe or mottlings).
</paragraph>
<paragraph id="8BE95672FFC7FFABFF3A7B00FC94FDD6" blockId="19.[151,1437,151,1934]" pageId="19" pageNumber="72">
At the larval stage, 
<taxonomicName id="4C562DF1FFC7FFABFE507B00FDC8FE96" box="[429,536,258,282]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC7FFABFE507B00FDC8FE96" box="[429,536,258,282]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFE507B00FE14FE96" box="[429,452,259,282]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
is morphologically similar to the other congeneric species, but has uniformly dark-brown dorsum with some small ochre flecks, and it can be distinguished from larval 
<taxonomicName id="4C562DF1FFC7FFABFB617B25FAFDFEB2" box="[1180,1325,294,318]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC7FFABFB617B25FAFDFEB2" box="[1180,1325,294,318]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFB617B25FB63FEB1" box="[1180,1203,294,317]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
, which is sympatric in one locality (
<figureCitation id="136D4AF7FFC7FFABFE3C7B4AFDC3FEED" box="[449,531,329,354]" captionStart="FIGURE 12" captionStartId="20.[151,250,975,997]" captionTargetBox="[180,1435,193,953]" captionTargetId="figure@20.[151,1435,193,954]" captionTargetPageId="20" captionText="FIGURE 12. Larvae of O. fuscus sp. nov. (A and B) and a sympatric O. japonicus larva (C) from Sanjo-shi, Niigata Prefecture (In preservation). Larvae of O. japonicus in life (D – F) collected from Tadami-machi, Fukushima Prefecture, near the type locality of O. fuscus sp. nov. Scale bar indicates 10 mm." httpUri="https://zenodo.org/record/287196/files/figure.png" pageId="19" pageNumber="72">Fig. 12</figureCitation>
C–F). In 
<taxonomicName id="4C562DF1FFC7FFABFD817B4AFCDCFEED" box="[636,780,329,353]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC7FFABFD817B4AFCDCFEED" box="[636,780,329,353]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFD817B4AFD43FEEC" box="[636,659,329,352]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
larvae of neighboring or sympatric populations, there are two morphotypes, striped and unstriped 
<typeStatus id="54EDE8D0FFC7FFABFD957B6EFD76FE09" box="[616,678,365,389]" pageId="19" pageNumber="72">types</typeStatus>
, and striped 
<typeStatus id="54EDE8D0FFC7FFABFCC17B6EFCBCFE09" box="[828,876,365,389]" pageId="19" pageNumber="72">type</typeStatus>
can easily be distinguished from 
<taxonomicName id="4C562DF1FFC7FFABFB097B6EFAB0FE09" box="[1268,1376,365,389]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC7FFABFB097B6EFAB0FE09" box="[1268,1376,365,389]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFB097B6EFADBFE08" box="[1268,1291,365,388]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
. The unstriped 
<taxonomicName id="4C562DF1FFC7FFABFEFB7B93FE45FE24" box="[262,405,400,424]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC7FFABFEFB7B93FE45FE24" box="[262,405,400,424]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFEFB7B93FECDFE2B" box="[262,285,400,423]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
larvae has a varying amount of black mottlings on yellowish-gray background and it differs from uniformly dark-brown 
<taxonomicName id="4C562DF1FFC7FFABFE2A7BB7FDEAFE40" box="[471,570,436,460]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fiscus">
<emphasis id="B9228A60FFC7FFABFE2A7BB7FDEAFE40" box="[471,570,436,460]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFE2A7BB7FE3EFE47" box="[471,494,436,459]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fiscus
</emphasis>
</taxonomicName>
. From the other species in the 
<taxonomicName id="4C562DF1FFC7FFABFC6A7BB7FBF7FE47" box="[919,1063,436,459]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC7FFABFC6A7BB7FBF7FE47" box="[919,1063,436,459]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFC6A7BB7FC7EFE47" box="[919,942,436,459]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
complex, 
<taxonomicName id="4C562DF1FFC7FFABFB607BB7FB2FFE40" box="[1181,1279,436,460]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC7FFABFB607BB7FB2FFE40" box="[1181,1279,436,460]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFB607BB7FB61FE47" box="[1181,1201,436,459]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
fuscus
</emphasis>
</taxonomicName>
larvae can be distinguished by the presence of various 
<typeStatus id="54EDE8D0FFC7FFABFD8A7BDBFD62FE7C" box="[631,690,472,496]" pageId="19" pageNumber="72">types</typeStatus>
of dorsal markings (distinct stripe, continuous or discontinuous series of blotches, or varying amount of mottlings). However, we would like to note that these diagnoses of 
<taxonomicName id="4C562DF1FFC7FFABFA7C7BF8FF0DFDBA" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC7FFABFA7C7BF8FF0DFDBA" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFA7C7BF8FA48FD9E" box="[1409,1432,507,530]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
are provisional, and future collection of more larval specimens and detailed comparisons are necessary to understand morphological characteristics of 
<taxonomicName id="4C562DF1FFC7FFABFD757841FD21FDD5" box="[648,753,577,601]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC7FFABFD757841FD21FDD5" box="[648,753,577,601]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFD757841FD4FFDD5" box="[648,671,578,601]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
larvae.
</paragraph>
<paragraph id="8BE95672FFC7FFABFF3A7866FC74FDF1" blockId="19.[151,1437,151,1934]" box="[199,932,613,638]" pageId="19" pageNumber="72">
<emphasis id="B9228A60FFC7FFABFF3A7866FE9EFDF2" bold="true" box="[199,334,613,638]" pageId="19" pageNumber="72">Karyotype.</emphasis>
No karyotypic information is available for 
<taxonomicName id="4C562DF1FFC7FFABFCC87866FC4DFDF1" box="[821,925,613,637]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC7FFABFCC87866FC4DFDF1" box="[821,925,613,637]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFCC87866FC9CFDF0" box="[821,844,613,636]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BE95672FFC7FFABFF3A788AFED5FC6A" blockId="19.[151,1437,151,1934]" pageId="19" pageNumber="72">
<emphasis id="B9228A60FFC7FFABFF3A788AFE0FFD2E" bold="true" box="[199,479,649,674]" pageId="19" pageNumber="72">Genetic characteristics.</emphasis>
<taxonomicName id="4C562DF1FFC7FFABFE18788AFD36FD2D" box="[485,742,649,673]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus" status="sp. nov.">
<emphasis id="B9228A60FFC7FFABFE18788AFD36FD2D" box="[485,742,649,673]" italics="true" pageId="19" pageNumber="72">Onychodactylus fuscus</emphasis>
</taxonomicName>
<emphasis id="B9228A60FFC7FFABFD10788AFC96FD2E" bold="true" box="[749,838,649,674]" pageId="19" pageNumber="72">
<taxonomicNameLabel id="A211371BFFC7FFABFD10788AFC96FD2E" box="[749,838,649,674]" pageId="19" pageNumber="72" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
forms a distinct clade, Subclade II-C, which is newly reported in this study (
<figureCitation id="136D4AF7FFC7FFABFE5E78AEFE3CFD49" box="[419,492,685,710]" captionStart="FIGURE 2" captionStartId="5.[151,250,1953,1975]" captionTargetBox="[254,1330,199,1928]" captionTargetId="figure@5.[242,1344,193,1932]" captionTargetPageId="5" captionText="FIGURE 2. The ML trees based on 1141 bp of mitochondrial cytochrome b gene, showing entire shape of the ML tree (A), detail of Subclade II-A (B), and detail of Clade III (C). Values at nodes indicate supports of 1000 bootstrap replicates. Numbers in parenthesis indicate population numbers shown in Appendix 3." httpUri="https://zenodo.org/record/287186/files/figure.png" pageId="19" pageNumber="72">Fig. 2</figureCitation>
). Based on the mitochondrial phylogeny, the new species is a sister taxon of 
<taxonomicName id="4C562DF1FFC7FFABFA7C78AEFECCFD65" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="intermedius">
<emphasis id="B9228A60FFC7FFABFA7C78AEFECCFD65" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFA7C78AEFA48FD48" box="[1409,1432,685,708]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. intermedius
</emphasis>
</taxonomicName>
described above (Subclade II-A), and is also closely related to 
<taxonomicName id="4C562DF1FFC7FFABFC0978D2FB78FD65" box="[1012,1192,721,745]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="tsukubaensis">
<emphasis id="B9228A60FFC7FFABFC0978D2FB78FD65" box="[1012,1192,721,745]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFC0978D2FBDBFD64" box="[1012,1035,721,744]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. tsukubaensis
</emphasis>
</taxonomicName>
(Subclade II-B) with mean pairwise p-distances between them for cytb gene of 4.15% (3.59–4.73%) and 5.51% (5.35–5.70%), respectively. Within the species, pairwise p-distances varied up to 1.75%, and two distinct haplotype groups were recognized although they showed no meaningful geographic genetic structure. The new species is sympatric in some localities with 
<taxonomicName id="4C562DF1FFC7FFABFE667962FDE4FCF5" box="[411,564,865,889]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC7FFABFE667962FDE4FCF5" box="[411,564,865,889]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFE667962FE62FCF4" box="[411,434,865,888]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
, but is genetically distinct from it, with a large p-distance of 8.18% (7.71–8.85%). Analysis of microsatellite loci also showed substantial difference between these sympatric species, and indicated presence of a high degree of reproductive isolation between them at least in these instances of sympatry.
</paragraph>
<paragraph id="8BE95672FFC7FFABFF3A79F2FC62FB6D" blockId="19.[151,1437,151,1934]" pageId="19" pageNumber="72">
<emphasis id="B9228A60FFC7FFABFF3A79F2FDE5FB86" bold="true" box="[199,565,1009,1034]" pageId="19" pageNumber="72">Fecundity and natural history.</emphasis>
Breeding habits of 
<taxonomicName id="4C562DF1FFC7FFABFCE979F2FCADFB85" box="[788,893,1009,1033]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus" status="sp. nov.">
<emphasis id="B9228A60FFC7FFABFCE979F2FCADFB85" box="[788,893,1009,1033]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFCE979F2FCFBFB84" box="[788,811,1009,1032]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A211371BFFC7FFABFC7979F2FC0EFB85" box="[900,990,1009,1034]" pageId="19" pageNumber="72" rank="species">
<emphasis id="B9228A60FFC7FFABFC7979F2FC07FB86" bold="true" box="[900,983,1009,1034]" pageId="19" pageNumber="72">sp. nov</emphasis>
.
</taxonomicNameLabel>
is not fully understood, but it is certain that the species breeds in late autumn to early winter. It is unknown if this species also breeds in summer as known in some congeners, or breeds biannually (summer and winter) as seen in 
<taxonomicName id="4C562DF1FFC7FFABFC187E3AFBA7FBDD" box="[997,1143,1081,1105]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC7FFABFC187E3AFBA7FBDD" box="[997,1143,1081,1105]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFC187E3AFC2CFBDC" box="[997,1020,1081,1104]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
in Ishikawa Pref. (
<bibRefCitation id="EFC72B83FFC7FFABFAA87E3AFF03FBF9" author="Akita" pageId="19" pageNumber="72" refString="Akita, Y. (1989) Breeding cycle of Onychodactylus japonicus on Mt. Hodatsu, with special reference to biannual spawning. In: Matsui, M., Hikida, T. &amp; Goris, R. C. (Eds.), Current Herpetology in East Asia. Herpetological Society of Japan, Kyoto, pp. 305." type="book chapter" year="1989">Akita, 1989</bibRefCitation>
). In the 
<typeStatus id="54EDE8D0FFC7FFABFEC17E5DFEBCFBFA" box="[316,364,1118,1142]" pageId="19" pageNumber="72">type</typeStatus>
locality, assemblage of breeding adults (seven males and two females) was observed on 
<date id="FFE870B2FFC7FFABFA737E5EFE88FB15" pageId="19" pageNumber="72" value="2013-11-09">9 November 2013</date>
, and newly laid egg sacs were confirmed on 
<date id="FFE870B2FFC7FFABFC5F7E82FB5DFB15" box="[930,1165,1153,1178]" pageId="19" pageNumber="72" value="2013-11-23">23 November 2013</date>
(
<collectingCountry id="F34116E2FFC7FFABFB5B7E82FB6BFB15" box="[1190,1211,1153,1177]" name="Italy" pageId="19" pageNumber="72">Y</collectingCountry>
. Nakano, personal communication). Egg sacs were strongly adhered under the rock by gelatinous stalks in complete darkness in the underground flowing water at a headstream (water temperature 9.3°C).
</paragraph>
<paragraph id="8BE95672FFC7FFABFF3A7EEEFE7CFA52" blockId="19.[151,1437,151,1934]" pageId="19" pageNumber="72">
The egg sacs of 
<taxonomicName id="4C562DF1FFC7FFABFE807EEEFE35FA89" box="[381,485,1261,1285]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC7FFABFE807EEEFE35FA89" box="[381,485,1261,1285]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFE807EEEFE44FA88" box="[381,404,1261,1284]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
are similar to those of 
<taxonomicName id="4C562DF1FFC7FFABFD1A7EEEFBDEFA89" authority="(Akita 1982)" baseAuthorityName="Akita" baseAuthorityYear="1982" box="[743,1038,1261,1286]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC7FFABFD1A7EEEFCA7FA89" box="[743,887,1261,1285]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFD1A7EEEFD2EFA88" box="[743,766,1261,1284]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. japonicus
</emphasis>
(
<bibRefCitation id="EFC72B83FFC7FFABFC787EEEFBD5FA89" author="Akita" box="[901,1029,1261,1286]" pageId="19" pageNumber="72" refString="Akita, Y. (1982) Notes on the egg-laying site of Onychodactylus japonicus on Mt. Hodatsu. Japanese Journal of Herpetology, 9, 111 - 117." type="journal article" year="1982">Akita 1982</bibRefCitation>
)
</taxonomicName>
and 
<taxonomicName id="4C562DF1FFC7FFABFBB97EEEFF2CFAA5" authority="Yoshikawa et al. 2013" authorityName="Yoshikawa et al." authorityYear="2013" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="tsukubaensis">
<emphasis id="B9228A60FFC7FFABFBB97EEEFB25FA89" box="[1092,1269,1261,1285]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFBB97EEEFB8BFA88" box="[1092,1115,1261,1284]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. tsukubaensis
</emphasis>
(Yoshikawa 
<emphasis id="B9228A60FFC7FFABFA757EEDFF64FAA5" italics="true" pageId="19" pageNumber="72">et al.</emphasis>
2013)
</taxonomicName>
in shape and texture (
<figureCitation id="136D4AF7FFC7FFABFE0E7F12FD95FAA5" box="[499,581,1297,1322]" captionStart="FIGURE 13. A" captionStartId="20.[151,250,1979,2001]" captionTargetBox="[151,1435,1108,1957]" captionTargetId="figure@20.[151,1436,1108,1957]" captionTargetPageId="20" captionText="FIGURE 13. A pair of egg sacs of O. fuscus sp. nov. in the type locality on 20 December 2013, about one month later from the deposition. Scale bar indicates 10 mm." httpUri="https://zenodo.org/record/287197/files/figure.png" pageId="19" pageNumber="72">Fig. 13</figureCitation>
), and were spindle in shape, with transparent, elastic, and strong outer layer. The egg sacs had very weak, longitudinal grooves on the surface, and short, filamentous structure on free ends. Among six pairs of egg sacs observed, length of sacs excluding stalk ranged 30.0– 
<quantity id="4CAEFB97FFC7FFABFB7F7F5AFB3CFAFD" box="[1154,1260,1369,1394]" metricMagnitude="-2" metricUnit="m" metricValue="3.62" pageId="19" pageNumber="72" unit="mm" value="36.2">36.2 mm</quantity>
(mean±1SD = 32.3±2.3, n = 12). In the egg sac, eggs were arranged in two rows, and the clutch size ranged from 13 to 16 (6– 
<specimenCount id="9D509DFBFFC7FFABFA737F7EFA4CFA1A" box="[1422,1436,1405,1430]" pageId="19" pageNumber="72" type="egg">9</specimenCount>
eggs per egg sac). Eggs were large, with diameters of 5.0– 
<quantity id="4CAEFB97FFC7FFABFCD47FA2FCAFFA35" box="[809,895,1441,1466]" metricMagnitude="-3" metricUnit="m" metricValue="6.5" pageId="19" pageNumber="72" unit="mm" value="6.5">6.5 mm</quantity>
(mean±1SD = 5.9±0.3, n = 87), and completely yellowish white in color.
</paragraph>
<paragraph id="8BE95672FFC7FFABFF3A7FEAFBE6F9C5" blockId="19.[151,1437,151,1934]" pageId="19" pageNumber="72">
Larval life history of 
<taxonomicName id="4C562DF1FFC7FFABFE3D7FEAFDFBF98D" box="[448,555,1513,1537]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus">
<emphasis id="B9228A60FFC7FFABFE3D7FEAFDFBF98D" box="[448,555,1513,1537]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFE3D7FEAFE07F98C" box="[448,471,1513,1536]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fuscus
</emphasis>
</taxonomicName>
, including the season and the size at hatching and metamorphosis, is poorly known. Our limited number of specimens suggest that larvae reach at least 
<quantity id="4CAEFB97FFC7FFABFC127C0EFB85F9A9" box="[1007,1109,1549,1573]" metricMagnitude="-2" metricUnit="m" metricValue="4.31" pageId="19" pageNumber="72" unit="mm" value="43.1">43.1 mm</quantity>
in SVL and 
<quantity id="4CAEFB97FFC7FFABFB157C0EFA9DF9A9" box="[1256,1357,1549,1573]" metricMagnitude="-2" metricUnit="m" metricValue="8.49" pageId="19" pageNumber="72" unit="mm" value="84.9">84.9 mm</quantity>
in TL. Metamorphosed juveniles would disperse to inhabit forest floor around the stream.
</paragraph>
<paragraph id="8BE95672FFC7FFABFF3A7C56FE47F972" blockId="19.[151,1437,151,1934]" pageId="19" pageNumber="72">
Life history of adults in the non-breeding season is unknown, but they are presumed to inhabit the cool and humid forest floor in the adjacent mountains, as is the case in the other 
<taxonomicName id="4C562DF1FFC7FFABFC4C7C7AFBB6F91D" box="[945,1126,1657,1681]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="genus">
<emphasis id="B9228A60FFC7FFABFC4C7C7AFBB6F91D" box="[945,1126,1657,1681]" italics="true" pageId="19" pageNumber="72">Onychodactylus</emphasis>
</taxonomicName>
species. The new species is sympatric with 
<taxonomicName id="4C562DF1FFC7FFABFEB37C9EFE32F939" box="[334,482,1693,1717]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC7FFABFEB37C9EFE32F939" box="[334,482,1693,1717]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFEB37C9EFEB5F938" box="[334,357,1693,1716]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
in one locality, and is probably so in its entire distributional range, but detailed distributional patterns and ecological features of these species, as well as mechanisms of their reproductive isolation are unknown.
</paragraph>
<paragraph id="8BE95672FFC7FFABFF3A7D0AFA57F801" blockId="19.[151,1437,151,1934]" pageId="19" pageNumber="72">
<emphasis id="B9228A60FFC7FFABFF3A7D0AFECAF8AE" bold="true" box="[199,282,1801,1826]" pageId="19" pageNumber="72">Range.</emphasis>
Known from two localities in western Tadami-machi, Fukushima Prefecture and southern Sanjo-shi, Niigata Prefecture. However, actual distributional range is probably wider. The new species is sympatric with 
<taxonomicName id="4C562DF1FFC7FFABFA7C7D2EFED4F8E5" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC7FFABFA7C7D2EFED4F8E5" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFA7C7D2EFA48F8C8" box="[1409,1432,1837,1860]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
in Sanjo-shi, Niigata Prefecture, where larvae of the two species are found together. In addition, we also found larvae of 
<taxonomicName id="4C562DF1FFC7FFABFEB47D76FE08F801" box="[329,472,1909,1933]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC7FFABFEB47D76FE08F801" box="[329,472,1909,1933]" italics="true" pageId="19" pageNumber="72">
<collectingCountry id="F34116E2FFC7FFABFEB47D76FEB0F800" box="[329,352,1909,1932]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. japonicus
</emphasis>
</taxonomicName>
(
<figureCitation id="136D4AF7FFC7FFABFE1A7D76FDE7F801" box="[487,567,1909,1934]" captionStart="FIGURE 12" captionStartId="20.[151,250,975,997]" captionTargetBox="[180,1435,193,953]" captionTargetId="figure@20.[151,1435,193,954]" captionTargetPageId="20" captionText="FIGURE 12. Larvae of O. fuscus sp. nov. (A and B) and a sympatric O. japonicus larva (C) from Sanjo-shi, Niigata Prefecture (In preservation). Larvae of O. japonicus in life (D – F) collected from Tadami-machi, Fukushima Prefecture, near the type locality of O. fuscus sp. nov. Scale bar indicates 10 mm." httpUri="https://zenodo.org/record/287196/files/figure.png" pageId="19" pageNumber="72">Fig. 12</figureCitation>
D–F) from a stream close (ca. 
<quantity id="4CAEFB97FFC7FFABFC707D76FC6FF802" box="[909,959,1909,1934]" metricMagnitude="3" metricUnit="m" metricValue="5.0" pageId="19" pageNumber="72" unit="km" value="5.0">5km</quantity>
distant) to the 
<typeStatus id="54EDE8D0FFC7FFABFB9A7D75FB47F802" box="[1127,1175,1910,1934]" pageId="19" pageNumber="72">type</typeStatus>
locality of 
<emphasis id="B9228A60FFC7FFABFAEA7D76FA57F801" box="[1303,1415,1909,1933]" italics="true" pageId="19" pageNumber="72">
<taxonomicName id="4C562DF1FFC7FFABFAEA7D76FA53F801" box="[1303,1411,1909,1933]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="19" pageNumber="72" phylum="Chordata" rank="species" species="fuscus">
<collectingCountry id="F34116E2FFC7FFABFAEA7D76FAFEF800" box="[1303,1326,1909,1932]" name="Iceland" pageId="19" pageNumber="72">O</collectingCountry>
. fuscus
</taxonomicName>
.
</emphasis>
</paragraph>
<caption id="DF2906FAFFC0FFACFF6A79CCFD02FBAE" httpUri="https://zenodo.org/record/287196/files/figure.png" pageId="20" pageNumber="73" targetBox="[180,1435,193,953]" targetPageId="20">
<paragraph id="8BE95672FFC0FFACFF6A79CCFD02FBAE" blockId="20.[151,1436,975,1058]" pageId="20" pageNumber="73">
<emphasis id="B9228A60FFC0FFACFF6A79CCFEC8FC69" bold="true" box="[151,280,975,997]" pageId="20" pageNumber="73">FIGURE 12</emphasis>
. Larvae of 
<taxonomicName id="4C562DF1FFC0FFACFE7179D3FE3AFC69" box="[396,490,975,997]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="20" pageNumber="73" phylum="Chordata" rank="species" species="fuscus" status="sp. nov.">
<emphasis id="B9228A60FFC0FFACFE7179D3FE3AFC69" box="[396,490,975,997]" italics="true" pageId="20" pageNumber="73">O. fuscus</emphasis>
</taxonomicName>
<emphasis id="B9228A60FFC0FFACFE0C79CCFD91FC69" bold="true" box="[497,577,975,997]" pageId="20" pageNumber="73">
<taxonomicNameLabel id="A211371BFFC0FFACFE0C79CCFD91FC69" box="[497,577,975,997]" pageId="20" pageNumber="73" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(A and B) and a sympatric 
<taxonomicName id="4C562DF1FFC0FFACFCA579D3FC09FC69" box="[856,985,976,997]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="20" pageNumber="73" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC0FFACFCA579D3FC09FC69" box="[856,985,976,997]" italics="true" pageId="20" pageNumber="73">O. japonicus</emphasis>
</taxonomicName>
larva (C) from Sanjo-shi, Niigata Prefecture (In preservation). Larvae of 
<taxonomicName id="4C562DF1FFC0FFACFE3F79ECFD97FB88" box="[450,583,1007,1028]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="20" pageNumber="73" phylum="Chordata" rank="species" species="japonicus">
<emphasis id="B9228A60FFC0FFACFE3F79ECFD97FB88" box="[450,583,1007,1028]" italics="true" pageId="20" pageNumber="73">O. japonicus</emphasis>
</taxonomicName>
in life (D–F) collected from Tadami-machi, Fukushima Prefecture, near the type locality of 
<taxonomicName id="4C562DF1FFC0FFACFEF97E0EFEB3FBAE" box="[260,355,1036,1058]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="20" pageNumber="73" phylum="Chordata" rank="species" species="fuscus" status="sp. nov.">
<emphasis id="B9228A60FFC0FFACFEF97E0EFEB3FBAE" box="[260,355,1036,1058]" italics="true" pageId="20" pageNumber="73">O. fuscus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A211371BFFC0FFACFE947E0FFE69FBAE" box="[361,441,1036,1058]" pageId="20" pageNumber="73" rank="species">
<emphasis id="B9228A60FFC0FFACFE947E0FFE63FBAE" bold="true" box="[361,435,1036,1058]" pageId="20" pageNumber="73">sp. nov</emphasis>
.
</taxonomicNameLabel>
Scale bar indicates 10 mm.
</paragraph>
</caption>
<caption id="DF2906FAFFC0FFACFF6A7DB8FDCFF87C" httpUri="https://zenodo.org/record/287197/files/figure.png" pageId="20" pageNumber="73" targetBox="[151,1435,1108,1957]" targetPageId="20">
<paragraph id="8BE95672FFC0FFACFF6A7DB8FDCFF87C" blockId="20.[151,1436,1979,2032]" pageId="20" pageNumber="73">
<emphasis id="B9228A60FFC0FFACFF6A7DB8FEC8F85D" bold="true" box="[151,280,1979,2001]" pageId="20" pageNumber="73">FIGURE 13</emphasis>
. A pair of egg sacs of 
<taxonomicName id="4C562DF1FFC0FFACFE067DBFFD89F85D" box="[507,601,1979,2001]" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="20" pageNumber="73" phylum="Chordata" rank="species" species="fuscus" status="sp. nov.">
<emphasis id="B9228A60FFC0FFACFE067DBFFD89F85D" box="[507,601,1979,2001]" italics="true" pageId="20" pageNumber="73">O. fuscus</emphasis>
</taxonomicName>
<emphasis id="B9228A60FFC0FFACFDA27DB8FD7FF85D" bold="true" box="[607,687,1979,2001]" pageId="20" pageNumber="73">
<taxonomicNameLabel id="A211371BFFC0FFACFDA27DB8FD7FF85D" box="[607,687,1979,2001]" pageId="20" pageNumber="73" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
in the type locality on 20 December 2013, about one month later from the deposition. Scale bar indicates 10 mm.
</paragraph>
</caption>
<paragraph id="8BE95672FFC1FFADFF3A7A94FD95FE0B" blockId="21.[151,1436,151,392]" pageId="21" pageNumber="74">
<emphasis id="B9228A60FFC1FFADFF3A7A94FEBEFF3C" bold="true" box="[199,366,151,176]" pageId="21" pageNumber="74">Conservation.</emphasis>
The new species is currently known only from a limited area in Fukushima and Niigata Prefectures, although potential distributional range is much wider. This species seems to be locally abundant and the habitat is generally well conserved, but some parts of the habitat is threatened by recent constructions of roads and dams. Because the known range is small, collection pressure for pets can also be a serious threat, as in 
<taxonomicName id="4C562DF1FFC1FFADFA7C7B07FEF6FECC" class="Amphibia" family="Hynobiidae" genus="Onychodactylus" kingdom="Animalia" order="Caudata" pageId="21" pageNumber="74" phylum="Chordata" rank="species" species="tsukubaensis">
<emphasis id="B9228A60FFC1FFADFA7C7B07FEF6FECC" italics="true" pageId="21" pageNumber="74">
<collectingCountry id="F34116E2FFC1FFADFA7C7B07FA48FE97" box="[1409,1432,260,283]" name="Iceland" pageId="21" pageNumber="74">O</collectingCountry>
. tsukubaensis
</emphasis>
</taxonomicName>
, which is now facing the problem of over collection (
<collectingCountry id="F34116E2FFC1FFADFC7C7B2BFC48FECC" box="[897,920,296,320]" name="Iceland" pageId="21" pageNumber="74">O</collectingCountry>
. Inaba, personal communication) shortly after recent description. We propose that the new species to be designated as Vulnerable in the Red Lists of IUCN and Ministry of the Environment of 
<collectingCountry id="F34116E2FFC1FFADFE007B73FD91FE0B" box="[509,577,368,391]" name="Japan" pageId="21" pageNumber="74">Japan</collectingCountry>
.
</paragraph>
</subSubSection>
</treatment>
</document>