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<mods:title id="C423E8E9D9F71B5CE902DEF8BCEB60A0">Description of a new subterranean breeding brown frog (Ranidae: Rana) from Japan</mods:title>
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<mods:namePart id="415081A6A80E92F169D2803D2E6F2F05">Matsui, Masafumi</mods:namePart>
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<emphasis id="B943EA8EFFC1FFDEF9A0669D5F1C2ABD" bold="true" box="[151,289,223,250]" italics="true" pageId="17" pageNumber="418">Rana kyoto</emphasis>
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<paragraph id="8B88369CFFC1FFDEF9A067415CD52B5A" blockId="17.[151,744,223,357]" box="[151,744,259,286]" pageId="17" pageNumber="418">(Suggested English name: Dwarf Tagos brown frog)</paragraph>
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<paragraph id="8B88369CFFC1FFDEF9A067655CA42B06" blockId="17.[151,744,223,357]" box="[151,665,295,322]" pageId="17" pageNumber="418">(Suggested Japanese name: Hime-tago-gaeru)</paragraph>
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(
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)
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<paragraph id="8B88369CFFC1FFDEF9A067D25F302BEF" blockId="17.[151,1437,400,546]" box="[151,269,400,424]" pageId="17" pageNumber="418">Synonymy:</paragraph>
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<taxonomicName id="4C374D1FFFC1FFDEF9A067ED5F232B81" ID-CoL="642HY" authorityName="Linnaeus" authorityYear="1758" box="[151,286,431,454]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="17" pageNumber="418" phylum="Chordata" rank="genus">
<emphasis id="B943EA8EFFC1FFDEF9A067ED5F232B81" box="[151,286,431,454]" italics="true" pageId="17" pageNumber="418">Rana t. tagoi</emphasis>
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:
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: p. 145
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;
<treatmentCitation id="0A96108DFFC1FFDEFBA267EC5D9A2B81" author="Tanaka, T. &amp; Matsui, M. &amp; Takenaka, O." box="[661,935,430,454]" page="756" pageId="17" pageNumber="418" year="1994">
<bibRefCitation id="EFA64B6DFFC1FFDEFBA267EC5D6A2B81" author="Tanaka, T. &amp; Matsui, M. &amp; Takenaka, O." box="[661,855,430,454]" pageId="17" pageNumber="418" pagination="753 - 757" refId="ref18525" refString="Tanaka, T., Matsui, M. &amp; Takenaka, O. (1994) Estimation of phylogenetic relationships among Japanese brown frogs from mitochondrial cytochrome b gene (Amphibia: Anura). Zoological Science, 11, 753 - 757." type="journal article" year="1994">
Tanaka
<emphasis id="B943EA8EFFC1FFDEFBD167ED5D292B81" box="[742,788,430,454]" italics="true" pageId="17" pageNumber="418">et al</emphasis>
. 1994
</bibRefCitation>
: p. 756
</treatmentCitation>
;
<treatmentCitation id="0A96108DFFC1FFDEFA8567ED5AA32B81" author="Eto, K. &amp; Matsui, M. &amp; Sugahara, T. &amp; Tanaka-Ueno, T." box="[946,1182,430,454]" page="663" pageId="17" pageNumber="418" year="2012">
<bibRefCitation id="EFA64B6DFFC1FFDEFA8567ED5A6D2B81" author="Eto, K. &amp; Matsui, M. &amp; Sugahara, T. &amp; Tanaka-Ueno, T." box="[946,1104,430,454]" pageId="17" pageNumber="418" pagination="662 - 671" refId="ref16740" refString="Eto, K., Matsui, M., Sugahara, T. &amp; Tanaka-Ueno, T. (2012) Highly complex mitochondrial DNA genealogy in an endemic Japanese subterranean breeding brown frog Rana tagoi (Amphibia, Anura, Ranidae). Zoological Science, 29, 662 - 671. https: // doi. org / 10.2108 / zsj. 29.662" type="journal article" year="2012">
Eto
<emphasis id="B943EA8EFFC1FFDEFAEA67ED5A362B81" box="[989,1035,430,454]" italics="true" pageId="17" pageNumber="418">et al</emphasis>
. 2012
</bibRefCitation>
: p. 663
</treatmentCitation>
;
<treatmentCitation id="0A96108DFFC1FFDEFD9D67EC5ECD2BA2" author="Sugihara, T. &amp; Matsui, M. &amp; Sugahara, T." page="233" pageId="17" pageNumber="418" year="2022">
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Sugihara
<emphasis id="B943EA8EFFC1FFDEFC3B67ED5B072B81" box="[1292,1338,430,454]" italics="true" pageId="17" pageNumber="418">et al</emphasis>
. 2022
</bibRefCitation>
: p. 233
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.
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<paragraph id="8B88369CFFC1FFDEF9A067AE5A082865" blockId="17.[151,1437,400,546]" pageId="17" pageNumber="418">
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<emphasis id="B943EA8EFFC1FFDEF9A067AE5EDD2844" box="[151,224,492,515]" italics="true" pageId="17" pageNumber="418">Rana t.</emphasis>
<treatmentCitation id="0A96108DFFC1FFDEF9D067AE5C4C2843" author="Daito, Y. &amp; Nakamizo, S." box="[231,625,492,516]" page="213" pageId="17" pageNumber="418" year="1993">
<emphasis id="B943EA8EFFC1FFDEF9D067AE5F262844" box="[231,283,492,515]" italics="true" pageId="17" pageNumber="418">tagoi</emphasis>
:
<bibRefCitation id="EFA64B6DFFC1FFDEF81F67AE5C182843" author="Daito, Y. &amp; Nakamizo, S." box="[296,549,492,516]" pageId="17" pageNumber="418" pagination="213 - 230" refId="ref16628" refString="Daito, Y. &amp; Nakamizo, S. (1993) Breeding habitat and call structure of Tago's brown frog (Rana tagoi tagoi) in Mt. Rokko. Bulletin of the Faculty of Education, Kobe University, 90, 213 - 230." type="journal article" year="1993">Daito &amp; Nakamizo 1993</bibRefCitation>
: p. 213
</treatmentCitation>
</taxonomicName>
;
<treatmentCitation id="0A96108DFFC1FFDEFB4B67AE5D4B2844" author="Daito, Y. &amp; Hiramoto, M. &amp; Yoshimoto, Y." box="[636,886,491,516]" page="523" pageId="17" pageNumber="418" year="1998">
<bibRefCitation id="EFA64B6DFFC1FFDEFB4B67AE5D142843" author="Daito, Y. &amp; Hiramoto, M. &amp; Yoshimoto, Y." box="[636,809,492,516]" pageId="17" pageNumber="418" pagination="523 - 530" refId="ref16675" refString="Daito, Y., Hiramoto, M. &amp; Yoshimoto, Y. (1998) Development of post- mating isolation in the Tago's brown frogs (Rana) II. Offspring between Rana tagoi tagoi and Oki population, Tago's brown frog of Oki Isl. Bulletin of the Faculty of Human Development, Kobe University, 5, 523 - 530." type="journal article" year="1998">
Daito
<emphasis id="B943EA8EFFC1FFDEFB8D67AE5CDA2843" box="[698,743,492,516]" italics="true" pageId="17" pageNumber="418">et al</emphasis>
. 1998
</bibRefCitation>
: p. 523
</treatmentCitation>
;
<treatmentCitation id="0A96108DFFC1FFDEFAB767AE5A992844" author="Maeda, N. &amp; Matsui, M." box="[896,1188,491,516]" page="58" pageId="17" pageNumber="418" year="1999">
<bibRefCitation id="EFA64B6DFFC1FFDEFAB767AE5A552843" author="Maeda, N. &amp; Matsui, M." box="[896,1128,492,516]" pageId="17" pageNumber="418" refId="ref17385" refString="Maeda, N. &amp; Matsui, M. (1999) Frogs and Toads of Japan, Revised Edition. Bun-ichi Sogo Shuppan, Tokyo, 223 pp." type="book" year="1999">Maeda &amp; Matsui 1999</bibRefCitation>
: p. 58
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(part);
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<bibRefCitation id="EFA64B6DFFC1FFDEFDC667AE5EC32865" author="Matsui, M. &amp; Maeda, M." pageId="17" pageNumber="418" refId="ref17513" refString="Matsui, M. &amp; Maeda, M. (2018) Encyclopedia of Japanese Frogs. Bun-ichi Sogo Shuppan, Tokyo, 271 pp." type="book" year="2018">Matsui &amp; Maeda 2018</bibRefCitation>
: p. 84
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(part);
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<bibRefCitation id="EFA64B6DFFC1FFDEF8B164495C702865" author="Eto, K. &amp; Matsui, M." box="[390,589,522,546]" pageId="17" pageNumber="418" pagination="231 - 239" refId="ref16874" refString="Eto, K. &amp; Matsui, M. (2014) Cytonuclear discordance and historical demography of two brown frogs, Rana tagoi and R. sakuraii (Amphibia: Ranidae). Molecular Phylogenetics and Evolution, 79, 231 - 239. https: // doi. org / 10.1016 / j. ympev. 2014.06.019" type="journal article" year="2014">Eto &amp; Matsui 2014</bibRefCitation>
: p. 231
</treatmentCitation>
(part).
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<bibRefCitation id="EFA64B6DFFC1FFDEFBD664495D8E2865" author="Eto, K. &amp; Matsui, M." box="[737,947,522,546]" pageId="17" pageNumber="418" pagination="62 - 63" refId="ref16990" refString="Eto, K. &amp; Matsui, M. (2021 a) Rana tagoi Okada, 1928. In: Herpetological Society of Japan (Ed.), Amphibians and Reptiles of Japan. Sunrise Publishing, Hikone, pp. 62 - 63." type="book chapter" year="2021">Eto &amp; Matsui 2021a</bibRefCitation>
: p. 62
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(part).
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<typeStatus id="548C883EFFC1FFDEF9A064135F35282C" box="[151,264,593,619]" pageId="17" pageNumber="418" type="holotype">Holotype</typeStatus>
:
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KUHE 48482, adult male from
<location id="8EE86047FFC1FFDEFBAC64125D05282C" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFBAC64125D05282C" box="[667,824,592,619]" country="Japan" county="City" latitude="35.139725" longLatPrecision="19" longitude="135.75333" municipality="Kitayama area" name="Takitani Pass" pageId="17" pageNumber="418" stateProvince="Kyoto">Takitani Pass</location>
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Kumogahata of
<collectingMunicipality id="6BECACE6FFC1FFDEFD3D64125A4A282D" box="[1034,1143,592,618]" pageId="17" pageNumber="418">Kitayama</collectingMunicipality>
</location>
area,
<collectingRegion id="49F3F87EFFC1FFDEFDF064135B30282C" box="[1223,1293,593,619]" country="Japan" name="Kyoto" pageId="17" pageNumber="418">Kyoto</collectingRegion>
<collectingCounty id="62E94E10FFC1FFDEFC2D64125B76282D" box="[1306,1355,592,618]" pageId="17" pageNumber="418">City</collectingCounty>
,
<collectingCountry id="F320760CFFC1FFDEFC6B64135BA1282C" box="[1372,1436,593,619]" name="Japan" pageId="17" pageNumber="418">Japan</collectingCountry>
(
<geoCoordinate id="EE03505BFFC1FFDEF9A864345F0D28C8" box="[159,304,629,657]" degrees="35" direction="north" minutes="08" orientation="latitude" pageId="17" pageNumber="418" precision="15" seconds="23" value="35.139725">35°0823″ N</geoCoordinate>
,
<geoCoordinate id="EE03505BFFC1FFDEF80C64345FF328D6" box="[315,462,630,657]" degrees="135" direction="east" minutes="45" orientation="longitude" pageId="17" pageNumber="418" precision="15" seconds="12" value="135.75333">135°4512″E</geoCoordinate>
,
<elevation id="001AD1AFFFC1FFDEF8EE64365CEC28C9" box="[473,721,628,655]" metricMagnitude="2" metricUnit="m" metricValue="6.87" pageId="17" pageNumber="418" unit="m" value="687.0">
<quantity id="4CCF9B79FFC1FFDEF8EE64365C1D28C8" box="[473,544,628,655]" metricMagnitude="2" metricUnit="m" metricValue="6.87" pageId="17" pageNumber="418" unit="m" value="687.0">687 m</quantity>
above sea level
</elevation>
[asl]), collected on
<date id="FF89105CFFC1FFDEFA8664365A0328C8" box="[945,1086,628,655]" pageId="17" pageNumber="418" value="2014-04-06">
<collectingDate id="EFCDE9B4FFC1FFDEFA8664365A0328C8" box="[945,1086,628,655]" pageId="17" pageNumber="418" value="2014-04-06">6 April 2014</collectingDate>
</date>
by Koshiro Eto.
</materialsCitation>
</paragraph>
<paragraph id="8B88369CFFC1FFDEF9F064DB5BA929E9" blockId="17.[151,1437,592,2023]" pageId="17" pageNumber="418">
<materialsCitation id="3B5F3CC1FFC1FFDEF9F064DB5BAD29E9" ID-GBIF-Occurrence="3967664302" collectingDate="1993-03-31" collectingDateMax="2017-03-25" collectingDateMin="1993-03-31" county="All" location="City" municipality="Kumogahata" pageId="17" pageNumber="418" specimenCount="10" specimenCount-female="3" specimenCount-male="7" stateProvince="Kyoto" typeStatus="paratype">
<emphasis id="B943EA8EFFC1FFDEF9F064DB5F7A28F4" bold="true" box="[199,327,665,691]" pageId="17" pageNumber="418">
<typeStatus id="548C883EFFC1FFDEF9F064DB5F7F28F4" box="[199,322,665,691]" pageId="17" pageNumber="418" type="paratype">Paratypes</typeStatus>
:
</emphasis>
<collectingCounty id="62E94E10FFC1FFDEF86764DA5F4E28F5" box="[336,371,664,690]" pageId="17" pageNumber="418">All</collectingCounty>
from
<collectingMunicipality id="6BECACE6FFC1FFDEF88B64DA5C7028F5" box="[444,589,664,690]" pageId="17" pageNumber="418">Kumogahata</collectingMunicipality>
,
<collectingRegion id="49F3F87EFFC1FFDEFB6E64DB5CA328F4" box="[601,670,665,691]" country="Japan" name="Kyoto" pageId="17" pageNumber="418">Kyoto</collectingRegion>
<location id="8EE86047FFC1FFDEFB9F64DA5CE428F5" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFB9F64DA5CE428F5" box="[680,729,664,690]" county="All" municipality="Kumogahata" name="City" pageId="17" pageNumber="418" stateProvince="Kyoto">City</location>
, near the type locality. KUHE S981985,
<specimenCount id="9D31FD15FFC1FFDEFD8D64DA5B0D28F5" box="[1210,1328,664,690]" pageId="17" pageNumber="418" type="male">five males</specimenCount>
collected on
<date id="FF89105CFFC1FFDEF98B64FF5F552890" box="[188,360,700,727]" pageId="17" pageNumber="418" value="1993-03-31">
<collectingDate id="EFCDE9B4FFC1FFDEF98B64FF5F552890" box="[188,360,700,727]" pageId="17" pageNumber="418" value="1993-03-31">31 March 1993</collectingDate>
</date>
by Takahiro Sugahara; KUHE 2709427096, 27098,
<specimenCount id="9D31FD15FFC1FFDEFAFD64FE5A7C2891" box="[970,1089,700,726]" pageId="17" pageNumber="418" type="male">four males</specimenCount>
collected on
<date id="FF89105CFFC1FFDEFDED64FF5B4A2890" box="[1242,1399,700,727]" pageId="17" pageNumber="418" value="2000-04-14">
<collectingDate id="EFCDE9B4FFC1FFDEFDED64FF5B4A2890" box="[1242,1399,700,727]" pageId="17" pageNumber="418" value="2000-04-14">14 April 2000</collectingDate>
</date>
by Tadafumi Sugihara; KUHE 42248,
<specimenCount id="9D31FD15FFC1FFDEFB1C64A35CAE28BD" box="[555,659,736,763]" pageId="17" pageNumber="418" type="male">one male</specimenCount>
collected on
<date id="FF89105CFFC1FFDEFA1864A35DC028BC" box="[815,1021,736,763]" pageId="17" pageNumber="418" value="2008-09-02">
<collectingDate id="EFCDE9B4FFC1FFDEFA1864A35DC028BC" box="[815,1021,736,763]" pageId="17" pageNumber="418" value="2008-09-02">2 September 2008</collectingDate>
</date>
by Koshiro Eto; KUHE 42034,
<specimenCount id="9D31FD15FFC1FFDEFC4364A35EDF2959" pageId="17" pageNumber="418" type="female">one female</specimenCount>
collected on
<date id="FF89105CFFC1FFDEF84865465C382958" box="[383,517,772,799]" pageId="17" pageNumber="418" value="2008-07-06">
<collectingDate id="EFCDE9B4FFC1FFDEF84865465C382958" box="[383,517,772,799]" pageId="17" pageNumber="418" value="2008-07-06">6 July 2008</collectingDate>
</date>
by Amir Hamidy; KUHE 42249,
<specimenCount id="9D31FD15FFC1FFDEFAB865475A362959" box="[911,1035,772,799]" pageId="17" pageNumber="418" type="female">one female</specimenCount>
collected on
<date id="FF89105CFFC1FFDEFD9F65475B4A2958" box="[1192,1399,772,799]" pageId="17" pageNumber="418" value="2008-09-02">
<collectingDate id="EFCDE9B4FFC1FFDEFD9F65475B4A2958" box="[1192,1399,772,799]" pageId="17" pageNumber="418" value="2008-09-02">2 September 2008</collectingDate>
</date>
by Koshiro Eto: KUHE 42273 and 42274,
<specimenCount id="9D31FD15FFC1FFDEFB7A656B5CE92905" box="[589,724,808,835]" pageId="17" pageNumber="418" type="female">two females</specimenCount>
collected on
<date id="FF89105CFFC1FFDEFA50656B5A032904" box="[871,1086,808,835]" pageId="17" pageNumber="418" value="2008-09-13">
<collectingDate id="EFCDE9B4FFC1FFDEFA50656B5A032904" box="[871,1086,808,835]" pageId="17" pageNumber="418" value="2008-09-13">13 September 2008</collectingDate>
</date>
by Koshiro Eto; KUHE 48481,
<specimenCount id="9D31FD15FFC1FFDEF9A0650F5EC12921" box="[151,252,844,871]" pageId="17" pageNumber="418" type="male">one male</specimenCount>
collected together with the
<typeStatus id="548C883EFFC1FFDEFB06650E5CA82921" box="[561,661,844,870]" pageId="17" pageNumber="418" type="holotype">holotype</typeStatus>
; KMNH 2020709VR-KE0218,
<specimenCount id="9D31FD15FFC1FFDEFACF650F5A632921" box="[1016,1118,844,871]" pageId="17" pageNumber="418" type="male">one male</specimenCount>
collected on
<date id="FF89105CFFC1FFDEFDC5650E5BA02920" box="[1266,1437,844,871]" pageId="17" pageNumber="418" value="2017-03-25">
<collectingDate id="EFCDE9B4FFC1FFDEFDC5650E5BA02920" box="[1266,1437,844,871]" pageId="17" pageNumber="418" value="2017-03-25">25 March 2017</collectingDate>
</date>
by Keita Niwa; KMNH 20220715-KE2140 (former KUHE 42271),
<specimenCount id="9D31FD15FFC1FFDEFAAC65335A3F29CD" box="[923,1026,880,907]" pageId="17" pageNumber="418" type="male">one male</specimenCount>
collected on
<date id="FF89105CFFC1FFDEFDAB65335B4A29CC" box="[1180,1399,880,907]" pageId="17" pageNumber="418" value="2008-09-13">
<collectingDate id="EFCDE9B4FFC1FFDEFDAB65335B4A29CC" box="[1180,1399,880,907]" pageId="17" pageNumber="418" value="2008-09-13">13 September 2008</collectingDate>
</date>
by Koshiro Eto; NSMT-H17097 (former KUHE 41266),
<specimenCount id="9D31FD15FFC1FFDEFBDB65D75D6C29E9" box="[748,849,916,943]" pageId="17" pageNumber="418" type="male">one male</specimenCount>
collected on
<date id="FF89105CFFC1FFDEFAD265D65A4D29E8" box="[997,1136,916,943]" pageId="17" pageNumber="418" value="2008-04-06">
<collectingDate id="EFCDE9B4FFC1FFDEFAD265D65A4D29E8" box="[997,1136,916,943]" pageId="17" pageNumber="418" value="2008-04-06">6 April 2008</collectingDate>
</date>
by Natsuhiko Yoshikawa
</materialsCitation>
.
</paragraph>
<paragraph id="8B88369CFFC1FFDEF9F065FA5C7D2FA8" blockId="17.[151,1437,592,2023]" pageId="17" pageNumber="418">
<emphasis id="B943EA8EFFC1FFDEF9F065FA5F802995" bold="true" box="[199,445,952,978]" pageId="17" pageNumber="418">Referred specimens:</emphasis>
<materialsCitation id="3B5F3CC1FFC1FFDEF8F165FA5C062FA8" ID-GBIF-Occurrence="3967664301" accessionNumber="H04218" country="Japan" county="All" location="Honshu Isl." municipality="Central-western" pageId="17" pageNumber="418" specimenCount="74" specimenCount-female="11" specimenCount-male="63" stateProvince="Fukui">
<collectingCounty id="62E94E10FFC1FFDEF8F165FA5FD72995" box="[454,490,952,978]" pageId="17" pageNumber="418">All</collectingCounty>
from
<collectingMunicipality id="6BECACE6FFC1FFDEFB0665FA5CD92995" box="[561,740,952,978]" pageId="17" pageNumber="418">Central-western</collectingMunicipality>
part of
<location id="8EE86047FFC1FFDEFA7665FA5DF82995" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFA7665FA5DF82995" box="[833,965,952,978]" country="Japan" county="All" municipality="Central-western" name="Honshu Isl." pageId="17" pageNumber="418" stateProvince="Fukui">Honshu Isl.</location>
,
<collectingCountry id="F320760CFFC1FFDEFAE465FB5A2E2994" box="[979,1043,953,979]" name="Japan" pageId="17" pageNumber="418">Japan</collectingCountry>
(around the
<location id="8EE86047FFC1FFDEFD9165FA5B7A2995" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFD9165FA5B7A2995" box="[1190,1351,952,978]" country="Japan" county="All" municipality="Central-western" name="Kinki District" pageId="17" pageNumber="418" stateProvince="Fukui">Kinki District</location>
).
<collectingRegion id="49F3F87EFFC1FFDEFC6C65FA5EEF29B1" country="Japan" name="Fukui" pageId="17" pageNumber="418">Fukui Pref.</collectingRegion>
: KUHE 44746,
<specimenCount id="9D31FD15FFC1FFDEF8BB659F5FCE29B1" box="[396,499,988,1015]" pageId="17" pageNumber="418" type="male">one male</specimenCount>
from
<location id="8EE86047FFC1FFDEFB0A659E5CA129B1" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFB0A659E5CA129B1" box="[573,668,988,1014]" country="Japan" county="All" municipality="Central-western" name="Natasho" pageId="17" pageNumber="418" stateProvince="Fukui">Natasho</location>
,
<location id="8EE86047FFC1FFDEFB9E659E5D3029B0" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFB9E659E5D3029B0" box="[681,781,988,1015]" country="Japan" county="All" municipality="Central-western" name="Oi Town" pageId="17" pageNumber="418" stateProvince="Fukui">Oi Town</location>
by
<location id="8EE86047FFC1FFDEFA0B659E5A1E29B1" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFA0B659E5A1E29B1" box="[828,1059,988,1014]" country="Japan" county="All" municipality="Central-western" name="Kazunori Kawauchi" pageId="17" pageNumber="418" stateProvince="Fukui">Kazunori Kawauchi</location>
; KUHE 45952,
<specimenCount id="9D31FD15FFC1FFDEFDE9659F5B6129B1" box="[1246,1372,988,1015]" pageId="17" pageNumber="418" type="female">one female</specimenCount>
from
<location id="8EE86047FFC1FFDEF9A062425EDE2E5D" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF9A062425EDE2E5D" box="[151,227,1024,1050]" country="Japan" county="All" municipality="Central-western" name="Shinjo" pageId="17" pageNumber="418" stateProvince="Fukui">Shinjo</location>
,
<location id="8EE86047FFC1FFDEF9D862425FAE2E5C" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF9D862425FAE2E5C" box="[239,403,1024,1051]" country="Japan" county="All" municipality="Central-western" name="Mihama Town" pageId="17" pageNumber="418" stateProvince="Fukui">Mihama Town</location>
by
<location id="8EE86047FFC1FFDEF8F762425C982E5D" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF8F762425C982E5D" box="[448,677,1024,1050]" country="Japan" county="All" municipality="Central-western" name="Kazunori Kawauchi" pageId="17" pageNumber="418" stateProvince="Fukui">Kazunori Kawauchi</location>
;
<collectingRegion id="49F3F87EFFC1FFDEFB8662425D0E2E5D" box="[689,819,1024,1050]" country="Japan" name="Shiga" pageId="17" pageNumber="418">Shiga Pref.</collectingRegion>
: KUHE T3392T3394 and T3396T3401,
<specimenCount id="9D31FD15FFC1FFDEFC2C62425BA12E5D" box="[1307,1436,1024,1050]" pageId="17" pageNumber="418" type="male">eight males</specimenCount>
and
<specimenCount id="9D31FD15FFC1FFDEF9F162675F7D2E79" box="[198,320,1060,1087]" pageId="17" pageNumber="418" type="female">one female</specimenCount>
from
<location id="8EE86047FFC1FFDEF8B562675C232E79" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF8B562675C232E79" box="[386,542,1060,1087]" country="Japan" county="All" municipality="Central-western" name="Mt. Hakodate" pageId="17" pageNumber="418" stateProvince="Fukui">Mt. Hakodate</location>
,
<location id="8EE86047FFC1FFDEFB1F62665CEB2E79" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFB1F62665CEB2E79" box="[552,726,1060,1086]" country="Japan" county="All" municipality="Central-western" name="Takashima City" pageId="17" pageNumber="418" stateProvince="Fukui">Takashima City</location>
by
<location id="8EE86047FFC1FFDEFBC962665D9E2E79" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFBC962665D9E2E79" box="[766,931,1060,1086]" country="Japan" county="All" municipality="Central-western" name="Shingo Tanabe" pageId="17" pageNumber="418" stateProvince="Fukui">Shingo Tanabe</location>
and
<location id="8EE86047FFC1FFDEFAEF62675AA02E79" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFAEF62675AA02E79" box="[984,1181,1060,1087]" country="Japan" county="All" municipality="Central-western" name="Kanto Nishikawa" pageId="17" pageNumber="418" stateProvince="Fukui">Kanto Nishikawa</location>
; KUHE 4141941422,
<specimenCount id="9D31FD15FFC1FFDEF9A0620A5F332E25" box="[151,270,1096,1122]" pageId="17" pageNumber="418" type="male">four males</specimenCount>
from
<location id="8EE86047FFC1FFDEF865620B5FFE2E25" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF865620B5FFE2E25" box="[338,451,1096,1123]" country="Japan" county="All" municipality="Central-western" name="Mt. Horai" pageId="17" pageNumber="418" stateProvince="Fukui">Mt. Horai</location>
,
<location id="8EE86047FFC1FFDEF8F9620B5C002E25" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF8F9620B5C002E25" box="[462,573,1096,1123]" country="Japan" county="All" municipality="Central-western" name="Otsu City" pageId="17" pageNumber="418" stateProvince="Fukui">Otsu City</location>
; KUHE 44155,
<specimenCount id="9D31FD15FFC1FFDEFBC7620B5D682E25" box="[752,853,1096,1123]" pageId="17" pageNumber="418" type="male">one male</specimenCount>
from
<location id="8EE86047FFC1FFDEFAAE620A5DC82E25" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFAAE620A5DC82E25" box="[921,1013,1096,1122]" country="Japan" county="All" municipality="Central-western" name="Kutsuki" pageId="17" pageNumber="418" stateProvince="Fukui">Kutsuki</location>
,
<location id="8EE86047FFC1FFDEFAC8620A5A8E2E25" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFAC8620A5A8E2E25" box="[1023,1203,1096,1122]" country="Japan" county="All" municipality="Central-western" name="Takashima City" pageId="17" pageNumber="418" stateProvince="Fukui">Takashima City</location>
;
<collectingRegion id="49F3F87EFFC1FFDEFD89620B5B782E25" box="[1214,1349,1096,1123]" country="Japan" name="Kyoto" pageId="17" pageNumber="418">Kyoto Pref.</collectingRegion>
: KUHE S967, S969, S971980, S986-990, S1005, S1009, S1011, and S1012,
<specimenCount id="9D31FD15FFC1FFDEFA92622F5A342EC1" box="[933,1033,1132,1159]" pageId="17" pageNumber="418" type="male">19 males</specimenCount>
and
<specimenCount id="9D31FD15FFC1FFDEFD77622F5AFA2EC1" box="[1088,1223,1132,1159]" pageId="17" pageNumber="418" type="female">two females</specimenCount>
from near the
<typeStatus id="548C883EFFC1FFDEFC5B622F5BA12EC0" box="[1388,1436,1133,1159]" pageId="17" pageNumber="418">type</typeStatus>
locality; KUHE 27099, 27100, 2864628653, 2865728670, 29274, 29277, 29336, 29343, 29344, 40143, 40144, 4022740235, 41265, 4282642828,
<specimenCount id="9D31FD15FFC1FFDEFB0E62F75CA12E89" box="[569,668,1204,1231]" pageId="17" pageNumber="418" type="male">34 males</specimenCount>
and
<specimenCount id="9D31FD15FFC1FFDEFBE662F75D532E89" box="[721,878,1204,1231]" pageId="17" pageNumber="418" type="female">seven females</specimenCount>
from near the
<typeStatus id="548C883EFFC1FFDEFD3862F75A022E88" box="[1039,1087,1205,1231]" pageId="17" pageNumber="418">type</typeStatus>
locality; KUHE 3597435988,
<specimenCount id="9D31FD15FFC1FFDEF9A0629A5F3D2EB5" box="[151,256,1240,1266]" pageId="17" pageNumber="418" type="male">six males</specimenCount>
and
<specimenCount id="9D31FD15FFC1FFDEF800629B5F822EB5" box="[311,447,1240,1267]" pageId="17" pageNumber="418" type="female">two females</specimenCount>
from
<location id="8EE86047FFC1FFDEFB34629A5C6B2EB5" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFB34629A5C6B2EB5" box="[515,598,1240,1266]" country="Japan" county="All" municipality="Central-western" name="Yasaka" pageId="17" pageNumber="418" stateProvince="Fukui">Yasaka</location>
,
<location id="8EE86047FFC1FFDEFB56629B5D3A2EB5" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFB56629B5D3A2EB5" box="[609,775,1240,1267]" country="Japan" county="All" municipality="Central-western" name="Kyotango City" pageId="17" pageNumber="418" stateProvince="Fukui">Kyotango City</location>
by
<location id="8EE86047FFC1FFDEFA07629A5DC12EB5" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFA07629A5DC12EB5" box="[816,1020,1240,1266]" country="Japan" county="All" municipality="Central-western" name="Atsushi Tominaga" pageId="17" pageNumber="418" stateProvince="Fukui">Atsushi Tominaga</location>
and
<location id="8EE86047FFC1FFDEFD04629A5AC12EB5" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFD04629A5AC12EB5" box="[1075,1276,1240,1266]" country="Japan" county="All" municipality="Central-western" name="Masafumi Matsui" pageId="17" pageNumber="418" stateProvince="Fukui">Masafumi Matsui</location>
; KUHE 4141,
<specimenCount id="9D31FD15FFC1FFDEF9A062BF5EC22F51" box="[151,255,1276,1303]" pageId="17" pageNumber="418" type="male">one male</specimenCount>
from
<location id="8EE86047FFC1FFDEF87062BE5FB82F51" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF87062BE5FB82F51" box="[327,389,1276,1302]" country="Japan" county="All" municipality="Central-western" name="Ashu" pageId="17" pageNumber="418" stateProvince="Fukui">Ashu</location>
,
<location id="8EE86047FFC1FFDEF8A462BF5C1C2F51" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF8A462BF5C1C2F51" box="[403,545,1276,1303]" country="Japan" county="All" municipality="Central-western" name="Nantan City" pageId="17" pageNumber="418" stateProvince="Fukui">Nantan City</location>
; KUHE 44604,
<specimenCount id="9D31FD15FFC1FFDEFBEA62BF5D672F51" box="[733,858,1276,1303]" pageId="17" pageNumber="418" type="female">one female</specimenCount>
from
<location id="8EE86047FFC1FFDEFA9462BE5DDC2F51" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFA9462BE5DDC2F51" box="[931,993,1276,1302]" country="Japan" county="All" municipality="Central-western" name="Ashu" pageId="17" pageNumber="418" stateProvince="Fukui">Ashu</location>
,
<location id="8EE86047FFC1FFDEFAD962BF5A452F51" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFAD962BF5A452F51" box="[1006,1144,1276,1303]" country="Japan" county="All" municipality="Central-western" name="Nantan City" pageId="17" pageNumber="418" stateProvince="Fukui">Nantan City</location>
by
<location id="8EE86047FFC1FFDEFD9F62BE5B7F2F50" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFD9F62BE5B7F2F50" box="[1192,1346,1276,1303]" country="Japan" county="All" municipality="Central-western" name="Makoto Kato" pageId="17" pageNumber="418" stateProvince="Fukui">Makoto Kato</location>
;
<collectingRegion id="49F3F87EFFC1FFDEFC6762BF5EEF2F7D" country="Japan" name="Hyogo" pageId="17" pageNumber="418">Hyogo Pref.</collectingRegion>
: KUHE T3451,
<specimenCount id="9D31FD15FFC1FFDEF8BC63635C352F7D" box="[395,520,1312,1339]" pageId="17" pageNumber="418" type="female">one female</specimenCount>
from
<location id="8EE86047FFC1FFDEFB6763635C822F7D" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFB6763635C822F7D" box="[592,703,1312,1339]" country="Japan" county="All" municipality="Central-western" name="Mt. Ooka" pageId="17" pageNumber="418" stateProvince="Fukui">Mt. Ooka</location>
,
<location id="8EE86047FFC1FFDEFBFC63625D582F7D" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFBFC63625D582F7D" box="[715,869,1312,1338]" country="Japan" county="All" municipality="Central-western" name="Toyooka City" pageId="17" pageNumber="418" stateProvince="Fukui">Toyooka City</location>
by
<location id="8EE86047FFC1FFDEFAA563625A032F7D" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFAA563625A032F7D" box="[914,1086,1312,1338]" country="Japan" county="All" municipality="Central-western" name="Shingo Tanabe" pageId="17" pageNumber="418" stateProvince="Fukui">Shingo Tanabe</location>
; KUHE 22647,
<specimenCount id="9D31FD15FFC1FFDEFDC163635B602F7D" box="[1270,1373,1312,1339]" pageId="17" pageNumber="418" type="male">one male</specimenCount>
from
<location id="8EE86047FFC1FFDEF9A063075F2B2F19" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF9A063075F2B2F19" box="[151,278,1348,1375]" country="Japan" county="All" municipality="Central-western" name="Mt. Rokko" pageId="17" pageNumber="418" stateProvince="Fukui">Mt. Rokko</location>
,
<location id="8EE86047FFC1FFDEF81463065FA52F19" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF81463065FA52F19" box="[291,408,1348,1374]" country="Japan" county="All" municipality="Central-western" name="Kobe City" pageId="17" pageNumber="418" stateProvince="Fukui">Kobe City</location>
by
<location id="8EE86047FFC1FFDEF8F263065CA32F19" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF8F263065CA32F19" box="[453,670,1348,1374]" country="Japan" county="All" municipality="Central-western" name="Yasuchika Misawa" pageId="17" pageNumber="418" stateProvince="Fukui">Yasuchika Misawa</location>
; KUHE 4539045394,
<specimenCount id="9D31FD15FFC1FFDEFA9B63065A182F19" box="[940,1061,1348,1374]" pageId="17" pageNumber="418" type="male">four males</specimenCount>
and
<specimenCount id="9D31FD15FFC1FFDEFD5763075AE02F19" box="[1120,1245,1348,1375]" pageId="17" pageNumber="418" type="female">one female</specimenCount>
from
<location id="8EE86047FFC1FFDEFC1163075BA52F19" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFC1163075BA52F19" box="[1318,1432,1348,1375]" country="Japan" county="All" municipality="Central-western" name="Mt. Tanjo" pageId="17" pageNumber="418" stateProvince="Fukui">Mt. Tanjo</location>
,
<location id="8EE86047FFC1FFDEF9A0632A5F372FC5" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF9A0632A5F372FC5" box="[151,266,1384,1410]" country="Japan" county="All" municipality="Central-western" name="Kobe City" pageId="17" pageNumber="418" stateProvince="Fukui">Kobe City</location>
by
<location id="8EE86047FFC1FFDEF804632A5C172FC5" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF804632A5C172FC5" box="[307,554,1384,1410]" country="Japan" county="All" municipality="Central-western" name="Natsuhiko Yoshikawa" pageId="17" pageNumber="418" stateProvince="Fukui">Natsuhiko Yoshikawa</location>
; KUHE 45852,
<specimenCount id="9D31FD15FFC1FFDEFBEC632B5D6B2FC5" box="[731,854,1384,1411]" pageId="17" pageNumber="418" type="female">one female</specimenCount>
from
<location id="8EE86047FFC1FFDEFAAE632A5A5D2FC5" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFAAE632A5A5D2FC5" box="[921,1120,1384,1410]" country="Japan" county="All" municipality="Central-western" name="Nishinomiya City" pageId="17" pageNumber="418" stateProvince="Fukui">Nishinomiya City</location>
by
<location id="8EE86047FFC1FFDEFDBF632A5B792FC5" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFDBF632A5B792FC5" box="[1160,1348,1384,1410]" country="Japan" county="All" municipality="Central-western" name="Takashi Matsuki" pageId="17" pageNumber="418" stateProvince="Fukui">Takashi Matsuki</location>
; KUHE 4935049352,
<specimenCount id="9D31FD15FFC1FFDEF87763CF5F8E2FE1" box="[320,435,1420,1447]" pageId="17" pageNumber="418" type="male">two males</specimenCount>
and
<specimenCount id="9D31FD15FFC1FFDEF8DB63CF5C552FE1" box="[492,616,1420,1447]" pageId="17" pageNumber="418" type="female">one female</specimenCount>
from
<location id="8EE86047FFC1FFDEFB9963CF5D1C2FE0" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFB9963CF5D1C2FE0" box="[686,801,1421,1447]" country="Japan" county="All" municipality="Central-western" name="Mt. Maya" pageId="17" pageNumber="418" stateProvince="Fukui">Mt. Maya</location>
,
<location id="8EE86047FFC1FFDEFA1B63CE5D982FE1" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFA1B63CE5D982FE1" box="[812,933,1420,1446]" country="Japan" county="All" municipality="Central-western" name="Kobe City" pageId="17" pageNumber="418" stateProvince="Fukui">Kobe City</location>
;
<collectingRegion id="49F3F87EFFC1FFDEFA8763CE5A022FE1" box="[944,1087,1420,1446]" country="Japan" name="Tottori" pageId="17" pageNumber="418">Tottori Pref.</collectingRegion>
: KUHE 34742 and 34743,
<specimenCount id="9D31FD15FFC1FFDEFC4563CF5EE52F8D" pageId="17" pageNumber="418" type="male">two males</specimenCount>
from
<location id="8EE86047FFC1FFDEF82C63F35F802F8C" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF82C63F35F802F8C" box="[283,445,1457,1483]" country="Japan" county="All" municipality="Central-western" name="Mt. Hyonosen" pageId="17" pageNumber="418" stateProvince="Fukui">Mt. Hyonosen</location>
,
<location id="8EE86047FFC1FFDEF8FF63F25C5E2F8C" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEF8FF63F25C5E2F8C" box="[456,611,1456,1483]" country="Japan" county="All" municipality="Central-western" name="Wakasa Town" pageId="17" pageNumber="418" stateProvince="Fukui">Wakasa Town</location>
by
<location id="8EE86047FFC1FFDEFBBC63F25D152F8D" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFBBC63F25D152F8D" box="[651,808,1456,1482]" country="Japan" county="All" municipality="Central-western" name="Sumio Okada" pageId="17" pageNumber="418" stateProvince="Fukui">Sumio Okada</location>
; NSMT H04218,
<specimenCount id="9D31FD15FFC1FFDEFADA63F35A5A2F8D" box="[1005,1127,1456,1483]" pageId="17" pageNumber="418" type="female">one female</specimenCount>
from
<location id="8EE86047FFC1FFDEFD9D63F35B7B2F8D" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC1FFDEFD9D63F35B7B2F8D" box="[1194,1350,1456,1483]" country="Japan" county="All" municipality="Central-western" name="Mt. Oginosen" pageId="17" pageNumber="418" stateProvince="Fukui">Mt. Oginosen</location>
,
<collectingRegion id="49F3F87EFFC1FFDEFC6763F25EF02FA9" country="Japan" name="Tottori" pageId="17" pageNumber="418">Tottori Pref.</collectingRegion>
/
<collectingRegion id="49F3F87EFFC1FFDEF9E263965F632FA9" box="[213,350,1492,1518]" country="Japan" name="Hyogo" pageId="17" pageNumber="418">Hyogo Pref.</collectingRegion>
by Shun-ichi Uéno
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C32D6517FFC1FFDEF9F063BB5FF52C1D" pageId="17" pageNumber="418" type="etymology">
<paragraph id="8B88369CFFC1FFDEF9F063BB5FF52C1D" blockId="17.[151,1437,592,2023]" pageId="17" pageNumber="418">
<emphasis id="B943EA8EFFC1FFDEF9F063BB5F6F2C54" bold="true" box="[199,338,1529,1555]" pageId="17" pageNumber="418">Etymology:</emphasis>
The specific epithet is derived from a Japanese city
<collectingRegion id="49F3F87EFFC1FFDEFAE763BB5A2B2C54" box="[976,1046,1529,1555]" country="Japan" name="Kyoto" pageId="17" pageNumber="418">Kyoto</collectingRegion>
(proper noun,
<taxonomicName id="4C374D1FFFC1FFDEFDFB63BA5B342C55" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[1228,1289,1528,1554]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="17" pageNumber="418" phylum="Chordata" rank="species" species="kyoto">
<emphasis id="B943EA8EFFC1FFDEFDFB63BA5B342C55" box="[1228,1289,1528,1554]" italics="true" pageId="17" pageNumber="418">kyôto</emphasis>
</taxonomicName>
), an ancient municipality in this country, where this species was first reported by T. Sugahara (
<bibRefCitation id="EFA64B6DFFC1FFDEFD08605E5AD02C70" author="Sugahara, T." box="[1087,1261,1564,1591]" pageId="17" pageNumber="418" pagination="145" refId="ref18214" refString="Sugahara, T. (1990) Reproductive ecology of Rana tagoi tagoi at Kitayama in Kyoto. Japanese Journal of Herpetology, 13, 145." type="journal article" year="1990">Sugahara 1990</bibRefCitation>
). The epithet is used as noun in apposition.
</paragraph>
</subSubSection>
<subSubSection id="C32D6517FFC1FFDEF9F060265C3A2DA1" pageId="17" pageNumber="418" type="diagnosis">
<paragraph id="8B88369CFFC1FFDEF9F060265C3A2DA1" blockId="17.[151,1437,592,2023]" pageId="17" pageNumber="418">
<emphasis id="B943EA8EFFC1FFDEF9F060265F792C39" bold="true" box="[199,324,1636,1662]" pageId="17" pageNumber="418">Diagnosis:</emphasis>
A small-sized form of the genus (adult males
<quantity id="4CCF9B79FFC1FFDEFA6660275DCA2C38" box="[849,1015,1637,1663]" metricMagnitude="-2" metricUnit="m" metricValue="3.635" metricValueMax="4.39" metricValueMin="2.88" pageId="17" pageNumber="418" unit="mm" value="36.35" valueMax="43.9" valueMin="28.8">28.843.9 mm</quantity>
, females
<quantity id="4CCF9B79FFC1FFDEFD5460275B3A2C38" box="[1123,1287,1637,1663]" metricMagnitude="-2" metricUnit="m" metricValue="3.7849999999999993" metricValueMax="4.83" metricValueMin="2.7399999999999998" pageId="17" pageNumber="418" unit="mm" value="37.849999999999994" valueMax="48.3" valueMin="27.4">27.448.3 mm</quantity>
in SVL after fixation); reddish-, dark-, or pale-brown dorsally; a dark mask covering tympanum; a dark shoulder chevron of inverted V shape present but indistinct; dorsal and lateral skin nearly smooth with minute granules; supratympanic and dorsolateral fold present; dark mottling present in throat, chest, ventrolateral area, and ventral side of thigh and tibia; heels overlapping when limbs are held at right angles to body; tibiotarsal articulation reaching eye or beyond; tip of digit blunt, slightly dilated but not grooved; toe web developed poorly; males with internal vocal sacs. The new species is similar to the other members of the
<taxonomicName id="4C374D1FFFC1FFDEFBED617F5D6A2D11" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[730,855,1853,1878]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="17" pageNumber="418" phylum="Chordata" rank="species" species="tagoi">
<emphasis id="B943EA8EFFC1FFDEFBED617F5D6A2D11" box="[730,855,1853,1878]" italics="true" pageId="17" pageNumber="418">Rana tagoi</emphasis>
</taxonomicName>
complex, as well as
<taxonomicName id="4C374D1FFFC1FFDEFD72617F5A8E2D11" authorityName="Boulenger" authorityYear="1909" box="[1093,1203,1853,1878]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="17" pageNumber="418" phylum="Chordata" rank="species" species="sauteri">
<emphasis id="B943EA8EFFC1FFDEFD72617F5A8E2D11" box="[1093,1203,1853,1878]" italics="true" pageId="17" pageNumber="418">R. sauteri</emphasis>
</taxonomicName>
and the members of the
<taxonomicName id="4C374D1FFFC1FFDEF9F561235F4F2D3D" authority="Smith" authorityName="Smith" authorityYear="1921" box="[194,370,1888,1914]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="17" pageNumber="418" phylum="Chordata" rank="species" species="johnsi">
<emphasis id="B943EA8EFFC1FFDEF9F561235F1B2D3D" box="[194,294,1888,1914]" italics="true" pageId="17" pageNumber="418">R. johnsi</emphasis>
Smith
</taxonomicName>
group (
<bibRefCitation id="EFA64B6DFFC1FFDEF8FC61235CBE2D3C" author="Wan, H. &amp; Lyu, Z. T. &amp; Qi, S. &amp; Zhao, J. &amp; Li, P. P. &amp; Wang, Y. Y." box="[459,643,1888,1915]" pageId="17" pageNumber="418" pagination="141 - 158" refId="ref18618" refString="Wan, H., Lyu, Z. T., Qi, S., Zhao, J., Li, P. P. &amp; Wang, Y. Y. (2020) A new species of the Rana japonica group (Anura, Ranidae, Rana) from China, with a taxonomic proposal for the R. johnsi group. ZooKeys, 2020 (942), 141 - 158. https: // doi. org / 10.3897 / zookeys. 942.46928" type="journal article" year="2020">
Wan
<emphasis id="B943EA8EFFC1FFDEFB3F61235C072D3D" box="[520,570,1888,1914]" italics="true" pageId="17" pageNumber="418">et al</emphasis>
. 2020
</bibRefCitation>
) but can be distinguished from them by the combination of following characters: small adult body size; less developed toe webs; digit characteristics; dense dark markings on ventral part of legs; small, poorly-pigmented larvae without sucker; low dominant frequency of advertisement call; and chromosome number of 2n = 26.
</paragraph>
</subSubSection>
<subSubSection id="C32D6517FFC2FFDAF9F066D45CC928EE" lastPageId="21" lastPageNumber="422" pageId="18" pageNumber="419" type="description">
<paragraph id="8B88369CFFC2FFDDF9F066D45B23287A" blockId="18.[151,1437,150,2013]" pageId="18" pageNumber="419">
<emphasis id="B943EA8EFFC2FFDDF9F066D45CC82AF7" bold="true" box="[199,757,150,177]" pageId="18" pageNumber="419">
Description of
<typeStatus id="548C883EFFC2FFDDF84466D55FEA2AF6" box="[371,471,151,177]" pageId="18" pageNumber="419" type="holotype">holotype</typeStatus>
(measurements in mm):
</emphasis>
A breeding male SVL 31.2; habitus moderately stout (
<figureCitation id="130C2A19FFC2FFDDFC7C66D55BB02AF6" box="[1355,1421,151,177]" captionStart="FIGURE 8" captionStartId="19.[151,250,1903,1927]" captionTargetBox="[157,1428,967,1880]" captionTargetId="figure-392@19.[151,1436,957,1880]" captionTargetPageId="19" captionText="FIGURE 8. Holotype of Rana kyoto sp. n.. Top left to right: dorsal, lateral, and ventral views of whole body; bottom left: photo in wild; and bottom right: ventral view of hand and foot. Scale bars indicate 10 mm for whole body, hand, and foot, whereas photo in wild is not scaled." figureDoi="http://doi.org/10.5281/zenodo.7333819" httpUri="https://zenodo.org/record/7333819/files/figure.png" pageId="18" pageNumber="419">Fig. 8</figureCitation>
); head as long as broad (HL 11.5, 36.9%SVL; HW 11.0, 35.4%SVL); snout rounded; eye large, eyelid length slightly greater than snout length (EL 4.9, 15.6%SVL; SL 4.1, 13.1%SVL); snout projecting beyond lower jaw, rounded in lateral profile; canthus rostralis rounded; lores slightly oblique, concave; nostril lateral, below canthus, nearer to corner of eye than to tip of snout (S-NL 2.2, 7.0%SVL; N-EL 1.6, 5.2%SVL); internarial distance (IND 3.7, 12.0%SVL) wider than interorbital distance (IOD 2.0, 6.5% SVL) and upper eyelid (UEW 3.2, 10.1% SVL) but narrower than intercanthal distance (5.3, 17.1% SVL); pupil elliptical; tympanum distinct, diameter (TD 1.1, 3.6% SVL) one-fourth that of eye, and separated from posterior corner of eye by more than half of tympanum diameter (T-EL 0.8, 2.4% SVL); vomerine teeth in oval, oblique series (right with four and left with five teeth), beginning from a line connecting centers of choanae and extending posteromedially, groups more widely separated from each other than from choanae, and anterior margin of series outlined by dark pigmentation; tongue notched posteriorly, without papilla; vocal sac bipartite, consisting of two laterally situated parts, with a pinhole-like opening near each jaw commissure.
</paragraph>
<paragraph id="8B88369CFFC2FFDDF9F064055C57288A" blockId="18.[151,1437,150,2013]" pageId="18" pageNumber="419">
Forelimb stout and moderately long (LAL 14.2, 45.6%SVL:
<figureCitation id="130C2A19FFC2FFDDFAB264055DF72826" box="[901,970,583,609]" captionStart="FIGURE 8" captionStartId="19.[151,250,1903,1927]" captionTargetBox="[157,1428,967,1880]" captionTargetId="figure-392@19.[151,1436,957,1880]" captionTargetPageId="19" captionText="FIGURE 8. Holotype of Rana kyoto sp. n.. Top left to right: dorsal, lateral, and ventral views of whole body; bottom left: photo in wild; and bottom right: ventral view of hand and foot. Scale bars indicate 10 mm for whole body, hand, and foot, whereas photo in wild is not scaled." figureDoi="http://doi.org/10.5281/zenodo.7333819" httpUri="https://zenodo.org/record/7333819/files/figure.png" pageId="18" pageNumber="419">Fig. 8</figureCitation>
); fingers unwebbed; third finger longest (4.5, 14.5%SVL); finger length formula: II&lt;I&lt;IV&lt;III; finger tips blunt, slightly dilated but not grooved; subarticular tubercles distinct; no fringes of skin along fingers; distinct nuptial pads on first finger; the pads divided into two parts, and covered with minute asperities.
</paragraph>
<paragraph id="8B88369CFFC2FFDDF9F064955A3E29E2" blockId="18.[151,1437,150,2013]" pageId="18" pageNumber="419">
Hindlimb moderately long (HLL 52.3, 167.6% SVL:
<figureCitation id="130C2A19FFC2FFDDFA2464955D6828B6" box="[787,853,727,753]" captionStart="FIGURE 6" captionStartId="11.[151,250,1885,1909]" captionTargetBox="[162,1421,379,1849]" captionTargetId="figure-99@11.[151,1436,345,1861]" captionTargetPageId="11" captionText="FIGURE 6. A comparison of measurement ratio to SVL in males. The ratio value with significant difference in Dunns multiple comparison tests are shown.Boxplots indicate range, lower quartile, median, and upper quartile.Asterisks show the combinations with significant difference in Kruskal-Wallis test." figureDoi="http://doi.org/10.5281/zenodo.7333813" httpUri="https://zenodo.org/record/7333813/files/figure.png" pageId="18" pageNumber="419">Fig. 6</figureCitation>
); tibia moderately long (TL 15.5, 49.6% SVL), heels overlapping when limbs are held at right angles to body; tibiotarsal articulation of adpressed limb reaching anterior margin of eye; foot (FL 16.5, 52.8%SVL) longer than tibia; toe length formula I&lt;II&lt;V&lt;III&lt;IV; toe tips similar to those of fingers; webbing developed poorly, formula: I 2
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II 13 III 2
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IV
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V; toes without lateral fringes; subarticular tubercles distinct; inner metatarsal tubercle oval, length (IMTL 1.6, 5.2%SVL) shorter than first toe (1TOEL 3.0, 9.5%SVL); outer metatarsal tubercle rounded and small.
</paragraph>
<paragraph id="8B88369CFFC2FFDDF9F065ED5D412E72" blockId="18.[151,1437,150,2013]" pageId="18" pageNumber="419">
Dorsal skin nearly smooth and both sides of body relatively granular (
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); chevron marking in scapular region indistinct; a distinct supratympanic fold from posterior margin of eye to axilla; a dorsolateral fold from supratympanic fold to groin; a junction of supratympanic and dorsolateral folds swelling outside; skin of both sides of body flabby; throat and chest smooth; abdomen weakly rugose.
</paragraph>
<paragraph id="8B88369CFFC2FFDDF9F0627D5B672FDA" blockId="18.[151,1437,150,2013]" pageId="18" pageNumber="419">Color in life brown dorsally with dark blotches; sides of body dark brown; dorsolateral fold brown with darker outline; dark brown blotches on head forming interrupted interorbital bar; lore with distinct black bar below canthus; labium with heavy, dusty dark pigmentations; distinct black marking below supratympanic fold, from behind eye, covering tympanum, and reaching above arm insertion; iris gold with reddish portions anteriorly and posteriorly; dorsal surface of limbs marked with alternating, dark brown crossbars, three on lower arm, three to four on thigh, four on tibia, and two to three on tarsus; throat and chest with heavy, dark pigmentation; the pigmentation relatively sparse in abdomen and almost absent on lower part; palm sparsely pigmented; ventral side of arm unpigmented in radial side but heavily pigmented in ulnar side; ventral side of thigh nearly unpigmented on proximal side but with dark blotches and dots on distal side; ventral side of tibia and tarsus with dark blotches; under side of foot dark brown. In preservative, coloration faded and became grayish but no obvious change in pattern was observed.</paragraph>
<paragraph id="8B88369CFFC2FFDDF9F063E45C832D9A" blockId="18.[151,1437,150,2013]" pageId="18" pageNumber="419">
<emphasis id="B943EA8EFFC2FFDDF9F063E45F7C2F87" bold="true" box="[199,321,1446,1472]" pageId="18" pageNumber="419">Variation:</emphasis>
A summary of morphometric data is shown in
<tableCitation id="C6B50327FFC2FFDDFA7863E55D9F2F86" box="[847,930,1447,1473]" captionStart="TABLE 1" captionStartId="6.[151,244,464,492]" captionTargetPageId="6" captionText="TABLE 1. Comparisons of SVL and the other measurements (means followed by ranges in parenthesis) of two Rana species." httpUri="http://table.plazi.org/id/DF486614FFD6FFC9F9A0679258282BAC" pageId="18" pageNumber="419" tableUuid="DF486614FFD6FFC9F9A0679258282BAC">Table 1</tableCitation>
together with those on the allied species
<emphasis id="B943EA8EFFC2FFDDFC5963E55EEC2FA3" italics="true" pageId="18" pageNumber="419">
R. t.
<taxonomicName id="4C374D1FFFC2FFDDF9A063895EEC2FA3" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[151,209,1483,1508]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="18" pageNumber="419" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
. Original measurements are available from Figshare (DOI: 10.6084/m9.figshare.21315384). Adult body size is apparently smaller in populations sympatric with
<emphasis id="B943EA8EFFC2FFDDFBE263AD5D7F2C4F" box="[725,834,1519,1544]" italics="true" pageId="18" pageNumber="419">
R. t.
<taxonomicName id="4C374D1FFFC2FFDDFA3F63AD5D7F2C4F" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[776,834,1519,1544]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="18" pageNumber="419" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
(male:
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; female:
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) than in solitary populations (36.0
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; 48.0
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). In the latter populations, females seem to be larger than males, although the tendency is not recognized in the former. When the hindlimb is bent forward along the body, tibiotarsal joint usually reaches between snout and anterior corner of eye in males, whereas it reaches between nostril and center of eye in females. Development of toe web varies from moderate (I
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2 II 1
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III 23 IV 31 V) to poor (I 2
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II 23 III
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4 IV
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2 V) in males, and from moderate (I
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II 12 III 23 IV 3
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V) to rudimentary (I 2
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II 23 III 24 IV
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2 V) in females. Generally, breeding males tend to have more developed web than males in the other season and females. Some have tubercles laying on the chevron marking in scapular region. Dorsal ground color varies from orange to dark brown. Individuals seem to change their dorsal color slightly from light to dark, and especially males in breeding season tend to become darker color than in the other seasons. Dark pigmentation on throat more or less sparse in some males, and almost absent in females. Development of ventral dark pigmentation also varies among individuals, and some have totally creamy ventrum. Most males have fine dots or blotches on ventral side of legs, but such a marking is less developed in females. Ground color of ventral side of legs varies from light yellow to pale pink.
</paragraph>
<paragraph id="8B88369CFFC3FFDCF9F066D55B622B22" blockId="19.[151,1437,151,933]" pageId="19" pageNumber="420">
<emphasis id="B943EA8EFFC3FFDCF9F066D55FB22AF6" bold="true" box="[199,399,151,177]" pageId="19" pageNumber="420">Eggs and larvae:</emphasis>
Diameter of fresh egg varies from 2.0 to
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before oviposition, and 3.1 to 3.4 at morula stage. Clutch size 3353, eggs yolky, black or dark-gray in animal hemisphere and white or cream in vegetal hemisphere. Larva at stage 33 monotone gray dorsally without obvious marking; ventrum largely unpigmented except for oral area, and yolky intestine can be seen through the skin; total length
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in preservation (5% formalin); head-body oval, depressed; eyes dorsolateral, not visible from below; tail long, body/tail ratio of 0.440.47; dental formula 1:1+1/1+1:2 (n = 3) in this stage. Frog SVL at metamorphosis 6.77.0 mm (n = 4).
</paragraph>
<paragraph id="8B88369CFFC3FFDCF9F0672D5D4E2BEA" blockId="19.[151,1437,151,933]" pageId="19" pageNumber="420">
<emphasis id="B943EA8EFFC3FFDCF9F0672D5F6D2BCE" bold="true" box="[199,336,367,393]" pageId="19" pageNumber="420">Karyotype:</emphasis>
One male (KUHE 44758) had diploid chromosome of 2n = 26, with five large and eight small pairs. One small pair had secondary constriction on the long arm.
</paragraph>
<paragraph id="8B88369CFFC3FFDCF9F067F45D2A29E2" blockId="19.[151,1437,151,933]" pageId="19" pageNumber="420">
<emphasis id="B943EA8EFFC3FFDCF9F067F45FFD2B97" bold="true" box="[199,448,438,464]" pageId="19" pageNumber="420">Call characteristics:</emphasis>
The advertisement call of the
<typeStatus id="548C883EFFC3FFDCFA7267F55D9A2B96" box="[837,935,439,465]" pageId="19" pageNumber="420" type="holotype">holotype</typeStatus>
(KUHE 48482:
<figureCitation id="130C2A19FFC3FFDCFD4D67F55AF82B96" box="[1146,1221,439,465]" captionStart="FIGURE 7" captionStartId="15.[151,250,1396,1420]" captionTargetBox="[159,1428,669,1364]" captionTargetId="figure-256@15.[151,1436,669,1373]" captionTargetPageId="15" captionText="FIGURE 7. Wave forms and sonagrams of advertisement calls with two notes. (A) call of holotype Rana kyoto sp. n. (sympatric population) recorded at a water temperature of 8.7°C and an air of 9.4°C; (B) Rana kyoto sp. n. from Kobe City, Hyogo Pref. (allopatric population), recorded at a water temperature of 13.6°C and an air of 21.6°C; (C) R. tagoi topotype recorded at an air temperature of 14.4°C; and (D) R. tagoi from Takashima City, Shiga Pref. (sympatric population), recorded at a water temperature of 11.5°C." figureDoi="http://doi.org/10.5281/zenodo.7333817" httpUri="https://zenodo.org/record/7333817/files/figure.png" pageId="19" pageNumber="420">Fig. 7</figureCitation>
) was recorded at the type locality at the water temperature of 8.7°C. The call recording was stored in Figshare (DOI: 10.6084/ m9.figshare.21315384). Of 20 calls measured, nine were single note call (type I), seven were two-notes call with longer first note (type II-B), and four were three-notes calls (type III-A (1) and III-C (3)). Mean (± SD) call duration (s) of single-, two-, and three-note calls were 0.38±0.11, 1.64±0.33, and 2.78±0.82, respectively. Single note calls contained 15.3±5.1 pulses in average. In two-notes calls, duration (s) and pulse number of the first note (0.81±0.25 and 42.6±11.9, respectively) was longer and larger than in the last one (0.32±0.05 and 13.6±3.4). Note duration (s) and pulse number, respectively, in three note calls were as follows: 0.34±0.02 and 19.3±
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the first note, 0.64±0.04 and 32.0±
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the middle note, and 0.40±0.06 and 15.7±
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the last note. A similar dominant frequency (Hz) was recorded among each note/call: 895.9±
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one-note call; 849.8 ± 19.7 and 925.0±
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the first and last notes of two-notes calls; and 847.9±23.3, 847.9±23.3, and 905.3±73.0 in the first, middle, and last notes of three-notes calls. Each note contained clear harmonics. Frequency modulation was not recognized in the most notes, but the middle note had modulation in one of the three-notes calls. For the variation of calls among individuals/localities, see RESULTS section and
<tableCitation id="C6B50327FFC3FFDCFB8B65C95D2C29E2" box="[700,785,906,933]" captionStart="TABLE 5" captionStartId="16.[151,239,471,497]" captionTargetPageId="16" captionText="TABLE 5. Bioacoustic characteristics of advertisement call of two Rana species. Asterisks indicate that the note contains continuous pulse(s)." httpUri="http://table.plazi.org/id/DF486614FFC0FFDFF9A0679558B92BB6" pageId="19" pageNumber="420" tableUuid="DF486614FFC0FFDFF9A0679558B92BB6">Table 5</tableCitation>
.
</paragraph>
<caption id="DF486614FFC3FFDCF9A0612D5F942D88" ID-DOI="http://doi.org/10.5281/zenodo.7333819" ID-Zenodo-Dep="7333819" httpUri="https://zenodo.org/record/7333819/files/figure.png" pageId="19" pageNumber="420" startId="19.[151,250,1903,1927]" targetBox="[157,1428,967,1880]" targetPageId="19">
<paragraph id="8B88369CFFC3FFDCF9A0612D5F942D88" blockId="19.[151,1437,1903,1999]" pageId="19" pageNumber="420">
<emphasis id="B943EA8EFFC3FFDCF9A0612D5F2F2DC0" bold="true" box="[151,274,1903,1927]" pageId="19" pageNumber="420">FIGURE 8.</emphasis>
Holotype of
<taxonomicName id="4C374D1FFFC3FFDCF8A061325C342DC0" authority="Eto &amp; Matsui &amp; Sugahara, 2022" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[407,521,1903,1927]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="19" pageNumber="420" phylum="Chordata" rank="species" species="kyoto" status="sp. nov.">
<emphasis id="B943EA8EFFC3FFDCF8A061325C342DC0" box="[407,521,1903,1927]" italics="true" pageId="19" pageNumber="420">Rana kyoto</emphasis>
</taxonomicName>
<emphasis id="B943EA8EFFC3FFDCFB38612D5C7A2DC0" bold="true" box="[527,583,1903,1927]" pageId="19" pageNumber="420">
<taxonomicNameLabel id="A27057F5FFC3FFDCFB38612D5C7A2DC0" box="[527,583,1903,1927]" pageId="19" pageNumber="420" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
. Top left to right: dorsal, lateral, and ventral views of whole body; bottom left: photo in wild; and bottom right: ventral view of hand and foot. Scale bars indicate 10 mm for whole body, hand, and foot, whereas photo in wild is not scaled.
</paragraph>
</caption>
<paragraph id="8B88369CFFC4FFDBF9F066D55C2D2A92" blockId="20.[151,1437,151,2013]" pageId="20" pageNumber="421">Breeding female often produces a call when it is handled. This call sounds higher pitch than mating call of the male, and could be a distress call.</paragraph>
<paragraph id="8B88369CFFC4FFDBF9F0669C5A1A2BB2" blockId="20.[151,1437,151,2013]" pageId="20" pageNumber="421">
<emphasis id="B943EA8EFFC4FFDBF9F0669C5F522ABF" bold="true" box="[199,367,222,248]" pageId="20" pageNumber="421">Comparisons:</emphasis>
Among brown frogs of the genus
<taxonomicName id="4C374D1FFFC4FFDBFBDC669D5DE82ABE" authority="Linnaeus 1758" authorityName="Linnaeus" authorityYear="1758" box="[747,981,222,249]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="genus">
<emphasis id="B943EA8EFFC4FFDBFBDC669D5D1B2ABF" box="[747,806,223,248]" italics="true" pageId="20" pageNumber="421">Rana</emphasis>
Linnaeus 1758
</taxonomicName>
,
<taxonomicName id="4C374D1FFFC4FFDBFAE8669D5A602ABE" authority="Eto &amp; Matsui &amp; Sugahara, 2022" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[991,1117,223,249]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="kyoto" status="sp. nov.">
<emphasis id="B943EA8EFFC4FFDBFAE8669D5A602ABE" box="[991,1117,223,249]" italics="true" pageId="20" pageNumber="421">Rana kyoto</emphasis>
</taxonomicName>
<emphasis id="B943EA8EFFC4FFDBFD54669D5A812ABE" bold="true" box="[1123,1212,223,249]" pageId="20" pageNumber="421">
<taxonomicNameLabel id="A27057F5FFC4FFDBFD54669D5A812ABE" box="[1123,1212,223,249]" pageId="20" pageNumber="421" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
resembles the
<taxonomicName id="4C374D1FFFC4FFDBFC56669D5EEC2B5B" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">
<emphasis id="B943EA8EFFC4FFDBFC56669D5EEC2B5B" italics="true" pageId="20" pageNumber="421">Rana tagoi</emphasis>
</taxonomicName>
complex,
<taxonomicName id="4C374D1FFFC4FFDBF87C67415F862B5B" authorityName="Boulenger" authorityYear="1909" box="[331,443,259,284]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="sauteri">
<emphasis id="B943EA8EFFC4FFDBF87C67415F862B5B" box="[331,443,259,284]" italics="true" pageId="20" pageNumber="421">R. sauteri</emphasis>
</taxonomicName>
,
<taxonomicName id="4C374D1FFFC4FFDBF8FC67415C0C2B5A" authorityName="Smith" authorityYear="1921" box="[459,561,259,285]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="johnsi">
<emphasis id="B943EA8EFFC4FFDBF8FC67415C0C2B5A" box="[459,561,259,285]" italics="true" pageId="20" pageNumber="421">R. johnsi</emphasis>
</taxonomicName>
and its close relatives. However, by its blunt digit tips without groove (tips grooved in
<taxonomicName id="4C374D1FFFC4FFDBF82067655FB92B07" authorityName="Boulenger" authorityYear="1909" box="[279,388,295,320]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="sauteri">
<emphasis id="B943EA8EFFC4FFDBF82067655FB92B07" box="[279,388,295,320]" italics="true" pageId="20" pageNumber="421">R. sauteri</emphasis>
</taxonomicName>
and
<taxonomicName id="4C374D1FFFC4FFDBF88D67655CDF2B06" authority=": Boulenger 1909" authorityName="Boulenger" authorityYear="1909" box="[442,738,295,321]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="johnsi">
<emphasis id="B943EA8EFFC4FFDBF88D67655C202B06" box="[442,541,295,321]" italics="true" pageId="20" pageNumber="421">R. johnsi</emphasis>
:
<bibRefCitation id="EFA64B6DFFC4FFDBFB1B67655CDF2B06" author="Boulenger, G. A." box="[556,738,295,321]" pageId="20" pageNumber="421" pagination="492 - 495" refId="ref16449" refString="Boulenger, G. A. (1909) Descriptions of four new frogs and a new snake discovered by Mr. H. Sauter in Formosa. Annals and Magazine of Natural History, 4 (24), 492 - 495. https: // doi. org / 10.1080 / 00222930908692704" type="journal article" year="1909">Boulenger 1909</bibRefCitation>
</taxonomicName>
;
<bibRefCitation id="EFA64B6DFFC4FFDBFBD967655D4E2B06" author="Smith, M. A." box="[750,883,295,321]" pageId="20" pageNumber="421" pagination="423 - 440" refId="ref18113" refString="Smith, M. A. (1921) New or little-known reptiles and batrachians from southern Annam (Indo-china). Proceedings of the Zoological Society of London, 91 (2), 423 - 440. https: // doi. org / 10.1111 / j. 1096 - 3642.1921. tb 03271. x" type="journal article" year="1921">Smith 1921</bibRefCitation>
;
<bibRefCitation id="EFA64B6DFFC4FFDBFA4867655A7B2B06" author="Chou, W. H. &amp; Lin, J. Y." box="[895,1094,295,321]" pageId="20" pageNumber="421" pagination="222 - 229" refId="ref16541" refString="Chou, W. H. &amp; Lin, J. Y. (1997) Description of a new species, Rana multidenticulata (Anura: Ranidae), from Taiwan. Zoological Studies, 36 (3), 222 - 229." type="journal article" year="1997">Chou &amp; Lin 1997</bibRefCitation>
) and small (TTL ca.
<quantity id="4CCF9B79FFC4FFDBFC0667655BBD2B06" box="[1329,1408,295,321]" metricMagnitude="-2" metricUnit="m" metricValue="2.0" pageId="20" pageNumber="421" unit="mm" value="20.0">20 mm</quantity>
in St. 35), endotrophic larva without sucker (abdominal sucker present in tadpole of
<taxonomicName id="4C374D1FFFC4FFDBFD1767095AB12B23" authorityName="Boulenger" authorityYear="1909" box="[1056,1164,331,356]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="sauteri">
<emphasis id="B943EA8EFFC4FFDBFD1767095AB12B23" box="[1056,1164,331,356]" italics="true" pageId="20" pageNumber="421">R. sauteri</emphasis>
</taxonomicName>
, and TTL can exceed
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in St. 2632 and can exceed
<quantity id="4CCF9B79FFC4FFDBFB27672D5C5D2BCE" box="[528,608,367,393]" metricMagnitude="-2" metricUnit="m" metricValue="4.0" pageId="20" pageNumber="421" unit="mm" value="40.0">40 mm</quantity>
in St. 3341:
<bibRefCitation id="EFA64B6DFFC4FFDBFA36672D5D962BCE" author="Lai, S. J. &amp; Kam, Y. C. &amp; Lin, Y. S." box="[769,939,367,393]" pageId="20" pageNumber="421" pagination="193 - 202" refId="ref17331" refString="Lai, S. J., Kam, Y. C. &amp; Lin, Y. S. (2003) Elevational variation in reproductive and life history traits of Sauter's frog Rana sauteri Boulenger, 1909 in Taiwan. Zoological Studies, 42 (1), 193 - 202." type="journal article" year="2003">
Lai
<emphasis id="B943EA8EFFC4FFDBFA19672D5D5A2BCE" box="[814,871,367,393]" italics="true" pageId="20" pageNumber="421">et al.</emphasis>
2003
</bibRefCitation>
), the new species can be distinguished from them and all the other congeners except for taxa belonging to the
<taxonomicName id="4C374D1FFFC4FFDBFA4F67D15DEC2BEB" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[888,977,403,428]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">
<emphasis id="B943EA8EFFC4FFDBFA4F67D15DEC2BEB" box="[888,977,403,428]" italics="true" pageId="20" pageNumber="421">R. tagoi</emphasis>
</taxonomicName>
complex (
<taxonomicName id="4C374D1FFFC4FFDBFD7E67D15A9E2BEB" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[1097,1187,403,428]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">
<emphasis id="B943EA8EFFC4FFDBFD7E67D15A9E2BEB" box="[1097,1187,403,428]" italics="true" pageId="20" pageNumber="421">R. tagoi</emphasis>
</taxonomicName>
sspp.,
<taxonomicName id="4C374D1FFFC4FFDBFDC767D15B7B2BEA" box="[1264,1350,403,429]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="neba">
<emphasis id="B943EA8EFFC4FFDBFDC767D15B7B2BEA" box="[1264,1350,403,429]" italics="true" pageId="20" pageNumber="421">R. neba</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C374D1FFFC4FFDBFCB367D15ECE2B96" authorityName="Matsui &amp; Matsui" authorityYear="1990" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="sakuraii">
<emphasis id="B943EA8EFFC4FFDBFCB367D15ECE2B96" italics="true" pageId="20" pageNumber="421">R. sakuraii</emphasis>
</taxonomicName>
). By its subterranean breeding habit, the new species is also distinguishable from all the other congeners except for the
<taxonomicName id="4C374D1FFFC4FFDBF80F67995FAC2BB3" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[312,401,475,500]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">
<emphasis id="B943EA8EFFC4FFDBF80F67995FAC2BB3" box="[312,401,475,500]" italics="true" pageId="20" pageNumber="421">R. tagoi</emphasis>
</taxonomicName>
complex and
<taxonomicName id="4C374D1FFFC4FFDBFB1867995D242BB2" authority="Boulenger" authorityName="Boulenger" authorityYear="1879" box="[559,793,475,501]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="iberica">
<emphasis id="B943EA8EFFC4FFDBFB1867995CA02BB2" box="[559,669,475,501]" italics="true" pageId="20" pageNumber="421">R. iberica</emphasis>
Boulenger
</taxonomicName>
(
<bibRefCitation id="EFA64B6DFFC4FFDBFA1067995A222BB2" author="Rosa, G. M. &amp; Penado, A." box="[807,1055,475,501]" pageId="20" pageNumber="421" pagination="15 - 29" refId="ref17754" refString="Rosa, G. M. &amp; Penado, A. (2013) Rana iberica (Boulenger, 1879) goes underground: subterranean habitat usage and new insights on natural history. Subterranean Biology, 11, 15 - 29. https: // doi. org / 10.3897 / subtbiol. 11.5170" type="journal article" year="2013">Rosa &amp; Penado, 2013</bibRefCitation>
)
</paragraph>
<paragraph id="8B88369CFFC4FFDBF9F067BD5B3B298E" blockId="20.[151,1437,151,2013]" pageId="20" pageNumber="421">
<taxonomicName id="4C374D1FFFC4FFDBF9F067BD5F7B285E" authority="Eto &amp; Matsui &amp; Sugahara, 2022" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[199,326,511,537]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="kyoto" status="sp. nov.">
<emphasis id="B943EA8EFFC4FFDBF9F067BD5F7B285E" box="[199,326,511,537]" italics="true" pageId="20" pageNumber="421">Rana kyoto</emphasis>
</taxonomicName>
<emphasis id="B943EA8EFFC4FFDBF87B67BD5F98285E" bold="true" box="[332,421,511,537]" pageId="20" pageNumber="421">
<taxonomicNameLabel id="A27057F5FFC4FFDBF87B67BD5F98285E" box="[332,421,511,537]" pageId="20" pageNumber="421" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is distinct by its small adult SVL (males ca.
<quantity id="4CCF9B79FFC4FFDBFAA067BD5A32285E" box="[919,1039,511,537]" metricMagnitude="-2" metricUnit="m" metricValue="3.25" metricValueMax="3.6" metricValueMin="2.9" pageId="20" pageNumber="421" unit="mm" value="32.5" valueMax="36.0" valueMin="29.0">2936 mm</quantity>
in sympatric area with
<taxonomicName id="4C374D1FFFC4FFDBFC2267BD5B53285F" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[1301,1390,511,536]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">
<emphasis id="B943EA8EFFC4FFDBFC2267BD5B53285F" box="[1301,1390,511,536]" italics="true" pageId="20" pageNumber="421">R. tagoi</emphasis>
</taxonomicName>
and
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in solitary area) compared to all the other members of the
<taxonomicName id="4C374D1FFFC4FFDBFAA064615DD2287B" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[919,1007,547,572]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">
<emphasis id="B943EA8EFFC4FFDBFAA064615DD2287B" box="[919,1007,547,572]" italics="true" pageId="20" pageNumber="421">R. tagoi</emphasis>
</taxonomicName>
complex, i.e.,
<emphasis id="B943EA8EFFC4FFDBFDA564615B3D287B" box="[1170,1280,547,572]" italics="true" pageId="20" pageNumber="421">
R. t.
<taxonomicName id="4C374D1FFFC4FFDBFDF164615B3D287B" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[1222,1280,547,572]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
(males
<quantity id="4CCF9B79FFC4FFDBFC6164605EFA2826" metricMagnitude="-2" metricUnit="m" metricValue="4.8" metricValueMax="5.8" metricValueMin="3.8" pageId="20" pageNumber="421" unit="mm" value="48.0" valueMax="58.0" valueMin="38.0">3858 mm</quantity>
),
<emphasis id="B943EA8EFFC4FFDBF9EB64055F522826" box="[220,367,583,609]" italics="true" pageId="20" pageNumber="421">R. t. okiensis</emphasis>
(
<quantity id="4CCF9B79FFC4FFDBF8B764055C3C2826" box="[384,513,583,609]" metricMagnitude="-2" metricUnit="m" metricValue="4.2" metricValueMax="4.6" metricValueMin="3.8" pageId="20" pageNumber="421" unit="mm" value="42.0" valueMax="46.0" valueMin="38.0">3846 mm</quantity>
),
<emphasis id="B943EA8EFFC4FFDBFB2164055CCC2826" box="[534,753,583,609]" italics="true" pageId="20" pageNumber="421">R. t. yakushimensis</emphasis>
(
<quantity id="4CCF9B79FFC4FFDBFA3564055DBF2826" box="[770,898,583,609]" metricMagnitude="-2" metricUnit="m" metricValue="4.25" metricValueMax="4.8" metricValueMin="3.7" pageId="20" pageNumber="421" unit="mm" value="42.5" valueMax="48.0" valueMin="37.0">3748 mm</quantity>
),
<taxonomicName id="4C374D1FFFC4FFDBFAA064055A292826" authorityName="Matsui &amp; Matsui" authorityYear="1990" box="[919,1044,583,609]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="sakuraii">
<emphasis id="B943EA8EFFC4FFDBFAA064055A292826" box="[919,1044,583,609]" italics="true" pageId="20" pageNumber="421">R. sakuraii</emphasis>
</taxonomicName>
(
<quantity id="4CCF9B79FFC4FFDBFD1264045A9B2826" box="[1061,1190,582,609]" metricMagnitude="-2" metricUnit="m" metricValue="4.7" metricValueMax="5.6" metricValueMin="3.8" pageId="20" pageNumber="421" unit="mm" value="47.0" valueMax="56.0" valueMin="38.0">3856 mm</quantity>
), and
<taxonomicName id="4C374D1FFFC4FFDBFDDB64055B7E2826" box="[1260,1347,583,609]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="neba">
<emphasis id="B943EA8EFFC4FFDBFDDB64055B7E2826" box="[1260,1347,583,609]" italics="true" pageId="20" pageNumber="421">R. neba</emphasis>
</taxonomicName>
(
<quantity id="4CCF9B79FFC4FFDBFC6264055EF928C2" metricMagnitude="-2" metricUnit="m" metricValue="4.25" metricValueMax="4.8" metricValueMin="3.7" pageId="20" pageNumber="421" unit="mm" value="42.5" valueMax="48.0" valueMin="37.0">3748 mm</quantity>
) (
<bibRefCitation id="EFA64B6DFFC4FFDBF9EC64295FE128C2" author="Matsui, M. &amp; Maeda, M." box="[219,476,619,645]" pageId="20" pageNumber="421" refId="ref17513" refString="Matsui, M. &amp; Maeda, M. (2018) Encyclopedia of Japanese Frogs. Bun-ichi Sogo Shuppan, Tokyo, 271 pp." type="book" year="2018">Matsui &amp; Maeda 2018</bibRefCitation>
; present study). In the populations sympatric with
<emphasis id="B943EA8EFFC4FFDBFD2564295ABC28C3" box="[1042,1153,619,644]" italics="true" pageId="20" pageNumber="421">
R. t.
<taxonomicName id="4C374D1FFFC4FFDBFD7064295ABC28C3" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[1095,1153,619,644]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
, the new species tends to show relatively narrower head, larger eye, and wider internarial distance than
<emphasis id="B943EA8EFFC4FFDBFAC864CD5A5028EF" box="[1023,1133,655,680]" italics="true" pageId="20" pageNumber="421">
R. t.
<taxonomicName id="4C374D1FFFC4FFDBFD0464CD5A5028EF" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[1075,1133,655,680]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
(
<figureCitation id="130C2A19FFC4FFDBFD4B64CD5AFD28EE" box="[1148,1216,655,681]" captionStart="FIGURE 6" captionStartId="11.[151,250,1885,1909]" captionTargetBox="[162,1421,379,1849]" captionTargetId="figure-99@11.[151,1436,345,1861]" captionTargetPageId="11" captionText="FIGURE 6. A comparison of measurement ratio to SVL in males. The ratio value with significant difference in Dunns multiple comparison tests are shown.Boxplots indicate range, lower quartile, median, and upper quartile.Asterisks show the combinations with significant difference in Kruskal-Wallis test." figureDoi="http://doi.org/10.5281/zenodo.7333813" httpUri="https://zenodo.org/record/7333813/files/figure.png" pageId="20" pageNumber="421">Fig. 6</figureCitation>
). In adult toe webs,
<taxonomicName id="4C374D1FFFC4FFDBF9A064F15F28288A" authority="Eto &amp; Matsui &amp; Sugahara, 2022" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[151,277,691,717]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="kyoto" status="sp. nov.">
<emphasis id="B943EA8EFFC4FFDBF9A064F15F28288A" box="[151,277,691,717]" italics="true" pageId="20" pageNumber="421">Rana kyoto</emphasis>
</taxonomicName>
<emphasis id="B943EA8EFFC4FFDBF82C64F15F4E288A" bold="true" box="[283,371,691,717]" pageId="20" pageNumber="421">
<taxonomicNameLabel id="A27057F5FFC4FFDBF82C64F15F4E288A" box="[283,371,691,717]" pageId="20" pageNumber="421" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
shows less developed webs (in most breeding males, the margins of innermost and/or outermost webs do not reach to the 1
<superScript id="7C429BD4FFC4FFDBF8F664955FF128A2" attach="left" box="[449,460,727,741]" fontSize="6" pageId="20" pageNumber="421">st</superScript>
phalanges of 1
<superScript id="7C429BD4FFC4FFDBFB4F64955CBE28A2" attach="left" box="[632,643,727,741]" fontSize="6" pageId="20" pageNumber="421">st</superScript>
and 5
<superScript id="7C429BD4FFC4FFDBFBFF64945CE828A3" attach="left" box="[712,725,726,740]" fontSize="6" pageId="20" pageNumber="421">th</superScript>
toes) than in
<emphasis id="B943EA8EFFC4FFDBFA4764955DDD28B7" box="[880,992,727,752]" italics="true" pageId="20" pageNumber="421">
R. t.
<taxonomicName id="4C374D1FFFC4FFDBFA9164955DDD28B7" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[934,992,727,752]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
,
<emphasis id="B943EA8EFFC4FFDBFAD964955A4328B6" box="[1006,1150,727,753]" italics="true" pageId="20" pageNumber="421">R. t. okiensis</emphasis>
,
<emphasis id="B943EA8EFFC4FFDBFDBA64955B5828B6" box="[1165,1381,727,753]" italics="true" pageId="20" pageNumber="421">R. t. yakushimensis</emphasis>
, and
<taxonomicName id="4C374D1FFFC4FFDBF9A064B95F2E2952" authorityName="Matsui &amp; Matsui" authorityYear="1990" box="[151,275,763,789]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="sakuraii">
<emphasis id="B943EA8EFFC4FFDBF9A064B95F2E2952" box="[151,275,763,789]" italics="true" pageId="20" pageNumber="421">R. sakuraii</emphasis>
</taxonomicName>
(in most breeding males, these webs reach to the 1
<superScript id="7C429BD4FFC4FFDBFA6C64B95D5B294E" attach="left" box="[859,870,763,777]" fontSize="6" pageId="20" pageNumber="421">st</superScript>
phalange or tip pad of each toe). By the presence of dark blotches/dots on ventral surface of legs, the new species is also distinguishable from
<emphasis id="B943EA8EFFC4FFDBFDE4655D5B75297F" box="[1235,1352,799,824]" italics="true" pageId="20" pageNumber="421">
R. t.
<taxonomicName id="4C374D1FFFC4FFDBFC39655D5B75297F" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[1294,1352,799,824]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
and
<taxonomicName id="4C374D1FFFC4FFDBFCB3655D5EF0291A" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="neba">
<emphasis id="B943EA8EFFC4FFDBFCB3655D5EF0291A" italics="true" pageId="20" pageNumber="421">R. neba</emphasis>
</taxonomicName>
(topotypes of both species usually lacking such markings). By the division of the nuptial pad of 1
<superScript id="7C429BD4FFC4FFDBFC2265015B1D2916" attach="left" box="[1301,1312,835,849]" fontSize="6" pageId="20" pageNumber="421">st</superScript>
finger into two parts separated by smooth skin, the breeding males of
<taxonomicName id="4C374D1FFFC4FFDBFA1865255DB129C6" authority="Eto &amp; Matsui &amp; Sugahara, 2022" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[815,908,871,897]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="kyoto" status="sp. nov.">
<emphasis id="B943EA8EFFC4FFDBFA1865255DB129C6" box="[815,908,871,897]" italics="true" pageId="20" pageNumber="421">R. kyoto</emphasis>
</taxonomicName>
<emphasis id="B943EA8EFFC4FFDBFAA465255DD029C6" bold="true" box="[915,1005,871,897]" pageId="20" pageNumber="421">
<taxonomicNameLabel id="A27057F5FFC4FFDBFAA465255DD029C6" box="[915,1005,871,897]" pageId="20" pageNumber="421" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
can be distinguished from
<taxonomicName id="4C374D1FFFC4FFDBFC1665255BA129C6" authorityName="Matsui &amp; Matsui" authorityYear="1990" box="[1313,1436,871,897]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="sakuraii">
<emphasis id="B943EA8EFFC4FFDBFC1665255BA129C6" box="[1313,1436,871,897]" italics="true" pageId="20" pageNumber="421">R. sakuraii</emphasis>
</taxonomicName>
(breeding male with multimodal but continuous nuptial pad on 1
<superScript id="7C429BD4FFC4FFDBFA4265C95DBD29DE" attach="left" box="[885,896,907,921]" fontSize="6" pageId="20" pageNumber="421">st</superScript>
finger). Also, in mature males, the new species can be distinguished from
<taxonomicName id="4C374D1FFFC4FFDBF8F765ED5C06298E" authorityName="Matsui &amp; Matsui" authorityYear="1990" box="[448,571,943,969]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="sakuraii">
<emphasis id="B943EA8EFFC4FFDBF8F765ED5C06298E" box="[448,571,943,969]" italics="true" pageId="20" pageNumber="421">R. sakuraii</emphasis>
</taxonomicName>
by the presence of a vocal sac (vocal sac absent in
<taxonomicName id="4C374D1FFFC4FFDBFD4D65ED5ACB298E" authorityName="Matsui &amp; Matsui" authorityYear="1990" box="[1146,1270,943,969]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="sakuraii">
<emphasis id="B943EA8EFFC4FFDBFD4D65ED5ACB298E" box="[1146,1270,943,969]" italics="true" pageId="20" pageNumber="421">R. sakuraii</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="8B88369CFFC4FFDBF9F065915DF82F86" blockId="20.[151,1437,151,2013]" pageId="20" pageNumber="421">
Mating call of
<taxonomicName id="4C374D1FFFC4FFDBF85065915FD929AA" authority="Eto &amp; Matsui &amp; Sugahara, 2022" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[359,484,979,1005]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="kyoto" status="sp. nov.">
<emphasis id="B943EA8EFFC4FFDBF85065915FD929AA" box="[359,484,979,1005]" italics="true" pageId="20" pageNumber="421">Rana kyoto</emphasis>
</taxonomicName>
<emphasis id="B943EA8EFFC4FFDBF8DE65915C7D29AA" bold="true" box="[489,576,979,1005]" pageId="20" pageNumber="421">
<taxonomicNameLabel id="A27057F5FFC4FFDBF8DE65915C7D29AA" box="[489,576,979,1005]" pageId="20" pageNumber="421" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is characterized by low dominant frequency (ca. 0.70.9 kHz in average), lack of obvious frequency modulation in each note, and few note number (14, mode: 1) with short last note (generally NL ratio of first/last notes is 3/2 or more) (
<figureCitation id="130C2A19FFC4FFDBFBB562595CFA2E72" box="[642,711,1051,1077]" captionStart="FIGURE 7" captionStartId="15.[151,250,1396,1420]" captionTargetBox="[159,1428,669,1364]" captionTargetId="figure-256@15.[151,1436,669,1373]" captionTargetPageId="15" captionText="FIGURE 7. Wave forms and sonagrams of advertisement calls with two notes. (A) call of holotype Rana kyoto sp. n. (sympatric population) recorded at a water temperature of 8.7°C and an air of 9.4°C; (B) Rana kyoto sp. n. from Kobe City, Hyogo Pref. (allopatric population), recorded at a water temperature of 13.6°C and an air of 21.6°C; (C) R. tagoi topotype recorded at an air temperature of 14.4°C; and (D) R. tagoi from Takashima City, Shiga Pref. (sympatric population), recorded at a water temperature of 11.5°C." figureDoi="http://doi.org/10.5281/zenodo.7333817" httpUri="https://zenodo.org/record/7333817/files/figure.png" pageId="20" pageNumber="421">Fig. 7</figureCitation>
;
<tableCitation id="C6B50327FFC4FFDBFBE262595D162E72" box="[725,811,1050,1077]" captionStart="TABLE 5" captionStartId="16.[151,239,471,497]" captionTargetPageId="16" captionText="TABLE 5. Bioacoustic characteristics of advertisement call of two Rana species. Asterisks indicate that the note contains continuous pulse(s)." httpUri="http://table.plazi.org/id/DF486614FFC0FFDFF9A0679558B92BB6" pageId="20" pageNumber="421" tableUuid="DF486614FFC0FFDFF9A0679558B92BB6">Table 5</tableCitation>
). Compared to the new species, calls of topotypic
<emphasis id="B943EA8EFFC4FFDBFC5A62595EEC2E1F" italics="true" pageId="20" pageNumber="421">
R. t.
<taxonomicName id="4C374D1FFFC4FFDBF9A0627D5EEC2E1F" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[151,209,1087,1112]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
show higher dominant frequency (ca. 0.91.2 kHz in average) with modulation at least within the first note in some cases, and higher number of notes (28, mode: 3) with last note longer than the first (NL ratio of first/last notes is 2/3 or less) (
<bibRefCitation id="EFA64B6DFFC4FFDBF8BB62C55C442EE6" author="Shimada, T. &amp; Sanda, M. &amp; Eto, K." box="[396,633,1159,1185]" pageId="20" pageNumber="421" pagination="54 - 65" refId="ref18050" refString="Shimada, T., Sanda, M. &amp; Eto, K. (2021) A comparison of mating calls recorded around the type localities of Rana tagoi and R. neba (Amphibia: Anura: Ranidae). Current Herpetology, 40, 54 - 65. https: // doi. org / 10.5358 / hsj. 40.54" type="journal article" year="2021">
Shimada
<emphasis id="B943EA8EFFC4FFDBF8CD62C55C132EE6" box="[506,558,1159,1185]" italics="true" pageId="20" pageNumber="421">et al</emphasis>
. 2021
</bibRefCitation>
;
<tableCitation id="C6B50327FFC4FFDBFBB062C55CE22EE6" box="[647,735,1158,1185]" captionStart="TABLE 5" captionStartId="16.[151,239,471,497]" captionTargetPageId="16" captionText="TABLE 5. Bioacoustic characteristics of advertisement call of two Rana species. Asterisks indicate that the note contains continuous pulse(s)." httpUri="http://table.plazi.org/id/DF486614FFC0FFDFF9A0679558B92BB6" pageId="20" pageNumber="421" tableUuid="DF486614FFC0FFDFF9A0679558B92BB6">Table 5</tableCitation>
). Calls of
<emphasis id="B943EA8EFFC4FFDBFA6B62C55DED2EE7" box="[860,976,1159,1184]" italics="true" pageId="20" pageNumber="421">
R. t.
<taxonomicName id="4C374D1FFFC4FFDBFAA162C55DED2EE7" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[918,976,1159,1184]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
in Kinki District also show a dominant frequency higher than the new species (ca. 0.81.4 kHz in average) with modulation at least within the first note in many cases, notes higher in number (112, mode: 2 and 4) and last note longer than the first (
<tableCitation id="C6B50327FFC4FFDBFD80628D5B332EAE" box="[1207,1294,1230,1257]" captionStart="TABLE 5" captionStartId="16.[151,239,471,497]" captionTargetPageId="16" captionText="TABLE 5. Bioacoustic characteristics of advertisement call of two Rana species. Asterisks indicate that the note contains continuous pulse(s)." httpUri="http://table.plazi.org/id/DF486614FFC0FFDFF9A0679558B92BB6" pageId="20" pageNumber="421" tableUuid="DF486614FFC0FFDFF9A0679558B92BB6">Table 5</tableCitation>
). Calls of
<taxonomicName id="4C374D1FFFC4FFDBFCB3628D5EF02F4A" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="neba">
<emphasis id="B943EA8EFFC4FFDBFCB3628D5EF02F4A" italics="true" pageId="20" pageNumber="421">R. neba</emphasis>
</taxonomicName>
show high dominant frequency (ca. 1.11.2 kHz in average) with obvious modulation within the first note, and more note number (37) with the last note similar to the first/middle ones in length (
<bibRefCitation id="EFA64B6DFFC4FFDBFD7D63555B0C2F76" author="Shimada, T. &amp; Sanda, M. &amp; Eto, K." box="[1098,1329,1303,1329]" pageId="20" pageNumber="421" pagination="54 - 65" refId="ref18050" refString="Shimada, T., Sanda, M. &amp; Eto, K. (2021) A comparison of mating calls recorded around the type localities of Rana tagoi and R. neba (Amphibia: Anura: Ranidae). Current Herpetology, 40, 54 - 65. https: // doi. org / 10.5358 / hsj. 40.54" type="journal article" year="2021">
Shimada
<emphasis id="B943EA8EFFC4FFDBFD8263555ADB2F76" box="[1205,1254,1303,1329]" italics="true" pageId="20" pageNumber="421">et al</emphasis>
. 2021
</bibRefCitation>
). Calls of
<taxonomicName id="4C374D1FFFC4FFDBF9A063795F2F2F12" authorityName="Matsui &amp; Matsui" authorityYear="1990" box="[151,274,1339,1365]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="sakuraii">
<emphasis id="B943EA8EFFC4FFDBF9A063795F2F2F12" box="[151,274,1339,1365]" italics="true" pageId="20" pageNumber="421">R. sakuraii</emphasis>
</taxonomicName>
(dominant frequency: 0.9/3.1 kHz; note number: 4; obvious modulation absent:
<bibRefCitation id="EFA64B6DFFC4FFDBFDA663795BAD2F12" author="Matsui, M. &amp; Maeda, M." box="[1169,1424,1339,1365]" pageId="20" pageNumber="421" refId="ref17513" refString="Matsui, M. &amp; Maeda, M. (2018) Encyclopedia of Japanese Frogs. Bun-ichi Sogo Shuppan, Tokyo, 271 pp." type="book" year="2018">Matsui &amp; Maeda 2018</bibRefCitation>
),
<emphasis id="B943EA8EFFC4FFDBF9A0631D5F522F3E" box="[151,367,1375,1401]" italics="true" pageId="20" pageNumber="421">R. t. yakushimensis</emphasis>
(1.0/1.7 kHz; 14; obvious modulation present:
<bibRefCitation id="EFA64B6DFFC4FFDBFAA4631D5AA82F3E" author="Matsui, M. &amp; Maeda, M." box="[915,1173,1375,1401]" pageId="20" pageNumber="421" refId="ref17513" refString="Matsui, M. &amp; Maeda, M. (2018) Encyclopedia of Japanese Frogs. Bun-ichi Sogo Shuppan, Tokyo, 271 pp." type="book" year="2018">Matsui &amp; Maeda 2018</bibRefCitation>
), and
<emphasis id="B943EA8EFFC4FFDBFDEE631D5B552F3E" box="[1241,1384,1375,1401]" italics="true" pageId="20" pageNumber="421">R. t. okiensis</emphasis>
(0.7 kHz; 46; modulation present:
<bibRefCitation id="EFA64B6DFFC4FFDBF8C363C15CCE2FDA" author="Matsui, M. &amp; Maeda, M." box="[500,755,1411,1437]" pageId="20" pageNumber="421" refId="ref17513" refString="Matsui, M. &amp; Maeda, M. (2018) Encyclopedia of Japanese Frogs. Bun-ichi Sogo Shuppan, Tokyo, 271 pp." type="book" year="2018">Matsui &amp; Maeda 2018</bibRefCitation>
) are not fully examined, but seem to be distinguishable from the call of
<taxonomicName id="4C374D1FFFC4FFDBF83963E55F572F86" authority="Eto &amp; Matsui &amp; Sugahara, 2022" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[270,362,1447,1473]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="kyoto" status="sp. nov.">
<emphasis id="B943EA8EFFC4FFDBF83963E55F572F86" box="[270,362,1447,1473]" italics="true" pageId="20" pageNumber="421">R. kyoto</emphasis>
</taxonomicName>
<emphasis id="B943EA8EFFC4FFDBF84663E55FF82F86" bold="true" box="[369,453,1447,1473]" pageId="20" pageNumber="421">
<taxonomicNameLabel id="A27057F5FFC4FFDBF84663E55FF82F86" box="[369,453,1447,1473]" pageId="20" pageNumber="421" rank="species">sp. nov</taxonomicNameLabel>
</emphasis>
by the combination of traits described above.
</paragraph>
<paragraph id="8B88369CFFC4FFDBF9F063895C732CA6" blockId="20.[151,1437,151,2013]" pageId="20" pageNumber="421">
In
<taxonomicName id="4C374D1FFFC4FFDBF9DE63895F752FA2" authority="Eto &amp; Matsui &amp; Sugahara, 2022" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[233,328,1483,1509]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="kyoto" status="sp. nov.">
<emphasis id="B943EA8EFFC4FFDBF9DE63895F752FA2" box="[233,328,1483,1509]" italics="true" pageId="20" pageNumber="421">R. kyoto</emphasis>
</taxonomicName>
<emphasis id="B943EA8EFFC4FFDBF86563895F932FA2" bold="true" box="[338,430,1483,1509]" pageId="20" pageNumber="421">
<taxonomicNameLabel id="A27057F5FFC4FFDBF86563895F932FA2" box="[338,430,1483,1509]" pageId="20" pageNumber="421" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, the animal hemisphere of eggs at early developmental stage is usually dark brown or almost black, whereas the animal hemisphere at early stage is pale gray to dark brown in
<emphasis id="B943EA8EFFC4FFDBFD4F63AD5ADB2C4F" box="[1144,1254,1519,1544]" italics="true" pageId="20" pageNumber="421">
R. t.
<taxonomicName id="4C374D1FFFC4FFDBFD9B63AD5ADB2C4F" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[1196,1254,1519,1544]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
and
<taxonomicName id="4C374D1FFFC4FFDBFC2C63AD5BA82C4E" authorityName="Matsui &amp; Matsui" authorityYear="1990" box="[1307,1429,1519,1545]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="sakuraii">
<emphasis id="B943EA8EFFC4FFDBFC2C63AD5BA82C4E" box="[1307,1429,1519,1545]" italics="true" pageId="20" pageNumber="421">R. sakuraii</emphasis>
</taxonomicName>
. Dorsal coloration of larva is also darker in the new species than in
<emphasis id="B943EA8EFFC4FFDBFAB660515DCD2C6B" box="[897,1008,1555,1580]" italics="true" pageId="20" pageNumber="421">
R. t.
<taxonomicName id="4C374D1FFFC4FFDBFA8160515DCD2C6B" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[950,1008,1555,1580]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
from Kinki District. At least in grown larvae (St. 3335), the new species tends to have more tooth rows (generally 1:1+1/1+1:2) than
<emphasis id="B943EA8EFFC4FFDBFDDD60755B602C17" box="[1258,1373,1591,1616]" italics="true" pageId="20" pageNumber="421">
R. t.
<taxonomicName id="4C374D1FFFC4FFDBFC1460755B602C17" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[1315,1373,1591,1616]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
from Kinki District at the similar stage (1:1+1/1+1:1) and tends to have fewer rows than those of
<taxonomicName id="4C374D1FFFC4FFDBFD9F60195B1E2C32" authorityName="Matsui &amp; Matsui" authorityYear="1990" box="[1192,1315,1627,1653]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="sakuraii">
<emphasis id="B943EA8EFFC4FFDBFD9F60195B1E2C32" box="[1192,1315,1627,1653]" italics="true" pageId="20" pageNumber="421">R. sakuraii</emphasis>
</taxonomicName>
(generally 1:2+2/2+2:1 or 1:2+2/1+1:1:
<bibRefCitation id="EFA64B6DFFC4FFDBF8DC603D5D102CDE" author="Kishimoto, K. &amp; Hayashi, F." box="[491,813,1663,1689]" pageId="20" pageNumber="421" pagination="173 - 183" refId="ref17170" refString="Kishimoto, K. &amp; Hayashi, F. (2020) Complete embryonic and larval stages of Rana sakuraii (Ranidae), a species that metamorphoses without feeding. Current Herpetology, 39, 173 - 183. https: // doi. org / 10.5358 / hsj. 39.173" type="journal article" year="2020">Kishimoto &amp; Hayashi 2020</bibRefCitation>
, shown as stages 2830 by their own staging method roughly correspond to St.
<quantity id="4CCF9B79FFC4FFDBF8FE60E05C0D2CFA" box="[457,560,1698,1725]" metricMagnitude="-1" metricUnit="m" metricValue="8.636" metricValueMax="8.89" metricValueMin="8.382" pageId="20" pageNumber="421" unit="in" value="34.0" valueMax="35.0" valueMin="33.0">3335 in</quantity>
this study), although all of them show variation in this trait (
<bibRefCitation id="EFA64B6DFFC4FFDBFC3360E15F0B2CA6" author="Kishimoto, K. &amp; Hayashi, F." pageId="20" pageNumber="421" pagination="173 - 183" refId="ref17170" refString="Kishimoto, K. &amp; Hayashi, F. (2020) Complete embryonic and larval stages of Rana sakuraii (Ranidae), a species that metamorphoses without feeding. Current Herpetology, 39, 173 - 183. https: // doi. org / 10.5358 / hsj. 39.173" type="journal article" year="2020">Kishimoto &amp; Hayashi 2020</bibRefCitation>
;
<bibRefCitation id="EFA64B6DFFC4FFDBF87660855C7F2CA6" author="Matsui, M. &amp; Maeda, M." box="[321,578,1735,1761]" pageId="20" pageNumber="421" refId="ref17513" refString="Matsui, M. &amp; Maeda, M. (2018) Encyclopedia of Japanese Frogs. Bun-ichi Sogo Shuppan, Tokyo, 271 pp." type="book" year="2018">Matsui &amp; Maeda 2018</bibRefCitation>
).
</paragraph>
<paragraph id="8B88369CFFC4FFDBF9F060A95AAA2D6E" blockId="20.[151,1437,151,2013]" pageId="20" pageNumber="421">
By its diploid chromosome 2n = 26,
<taxonomicName id="4C374D1FFFC4FFDBFB5560A95C822D42" authority="Eto &amp; Matsui &amp; Sugahara, 2022" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[610,703,1771,1797]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="kyoto" status="sp. nov.">
<emphasis id="B943EA8EFFC4FFDBFB5560A95C822D42" box="[610,703,1771,1797]" italics="true" pageId="20" pageNumber="421">R. kyoto</emphasis>
</taxonomicName>
<emphasis id="B943EA8EFFC4FFDBFBF260A95D222D42" bold="true" box="[709,799,1771,1797]" pageId="20" pageNumber="421">
<taxonomicNameLabel id="A27057F5FFC4FFDBFBF260A95D222D42" box="[709,799,1771,1797]" pageId="20" pageNumber="421" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is separated from
<taxonomicName id="4C374D1FFFC4FFDBFAD960A95A7E2D42" box="[1006,1091,1771,1797]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="neba">
<emphasis id="B943EA8EFFC4FFDBFAD960A95A7E2D42" box="[1006,1091,1771,1797]" italics="true" pageId="20" pageNumber="421">R. neba</emphasis>
</taxonomicName>
(2n = 28 following
<bibRefCitation id="EFA64B6DFFC4FFDBFC1260A95ECB2D6E" author="Ryuzaki, M. &amp; Hasegawa, Y. &amp; Kuramoto, M." pageId="20" pageNumber="421" pagination="49 - 58" refId="ref17875" refString="Ryuzaki, M., Hasegawa, Y. &amp; Kuramoto, M. (2014) A new brown frog of the genus Rana from Japan (Anura: Ranidae) revealed by cytological and bioacoustic studies. Alytes, 31, 49 - 58." type="journal article" year="2014">
Ryuzaki
<emphasis id="B943EA8EFFC4FFDBFCBF60A95E902D6E" italics="true" pageId="20" pageNumber="421">et al</emphasis>
. 2014
</bibRefCitation>
, although the presence of variation in the latter is suspected:
<bibRefCitation id="EFA64B6DFFC4FFDBFA96614D5AB62D6E" author="Eto, K. &amp; Matsui, M." box="[929,1163,1807,1833]" pageId="20" pageNumber="421" pagination="65" refId="ref17037" refString="Eto, K. &amp; Matsui, M. (2021 b) Rana neba Ryuzaki, Hasegawa et Kuramoto, 2014. In: Herpetological Society of Japan (Ed.), Amphibians and Reptiles of Japan. Sunrise Publishing, Hikone, pp. 65." type="book chapter" year="2021">Eto &amp; Matsui 2021b</bibRefCitation>
).
</paragraph>
<paragraph id="8B88369CFFC4FFDBF9F061715A3C2DFE" blockId="20.[151,1437,151,2013]" pageId="20" pageNumber="421">
<emphasis id="B943EA8EFFC4FFDBF9F061715F202D0A" bold="true" box="[199,285,1843,1869]" pageId="20" pageNumber="421">Range:</emphasis>
Known from northern part of Kinki District, central-western part of Honshu Island,
<collectingCountry id="F320760CFFC4FFDBFC3361715B7A2D0A" box="[1284,1351,1843,1869]" name="Japan" pageId="20" pageNumber="421">Japan</collectingCountry>
.
<collectingRegion id="49F3F87EFFC4FFDBFC6161715F322D36" country="Japan" name="Kyoto" pageId="20" pageNumber="421">Kyoto Prefecture</collectingRegion>
:
<collectorName id="26C2534AFFC4FFDBF82E61155FFD2D36" box="[281,448,1879,1905]" pageId="20" pageNumber="421">Kyotango City</collectorName>
,
<collectingRegion id="49F3F87EFFC4FFDBF8FB61155C2F2D36" box="[460,530,1879,1905]" country="Japan" name="Kyoto" pageId="20" pageNumber="421">Kyoto</collectingRegion>
<location id="8EE86047FFC4FFDBFB2E61155C772D36" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC4FFDBFB2E61155C772D36" box="[537,586,1879,1905]" country="Japan" name="City" pageId="20" pageNumber="421" stateProvince="Kyoto">City</location>
,
<collectorName id="26C2534AFFC4FFDBFB6161155CDF2D36" box="[598,738,1879,1905]" pageId="20" pageNumber="421">Nantan City</collectorName>
;
<collectingRegion id="49F3F87EFFC4FFDBFBDA61155D932D36" box="[749,942,1879,1905]" country="Japan" name="Fukui" pageId="20" pageNumber="421">Fukui Prefecture</collectingRegion>
:
<collectingMunicipality id="6BECACE6FFC4FFDBFA8F61155A632D36" box="[952,1118,1879,1905]" pageId="20" pageNumber="421">Mihama Town</collectingMunicipality>
,
<location id="8EE86047FFC4FFDBFD5E61155AF22D36" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC4FFDBFD5E61155AF22D36" box="[1129,1231,1879,1905]" country="Japan" municipality="Mihama Town" name="Oi Town" pageId="20" pageNumber="421" stateProvince="Fukui">Oi Town</location>
;
<collectingRegion id="49F3F87EFFC4FFDBFDED61155BA42D36" box="[1242,1433,1879,1905]" country="Japan" name="Shiga" pageId="20" pageNumber="421">Shiga Prefecture</collectingRegion>
: Otsu City,
<collectorName id="26C2534AFFC4FFDBF83B61395F832DD2" box="[268,446,1915,1941]" pageId="20" pageNumber="421">Takashima City</collectorName>
;
<collectingRegion id="49F3F87EFFC4FFDBF8FF61395CAC2DD2" box="[456,657,1915,1941]" country="Japan" name="Hyogo" pageId="20" pageNumber="421">Hyogo Prefecture</collectingRegion>
:
<collectingCounty id="62E94E10FFC4FFDBFBAF61395D2B2DD2" box="[664,790,1915,1941]" pageId="20" pageNumber="421">Asago City</collectingCounty>
,
<collectingMunicipality id="6BECACE6FFC4FFDBFA1761395DAE2DD2" box="[800,915,1915,1941]" pageId="20" pageNumber="421">Kobe City</collectingMunicipality>
,
<location id="8EE86047FFC4FFDBFAAA61395A592DD2" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC4FFDBFAAA61395A592DD2" box="[925,1124,1915,1941]" country="Japan" county="Asago City" municipality="Kobe City" name="Nishinomiya City" pageId="20" pageNumber="421" stateProvince="Hyogo">Nishinomiya City</location>
,
<location id="8EE86047FFC4FFDBFD5961395B292DD2" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC4FFDBFD5961395B292DD2" box="[1134,1300,1915,1941]" country="Japan" county="Asago City" municipality="Kobe City" name="Sasayama City" pageId="20" pageNumber="421" stateProvince="Hyogo">Sasayama City</location>
,
<location id="8EE86047FFC4FFDBFC2961395EE42DFE" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC4FFDBFC2961395EE42DFE" country="Japan" county="Asago City" municipality="Kobe City" name="Shin'onsen Town" pageId="20" pageNumber="421" stateProvince="Hyogo">Shinonsen Town</location>
,
<collectorName id="26C2534AFFC4FFDBF9D261DD5F662DFE" box="[229,347,1951,1977]" pageId="20" pageNumber="421">Shiso City</collectorName>
,
<collectorName id="26C2534AFFC4FFDBF85161DD5FC22DFE" box="[358,511,1951,1977]" pageId="20" pageNumber="421">Toyooka City</collectorName>
,
<collectorName id="26C2534AFFC4FFDBFB3D61DD5C432DFE" box="[522,638,1951,1977]" pageId="20" pageNumber="421">Yabu City</collectorName>
;
<collectingRegion id="49F3F87EFFC4FFDBFBBE61DD5D692DFE" box="[649,852,1951,1977]" country="Japan" name="Tottori" pageId="20" pageNumber="421">Tottori Prefecture</collectingRegion>
:
<location id="8EE86047FFC4FFDBFA6A61DD5DC02DFE" LSID="urn:lsid:plazi:treatment:039E878AFFC1FFDAF9A0669D5CC928EE:8EE86047FFC4FFDBFA6A61DD5DC02DFE" box="[861,1021,1951,1977]" country="Japan" name="Wakasa Town" pageId="20" pageNumber="421" stateProvince="Tottori">Wakasa Town</location>
.
</paragraph>
<paragraph id="8B88369CFFC4FFDAF9F061815CC928EE" blockId="20.[151,1437,151,2013]" lastBlockId="21.[151,1437,151,681]" lastPageId="21" lastPageNumber="422" pageId="20" pageNumber="421">
<emphasis id="B943EA8EFFC4FFDBF9F061815FB72D9B" bold="true" box="[199,394,1986,2013]" pageId="20" pageNumber="421">Natural history:</emphasis>
Inhabits montane forest. Topotypic
<taxonomicName id="4C374D1FFFC4FFDBFA1261815D9B2D9A" authority="Eto &amp; Matsui &amp; Sugahara, 2022" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[805,934,1987,2013]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="20" pageNumber="421" phylum="Chordata" rank="species" species="kyoto" status="sp. nov.">
<emphasis id="B943EA8EFFC4FFDBFA1261815D9B2D9A" box="[805,934,1987,2013]" italics="true" pageId="20" pageNumber="421">Rana kyoto</emphasis>
</taxonomicName>
<emphasis id="B943EA8EFFC4FFDBFA9961815A352D9A" bold="true" box="[942,1032,1987,2013]" pageId="20" pageNumber="421">
<taxonomicNameLabel id="A27057F5FFC4FFDBFA9961815A352D9A" box="[942,1032,1987,2013]" pageId="20" pageNumber="421" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
breeds in early spring, early to late April (
<bibRefCitation id="EFA64B6DFFC5FFDAF9D466D55C372AF6" author="Sugahara, T. &amp; Matsui, M." box="[227,522,151,177]" pageId="21" pageNumber="422" pagination="68 - 69" refId="ref18425" refString="Sugahara, T. &amp; Matsui, M. (1997) Breeding seasons and sites in two types of Rana tagoi tagoi from Kumogahata, Kyoto. Japanese Journal of Herpetology, 17, 68 - 69." type="journal article" year="1997">Sugahara &amp; Matsui 1997</bibRefCitation>
), whereas it breeds from late April to late May around Kobe City,
<collectingRegion id="49F3F87EFFC5FFDAFC2666D55BA02AF6" box="[1297,1437,151,177]" country="Japan" name="Hyogo" pageId="21" pageNumber="422">Hyogo Pref.</collectingRegion>
(
<bibRefCitation id="EFA64B6DFFC5FFDAF9A866F95F872A92" author="Daito, Y. &amp; Nakamizo, S." box="[159,442,187,213]" pageId="21" pageNumber="422" pagination="213 - 230" refId="ref16628" refString="Daito, Y. &amp; Nakamizo, S. (1993) Breeding habitat and call structure of Tago's brown frog (Rana tagoi tagoi) in Mt. Rokko. Bulletin of the Faculty of Education, Kobe University, 90, 213 - 230." type="journal article" year="1993">Daito &amp; Nakamizo 1993</bibRefCitation>
; Eto, unpublished data), the only known solitary range of this species without any close relatives.
<taxonomicName id="4C374D1FFFC5FFDAF832669D5FB92ABE" authority="Eto &amp; Matsui &amp; Sugahara, 2022" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[261,388,223,249]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="21" pageNumber="422" phylum="Chordata" rank="species" species="kyoto" status="sp. nov.">
<emphasis id="B943EA8EFFC5FFDAF832669D5FB92ABE" box="[261,388,223,249]" italics="true" pageId="21" pageNumber="422">Rana kyoto</emphasis>
</taxonomicName>
<emphasis id="B943EA8EFFC5FFDAF8BB669D5FDB2ABE" bold="true" box="[396,486,223,249]" pageId="21" pageNumber="422">
<taxonomicNameLabel id="A27057F5FFC5FFDAF8BB669D5FDB2ABE" box="[396,486,223,249]" pageId="21" pageNumber="422" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is a subterranean breeding species, and the males call inside crevices of rock bed or under rocks near springs or streams, with their bodies almost invisible from the outside while calling. Two or more males often call in a same burrow. Such a burrow is partially filled by water exuded from the back, and males sit inside soaking their bodies in the water. Females lay eggs inside such a burrow, and hatched tadpoles stay inside or around the burrow almost two months until they complete metamorphosis. Larvae can metamorphose even without feeding in the captivity, although it may feed detritus in the wild. Metamorphosis begins at mid June in the
<typeStatus id="548C883EFFC5FFDAFC5B67D15BA12BEA" box="[1388,1436,403,429]" pageId="21" pageNumber="422">type</typeStatus>
locality (
<bibRefCitation id="EFA64B6DFFC5FFDAF9CD67F55FD92B96" author="Sugihara, T. &amp; Matsui, M. &amp; Sugahara, T." box="[250,484,439,465]" pageId="21" pageNumber="422" pagination="230 - 237" refId="ref18464" refString="Sugihara, T., Matsui, M. &amp; Sugahara, T. (2022) A comparison of growth patterns between large- and small-sized Rana tagoi (Amphibia: Ranidae) from Kitayama, Kyoto, Japan. Current Herpetology, 41, 230 - 237. https: // doi. org / 10.5358 / hsj. 41.230" type="journal article" year="2022">
Sugihara
<emphasis id="B943EA8EFFC5FFDAF85067F55FA52B96" box="[359,408,439,465]" italics="true" pageId="21" pageNumber="422">et al</emphasis>
. 2022
</bibRefCitation>
). In many known habitats, the new species occurs syntopically with its close relative
<emphasis id="B943EA8EFFC5FFDAF9A067995F3B2BB3" box="[151,262,475,500]" italics="true" pageId="21" pageNumber="422">
R. t.
<taxonomicName id="4C374D1FFFC5FFDAF9FB67995F3B2BB3" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[204,262,475,500]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="21" pageNumber="422" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
. Their breeding sites are close to each other in the
<typeStatus id="548C883EFFC5FFDAFA7567995D4F2BB2" box="[834,882,475,501]" pageId="21" pageNumber="422">type</typeStatus>
locality and both of them use subterranean water current for oviposition site; however, it seems they prefer different micro habitat and generally they do not share same burrow (
<bibRefCitation id="EFA64B6DFFC5FFDAF80D64615C60287A" author="Sugahara, T. &amp; Matsui, M." box="[314,605,547,573]" pageId="21" pageNumber="422" pagination="68 - 69" refId="ref18425" refString="Sugahara, T. &amp; Matsui, M. (1997) Breeding seasons and sites in two types of Rana tagoi tagoi from Kumogahata, Kyoto. Japanese Journal of Herpetology, 17, 68 - 69." type="journal article" year="1997">Sugahara &amp; Matsui 1997</bibRefCitation>
). In the
<typeStatus id="548C883EFFC5FFDAFB8B64615CD1287A" box="[700,748,547,573]" pageId="21" pageNumber="422">type</typeStatus>
locality of
<taxonomicName id="4C374D1FFFC5FFDAFA4664615DF2287A" authority="Eto &amp; Matsui &amp; Sugahara, 2022" authorityName="Eto &amp; Matsui &amp; Sugahara" authorityYear="2022" box="[881,975,547,573]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="21" pageNumber="422" phylum="Chordata" rank="species" species="kyoto" status="sp. nov.">
<emphasis id="B943EA8EFFC5FFDAFA4664615DF2287A" box="[881,975,547,573]" italics="true" pageId="21" pageNumber="422">R. kyoto</emphasis>
</taxonomicName>
<emphasis id="B943EA8EFFC5FFDAFAE064615A0F287A" bold="true" box="[983,1074,547,573]" pageId="21" pageNumber="422">
<taxonomicNameLabel id="A27057F5FFC5FFDAFAE064615A0F287A" box="[983,1074,547,573]" pageId="21" pageNumber="422" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
,
<emphasis id="B943EA8EFFC5FFDAFD7564615A89287B" box="[1090,1204,547,572]" italics="true" pageId="21" pageNumber="422">
R. t.
<taxonomicName id="4C374D1FFFC5FFDAFD4D64615A89287B" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[1146,1204,547,572]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="21" pageNumber="422" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
starts calling at mid May, which lasts until early June (
<bibRefCitation id="EFA64B6DFFC5FFDAFB2B64055D032826" author="Sugahara, T. &amp; Matsui, M." box="[540,830,583,609]" pageId="21" pageNumber="422" pagination="68 - 69" refId="ref18425" refString="Sugahara, T. &amp; Matsui, M. (1997) Breeding seasons and sites in two types of Rana tagoi tagoi from Kumogahata, Kyoto. Japanese Journal of Herpetology, 17, 68 - 69." type="journal article" year="1997">Sugahara &amp; Matsui 1997</bibRefCitation>
). They begin metamorphosis at mid July (
<bibRefCitation id="EFA64B6DFFC5FFDAFC2C64055EC328C2" author="Sugihara, T. &amp; Matsui, M. &amp; Sugahara, T." pageId="21" pageNumber="422" pagination="230 - 237" refId="ref18464" refString="Sugihara, T., Matsui, M. &amp; Sugahara, T. (2022) A comparison of growth patterns between large- and small-sized Rana tagoi (Amphibia: Ranidae) from Kitayama, Kyoto, Japan. Current Herpetology, 41, 230 - 237. https: // doi. org / 10.5358 / hsj. 41.230" type="journal article" year="2022">
Sugihara
<emphasis id="B943EA8EFFC5FFDAFCBF64055E9028C2" italics="true" pageId="21" pageNumber="422">et al</emphasis>
., 2022
</bibRefCitation>
). In addition to
<emphasis id="B943EA8EFFC5FFDAF89F64295C2828C3" box="[424,533,619,644]" italics="true" pageId="21" pageNumber="422">
R. t.
<taxonomicName id="4C374D1FFFC5FFDAF8EC64295C2828C3" authorityName=": Daito &amp; Nakamizo" authorityYear="1993" box="[475,533,619,644]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="21" pageNumber="422" phylum="Chordata" rank="species" species="tagoi">tagoi</taxonomicName>
</emphasis>
, another close relative
<taxonomicName id="4C374D1FFFC5FFDAFA2464295DB028C2" authorityName="Matsui &amp; Matsui" authorityYear="1990" box="[787,909,619,645]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="21" pageNumber="422" phylum="Chordata" rank="species" species="sakuraii">
<emphasis id="B943EA8EFFC5FFDAFA2464295DB028C2" box="[787,909,619,645]" italics="true" pageId="21" pageNumber="422">R. sakuraii</emphasis>
</taxonomicName>
also co-occurs in several areas such as northern part of
<collectingRegion id="49F3F87EFFC5FFDAF9DF64CD5F5628EE" box="[232,363,655,681]" country="Japan" name="Kyoto" pageId="21" pageNumber="422">Kyoto Pref.</collectingRegion>
and Hira Mountains of
<collectingRegion id="49F3F87EFFC5FFDAFB4064CD5CC928EE" box="[631,756,655,681]" country="Japan" name="Shiga" pageId="21" pageNumber="422">Shiga Pref.</collectingRegion>
</paragraph>
</subSubSection>
</treatment>
</document>