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<mods:title id="4B293614811035BC2B78BDF6A36E21FE">Revisiting the phylogenetic predicament of the genus Huia (Amphibia: Ranidae) using molecular data and tadpole morphology</mods:title>
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<paragraph id="8BE736BACF68FFC8FEDAE543FDC6FDDA" blockId="14.[309,596,527,552]" box="[309,596,527,552]" pageId="14" pageNumber="687">
<heading id="D0AF81D6CF68FFC8FEDAE543FDC6FDDA" box="[309,596,527,552]" centered="true" fontSize="9" level="2" pageId="14" pageNumber="687" reason="2">
GENUS
<taxonomicName id="4C584D39CF68FFC8FE63E55CFDC6FDDA" ID-CoL="4Z6D" authority="YANG, 1991" authorityName="Yang" authorityYear="1991" box="[396,596,527,552]" class="Amphibia" family="Ranidae" genus="Huia" kingdom="Animalia" order="Anura" pageId="14" pageNumber="687" phylum="Chordata" rank="genus">
<emphasis id="B92CEAA8CF68FFC8FE63E55CFE5AFDD5" box="[396,456,528,551]" italics="true" pageId="14" pageNumber="687">HUIA</emphasis>
<bibRefCitation id="EFC94B4BCF68FFC8FE21E543FDC6FDDA" author="Yang DT" box="[462,596,527,552]" pageId="14" pageNumber="687" pagination="1 - 42" refId="ref15214" refString="Yang DT. 1991. Phylogenetic systematics of the Amolops group of ranid frogs of southeastern Asia and the Greater Sunda Islands. Fieldiana Zoology 63: 1 - 42." type="journal article" year="1991">YANG, 1991</bibRefCitation>
</taxonomicName>
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<subSubSection id="C3426531CF68FFC8FE7EE577FC4BFEA4" pageId="14" pageNumber="687" type="description">
<paragraph id="8BE736BACF68FFC8FE7EE577FE6AFDA6" blockId="14.[401,504,571,596]" box="[401,504,571,596]" pageId="14" pageNumber="687">
(
<figureCitation id="13632A3FCF68FFC8FE75E577FE62FDA6" box="[410,496,571,596]" captionStart="Figure 6" captionStartId="15.[163,241,1574,1596]" captionTargetBox="[165,1441,197,1532]" captionTargetId="figure-26@15.[163,1443,195,1534]" captionTargetPageId="15" captionText="Figure 6. A, Huia cavitympanum male, Taman Nasional Bukit Baka Bukit Raya, Provinsi Kalimantan Barat, Indonesia. B, Wijayarana (cladeA); UA20140663, male, PadangAro, Taman Nasional Kerinci-Seblat, Provinsi Sumatera Barat, Indonesia. C, Wijayarana (clade B); UA20150202, male, Wiyono Waterfall, Provinsi Lampung, Indonesia. D, Wijayarana (clade C); UA2015046465, male and female in amplexus, Batang Karangan, Provinsi Sumatera Barat, Indonesia. E, Wijayarana (clade E); UA20150032, male, Banyumas, Provinsi Jawa Tengah, Indonesia. F, Wijayarana (clade F); UA20150049; Palutungan, Provinsi Jawa Barat, Indonesia. Size not to scale. Photographs by U.A." pageId="14" pageNumber="687">FIG. 6A</figureCitation>
)
</paragraph>
<paragraph id="8BE736BACF68FFC8FF7EE528FE89FD25" blockId="14.[145,762,612,727]" pageId="14" pageNumber="687">
<emphasis id="B92CEAA8CF68FFC8FF7EE528FEA0FD89" box="[145,306,612,635]" italics="true" pageId="14" pageNumber="687">Type species:</emphasis>
<taxonomicName id="4C584D39CF68FFC8FED5E52AFD60FD88" authority="Boulenger, 1893" authorityName="Boulenger" authorityYear="1893" box="[314,754,613,635]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="14" pageNumber="687" phylum="Chordata" rank="species" species="cavitympanum">
<emphasis id="B92CEAA8CF68FFC8FED5E52AFDBDFD89" box="[314,559,614,635]" italics="true" pageId="14" pageNumber="687">Rana cavitympanum</emphasis>
<materialsCitation id="3B303CE7CF68FFC8FDD8E529FD60FD88" box="[567,754,613,635]" collectionCode="BMNH" location="Proceedings" municipality="Zoological Society of London" pageId="14" pageNumber="687" specimenCode="BMNH 1893.5" specimenCount="1" typeStatus="holotype">Boulenger, 1893</materialsCitation>
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,
<emphasis id="B92CEAA8CF68FFC8FF7EE5C8FD28FD6B" box="[145,698,644,665]" italics="true" pageId="14" pageNumber="687">
<location id="8E876061CF68FFC8FF7EE5C8FE8EFD6B" LSID="urn:lsid:plazi:treatment:03F187ACCF68FFC8FEDAE543FC4BFEA4:8E876061CF68FFC8FF7EE5C8FE8EFD6B" box="[145,284,644,665]" municipality="Zoological Society of London" name="Proceedings" pageId="14" pageNumber="687">Proceedings</location>
of the
<collectingMunicipality id="6B83ACC0CF68FFC8FE83E5C8FD28FD6B" box="[364,698,644,665]" pageId="14" pageNumber="687">Zoological Society of London</collectingMunicipality>
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: 525.
<typeStatus id="54E38818CF68FFC8FF7EE5EEFF6FFD4A" box="[145,253,674,696]" pageId="14" pageNumber="687" type="holotype">Holotype</typeStatus>
:
<specimenCode id="DBFE9EC1CF68FFC8FEE7E5EFFE20FD4A" box="[264,434,674,696]" collectionCode="BMNH" pageId="14" pageNumber="687">BMNH 1893.5</specimenCode>
.30.26 (BMNH 1947.2.4.16); adult male).
</paragraph>
<paragraph id="8BE736BACF68FFC8FF7EE454FDC2FCBF" blockId="14.[145,761,791,846]" pageId="14" pageNumber="687">
<materialsCitation id="3B303CE7CF68FFC8FF7EE454FECBFCBC" country="Malaysia" location="Kinabalu" pageId="14" pageNumber="687" specimenCount="1" stateProvince="Sabah" typeStatus="holotype">
<emphasis id="B92CEAA8CF68FFC8FF7EE454FEABFCDD" box="[145,313,791,815]" italics="true" pageId="14" pageNumber="687">
<typeStatus id="54E38818CF68FFC8FF7EE454FF5EFCDD" box="[145,204,792,815]" pageId="14" pageNumber="687">Type</typeStatus>
locality:
</emphasis>
<taxonomicName id="4C584D39CF68FFC8FEACE455FE55FCDD" authority="Baloo" authorityName="Baloo" box="[323,455,793,815]" class="Echinoidea" family="Schizasteridae" genus="Kina" kingdom="Animalia" order="Spatangoida" pageId="14" pageNumber="687" phylum="Echinodermata" rank="genus">Kina Baloo</taxonomicName>
, North Borneo (
<location id="8E876061CF68FFC8FD69E455FD67FCDD" LSID="urn:lsid:plazi:treatment:03F187ACCF68FFC8FEDAE543FC4BFEA4:8E876061CF68FFC8FD69E455FD67FCDD" box="[646,757,793,815]" country="Malaysia" name="Kinabalu" pageId="14" pageNumber="687" stateProvince="Sabah">Kinabalu</location>
,
<collectingRegion id="499CF858CF68FFC8FF7EE474FF48FCBC" box="[145,218,824,846]" country="Malaysia" name="Sabah" pageId="14" pageNumber="687">Sabah</collectingRegion>
,
<collectingCountry id="F34F762ACF68FFC8FF0AE474FEC2FCBC" box="[229,336,824,846]" name="Malaysia" pageId="14" pageNumber="687">Malaysia</collectingCountry>
)
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. Collector: A. Everett.
</paragraph>
<paragraph id="8BE736BACF68FFC8FF7EE4C2FD80FC11" blockId="14.[145,761,910,995]" pageId="14" pageNumber="687">
<emphasis id="B92CEAA8CF68FFC8FF7EE4C2FEC0FC54" box="[145,338,910,934]" italics="true" pageId="14" pageNumber="687">Common name:</emphasis>
Sabah
<taxonomicName id="4C584D39CF68FFC8FE46E4DCFE70FC54" authorityName="Yang" authorityYear="1991" box="[425,482,912,934]" class="Amphibia" family="Ranidae" genus="Huia" kingdom="Animalia" order="Anura" pageId="14" pageNumber="687" phylum="Chordata" rank="genus">Huia</taxonomicName>
frog (English) following
<bibRefCitation id="EFC94B4BCF68FFC8FF7EE4E2FE31FC36" author="Frank N &amp; Ramus E" box="[145,419,942,964]" pageId="14" pageNumber="687" refId="ref13385" refString="Frank N, Ramus E. 1995. A complete guide to scientific and common names of reptiles and amphibians of the world. NG Publishing, Inc." type="book" year="1995">Frank &amp; Ramus (1995)</bibRefCitation>
and
<bibRefCitation id="EFC94B4BCF68FFC8FE0FE4E3FDE1FC37" author="Frost DR" box="[480,627,943,965]" pageId="14" pageNumber="687" refId="ref13414" refString="Frost DR. 2019. Amphibian species of the world: an online reference. Version 6. 0. Available at: https: // amphibiansoftheworld. amnh. org" type="url" year="2019">Frost (2019)</bibRefCitation>
; Kongkang Jeram Sabah (Bahasa
<collectingCountry id="F34F762ACF68FFC8FE7CE481FD95FC11" box="[403,519,973,995]" name="Indonesia" pageId="14" pageNumber="687">Indonesia</collectingCountry>
).
</paragraph>
<paragraph id="8BE736BACF68FFC8FF7EE36FFE6DFB26" blockId="14.[145,761,1059,1236]" pageId="14" pageNumber="687">
<emphasis id="B92CEAA8CF68FFC8FF7EE36FFD52FBC9" box="[145,704,1059,1083]" italics="true" pageId="14" pageNumber="687">
Phylogenetic definition and content:
<taxonomicName id="4C584D39CF68FFC8FD6CE36AFD52FBC9" authorityName="Yang" authorityYear="1991" box="[643,704,1062,1083]" class="Amphibia" family="Ranidae" genus="Huia" kingdom="Animalia" order="Anura" pageId="14" pageNumber="687" phylum="Chordata" rank="genus">Huia</taxonomicName>
</emphasis>
is a distinct and independently evolving genus, which shares a most recent common ancestor with the genus
<taxonomicName id="4C584D39CF68FFC8FF7EE3CEFEBFFB65" authorityName="Yang" authorityYear="1991" box="[145,301,1154,1175]" class="Amphibia" family="Ranidae" genus="Meristogenys" kingdom="Animalia" order="Anura" pageId="14" pageNumber="687" phylum="Chordata" rank="genus">
<emphasis id="B92CEAA8CF68FFC8FF7EE3CEFEBFFB65" box="[145,301,1154,1175]" italics="true" pageId="14" pageNumber="687">Meristogenys</emphasis>
</taxonomicName>
(
<figureCitation id="13632A3FCF68FFC8FEAFE3CDFE16FB64" box="[320,388,1153,1175]" captionStart="Figure 2" captionStartId="5.[163,245,1601,1623]" captionTargetBox="[166,1446,198,1558]" captionTargetId="figure-27@5.[163,1449,195,1561]" captionTargetPageId="5" captionText="Figure 2. Maximum likelihood tree (concatenated dataset) depicting phylogenetic relationship of frogs previously recognized as Huia (coloured branches, which also match with the island/locality of the samples; clades AH) from Sumatra, Java, Borneo and the mainland to other closely related species within the family Ranidae. Black circles represent highly supported nodes [bootstrap support (BS)&gt; 70] for unphased and phased dataset, whereas low support values are shown with a slash (/) for unphased dataset (BS)/phased dataset 1 (BS1)/phased dataset 2 (BS2). Tadpole samples are represented by _Tad. Topotypic samples are represented by numbers *1*4 and the locality shown on the map. Pink triangles, orange circles, blue stars and purple squares on the map represent the sampling sites." pageId="14" pageNumber="687">Fig. 2</figureCitation>
). It is currently understood to be monotypic, containing
<taxonomicName id="4C584D39CF68FFC8FE54E3ECFD18FB47" authorityName="Boulenger" authorityYear="1893" box="[443,650,1184,1205]" class="Amphibia" family="Ranidae" genus="Rana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="14" pageNumber="687" phylum="Chordata" rank="species" species="cavitympanum">
<emphasis id="B92CEAA8CF68FFC8FE54E3ECFD18FB47" box="[443,650,1184,1205]" italics="true" pageId="14" pageNumber="687">H. cavitympanum</emphasis>
</taxonomicName>
, which is endemic to the island of Borneo.
</paragraph>
<paragraph id="8BE736BACF68FFC8FF7EE259FC4BFEA4" blockId="14.[145,762,1301,1907]" lastBlockId="14.[809,1425,197,342]" pageId="14" pageNumber="687">
<emphasis id="B92CEAA8CF68FFC8FF7EE259FE85FADE" box="[145,279,1301,1324]" italics="true" pageId="14" pageNumber="687">Diagnosis:</emphasis>
(Tadpole characters presented here are based on observation by U.A., whereas morphological characters of adults are taken from published studies (e.g.
<bibRefCitation id="EFC94B4BCF68FFC8FF2BE23FFED0FA7A" author="Inger RF" box="[196,322,1394,1416]" pageId="14" pageNumber="687" pagination="1 - 402" refId="ref13706" refString="Inger RF. 1966. The systematics and zoogeography of the Amphibia of Borneo. Fieldiana Zoology 52: 1 - 402." type="journal article" year="1966">Inger, 1966</bibRefCitation>
;
<bibRefCitation id="EFC94B4BCF68FFC8FEA3E23FFE55FA7A" author="Yang DT" box="[332,455,1395,1416]" pageId="14" pageNumber="687" pagination="1 - 42" refId="ref15214" refString="Yang DT. 1991. Phylogenetic systematics of the Amolops group of ranid frogs of southeastern Asia and the Greater Sunda Islands. Fieldiana Zoology 63: 1 - 42." type="journal article" year="1991">Yang, 1991</bibRefCitation>
;
<bibRefCitation id="EFC94B4BCF68FFC8FE3CE23EFD79FA7A" author="Manthey U &amp; Denzer W" box="[467,747,1394,1416]" pageId="14" pageNumber="687" pagination="31 - 48" refId="ref14177" refString="Manthey U, Denzer W. 2014. Sudostasiatische Anuren im Fokus, Spezies der Gattung Huia (sensu lato) Yang, 1991 (Amphibia: Anura: Ranidae). Sauria 36: 31 - 48." type="journal article" year="2014">Manthey &amp; Denzer, 2014</bibRefCitation>
). Larvae gastromyzophorous; upper beak Λ- shaped, lower beak V-shaped, both undivided; LTRF 1012((3 4)(1012))/56(02); postorbital glands present; an oblique cluster of glands immediately below the spiracular tube, running upward and backward present; prespiracular glands present in some individuals (
<bibRefCitation id="EFC94B4BCF68FFC8FECFE106FE0DF9AD" author="Inger RF" box="[288,415,1609,1631]" pageId="14" pageNumber="687" pagination="1 - 89" refId="ref13729" refString="Inger RF. 1985. Tadpoles of the forested regions of Borneo. Fieldiana Zoology 26: 1 - 89." type="journal article" year="1985">Inger, 1985</bibRefCitation>
); glands on the tail fin absent. Adults medium in size, body stocky; snout rounded; tympanum clear, deep in cavity; supra- and posttympanic folds present, dorsolateral folds present; ratio of tibia length to snoutvent length&gt; 70%; outer metatarsal tubercle present or absent (absent according to
<bibRefCitation id="EFC94B4BCF68FFC8FEC5E04EFE3FF8E4" author="Yang DT" box="[298,429,1793,1815]" pageId="14" pageNumber="687" pagination="1 - 42" refId="ref15214" refString="Yang DT. 1991. Phylogenetic systematics of the Amolops group of ranid frogs of southeastern Asia and the Greater Sunda Islands. Fieldiana Zoology 63: 1 - 42." type="journal article" year="1991">Yang, 1991</bibRefCitation>
), inner metatarsal tubercle present, small; length of Finger I = Finger II; length of metacarpal of Finger I 54% of Finger II; width of crossbar terminal phalanx of Finger III 50% length; male with nuptial pad and paired lateral vocal sacs; humeral gland absent. In life, dorsum generally dark brown; flanks light brown with markings; head light brown with scattered dark brown markings; lips with dark markings.
</paragraph>
</subSubSection>
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