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<document ID-DOI="http://dx.doi.org/10.3897/vz.73.e102475" ID-Pensoft-Pub="2625-8498-73-931" ID-Pensoft-UUID="EE261ADB23925B95AD23FA3F71DF2FBA" ID-ZooBank="6723C6BC034B49B68F26806978782527" ModsDocID="2625-8498-73-931" checkinTime="1696476018141" checkinUser="pensoft" docAuthor="Sheridan, Jennifer A., Phimmachak, Somphouthone, Sivongxay, Niane &amp; Stuart, Bryan L." docDate="2023" docId="5EB04F147E28594F84C4897B2938A2B6" docLanguage="en" docName="VertZool 73: 931-956" docOrigin="Vertebrate Zoology 73" docPubDate="2023-10-04" docSource="http://dx.doi.org/10.3897/vz.73.e102475" docTitle="Amolops attiguus Sheridan &amp; Phimmachak &amp; Sivongxay &amp; Stuart 2023, sp. nov." docType="treatment" docUuid="27659B24-8B32-4D0A-B0B9-AECBEFB954AC" docUuidSource="ZooBank" docVersion="2" id="EE261ADB23925B95AD23FA3F71DF2FBA" lastPageNumber="931" masterDocId="EE261ADB23925B95AD23FA3F71DF2FBA" masterDocTitle="Systematics of the Lao torrent frog, Amolops cremnobatus Inger &amp; Kottelat, 1998 (Anura: Ranidae), with descriptions of four new species" masterLastPageNumber="956" masterPageNumber="931" pageNumber="931" updateTime="1696476320615" updateUser="ExternalLinkService">
<mods:mods id="B781B0F9267B716748D6D054C9A42F24" xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:title id="C453C1155BA54FA18B5DF82994EFE580">Systematics of the Lao torrent frog, Amolops cremnobatus Inger &amp; Kottelat, 1998 (Anura: Ranidae), with descriptions of four new species</mods:title>
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<mods:name id="E08B3C8294010F8B1C64459396EF2230" type="personal">
<mods:role id="C8AFDFA991961C2684C7F52AFFC46FFD">
<mods:roleTerm id="0292A0BB5997E2BD8BCED8A5E9291F32">Author</mods:roleTerm>
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<mods:namePart id="5D00E6ACFA82D462C824D94DA1F5AB42">Sheridan, Jennifer A.</mods:namePart>
<mods:affiliation id="1EBE940A450E15A54E1A8866461ED580">Section of Amphibians &amp; Reptiles, Carnegie Museum of Natural History, 4400 Forbes Ave, Pittsburgh, PA 15213 USA</mods:affiliation>
<mods:nameIdentifier id="6AF560B67584299B0078B5377D673A4E" type="email">jasheridan@gmail.com</mods:nameIdentifier>
</mods:name>
<mods:name id="4297FB0A4FD6C0158B1938689B87E4BE" type="personal">
<mods:role id="23A529448BBB2921FBB8A8982BD222EF">
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<mods:namePart id="F1F8F4CC33DE9BE9A351129C463CB3DC">Phimmachak, Somphouthone</mods:namePart>
<mods:affiliation id="7AB3D55C64B648573DA8E449637DC956">Department of Biology, Faculty of Natural Sciences, National University of Laos, Dongdok Campus, P. O. Box 7322, Vientiane, Laos PDR</mods:affiliation>
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<mods:name id="266D398DE142BE8C6F4AEB4FDB236572" type="personal">
<mods:role id="C78495EFB1445F2BB5BF48FAC02CC1C0">
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<mods:namePart id="A08EA81EE2494A902640F8AFF2923E3C">Sivongxay, Niane</mods:namePart>
<mods:affiliation id="BF7FF2F35AA04A8FD24385050DB9C342">SEAMEO Regional Centre for Community Education Development, Ministry of Education and Sports, Dongdok, Xaithany District, P. O. Box 67, Vientiane, Lao PDR</mods:affiliation>
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<mods:name id="E51E30D80B712EBCEDD6F602BC1CD343" type="personal">
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<mods:namePart id="DD1A6AAC7B9CAABAAA0265CBEDE52AE5">Stuart, Bryan L.</mods:namePart>
<mods:affiliation id="3AA44AEDF8CE1866AAAAC6C37A41DEBD">North Carolina Museum of Natural Sciences, 11 West Jones St, Raleigh, NC 27601 USA</mods:affiliation>
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<mods:typeOfResource id="E3B985D9E83BFB0AACD15D0E96F9C1C3">text</mods:typeOfResource>
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<mods:title id="5843D0A82270DE601676F11AC5484B95">Vertebrate Zoology</mods:title>
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<mods:part id="0874649CFF2BB0F46F8C402E4BF1EF84">
<mods:date id="E1B8A9AF307806BDF112C77016DE25BA">2023</mods:date>
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<mods:number id="A0448E62DDAE2C473D15DE00A648E8CD">2023-10-04</mods:number>
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<mods:number id="8CD336692B21902C02B1A127D0606115">73</mods:number>
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<mods:classification id="EA27C16ACE40E419BF29A477C4F75E03">journal article</mods:classification>
<mods:identifier id="607B28D8F9ACA821B737BE2D7307F529" type="DOI">http://dx.doi.org/10.3897/vz.73.e102475</mods:identifier>
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<subSection id="493E9FEEAC042E0BECDDAF2E9EC42E9D" pageId="0" pageNumber="931" type="taxonomy">
<treatment id="5EB04F147E28594F84C4897B2938A2B6" LSID="urn:lsid:zoobank.org:act:27659B24-8B32-4D0A-B0B9-AECBEFB954AC" httpUri="http://treatment.plazi.org/id/5EB04F147E28594F84C4897B2938A2B6" lastPageNumber="931" pageId="0" pageNumber="931">
<subSubSection id="AA1662D3D09DA1DCE4560F4561B51096" pageId="0" pageNumber="931" type="nomenclature">
<paragraph id="4798561F9081B1CE7F39D9054EE5BDA6" pageId="0" pageNumber="931">
<taxonomicName id="C69B469480CB77ABB6F6E9173E31109A" LSID="https://zoobank.org/27659B24-8B32-4D0A-B0B9-AECBEFB954AC" authority="Sheridan &amp; Phimmachak &amp; Sivongxay &amp; Stuart, 2023" authorityName="Sheridan &amp; Phimmachak &amp; Sivongxay &amp; Stuart" authorityYear="2023" class="Amphibia" family="Ranidae" genus="Amolops" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Amolops attiguus" order="Anura" pageId="0" pageNumber="931" phylum="Chordata" rank="species" species="attiguus" status="sp. nov.">Amolops attiguus</taxonomicName>
<taxonomicNameLabel id="E762434F5743CE7B1F086D9C19D14F1C" pageId="0" pageNumber="931">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="CF0491DE9B210B965E564D17D232732E" pageId="0" pageNumber="931" type="description">
<paragraph id="4CF2BFE21BA4F4ADF5C0BBB1B3352613" pageId="0" pageNumber="931">
<figureCitation id="59B1A449EB88D5E24E99727812BC3966" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Amolops attiguus sp. nov. NCSM 80907 in life (top row) and in preservative (bottom row)." figureDoi="10.3897/vz.73.e102475.figure12" httpUri="https://binary.pensoft.net/fig/916375" pageId="0" pageNumber="931">Figure 12 &quot;Clade E&quot;</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="FFB795CF5135CA235D1CF3E648AE07C6" pageId="0" pageNumber="931" type="holotype">
<paragraph id="D61659AA633EAAADA4755428A232F950" pageId="0" pageNumber="931">Holotype.</paragraph>
<paragraph id="3600917D4541EA56D5CFE59ADD1EC3D7" pageId="0" pageNumber="931">
NCSM 80907 (field number BLS 15852), adult male, Laos, Bolikhamxay Province, Viengthong District, Nam Kading National Protected Area, Nam Xouang,
<geoCoordinate id="8CF7E6B6E7ED5BD40C514C769183754F" degrees="18.42676" direction="north" orientation="latitude" precision="1" value="18.42676">18.42676°N</geoCoordinate>
,
<geoCoordinate id="54ABE44C31310FAAB12C2B0A8DC8D47A" degrees="104.39136" direction="east" orientation="longitude" precision="1" value="104.39136">104.39136°E</geoCoordinate>
, 343 m elev., coll. 2200 h on wet rock face next to series of waterfalls over exposed sandstone bedrock with potholes on 1 March 2013 by Bryan L. Stuart, Jennifer A. Sheridan, Sengvilay Seateun, and Niane Sivongxay.
</paragraph>
<caption id="B6CE9B09BA728217D6B8D2820D033287" doi="10.3897/vz.73.e102475.figure12" httpUri="https://binary.pensoft.net/fig/916375" pageId="0" pageNumber="931" start="Figure 12" startId="F12">
<paragraph id="ADE1389EB14201465F5D32C6D2A076DD" pageId="0" pageNumber="931">
<emphasis id="072625039CF03644D1C1E9E0D8F0752C" bold="true" pageId="0" pageNumber="931">Figure 12.</emphasis>
<taxonomicName id="98AB7DB48E0F901BB24049C7081B56E5" authorityName="Sheridan &amp; Phimmachak &amp; Sivongxay &amp; Stuart" authorityYear="2023" class="Amphibia" family="Ranidae" genus="Amolops" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Amolops attiguus" order="Caudata" pageId="0" pageNumber="931" phylum="Chordata" rank="species" species="attiguus">
<emphasis id="BB079041A34FF553C55ED546165D91A6" italics="true" pageId="0" pageNumber="931">Amolops attiguus</emphasis>
</taxonomicName>
<emphasis id="2800555232728A5031D56208B143C254" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
NCSM 80907 in life (top row) and in preservative (bottom row).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="A87341959DCE0459002E6A34E5182E8C" pageId="0" pageNumber="931" type="paratypes">
<paragraph id="0F30D2DB8C55ADF97B962C459C6F169C" pageId="0" pageNumber="931">Paratypes.</paragraph>
<paragraph id="DC771A9BB129E8DA2FF2A88CB1E2ABB7" pageId="0" pageNumber="931">
Laos, Xieng Khouang, Mork District, tributary of Nam Mork: NCSM 80761 (one adult male),
<geoCoordinate id="434A5EA310A5337873B0D8CC3C8CCEC0" degrees="19.07075" direction="north" orientation="latitude" precision="1" value="19.07075">19.07075°N</geoCoordinate>
,
<geoCoordinate id="DE62B2A74073EC41452129DD432D1E45" degrees="103.95870" direction="east" orientation="longitude" precision="1" value="103.9587">103.95870°E</geoCoordinate>
, 454 m elev., coll. 12 April 2012 by Somphouthone Phimmachak. - Vietnam, Nghe An Province, Anh Son District, Pu Mat National Park, Cao Ven: AMNH A 191846 (one adult male),
<geoCoordinate id="BD0D0FDF1D20C942386F0BFA07AD6CE0" degrees="18.81050" direction="north" orientation="latitude" precision="1" value="18.8105">18.81050°N</geoCoordinate>
,
<geoCoordinate id="EC918D78729F2910FCE6EE044F169929" degrees="104.95150" direction="east" orientation="longitude" precision="1" value="104.9515">104.95150°E</geoCoordinate>
, 273 m elev., coll. 16 May 2010 by David A. Kizirian, Truong Quang Nguyen, and Sinh Van Nguyen; AMNH A 191848 (one adult male), AMNH A 191849, NCSM 79166 (two adult females), coll. 18 May 2010 by David A. Kizirian, Truong Quang Nguyen, and Sinh Van Nguyen.
</paragraph>
</subSubSection>
<subSubSection id="AEF6720757B2C78BF767E2F1BBC07E8E" pageId="0" pageNumber="931" type="etymology">
<paragraph id="722BA891EDE554DC12D7D44FAC984461" pageId="0" pageNumber="931">Etymology.</paragraph>
<paragraph id="FC7013AE45B41E672697B4C75AB939CE" pageId="0" pageNumber="931">
The specific epithet taken from
<emphasis id="26E5E9266BD3AE46AB62D9444A64F530" italics="true" pageId="0" pageNumber="931">attigua</emphasis>
(L.) for neighboring or adjacent, in reference to the new
<normalizedToken id="ED1D23F383D0BA54C3B200581EA62ED2" originalValue="species">species'</normalizedToken>
nested geographic distribution between its morphologically similar relatives
<taxonomicName id="DCDBC8ADEFF90654FB8368457D07F76E" lsidName="A. tanfuilianae" pageId="0" pageNumber="931" rank="species" species="tanfuilianae">
<emphasis id="343C29A1AE84E59E45FACF26DC14CBFD" italics="true" pageId="0" pageNumber="931">A. tanfuilianae</emphasis>
</taxonomicName>
<emphasis id="0DCF7462B8240ED4E3DA3BE94E92DAB4" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
and
<taxonomicName id="41A251D4F9077FB40E0AE5B0C2416200" lsidName="A. kottelati" pageId="0" pageNumber="931" rank="species" species="kottelati">
<emphasis id="79C3BA9F5FFD48FE3647EBA4C3B61E39" italics="true" pageId="0" pageNumber="931">A. kottelati</emphasis>
</taxonomicName>
<emphasis id="4004B94B4F49FC5403C26F4CEFA17ECB" bold="true" pageId="0" pageNumber="931">sp. nov</emphasis>
. (Fig.
<figureCitation id="92DC412912078DAD1A108225642E829B" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Map of specimen sampling locations of Amolops cremnobatus (orange stars), A. tanfuilianae sp. nov. (blue circles), A. sengae sp. nov. (green diamonds), A. kottelati sp. nov. (red triangles), and A. attiguus sp. nov. (purple squares). Filled symbols are adults and open symbols are tadpoles. Symbols with black dots are sequenced individuals, and symbols without black dots are unsequenced individuals. The sample in Thailand is provisionally referred to A. sengae sp. nov. (see text)." figureDoi="10.3897/vz.73.e102475.figure1" httpUri="https://binary.pensoft.net/fig/916364" pageId="0" pageNumber="931">1</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="306CEFD68D61B49E43FE51AD650D18E5" pageId="0" pageNumber="931" type="suggested common names">
<paragraph id="0BEFAFC48EB1B1BECE1F3B0128F3544B" pageId="0" pageNumber="931">Suggested Common Names.</paragraph>
<paragraph id="9BFE6F850724B7D9886654188E932ACA" pageId="0" pageNumber="931">
Similar Lao torrent frog (English),
<normalizedToken id="ED38F0D645E38E12D4013299112F7247" originalValue="ຂຽດເກາະຜາລາວແອດທ">ຂຽດເກາະຜາລາວແອດທ</normalizedToken>
ີກຸສ (Khiat Korpha Lao Sumphan; Lao).
</paragraph>
</subSubSection>
<subSubSection id="E0F69276D8E468052EFF26FDB9A7C837" pageId="0" pageNumber="931" type="diagnosis">
<paragraph id="D8D08F060C42CAAEBA463D869852C631" pageId="0" pageNumber="931">Diagnosis.</paragraph>
<paragraph id="A5531AC881A0816206D3677DBEA7B7E5" pageId="0" pageNumber="931">
A member of the
<taxonomicName id="B72CBBEAB46A16508504322E917A8D54" baseAuthorityName="Boulenger" baseAuthorityYear="1899" class="Amphibia" family="Ranidae" genus="Amolops" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Amolops larutensis" order="Caudata" pageId="0" pageNumber="931" phylum="Chordata" rank="species" species="larutensis">
<emphasis id="A3B851C758FDDC64A4F5463F2E6F1B61" italics="true" pageId="0" pageNumber="931">Amolops larutensis</emphasis>
</taxonomicName>
group having the combination of 2-3 vomerine teeth, sometimes faint or absent; relative finger lengths I &lt;IV&lt;II &lt;III; mean
<normalizedToken id="1BED603DAE7C14DDE9527B887FC31B1D" originalValue="±">+/-</normalizedToken>
SE SVL of adult males 30.2
<normalizedToken id="67F2C956E156645B5CD56ABD907F6407" originalValue="±">+/-</normalizedToken>
0.9 mm (range 28.9-31.3 mm; n = 5) and of females 39.2-39.4 mm (n = 2); EYE 4.7
<normalizedToken id="0CADB6F33856DB652AC18371AC9B0D83" originalValue="±">+/-</normalizedToken>
0.3 mm (range 4.1-4.8 mm) in males and 5.3-6.1 mm in females; TEY 0.7
<normalizedToken id="AC14C8998CC7742FF6798238AC4C68B8" originalValue="±">+/-</normalizedToken>
0.1 mm (range 0.5-0.9 mm) in males and 1.1-1.3 mm in females; SNT 4.1
<normalizedToken id="CD00B940B48A6EE5D190511909AADF03" originalValue="±">+/-</normalizedToken>
0.1 mm (range 4.5-4.7 mm) in males and 5.4-5.5 mm in females; and FTL 15.8
<normalizedToken id="586C5378595E83FC6440F35F31E49018" originalValue="±">+/-</normalizedToken>
0.7 mm (range 15.0-16.8 mm) in males and 18.6-18.7mm in females.
</paragraph>
</subSubSection>
<subSubSection id="24415CF1EF961264380BEDAB4F890A2D" pageId="0" pageNumber="931" type="description">
<paragraph id="DC05FE90A82DCB5709E660C6E3205FA3" pageId="0" pageNumber="931">Description of the holotype.</paragraph>
<paragraph id="5FC312B505FA1764EC1FEF72DFF8E1B9" pageId="0" pageNumber="931">Habitus moderately slender. Head length subequal to head width. Snout weakly pointed in dorsal view. Snout projecting slightly beyond lower jaw in lateral view, sloping obliquely back to lip. Nostril lateral, nearer to tip of snout than to eye. Canthus rostralis distinct. Lores obliquely and slightly concave. Eye diameter approximately equal to snout. No pineal body visible. Tympanum distinct, round, approximately one-third of eye diameter, slightly depressed relative to skin of temporal region, tympanic rim elevated relative to tympanum. Rictal glands slightly oval and pronounced. Vomerine teeth obliquely angled, closer to each other than to choanae, and extremely small, with 2-3 teeth each. Tongue cordiform, notched posteriorly. Vocal slit opening near corner of jaw. No gular pouch, but with some extra skin at edge of jaw.</paragraph>
<paragraph id="C7B95BF3D7FD9B76F0DC856C42EEAA2E" pageId="0" pageNumber="931">Forelimb moderately slender. Tips of all four fingers expanded into wide discs with circummarginal grooves. Fingers slender. Relative finger lengths I &lt;IV&lt;II &lt;III. Minimal basal webbing on hands. Subarticular tubercles conspicuous, surfaces rounded, formula 1, 1, 2, 2. One supernumary tubercle at base of fingers 2-4. Irregularly-shaped palmar and oval thenar tubercles present. Well-developed nuptial pad on Finger I, on dorsal surface to level of distal end of subarticular tubercle.</paragraph>
<paragraph id="0810B9FC2139B5334897D353E7F20721" pageId="0" pageNumber="931">Hindlimbs moderately long and slender. Toes slender. Tips of toes expanded into wide discs with circummarginal grooves. Web on all toes to base of disc. Subarticular tubercles conspicuous, surfaces rounded, formula 1, 1, 2, 2, 2. Inner metatarsal tubercle oval. No outer metatarsal tubercle. Discs of toes smaller than that of finger, but larger than or equal to that of outermost finger. Outermost toe disc slightly wider than toe. Widest toe disc narrower than all finger discs except for that of thumb. Subarticular tubercles conspicuous, surfaces rounded; formula 1, 1, 2, 2, 2. Inner metatarsal tubercle oval; outer metatarsal tubercle barely visible, small, round.</paragraph>
<paragraph id="61EE07437B01AFA799D1457BAA4EA933" pageId="0" pageNumber="931">Dorsal surface with weakly spinose glands. No supra-tympanic fold. Glandular dorsolateral fold. Flanks glandular. Spinose glands behind tympanum above arm. Glands around vent. Ventral surfaces weakly granular.</paragraph>
<paragraph id="E1187157103050CDB873B40BD3945A9B" pageId="0" pageNumber="931">Measurements of holotype (mm): SVL 30.4, HDL 10.3, HDW 11.0, SNT 4.7, EYE 4.6, IOD 3.2, IND 3.6, TMP 1.7, TEY 0.8, SHK 19.1, TGH 17.2, HND 9.7, FTL 16.8.</paragraph>
</subSubSection>
<subSubSection id="F6011B5188EC02DFA8F31533BA10A485" pageId="0" pageNumber="931" type="coloration">
<paragraph id="9668C6B9C2EAAB5A9965FB97F7D04B77" pageId="0" pageNumber="931">Coloration.</paragraph>
<paragraph id="B0E394AEFE65B915C10D1449F3893698" pageId="0" pageNumber="931">In preservative, dorsal coloration black-brown with paler markings. Pale bars on dorsal surface of thighs and arms. Posterior surface of thighs with irregular pale mottling or reticulation on dark surface. Ventral surfaces cream with very fine dark mottling. Ventral surface of feet dark grey-brown, ventral surface of hands dark beige.</paragraph>
<paragraph id="2A08AE3C3C4311E9E52E28F08CFC290C" pageId="0" pageNumber="931">
In life (from photos of NCSM 80906, Fig.
<figureCitation id="0BFFC824E6074FB7981263EA9EA2AFC9" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Map of specimen sampling locations of Amolops cremnobatus (orange stars), A. tanfuilianae sp. nov. (blue circles), A. sengae sp. nov. (green diamonds), A. kottelati sp. nov. (red triangles), and A. attiguus sp. nov. (purple squares). Filled symbols are adults and open symbols are tadpoles. Symbols with black dots are sequenced individuals, and symbols without black dots are unsequenced individuals. The sample in Thailand is provisionally referred to A. sengae sp. nov. (see text)." figureDoi="10.3897/vz.73.e102475.figure1" httpUri="https://binary.pensoft.net/fig/916364" pageId="0" pageNumber="931">1</figureCitation>
), dorsal surface greenish-grey with bright green markings. Dorsal surface of legs paler in color than dorsum. Eye mottled gray on lower one-third, mottled yellowish on upper one-third, with orange on anterior and posterior portions. No ventral photos exist for this specimen in life, but photos of this species recently euthanized (NCSM 80907, Fig.
<figureCitation id="0BB703887C992BDC259AB596AEAAC996" captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Amolops attiguus sp. nov. NCSM 80907 in life (top row) and in preservative (bottom row)." figureDoi="10.3897/vz.73.e102475.figure12" httpUri="https://binary.pensoft.net/fig/916375" pageId="0" pageNumber="931">12</figureCitation>
) show chin and chest to be creamy white with dark mottling, belly to be white with very fine mottling, and underside of arms and thighs to be translucent.
</paragraph>
</subSubSection>
<subSubSection id="090CA23FF6FA0F4E86CB430F99B41A16" pageId="0" pageNumber="931" type="sexual dimorphism">
<paragraph id="B98C62C2AE797D05A424B7F439BBEC10" pageId="0" pageNumber="931">Sexual dimorphism.</paragraph>
<paragraph id="3FBE1CEC31FA7359A4B22E290C1EE255" pageId="0" pageNumber="931">
Males (n = 5) differ from females (n = 2) in being smaller (30.2
<normalizedToken id="DA3497FF24D3ACD3A40DC3D7E46833E8" originalValue="±">+/-</normalizedToken>
0.9 mm SVL vs 39.2-39.4 mm; Table
<tableCitation id="0AFC66151E53F07F06391F8FA1C0F9FB" captionStart="Table 1" captionStartId="T1" captionText="Table 1. Mean + / - SD (range) of adult body measurements (mm) of Amolops examined in the present study." httpUri="http://table.plazi.org/id/229B758EC90EA927C14B047ECBBC36DC" pageId="0" pageNumber="931" tableUuid="229B758EC90EA927C14B047ECBBC36DC">1</tableCitation>
), and in possessing paired vocal slits and distinct nuptial pads at base of first finger.
</paragraph>
</subSubSection>
<subSubSection id="FE5F982825F37D5BC0E07F2234A76BCD" pageId="0" pageNumber="931" type="variation">
<paragraph id="874194CEC6330135C727A81316990E1A" pageId="0" pageNumber="931">Variation.</paragraph>
<paragraph id="FE4A02C9C51A2820C9334AAECC9C1355" pageId="0" pageNumber="931">Pineal body distinguishable in NCSM 80761, and clear and obvious in AMNH A 191845, 191846, 191848, and 191849, but not apparent in NCSM 79166. Rictal glands indistinct, bordering on absent in all paratypes. Vomerine teeth totally absent in AMNH A 191847; almost absent in AMNH A 191846 and A 191848; extremely small in NCSM 80761, and a bit more prominent in AMNH A 191845. Females have oval thenar tubercle. Outermost toe disc variable: in some individuals it is barely wider than toe, but in others appears wider.</paragraph>
<paragraph id="6828910E7677551B40A6E03193596526" pageId="0" pageNumber="931">Male AMNH A 191845 very slightly spinose. Dorsal surface of females glandular but not spinose. Glandular dorsolateral fold weak in females and in AMNH A 191845, 191846, and 191848. Spinose glands behind tympanum above arm, and in NCSM 79166 (female), coming down to skin in front of arm, though this feature is absent in AMNH A 191845, 191846, and 191848. In NCSM 79166, the pale dorsal markings are extremely sparse, giving the frog a dark appearance overall with few markings, while in the others, the dorsal surfaces are much more mottled with the paler markings. Pale bars on dorsal surface of thighs and arms less apparent in NCSM 79166 than in other individuals. Ventral coloration of NCSM 79166 creamy on the belly, and white on chin and chest with dark mottling. AMNH specimens A 191845, 19148, and 19149 have white chin and chest with yellow-cream belly (no dark mottling). AMNH A 19146 has white chin and chest with some dark mottling, and yellow-cream belly. Ventral surfaces of feet are dark brown to black, and ventral surfaces of hands are dark beige to black.</paragraph>
</subSubSection>
<subSubSection id="6D480055F3877030708877EFB8BD66DD" pageId="0" pageNumber="931" type="distribution">
<paragraph id="2A43510AE498E023330BE14CAF6F9D71" pageId="0" pageNumber="931">Distribution and natural history.</paragraph>
<paragraph id="4CD478F1385CA67A25204F7700E0966E" pageId="0" pageNumber="931">This species is known from clear streams with torrents in eastern Bolikhamxay and Xieng Khouang Provinces of northern Laos and southern Nghe An Province of northern Vietnam. Larvae of the new species remain unknown.</paragraph>
</subSubSection>
<subSubSection id="CB7D70E328E3E3DEFE2395004B146A83" pageId="0" pageNumber="931" type="comparisons">
<paragraph id="684240653A640233A829FE904768639A" pageId="0" pageNumber="931">Comparisons.</paragraph>
<paragraph id="1913EA17744B60D6FBA5A5796FF9A1A0" pageId="0" pageNumber="931">
<taxonomicName id="ADF3BD79072BCE99AC5875A5FF3FC452" authorityName="Sheridan &amp; Phimmachak &amp; Sivongxay &amp; Stuart" authorityYear="2023" class="Amphibia" family="Ranidae" genus="Amolops" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Amolops attiguus" order="Caudata" pageId="0" pageNumber="931" phylum="Chordata" rank="species" species="attiguus">
<emphasis id="62CDC12DF143A6E8D76CCBE0396F1468" italics="true" pageId="0" pageNumber="931">Amolops attiguus</emphasis>
</taxonomicName>
<emphasis id="FD1151528ED4311221B897B7809D00D7" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
differs from
<taxonomicName id="BB8FA81786A970D67051B2B126223D3D" lsidName="A. tanfuilianae" pageId="0" pageNumber="931" rank="species" species="tanfuilianae">
<emphasis id="2E9AE0E969C6D50986D3AC4FCF244896" italics="true" pageId="0" pageNumber="931">A. tanfuilianae</emphasis>
</taxonomicName>
<emphasis id="40F9C3360A4618230987E30BCE5E66F9" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
and
<taxonomicName id="5DD071FC4161DAD6D15C0018B0B8FF0E" lsidName="A. kottelati" pageId="0" pageNumber="931" rank="species" species="kottelati">
<emphasis id="3FEFB54C72BC035B0D49ED359DA73E28" italics="true" pageId="0" pageNumber="931">A. kottelati</emphasis>
</taxonomicName>
<emphasis id="52C2FC7F235512EDD59F61C766D38818" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
by having a smaller number (2-3) of vomerine teeth (3-4 in
<taxonomicName id="24F1B333E41E613309794DE967FD95B4" lsidName="A. tanfuilianae" pageId="0" pageNumber="931" rank="species" species="tanfuilianae">
<emphasis id="C4A0ED3AC396DDC74B9B65AE47E3A081" italics="true" pageId="0" pageNumber="931">A. tanfuilianae</emphasis>
</taxonomicName>
<emphasis id="7088BF1F2DCF75B55124273A327452C1" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
and
<taxonomicName id="62E416A151428A31013B0E7FC67FBCA6" lsidName="A. kottelati" pageId="0" pageNumber="931" rank="species" species="kottelati">
<emphasis id="729029E0E5B3B3C2C39CD746EDD50614" italics="true" pageId="0" pageNumber="931">A. kottelati</emphasis>
</taxonomicName>
<emphasis id="B96723FCB0B3B5A22E0DB4871FB40D17" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
).
<taxonomicName id="9A8CDDEA58061A2E4B0B41118FBF89D5" authorityName="Sheridan &amp; Phimmachak &amp; Sivongxay &amp; Stuart" authorityYear="2023" class="Amphibia" family="Ranidae" genus="Amolops" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Amolops attiguus" order="Caudata" pageId="0" pageNumber="931" phylum="Chordata" rank="species" species="attiguus">
<emphasis id="A857CD7DCBA95CD5A437D299A18E306D" italics="true" pageId="0" pageNumber="931">Amolops attiguus</emphasis>
</taxonomicName>
<emphasis id="61ACA496EF0274367F06384A90DE5298" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
differs from
<taxonomicName id="F37D519F98232083A9602CAE22582AF5" lsidName="A. cremnobatus" pageId="0" pageNumber="931" rank="species" species="cremnobatus">
<emphasis id="93B358B129EFC1C668A01BF9BDDCDF22" italics="true" pageId="0" pageNumber="931">A. cremnobatus</emphasis>
</taxonomicName>
and further differs from
<taxonomicName id="11D193C62683B215C44DF2A242BA5779" lsidName="A. tanfuilianae" pageId="0" pageNumber="931" rank="species" species="tanfuilianae">
<emphasis id="612FE5EBB37D864B35320373BD9F8224" italics="true" pageId="0" pageNumber="931">A. tanfuilianae</emphasis>
</taxonomicName>
<emphasis id="BC0F40FFD97B6F4707776F7DDEB91A03" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
by having relative finger lengths I&lt;IV&lt;II&lt;III (I&lt;II&lt;IV&lt;III in
<taxonomicName id="E7A0FE9B894E4CDDE38A6E58F4031600" lsidName="A. cremnobatus" pageId="0" pageNumber="931" rank="species" species="cremnobatus">
<emphasis id="D4BD22F181E4A759308246D5896B572E" italics="true" pageId="0" pageNumber="931">A. cremnobatus</emphasis>
</taxonomicName>
and
<taxonomicName id="55DCF261047AC8F5DF699E4108205C39" lsidName="A. tanfuilianae" pageId="0" pageNumber="931" rank="species" species="tanfuilianae">
<emphasis id="325176829715606D8D7225CE550C12BA" italics="true" pageId="0" pageNumber="931">A. tanfuilianae</emphasis>
</taxonomicName>
<emphasis id="9C9F889286B65A4577AC362D5BFA82F9" bold="true" pageId="0" pageNumber="931">sp. nov.)</emphasis>
.
<taxonomicName id="BE5708C8A8E075E3810EE74B8A959FFC" authorityName="Sheridan &amp; Phimmachak &amp; Sivongxay &amp; Stuart" authorityYear="2023" class="Amphibia" family="Ranidae" genus="Amolops" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Amolops attiguus" order="Caudata" pageId="0" pageNumber="931" phylum="Chordata" rank="species" species="attiguus">
<emphasis id="8B363C890C31EBD18A860DBB1DA82A23" italics="true" pageId="0" pageNumber="931">Amolops attiguus</emphasis>
</taxonomicName>
<emphasis id="65B3F2A18F4483B8D0B76FFE32410ABE" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
further differs from
<taxonomicName id="274B850DE38E8F2DDB5462F7950199DF" lsidName="A. sengae" pageId="0" pageNumber="931" rank="species" species="sengae">
<emphasis id="9C587F31AA7EB95417FD3B60C05F3A40" italics="true" pageId="0" pageNumber="931">A. sengae</emphasis>
</taxonomicName>
<emphasis id="B3B7A82B461C81ACF3EE1294BEB74610" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
by having spinose glands above the arm usually present (absent in
<taxonomicName id="75C04AD87D69C1009C562AEA915D7599" lsidName="A. sengae" pageId="0" pageNumber="931" rank="species" species="sengae">
<emphasis id="2681C3498BC3F7AA60E24C07F0CC126B" italics="true" pageId="0" pageNumber="931">A. sengae</emphasis>
</taxonomicName>
<emphasis id="0C676C60464C66AB517467DC17A20C1F" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
).
<taxonomicName id="70F0EB98A3556C63AD2A93FA6837112F" authorityName="Sheridan &amp; Phimmachak &amp; Sivongxay &amp; Stuart" authorityYear="2023" class="Amphibia" family="Ranidae" genus="Amolops" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Amolops attiguus" order="Caudata" pageId="0" pageNumber="931" phylum="Chordata" rank="species" species="attiguus">
<emphasis id="6F1EC2B112D57DBBAF999E72E86FCF3B" italics="true" pageId="0" pageNumber="931">Amolops attiguus</emphasis>
</taxonomicName>
<emphasis id="892CFAE9A680B01D81BEB37805489CB5" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
further differs from
<taxonomicName id="30CAA4142BBB69F07211311C6A8FD39D" lsidName="A. cremnobatus" pageId="0" pageNumber="931" rank="species" species="cremnobatus">
<emphasis id="99C163FD6203E1B7BF8667A96B288C2E" italics="true" pageId="0" pageNumber="931">A. cremnobatus</emphasis>
</taxonomicName>
and
<taxonomicName id="55137447E80A332A63E248577DDD6E69" lsidName="A. sengae" pageId="0" pageNumber="931" rank="species" species="sengae">
<emphasis id="FC65FA411FECA3E608BC87ECB5C190A8" italics="true" pageId="0" pageNumber="931">A. sengae</emphasis>
</taxonomicName>
<emphasis id="92A4DD8DC0D2FAA65BAEBF2D5117112D" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
by having larger EYE and smaller TEY (Fig.
<figureCitation id="A97C51F4104B2EE55A00B089A7FAF7D0" captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Box and whisker plots of measurements (mm) of morphological features of males of Amolops cremnobatus (B; n = 12), A. sengae sp. nov. (C; n = 5), and A. attiguus sp. nov. (E; n = 5). Measurements are snout-vent length (SVL), snout length (SNT), eye diameter (EYE), tympanum-eye distance (TEY), tarsus length (SHK), and foot length (FTL). Boxes show interquartile range and median (thick black line). Whiskers represent range up to 1.5 x interquartile range (IQR); dots show range beyond 1.5 x IQR. Clusters with the same symbol (* and †) are not significantly different, as determined by paired two-tailed t-tests following ANOVA." figureDoi="10.3897/vz.73.e102475.figure11" httpUri="https://binary.pensoft.net/fig/916374" pageId="0" pageNumber="931">11</figureCitation>
).
<taxonomicName id="5A37D7F276CB6708E8A5CA599A62C31E" authorityName="Sheridan &amp; Phimmachak &amp; Sivongxay &amp; Stuart" authorityYear="2023" class="Amphibia" family="Ranidae" genus="Amolops" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Amolops attiguus" order="Caudata" pageId="0" pageNumber="931" phylum="Chordata" rank="species" species="attiguus">
<emphasis id="8902376F88D19481131C48061F184B36" italics="true" pageId="0" pageNumber="931">Amolops attiguus</emphasis>
</taxonomicName>
<emphasis id="A269F9E4492DE9D66227A6565ABEE020" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
further differs from
<taxonomicName id="02D14BD2BEDF7621EA604253F02F485F" lsidName="A. sengae" pageId="0" pageNumber="931" rank="species" species="sengae">
<emphasis id="F6249824937EC7D00C50E4D281E23824" italics="true" pageId="0" pageNumber="931">A. sengae</emphasis>
</taxonomicName>
<emphasis id="3FC2AD156B2061582035EF9F9CF32CCD" bold="true" pageId="0" pageNumber="931">sp. nov.</emphasis>
by having smaller SNT and FTL (Fig.
<figureCitation id="20777E83CBA5E1D09AC53761632C25EC" captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Box and whisker plots of measurements (mm) of morphological features of males of Amolops cremnobatus (B; n = 12), A. sengae sp. nov. (C; n = 5), and A. attiguus sp. nov. (E; n = 5). Measurements are snout-vent length (SVL), snout length (SNT), eye diameter (EYE), tympanum-eye distance (TEY), tarsus length (SHK), and foot length (FTL). Boxes show interquartile range and median (thick black line). Whiskers represent range up to 1.5 x interquartile range (IQR); dots show range beyond 1.5 x IQR. Clusters with the same symbol (* and †) are not significantly different, as determined by paired two-tailed t-tests following ANOVA." figureDoi="10.3897/vz.73.e102475.figure11" httpUri="https://binary.pensoft.net/fig/916374" pageId="0" pageNumber="931">11</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</subSection>
</document>