treatments-xml/data/C2/0D/87/C20D878E6878FFB165ED6565FCECF98A.xml

<document ID-DOI="10.1093/zoolinnean/zlac038" ID-GBIF-Dataset="6ae97081-de1d-4297-a576-fa4d72beea43" ID-ISSN="0024-4082" ID-Zenodo-Dep="7695335" ID-ZooBank="A09E3E61-12DE-43F6-A427-922EE0381F58" checkinTime="1677744031252" checkinUser="plazi" docAuthor="Dufresnes, Christophe &amp; Hernandez, Axel" docDate="2023" docId="C20D878E6878FFB165ED6565FCECF98A" docLanguage="en" docName="zlac038.pdf" docOrigin="Zoological Journal of the Linnean Society 197" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-.journal_article" docStyleVersion="7" docTitle="Echinotriton (Echinotriton) raffaellii Dufresnes &amp; Hernandez, 2023, SP. NOV." docType="treatment" docVersion="2" lastPageNumber="636" masterDocId="3E34FFF66875FFA165546331FFE0FF9C" masterDocTitle="Towards completing the crocodile newts’ puzzle with all-inclusive phylogeographic resources" masterLastPageNumber="640" masterPageNumber="620" pageNumber="633" updateTime="1677840223197" updateUser="ExternalLinkService">
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<mods:title>Towards completing the crocodile newts’ puzzle with all-inclusive phylogeographic resources</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Dufresnes, Christophe</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Hernandez, Axel</mods:namePart>
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<mods:title>Zoological Journal of the Linnean Society</mods:title>
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<mods:date>2023</mods:date>
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<mods:number>2022-06-02</mods:number>
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<mods:identifier type="DOI">10.1093/zoolinnean/zlac038</mods:identifier>
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<paragraph blockId="13.[185,721,1619,1675]" pageId="13" pageNumber="633">
<heading centered="true" fontSize="9" level="2" pageId="13" pageNumber="633" reason="3">
<taxonomicName authorityName="Dufresnes &amp; Hernandez" authorityYear="2023" box="[185,721,1619,1643]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="634" phylum="Chordata" rank="species" species="raffaellii" subGenus="Echinotriton">
<emphasis bold="true" box="[185,368,1620,1643]" italics="true" pageId="13" pageNumber="633">ECHINOTRITON</emphasis>
<emphasis bold="true" box="[376,579,1619,1643]" pageId="13" pageNumber="633">
(
<emphasis bold="true" box="[385,569,1620,1643]" italics="true" pageId="13" pageNumber="633">ECHINOTRITON</emphasis>
)
</emphasis>
<emphasis bold="true" box="[586,721,1623,1642]" italics="true" pageId="13" pageNumber="633">RAFFAELLII</emphasis>
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HERNANDEZ &amp; DUFRESNES 
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<taxonomicNameLabel box="[562,661,1650,1674]" pageId="13" pageNumber="633" rank="species">SP. NOV.</taxonomicNameLabel>
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</paragraph>
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<subSubSection pageId="13" pageNumber="633" type="description">
<paragraph blockId="13.[410,495,1689,1714]" box="[410,495,1689,1714]" pageId="13" pageNumber="633">
(
<figureCitation box="[419,487,1689,1714]" captionStart="Figure 9" captionStartId="14.[163,243,1616,1638]" captionTargetBox="[338,1260,195,1563]" captionTargetId="figure-103@14.[348,1267,697,1577]" captionTargetPageId="14" captionText="Figure 9. Appearance of Echinotriton (Echinotriton) raffaellii sp. nov. Top: live individual observed at the type locality (credits: AH); bottom: the holotype MVZ:Herp:232187 (credits: MVZ)." figureDoi="http://doi.org/10.5281/zenodo.7695361" httpUri="https://zenodo.org/record/7695361/files/figure.png" pageId="13" pageNumber="633">FIG. 9</figureCitation>
)
</paragraph>
<paragraph blockId="13.[145,762,1761,1814]" pageId="13" pageNumber="633">
<emphasis box="[145,479,1761,1783]" italics="true" pageId="13" pageNumber="633">Z o o b a n k r e g i s t r a t i o n:</emphasis>
u r n: l s i d: z o o b a n k. org:act: 
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.
</paragraph>
</subSubSection>
<subSubSection pageId="13" pageNumber="633" type="diagnosis">
<paragraph blockId="13.[145,761,1854,1907]" lastBlockId="13.[809,1425,197,1108]" pageId="13" pageNumber="633">
<emphasis box="[145,415,1854,1875]" italics="true" pageId="13" pageNumber="633">Identity and diagnosis:</emphasis>
A robust newt characterized by a series of 12–13 conspicuous spiny costal warts with one to three extra rows of smaller warts next to the vertebral ridge (
<figureCitation box="[1045,1111,228,250]" captionStart="Figure 9" captionStartId="14.[163,243,1616,1638]" captionTargetBox="[338,1260,195,1563]" captionTargetId="figure-103@14.[348,1267,697,1577]" captionTargetPageId="14" captionText="Figure 9. Appearance of Echinotriton (Echinotriton) raffaellii sp. nov. Top: live individual observed at the type locality (credits: AH); bottom: the holotype MVZ:Herp:232187 (credits: MVZ)." figureDoi="http://doi.org/10.5281/zenodo.7695361" httpUri="https://zenodo.org/record/7695361/files/figure.png" pageId="13" pageNumber="633">Fig. 9</figureCitation>
). Background coloration is uniformly black, except for the underside of the tail, the cloacal region, the feet soles and the tips of the costal warts, which are yellow-orange to reddish.The fifth toe is rudimentary. The vomero-palatine teeth are V-shaped in two longitudinal series that meet in front. 
<taxonomicName authorityName="Dufresnes &amp; Hernandez" authorityYear="2023" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="633" phylum="Chordata" rank="species" species="raffaellii">
<emphasis italics="true" pageId="13" pageNumber="633">Echinotriton raffaellii</emphasis>
</taxonomicName>
is the sister-species to 
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1892" box="[1153,1292,412,433]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="633" phylum="Chordata" rank="species" species="andersoni">
<emphasis box="[1153,1292,412,433]" italics="true" pageId="13" pageNumber="633">E. andersoni</emphasis>
</taxonomicName>
, with which it was previously confounded. Specifically, it corresponds to the 
<taxonomicName box="[882,1025,473,494]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="633" phylum="Chordata" rank="genus">
<emphasis box="[882,1025,473,494]" italics="true" pageId="13" pageNumber="633">Echinotriton</emphasis>
</taxonomicName>
populations inhabiting the Amami Island group (yellow, orange and red squares in 
<figureCitation box="[1348,1411,504,526]" captionStart="Figure 2" captionStartId="5.[144,225,1312,1334]" captionTargetBox="[145,1425,196,1272]" captionTargetId="graphics-288@5.[712,847,726,743]" captionTargetPageId="5" captionText="Figure 2. Phylogeography of crocodile newts part I: genus Echinotriton and subgenus Tylototriton. The phylogenetic position and geographic distribution of each lineage is detailed by coloured symbols on the tree and the corresponding maps. Type localities of described taxa are indicated by stars. For T. Ʋerrucosus, we highlighted the paraphyletic position of sequences attributed to T. shanjing, as well as its type locality with asterisks." figureDoi="http://doi.org/10.5281/zenodo.7695343" httpUri="https://zenodo.org/record/7695343/files/figure.png" pageId="13" pageNumber="633">Fig. 2</figureCitation>
), often referred to as the ‘Amami clade of 
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1892" box="[1250,1391,534,556]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="633" phylum="Chordata" rank="species" species="andersoni">
<emphasis box="[1250,1391,534,556]" italics="true" pageId="13" pageNumber="633">E. andersoni</emphasis>
</taxonomicName>
’ in the literature. True 
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1892" box="[1041,1187,565,587]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="633" phylum="Chordata" rank="species" species="andersoni">
<emphasis box="[1041,1187,565,587]" italics="true" pageId="13" pageNumber="633">E. andersoni</emphasis>
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should be restricted to the populations inhabiting the 
<collectingRegion box="[1230,1338,596,618]" country="Japan" name="Okinawa" pageId="13" pageNumber="633">Okinawa</collectingRegion>
Island group (pink and purple squares in 
<figureCitation box="[1219,1284,626,648]" captionStart="Figure 2" captionStartId="5.[144,225,1312,1334]" captionTargetBox="[145,1425,196,1272]" captionTargetId="graphics-288@5.[712,847,726,743]" captionTargetPageId="5" captionText="Figure 2. Phylogeography of crocodile newts part I: genus Echinotriton and subgenus Tylototriton. The phylogenetic position and geographic distribution of each lineage is detailed by coloured symbols on the tree and the corresponding maps. Type localities of described taxa are indicated by stars. For T. Ʋerrucosus, we highlighted the paraphyletic position of sequences attributed to T. shanjing, as well as its type locality with asterisks." figureDoi="http://doi.org/10.5281/zenodo.7695343" httpUri="https://zenodo.org/record/7695343/files/figure.png" pageId="13" pageNumber="633">Fig. 2</figureCitation>
), as per the 
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locality of this taxon (‘ 
<collectingRegion box="[1132,1241,657,679]" country="Japan" name="Okinawa" pageId="13" pageNumber="633">Okinawa</collectingRegion>
, Japan’). Based on our mitochondrial phylogeny, 
<taxonomicName authorityName="Dufresnes &amp; Hernandez" authorityYear="2023" box="[1186,1318,688,709]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="633" phylum="Chordata" rank="species" species="raffaellii">
<emphasis box="[1186,1318,688,709]" italics="true" pageId="13" pageNumber="633">E. raffaellii</emphasis>
</taxonomicName>
diverged from 
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1892" box="[872,1022,718,740]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="633" phylum="Chordata" rank="species" species="andersoni">
<emphasis box="[872,1022,718,740]" italics="true" pageId="13" pageNumber="633">E. andersoni</emphasis>
</taxonomicName>
in the Late Miocene (~7 Mya). It features 2.7% sequence differentiation at 16S, 6.4% at 
<emphasis box="[809,862,780,801]" italics="true" pageId="13" pageNumber="633">ND2</emphasis>
and 7.9% at 
<emphasis box="[1014,1067,780,801]" italics="true" pageId="13" pageNumber="633">Cytb</emphasis>
. In addition, 
<taxonomicName authorityName="Dufresnes &amp; Hernandez" authorityYear="2023" box="[1222,1355,780,801]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="633" phylum="Chordata" rank="species" species="raffaellii">
<emphasis box="[1222,1355,780,801]" italics="true" pageId="13" pageNumber="633">E. raffaellii</emphasis>
</taxonomicName>
bears private nuclear alleles at intron loci (
<figureCitation box="[1222,1283,810,832]" captionStart="Figure 5" captionStartId="8.[162,240,1620,1642]" captionTargetBox="[243,1364,197,1579]" captionTargetId="figure-104@8.[241,1364,195,1580]" captionTargetPageId="8" captionText="Figure 5. Nuclear phylogeny of 27 crocodile newt species based on 3.2 kb combining four nuclear introns (POM-C, RAG1, BDNF and NCX1). Species are symbol-coded by genera and colour-coded by subclades as in the mitochondrial phylogeny (Fig. 1). Photos: E. maxiquadratus and T. pseudoƲerrucosus (credits: AH)." figureDoi="http://doi.org/10.5281/zenodo.7695353" httpUri="https://zenodo.org/record/7695353/files/figure.png" pageId="13" pageNumber="633">Fig. 5</figureCitation>
) and unique microsatellite profiles that suggest acute nuclear differentiation without recent genetic introgression (
<bibRefCitation author="Igawa T &amp; Sugawara H &amp; Honda M &amp; Tominaga A &amp; Oumi S &amp; Katsuren S &amp; Ota H &amp; Matsui M &amp; Sumida M" box="[818,1035,902,924]" pageId="13" pageNumber="633" pagination="13 - 26" refId="ref12954" refString="Igawa T, Sugawara H, Honda M, Tominaga A, Oumi S, Katsuren S, Ota H, Matsui M, Sumida M. 2020. Detecting inter- and intra-island genetic diversity: population structure of the endangered crocodile newt, Echinotriton andersoni, in the Ryukyus. ConserVation Genetics 21: 13 - 26." type="journal article" year="2020">
Igawa 
<emphasis box="[900,961,902,924]" italics="true" pageId="13" pageNumber="633">et al.</emphasis>
, 2020
</bibRefCitation>
). Besides molecular characters, 
<taxonomicName authorityName="Dufresnes &amp; Hernandez" authorityYear="2023" box="[809,941,933,955]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="633" phylum="Chordata" rank="species" species="raffaellii">
<emphasis box="[809,941,933,955]" italics="true" pageId="13" pageNumber="633">E. raffaellii</emphasis>
</taxonomicName>
differs from other 
<taxonomicName box="[1158,1301,933,954]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="633" phylum="Chordata" rank="genus">
<emphasis box="[1158,1301,933,954]" italics="true" pageId="13" pageNumber="633">Echinotriton</emphasis>
</taxonomicName>
(including 
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1892" box="[809,960,964,985]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="13" pageNumber="633" phylum="Chordata" rank="species" species="andersoni">
<emphasis box="[809,960,964,985]" italics="true" pageId="13" pageNumber="633">E. andersoni</emphasis>
</taxonomicName>
) by a moderately smaller size (both in TTL and SVL) and a shorter tail. The head is more distinctively triangular. The body is covered by brightly coloured, anteriorly directed projections, and the costal spiny warts feature bright yellow-orange/reddish tips.
</paragraph>
</subSubSection>
<subSubSection pageId="13" pageNumber="633" type="materials_examined">
<paragraph blockId="13.[809,1425,1149,1386]" pageId="13" pageNumber="633">
<materialsCitation collectingDate="2001-01" country="Japan" county="The" location="Herp" municipality="Berkeley" pageId="13" pageNumber="633" specimenCount="1" stateProvince="Kagoshima" typeStatus="holotype">
<emphasis box="[809,1159,1149,1170]" italics="true" pageId="13" pageNumber="633">
<typeStatus box="[809,916,1149,1170]" pageId="13" pageNumber="633">Holotype</typeStatus>
: MVZ: 
<location LSID="urn:lsid:plazi:treatment:C20D878E6878FFB165ED6565FCECF98A:4F7B60436878FFAC66B9674CFBC7FB0E" box="[1005,1063,1149,1170]" country="Japan" county="The" municipality="Berkeley" name="Herp" pageId="13" pageNumber="633" stateProvince="Kagoshima">Herp</location>
:232187
</emphasis>
, adult male collected in 
<location LSID="urn:lsid:plazi:treatment:C20D878E6878FFB165ED6565FCECF98A:4F7B60436878FFAC661E67ADFC11FB2E" box="[842,1009,1180,1202]" country="Japan" county="The" municipality="Berkeley" name="Tokunoshima" pageId="13" pageNumber="633" stateProvince="Kagoshima">Tokunoshima</location>
, 
<location LSID="urn:lsid:plazi:treatment:C20D878E6878FFB165ED6565FCECF98A:4F7B60436878FFAC66AB67ADFB1BFB2E" box="[1023,1275,1180,1202]" country="Japan" county="The" municipality="Berkeley" name="Ryukyu Archipelago" pageId="13" pageNumber="633" stateProvince="Kagoshima">Ryukyu Archipelago</location>
, 
<collectingRegion country="Japan" name="Kagoshima" pageId="13" pageNumber="633">Kagoshima Prefecture</collectingRegion>
, 
<collectingCountry box="[942,1014,1211,1232]" name="Japan" pageId="13" pageNumber="633">Japan</collectingCountry>
, by 
<location LSID="urn:lsid:plazi:treatment:C20D878E6878FFB165ED6565FCECF98A:4F7B60436878FFAC6177678BFAECFB4C" box="[1059,1292,1210,1232]" country="Japan" county="The" municipality="Berkeley" name="Theodore Papenfuss" pageId="13" pageNumber="633" stateProvince="Kagoshima">Theodore Papenfuss</location>
in 
<date pageId="13" pageNumber="633" value="2001-01">
<collectingDate pageId="13" pageNumber="633" value="2001-01">January 2001</collectingDate>
</date>
(depicted in 
<figureCitation box="[1006,1068,1241,1263]" captionStart="Figure 9" captionStartId="14.[163,243,1616,1638]" captionTargetBox="[338,1260,195,1563]" captionTargetId="figure-103@14.[348,1267,697,1577]" captionTargetPageId="14" captionText="Figure 9. Appearance of Echinotriton (Echinotriton) raffaellii sp. nov. Top: live individual observed at the type locality (credits: AH); bottom: the holotype MVZ:Herp:232187 (credits: MVZ)." figureDoi="http://doi.org/10.5281/zenodo.7695361" httpUri="https://zenodo.org/record/7695361/files/figure.png" pageId="13" pageNumber="633">Fig. 9</figureCitation>
), and curated at the 
<location LSID="urn:lsid:plazi:treatment:C20D878E6878FFB165ED6565FCECF98A:4F7B60436878FFAC604067E8FBE0FA92" country="Japan" county="The" municipality="Berkeley" name="Museum of Vertebrate Zoology" pageId="13" pageNumber="633" stateProvince="Kagoshima">Museum of Vertebrate Zoology</location>
, 
<location LSID="urn:lsid:plazi:treatment:C20D878E6878FFB165ED6565FCECF98A:4F7B60436878FFAC615E67C9FB64FA92" box="[1034,1156,1272,1294]" country="Japan" county="The" municipality="Berkeley" name="University" pageId="13" pageNumber="633" stateProvince="Kagoshima">University</location>
of California, 
<collectingMunicipality box="[1320,1421,1272,1294]" pageId="13" pageNumber="633">Berkeley</collectingMunicipality>
, USA. 
<collectingCounty box="[873,917,1302,1324]" pageId="13" pageNumber="633">The</collectingCounty>
mitogenome of this specimen was sequenced (Weisrock 
<emphasis box="[926,980,1333,1355]" italics="true" pageId="13" pageNumber="633">et al.</emphasis>
, 2006) and is featured in our phylogeny (red square in 
<figureCitation box="[975,1038,1364,1386]" captionStart="Figure 2" captionStartId="5.[144,225,1312,1334]" captionTargetBox="[145,1425,196,1272]" captionTargetId="graphics-288@5.[712,847,726,743]" captionTargetPageId="5" captionText="Figure 2. Phylogeography of crocodile newts part I: genus Echinotriton and subgenus Tylototriton. The phylogenetic position and geographic distribution of each lineage is detailed by coloured symbols on the tree and the corresponding maps. Type localities of described taxa are indicated by stars. For T. Ʋerrucosus, we highlighted the paraphyletic position of sequences attributed to T. shanjing, as well as its type locality with asterisks." figureDoi="http://doi.org/10.5281/zenodo.7695343" httpUri="https://zenodo.org/record/7695343/files/figure.png" pageId="13" pageNumber="633">Fig. 2</figureCitation>
).
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection lastPageId="14" lastPageNumber="634" pageId="13" pageNumber="633" type="description">
<paragraph blockId="13.[809,1425,1426,1908]" lastBlockId="14.[163,780,1722,1806]" lastPageId="14" lastPageNumber="634" pageId="13" pageNumber="633">
<emphasis box="[809,1114,1426,1448]" italics="true" pageId="13" pageNumber="633">
Description of the 
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:
</emphasis>
Good state of preservation in spite of the torn ventral cavity (
<figureCitation box="[1217,1283,1457,1479]" captionStart="Figure 9" captionStartId="14.[163,243,1616,1638]" captionTargetBox="[338,1260,195,1563]" captionTargetId="figure-103@14.[348,1267,697,1577]" captionTargetPageId="14" captionText="Figure 9. Appearance of Echinotriton (Echinotriton) raffaellii sp. nov. Top: live individual observed at the type locality (credits: AH); bottom: the holotype MVZ:Herp:232187 (credits: MVZ)." figureDoi="http://doi.org/10.5281/zenodo.7695361" httpUri="https://zenodo.org/record/7695361/files/figure.png" pageId="13" pageNumber="633">Fig. 9</figureCitation>
). Slim newt (TTL: 
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) with a strongly granulated skin and a laterally compressed tail (TAL: 46.0 mm), 1.5 × times shorter than the body (SVL: 
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); distinctive triangular head (HW: 
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, HL: 
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, HD: 
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), much larger than body (CW: 
<quantity box="[1248,1348,1610,1631]" metricMagnitude="-2" metricUnit="m" metricValue="1.23" pageId="13" pageNumber="633" unit="mm" value="12.3">12.3 mm</quantity>
); short snout (SL: 
<quantity box="[936,1025,1641,1663]" metricMagnitude="-3" metricUnit="m" metricValue="6.1" pageId="13" pageNumber="633" unit="mm" value="6.1">6.1 mm</quantity>
), with nostrils slightly closer from each other’s (IN: 
<quantity box="[1019,1111,1672,1694]" metricMagnitude="-3" metricUnit="m" metricValue="4.6" pageId="13" pageNumber="633" unit="mm" value="4.6">4.6 mm</quantity>
) than from the eyes (ON: 
<quantity box="[809,900,1702,1724]" metricMagnitude="-3" metricUnit="m" metricValue="5.1" pageId="13" pageNumber="633" unit="mm" value="5.1">5.1 mm</quantity>
); large eyes (ED: 
<quantity box="[1110,1199,1702,1723]" metricMagnitude="-3" metricUnit="m" metricValue="4.4" pageId="13" pageNumber="633" unit="mm" value="4.4">4.4 mm</quantity>
) laterally disposed (IC: 
<quantity box="[863,956,1733,1754]" metricMagnitude="-3" metricUnit="m" metricValue="7.7" pageId="13" pageNumber="633" unit="mm" value="7.7">7.7 mm</quantity>
); vertebral ridge prominent; 12 pointy lateral glandular warts, disposed from axilla to the tail basis; forelimbs (AL: 
<quantity box="[1043,1140,1794,1815]" metricMagnitude="-2" metricUnit="m" metricValue="1.42" pageId="13" pageNumber="633" unit="mm" value="14.2">14.2 mm</quantity>
) of similar size than hind limbs (PL: 
<quantity box="[931,1030,1825,1846]" metricMagnitude="-2" metricUnit="m" metricValue="1.41" pageId="13" pageNumber="633" unit="mm" value="14.1">14.1 mm</quantity>
) and widely spaced (AX: 
<quantity box="[1313,1414,1825,1847]" metricMagnitude="-2" metricUnit="m" metricValue="3.86" pageId="13" pageNumber="633" unit="mm" value="38.6">38.6 mm</quantity>
); four fingers and five toes (fifth one rudimentary), with no visible webbing; cloaca distinct, with longitudinal vent slit; coloration includes a dark mate background, with orange colours only at the lateral warts, fingers and toes, the cloaca region and the underside of the tail.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7695361" ID-Zenodo-Dep="7695361" httpUri="https://zenodo.org/record/7695361/files/figure.png" pageId="14" pageNumber="634" startId="14.[163,243,1616,1638]" targetBox="[338,1260,195,1563]" targetPageId="14">
<paragraph blockId="14.[163,1443,1616,1668]" pageId="14" pageNumber="634">
<emphasis bold="true" box="[163,270,1616,1638]" pageId="14" pageNumber="634">Figure 9.</emphasis>
Appearance of 
<taxonomicName authority="Dufresnes &amp; Hernandez, 2023" authorityName="Dufresnes &amp; Hernandez" authorityYear="2023" box="[441,834,1617,1639]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="14" pageNumber="634" phylum="Chordata" rank="species" species="raffaellii" status="sp. nov." subGenus="Echinotriton">
<emphasis box="[441,834,1617,1639]" italics="true" pageId="14" pageNumber="634">Echinotriton (Echinotriton) raffaellii</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[841,920,1617,1639]" pageId="14" pageNumber="634" rank="species">sp. nov.</taxonomicNameLabel>
Top: live individual observed at the type locality (credits: AH); bottom: the holotype 
<emphasis box="[527,720,1646,1668]" italics="true" pageId="14" pageNumber="634">MVZ:Herp:232187</emphasis>
(credits: MVZ).
</paragraph>
</caption>
<paragraph blockId="14.[163,778,1846,1899]" lastBlockId="14.[827,1443,1722,1806]" pageId="14" pageNumber="634">
<materialsCitation collectorName="C. B. Fleck" location="Cantonal Museum of Zoology" municipality="Zoology of Lausanne" pageId="14" pageNumber="634" specimenCount="1" typeStatus="paratype">
<emphasis box="[163,276,1847,1868]" italics="true" pageId="14" pageNumber="634">
<typeStatus box="[163,270,1847,1868]" pageId="14" pageNumber="634">Paratype</typeStatus>
:
</emphasis>
MZL–46961, specimen born in captivity, descending from adults collected on Tokunoshima by 
<collectorName box="[863,992,1722,1744]" pageId="14" pageNumber="634">C. B. Fleck</collectorName>
and which identity was confirmed by 16S barcoding. Curated at the 
<location LSID="urn:lsid:plazi:treatment:C20D878E6878FFB165ED6565FCECF98A:4F7B6043687BFFAF61FE65E8FC71F892" municipality="Zoology of Lausanne" name="Cantonal Museum of Zoology" pageId="14" pageNumber="634">
Cantonal Museum of 
<collectingMunicipality box="[827,913,1784,1806]" pageId="14" pageNumber="634">Zoology</collectingMunicipality>
</location>
of Lausanne
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection lastPageId="16" lastPageNumber="636" pageId="14" pageNumber="634" type="etymology">
<paragraph blockId="14.[827,1443,1846,1899]" lastBlockId="16.[163,778,197,250]" lastPageId="16" lastPageNumber="636" pageId="14" pageNumber="634">
<emphasis box="[827,966,1846,1867]" italics="true" pageId="14" pageNumber="634">Etymology:</emphasis>
We coin the new name 
<taxonomicName authorityName="Dufresnes &amp; Hernandez" authorityYear="2023" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="14" pageNumber="634" phylum="Chordata" rank="species" species="raffaellii" subGenus="Echinotriton">
<emphasis italics="true" pageId="14" pageNumber="634">Echinotriton (Echinotriton) raffaellii</emphasis>
</taxonomicName>
as a tribute to Jean Raffaëlli, French naturalist and world expert of newts and salamanders.
</paragraph>
</subSubSection>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7695363" ID-Zenodo-Dep="7695363" httpUri="https://zenodo.org/record/7695363/files/figure.png" pageId="15" pageNumber="635" startId="15.[145,226,1775,1797]" targetBox="[147,1372,159,1736]" targetPageId="15">
<paragraph blockId="15.[145,1425,1775,1885]" pageId="15" pageNumber="635">
<emphasis bold="true" box="[145,267,1775,1797]" pageId="15" pageNumber="635">Figure 10.</emphasis>
Appearance of 
<taxonomicName authority="Dufresnes &amp; Hernandez, 2023" authorityName="Dufresnes &amp; Hernandez" authorityYear="2023" box="[442,776,1775,1797]" class="Amphibia" family="Salamandridae" genus="Tylototriton" kingdom="Animalia" order="Caudata" pageId="15" pageNumber="635" phylum="Chordata" rank="species" species="houi" status="sp. nov." subGenus="Tylototriton">
<emphasis box="[442,776,1775,1797]" italics="true" pageId="15" pageNumber="635">Tylototriton (Tylototriton) houi</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[784,864,1775,1797]" pageId="15" pageNumber="635" rank="species">sp. nov.</taxonomicNameLabel>
Top: live individual observed near the type locality (credits: Mian Hou); middle: the holotype 
<emphasis box="[591,714,1805,1827]" italics="true" pageId="15" pageNumber="635">MZL-46960</emphasis>
curated at the Cantonal Museum of Zoology of Lausanne (credits: CD); bottom: the type locality in Jade Dragon Snow Mountain (Hengduan massif in northern Yunnan) and observation of a larvae of the new species (credits: AH).
</paragraph>
</caption>
<subSubSection pageId="16" pageNumber="636" type="vernacular_names">
<paragraph blockId="16.[163,778,285,338]" pageId="16" pageNumber="636">
<emphasis box="[163,368,285,307]" italics="true" pageId="16" pageNumber="636">Common names:</emphasis>
Raffaëlli’s spiny crocodile newt (English), 
<taxonomicName authority="de Raffaelli" authorityName="de Raffaelli" box="[280,561,316,338]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="16" pageNumber="636" phylum="Chordata" rank="genus">Echinotriton de Raffaëlli</taxonomicName>
(French).
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="636" type="distribution">
<paragraph blockId="16.[163,779,378,676]" pageId="16" pageNumber="636">
<emphasis box="[163,779,378,400]" italics="true" pageId="16" pageNumber="636">
DiƲersity and distribution: 
<taxonomicName authorityName="Dufresnes &amp; Hernandez" authorityYear="2023" box="[479,779,378,400]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="16" pageNumber="636" phylum="Chordata" rank="species" species="raffaellii" subGenus="Echinotriton">Echinotriton (E.) raffaellii</taxonomicName>
</emphasis>
<materialsCitation country="Japan" county="Tokuno" location="Amami Island" municipality="Ryukyu Archipelago" pageId="16" pageNumber="636" specimenCount="1">
is only known from three islands of the 
<location LSID="urn:lsid:plazi:treatment:C20D878E6878FFB165ED6565FCECF98A:4F7B60436865FFB1673C62A8FCEAFE33" box="[616,778,409,431]" country="Japan" county="Tokuno" municipality="Ryukyu Archipelago" name="Amami Island" pageId="16" pageNumber="636">Amami Island</location>
group in the northern part of the 
<collectingMunicipality box="[542,774,440,462]" pageId="16" pageNumber="636">Ryukyu Archipelago</collectingMunicipality>
, 
<collectingCountry box="[163,238,471,492]" name="Japan" pageId="16" pageNumber="636">Japan</collectingCountry>
: 
<collectingCounty box="[250,338,470,492]" pageId="16" pageNumber="636">Tokuno</collectingCounty>
(the 
<typeStatus box="[404,455,471,492]" pageId="16" pageNumber="636">type</typeStatus>
locality), Amami and Uke. Genetic analyses suggest mitochondrial and nuclear differentiation between these populations (
<bibRefCitation author="Igawa T &amp; Sugawara H &amp; Honda M &amp; Tominaga A &amp; Oumi S &amp; Katsuren S &amp; Ota H &amp; Matsui M &amp; Sumida M" pageId="16" pageNumber="636" pagination="13 - 26" refId="ref12954" refString="Igawa T, Sugawara H, Honda M, Tominaga A, Oumi S, Katsuren S, Ota H, Matsui M, Sumida M. 2020. Detecting inter- and intra-island genetic diversity: population structure of the endangered crocodile newt, Echinotriton andersoni, in the Ryukyus. ConserVation Genetics 21: 13 - 26." type="journal article" year="2020">
Igawa 
<emphasis box="[163,224,562,584]" italics="true" pageId="16" pageNumber="636">et al.</emphasis>
, 2020
</bibRefCitation>
), reflecting recent disconnections after the disappearance of land bridges that connected the islands during the last glaciation (
<bibRefCitation author="Dufresnes C &amp; Litvinchuk SN" pageId="16" pageNumber="636" pagination="695 - 760" refId="ref12051" refString="Dufresnes C, Litvinchuk SN. 2022. Diversity, distribution and molecular species delimitation in frogs and toads from the Eastern Palearctic. Zoological Journal of the Linnean Society 195: 695 - 760." type="journal article" year="2022">Dufresnes &amp; Litvinchuk, 2022</bibRefCitation>
: fig. 45)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="636" type="biology_ecology">
<paragraph blockId="16.[163,779,712,1071]" pageId="16" pageNumber="636">
<emphasis box="[163,363,712,733]" italics="true" pageId="16" pageNumber="636">Natural history:</emphasis>
The species inhabits a mosaic of grasslands, forests and sugarcane plantations (
<quantity box="[684,779,743,764]" metricMagnitude="2" metricUnit="m" metricValue="1.885" metricValueMax="3.7399999999999998" metricValueMin="0.03" pageId="16" pageNumber="636" unit="m" value="188.5" valueMax="374.0" valueMin="3.0">3–374 m</quantity>
a.s.l.), where it is usually found near water bodies, hidden under rocks or in leaf litter. Breeding occurs from February to June and is exclusively terrestrial. During courtship, the male takes a semicircular posture while rubbing the flanks of the female with its head, then deposits a spermatophore on the ground that the female eventually picks up. Clutches of eggs are then spawned under pine needles, dead leaves and in humus close to ponds and small streams, where hatching larvae are carried away by heavy rains.
</paragraph>
</subSubSection>
<subSubSection pageId="16" pageNumber="636" type="conservation">
<paragraph blockId="16.[163,780,1106,1558]" pageId="16" pageNumber="636">
<emphasis box="[163,315,1107,1128]" italics="true" pageId="16" pageNumber="636">ConserƲation:</emphasis>
Japanese 
<taxonomicName box="[437,576,1106,1127]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="16" pageNumber="636" phylum="Chordata" rank="genus">
<emphasis box="[437,576,1106,1127]" italics="true" pageId="16" pageNumber="636">Echinotriton</emphasis>
</taxonomicName>
(as ‘ 
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1892" box="[625,766,1106,1128]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="16" pageNumber="636" phylum="Chordata" rank="species" species="andersoni">
<emphasis box="[625,766,1106,1128]" italics="true" pageId="16" pageNumber="636">E. andersoni</emphasis>
</taxonomicName>
’) are categorized as ‘Endangered’ (EN) class B 
<quantity box="[691,733,1137,1159]" metricMagnitude="-2" metricUnit="m" metricValue="2.54" pageId="16" pageNumber="636" unit="in" value="1.0">1 in</quantity>
the IUCN Red List and as ‘Vulnerable’ (VU) at the national level (
<bibRefCitation author="Environment Agency of Japan" box="[229,631,1198,1220]" pageId="16" pageNumber="636" refId="ref12374" refString="Environment Agency of Japan. 2000. Kankyo hakusho 1999. Vols 1 - 2. Tokyo: Environment Agency." type="book" year="2000">
Environment Agency of 
<collectingCountry box="[497,567,1199,1220]" name="Japan" pageId="16" pageNumber="636">Japan</collectingCountry>
, 2000
</bibRefCitation>
). The species further appears on CITES appendix II – international level (
<bibRefCitation author="CITES" box="[237,390,1260,1281]" pageId="16" pageNumber="636" refId="ref11932" refString="CITES. 2021. ConVention on International Trade in Endangered Species of aeild fauna and flora - Appendices I, II and III. Available at https: // cites. org / eng / app / appendices. php. Accessed November 2021." type="url" year="2021">CITES, 2021</bibRefCitation>
). The prefectures of 
<collectingRegion box="[645,778,1260,1282]" country="Japan" name="Kagoshima" pageId="16" pageNumber="636">Kagoshima</collectingRegion>
and 
<collectingRegion box="[213,316,1290,1312]" country="Japan" name="Okinawa" pageId="16" pageNumber="636">Okinawa</collectingRegion>
have afforded them the rank of ‘natural monument’. Hence, 
<taxonomicName authorityName="Dufresnes &amp; Hernandez" authorityYear="2023" box="[393,576,1321,1343]" class="Amphibia" family="Salamandridae" genus="Echinotriton" kingdom="Animalia" order="Caudata" pageId="16" pageNumber="636" phylum="Chordata" rank="species" species="raffaellii" subGenus="Echinotriton">
<emphasis box="[393,576,1321,1343]" italics="true" pageId="16" pageNumber="636">E. (E.) raffaellii</emphasis>
</taxonomicName>
is undoubtedly a rare and endangered species. The major threats include: international trafficking, which involves illegal poaching and exports to 
<collectingCountry box="[348,423,1413,1435]" name="China" pageId="16" pageNumber="636">China</collectingCountry>
, 
<collectingCountry box="[437,575,1413,1434]" name="Hong Kong" pageId="16" pageNumber="636">Hong Kong</collectingCountry>
, Europe and the 
<collectingCountry box="[163,318,1444,1466]" name="United States of America" pageId="16" pageNumber="636">United States</collectingCountry>
; predation by the invasive Java mongoose 
<emphasis box="[163,338,1475,1496]" italics="true" pageId="16" pageNumber="636">
<taxonomicName authorityName="Hodgson" authorityYear="1841" box="[163,224,1475,1496]" class="Mammalia" family="Herpestidae" genus="Urva" higherTaxonomySource="GBIF" kingdom="Animalia" order="Carnivora" pageId="16" pageNumber="636" phylum="Chordata" rank="genus">UrƲa</taxonomicName>
jaƲanica
</emphasis>
(Geoffroy Saint-Hilaire, 1818); and deforestation of the Ryukyus for sugarcane plantations and urbanization since the 1960s (especially on Tokuno).
</paragraph>
</subSubSection>
</treatment>
</document>