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<mods:title id="758C8798B196188C7C630CAC80D49FF5">Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand</mods:title>
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<mods:affiliation id="6B30DFAC2B577CF2DC6DE7C27454B6C6">Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand somsak. pan @ chula. ac. th</mods:affiliation>
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<taxonomicName id="0DC53FBAA4614B5FFD3E81D76768BC4B" ID-CoL="7TBDM" ID-ENA="2729312" authority="(Panha, 1997)" baseAuthorityName="Panha" baseAuthorityYear="1997" box="[676,1049,495,520]" class="Gastropoda" family="Ariophantidae" genus="Taphrenalla" kingdom="Animalia" order="Stylommatophora" pageId="28" pageNumber="49" phylum="Mollusca" rank="species" species="asamurai">
<emphasis id="F8B1982BA4614B5FFD3E81D760FDBC4B" bold="true" box="[676,908,495,520]" italics="true" pageId="28" pageNumber="49">Taphrenalla asamurai</emphasis>
(Panha, 1997)
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<paragraph id="CA7A4439A4614B5FFD3E822C60BABC6F" blockId="28.[676,1049,495,556]" box="[676,971,532,556]" pageId="28" pageNumber="49">
(
<figureCitation id="52FE58BCA4614B5FFD30822C61ACBC6F" box="[682,733,532,556]" captionStart="Figure 1" captionStartId="3.[98,166,1062,1084]" captionTargetBox="[102,1153,164,1043]" captionTargetId="figure-208@3.[102,1153,164,1043]" captionTargetPageId="3" captionText="Figure 1 Map showing the sampling sites of the 11 Taphrenalla species along Southern Thailand.Each colour of the distribution range follows the phylogenetic tree of all Taphrenalla species in fig.3." figureDoi="http://doi.org/10.5281/zenodo.8343291" httpUri="https://zenodo.org/record/8343291/files/figure.png" pageId="28" pageNumber="49">figs 1</figureCitation>
,
<figureCitation id="52FE58BCA4614B5FFD7D822C6182BC6F" box="[743,755,532,556]" captionStart="Figure 3" captionStartId="13.[98,166,1547,1569]" captionTargetBox="[103,1152,168,1525]" captionTargetId="figure-2@13.[99,1155,164,1528]" captionTargetPageId="13" captionText="Figure 3 A representative ML tree showing only the Taphrenalla clade.The synopsis of the radial groove on shell sculpture, genitalia showing location of dart apparatus, and live specimens of Taphrenalla spp. Results of species delimitation analyses based on the COI gene using ABGD and PTP.Black circles represent nodes with both ML (when Ē 70%) and BI (when Ē 0.95) support.White circles represent nodes with ML BS support but no BI PP support (when &lt;0.95). Each clade colour refers to the distribution range in fig. 1." figureDoi="http://doi.org/10.5281/zenodo.8343295" httpUri="https://zenodo.org/record/8343295/files/figure.png" pageId="28" pageNumber="49">3</figureCitation>
, 5, 6A, 7AC, table 6)
</paragraph>
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<paragraph id="CA7A4439A4614B5FFD3E8264600ABCF8" blockId="28.[676,1187,603,843]" pageId="28" pageNumber="49">
<taxonomicName id="0DC53FBAA4614B5FFD3E826467D1BC30" ID-CoL="7RZKX" authority="Panha, 1997: 101 - 105" authorityName="Panha" authorityPageNumber="101 - 105" authorityYear="1997" box="[676,1184,603,628]" class="Gastropoda" family="Ariophantidae" genus="Macrochlamys" kingdom="Animalia" order="Stylommatophora" pageId="28" pageNumber="49" phylum="Mollusca" rank="species" species="asamurai">
<emphasis id="F8B1982BA4614B5FFD3E826460D6BC30" box="[676,935,604,628]" italics="true" pageId="28" pageNumber="49">Macrochlamys asamurai</emphasis>
Panha, 1997: 101105
</taxonomicName>
,
<figureCitation id="52FE58BCA4614B5FFD3E82476068BCD4" box="[676,793,639,664]" captionStart="Figure 2" captionStartId="12.[129,197,1607,1629]" captionTargetBox="[135,1179,167,1586]" captionTargetId="figure-1@12.[132,1183,164,1588]" captionTargetPageId="12" captionText="Figure 2 Representative ML tree of the Taphrenalla gen.nov., Macrochlamys, Sarika, Cryptozona, Hemiplecta, Megaustenia, and Durgella based on the combined partial sequences of the COI, 16S, and 28S genes. The code in the parenthesis next to the species name refers to voucher numbers of the analysed samples.Numbers on the branches indicate the ML BS and BI PP values." figureDoi="http://doi.org/10.5281/zenodo.8343293" httpUri="https://zenodo.org/record/8343293/files/figure.png" pageId="28" pageNumber="49">figs 2a, 2b</figureCitation>
<figureCitation id="52FE58BCA4614B5FFC8082476053BCD4" box="[794,802,639,663]" captionStart="Figure 1" captionStartId="3.[98,166,1062,1084]" captionTargetBox="[102,1153,164,1043]" captionTargetId="figure-208@3.[102,1153,164,1043]" captionTargetPageId="3" captionText="Figure 1 Map showing the sampling sites of the 11 Taphrenalla species along Southern Thailand.Each colour of the distribution range follows the phylogenetic tree of all Taphrenalla species in fig.3." figureDoi="http://doi.org/10.5281/zenodo.8343291" httpUri="https://zenodo.org/record/8343291/files/figure.png" pageId="28" pageNumber="49">1</figureCitation>
,
<figureCitation id="52FE58BCA4614B5FFCAE82B8604EBCDB" box="[820,831,640,664]" captionStart="Figure 3" captionStartId="13.[98,166,1547,1569]" captionTargetBox="[103,1152,168,1525]" captionTargetId="figure-2@13.[99,1155,164,1528]" captionTargetPageId="13" captionText="Figure 3 A representative ML tree showing only the Taphrenalla clade.The synopsis of the radial groove on shell sculpture, genitalia showing location of dart apparatus, and live specimens of Taphrenalla spp. Results of species delimitation analyses based on the COI gene using ABGD and PTP.Black circles represent nodes with both ML (when Ē 70%) and BI (when Ē 0.95) support.White circles represent nodes with ML BS support but no BI PP support (when &lt;0.95). Each clade colour refers to the distribution range in fig. 1." figureDoi="http://doi.org/10.5281/zenodo.8343295" httpUri="https://zenodo.org/record/8343295/files/figure.png" pageId="28" pageNumber="49">3</figureCitation>
4.
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locality: Takun Village,
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,
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.
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<taxonomicName id="0DC53FBAA4614B5FFD3E82FF67EEBC9C" ID-CoL="7RZKX" authority=": Maneevong, 2000: 26 - 27" authorityName="Maneevong" authorityPageNumber="26 - 27" authorityYear="2000" box="[676,1183,711,736]" class="Gastropoda" family="Ariophantidae" genus="Macrochlamys" kingdom="Animalia" order="Stylommatophora" pageId="28" pageNumber="49" phylum="Mollusca" rank="species" species="asamurai">
<emphasis id="F8B1982BA4614B5FFD3E82FF60EBBC9C" box="[676,922,711,735]" italics="true" pageId="28" pageNumber="49">Macrochlamys asamurai</emphasis>
: Maneevong, 2000: 2627
</taxonomicName>
, fig. 4-1.
</paragraph>
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<taxonomicName id="0DC53FBAA4614B5FFD3E833767EEBD64" ID-CoL="7T3F2" authority=": Sutcharit &amp; Panha, 2008: 95 - 100" authorityName="Sutcharit &amp; Panha" authorityPageNumber="95 - 100" authorityYear="2008" box="[676,1183,783,807]" class="Gastropoda" family="Ariophantidae" genus="Sarika" kingdom="Animalia" order="Stylommatophora" pageId="28" pageNumber="49" phylum="Mollusca" rank="species" species="asamurai">
<emphasis id="F8B1982BA4614B5FFD3E83376038BD64" box="[676,841,783,807]" italics="true" pageId="28" pageNumber="49">Sarika asamurai</emphasis>
: Sutcharit &amp; Panha, 2008: 95100
</taxonomicName>
,
<figureCitation id="52FE58BCA4614B5FFD3E830B618ABD08" box="[676,763,819,843]" captionStart="Figure 2" captionStartId="12.[129,197,1607,1629]" captionTargetBox="[135,1179,167,1586]" captionTargetId="figure-1@12.[132,1183,164,1588]" captionTargetPageId="12" captionText="Figure 2 Representative ML tree of the Taphrenalla gen.nov., Macrochlamys, Sarika, Cryptozona, Hemiplecta, Megaustenia, and Durgella based on the combined partial sequences of the COI, 16S, and 28S genes. The code in the parenthesis next to the species name refers to voucher numbers of the analysed samples.Numbers on the branches indicate the ML BS and BI PP values." figureDoi="http://doi.org/10.5281/zenodo.8343293" httpUri="https://zenodo.org/record/8343293/files/figure.png" pageId="28" pageNumber="49">figs 2c, d</figureCitation>
,
<figureCitation id="52FE58BCA4614B5FFC9F830B605DBD08" box="[773,812,819,843]" captionStart="Figure 3" captionStartId="13.[98,166,1547,1569]" captionTargetBox="[103,1152,168,1525]" captionTargetId="figure-2@13.[99,1155,164,1528]" captionTargetPageId="13" captionText="Figure 3 A representative ML tree showing only the Taphrenalla clade.The synopsis of the radial groove on shell sculpture, genitalia showing location of dart apparatus, and live specimens of Taphrenalla spp. Results of species delimitation analyses based on the COI gene using ABGD and PTP.Black circles represent nodes with both ML (when Ē 70%) and BI (when Ē 0.95) support.White circles represent nodes with ML BS support but no BI PP support (when &lt;0.95). Each clade colour refers to the distribution range in fig. 1." figureDoi="http://doi.org/10.5281/zenodo.8343295" httpUri="https://zenodo.org/record/8343295/files/figure.png" pageId="28" pageNumber="49">3fi</figureCitation>
, 4df.
</paragraph>
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<emphasis id="F8B1982BA4614B5FFD3E83436018BDD0" box="[676,873,891,915]" italics="true" pageId="28" pageNumber="49">Material examined.</emphasis>
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material: not examined. Non-type material: Wat Tham Wararam,
<collectingRegion id="08018ADBA4614B5FFBD783A76183BD99" pageId="28" pageNumber="49">Phanom District</collectingRegion>
,
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<collectingRegion id="08018ADBA4614B5FFCA483FA600BBD99" box="[830,890,962,986]" pageId="28" pageNumber="49">Thani</collectingRegion>
(98°5307.3”N 98°4001.0”E):
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7153 (fig. 6A; 26 shells and
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in ethanol)
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,
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<collectionCode id="ACD4DCFCA4614B5FFC928432603DBA61" box="[776,844,1034,1058]" collectionName="United Kingdom, Cambridge, University, Museum of Zoology" pageId="28" pageNumber="49">CUMZ</collectionCode>
7154 (
<specimenCount id="DCC38FB0A4614B5FFC1584326750BA61" box="[911,1057,1034,1058]" pageId="28" pageNumber="49" type="generic">six specimens</specimenCount>
in ethanol)
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,
<materialsCitation id="7AAD4E64A4614B5FFD3E841661BEBA29" ID-GBIF-Occurrence="4412265342" collectionCode="CUMZ" country="India" pageId="28" pageNumber="49" stateProvince="Thani">
<collectionCode id="ACD4DCFCA4614B5FFD3E84166199BA05" box="[676,744,1070,1094]" collectionName="United Kingdom, Cambridge, University, Museum of Zoology" pageId="28" pageNumber="49">CUMZ</collectionCode>
7155 (14 shells and
<specimenCount id="DCC38FB0A4614B5FFC308416673EBA05" box="[938,1103,1070,1094]" pageId="28" pageNumber="49" type="generic">eight specimens</specimenCount>
in ethanol)
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,
<materialsCitation id="7AAD4E64A4614B5FFD43846A608FBA29" ID-GBIF-Occurrence="4412265406" box="[729,1022,1106,1130]" collectionCode="CUMZ" country="India" pageId="28" pageNumber="49" stateProvince="Thani">
and
<collectionCode id="ACD4DCFCA4614B5FFC9C846A603BBA29" box="[774,842,1106,1130]" collectionName="United Kingdom, Cambridge, University, Museum of Zoology" pageId="28" pageNumber="49">CUMZ</collectionCode>
7156 (four shells)
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.
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Limestone outcrops at Khlong Sok,
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(
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):
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7157 (three shells)
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and
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7158 (one shell)
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.
<materialsCitation id="7AAD4E64A4614B5FFBAD848660D3BAB9" ID-GBIF-Occurrence="4412265351" country="India" pageId="28" pageNumber="49" stateProvince="Phanom District">Limestone outcrops at Khlong Sok</materialsCitation>
,
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,
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(
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<geoCoordinate id="AFF122FEA4614B5FFC1B853E6773BB5D" box="[897,1026,1286,1310]" degrees="98" direction="east" minutes="44" orientation="longitude" pageId="28" pageNumber="49" precision="1" seconds="18.0" value="98.73833">98°4418.0”E</geoCoordinate>
)
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:
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<collectionCode id="ACD4DCFCA4614B5FFBBA853E6715BB5D" box="[1056,1124,1286,1310]" collectionName="United Kingdom, Cambridge, University, Museum of Zoology" pageId="28" pageNumber="49">CUMZ</collectionCode>
7159 (31 shells and
<specimenCount id="DCC38FB0A4614B5FFCD7851260ADBB02" box="[845,988,1321,1346]" pageId="28" pageNumber="49" type="generic">12 specimens</specimenCount>
in ethanol).
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Wat Tham Phanthurat,
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,
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<collectingRegion id="08018ADBA4614B5FFBFD857567D2BB26" box="[1127,1187,1357,1381]" pageId="28" pageNumber="49">Thani</collectingRegion>
(
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<geoCoordinate id="AFF122FEA4614B5FFCB9854960D0BBCA" box="[803,929,1393,1417]" degrees="98" direction="east" minutes="31" orientation="longitude" pageId="28" pageNumber="49" precision="1" seconds="30.2" value="98.52506">98°3130.2”E</geoCoordinate>
):
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7160 (
<specimenCount id="DCC38FB0A4614B5FFBAB85496193BBEE" pageId="28" pageNumber="49" type="generic">three specimens</specimenCount>
in ethanol)
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.
</paragraph>
</subSubSection>
<subSubSection id="82DF17B2A4614B5DFD3E85E56132BAB8" lastPageId="30" lastPageNumber="51" pageId="28" pageNumber="49" type="discussion">
<paragraph id="CA7A4439A4614B5DFD3E85E56132BAB8" blockId="28.[676,1187,1501,1705]" lastBlockId="30.[129,640,1035,1275]" lastPageId="30" lastPageNumber="51" pageId="28" pageNumber="49">
<emphasis id="F8B1982BA4614B5FFD3E85E5609EBBB6" box="[676,1007,1501,1525]" italics="true" pageId="28" pageNumber="49">
Remarks.
<taxonomicName id="0DC53FBAA4614B5FFC9385E5609EBBB6" baseAuthorityName="Panha" baseAuthorityYear="1997" box="[777,1007,1501,1525]" class="Gastropoda" family="Ariophantidae" genus="Taphrenalla" kingdom="Animalia" order="Stylommatophora" pageId="28" pageNumber="49" phylum="Mollusca" rank="species" species="asamurai">Taphrenalla asamurai</taxonomicName>
</emphasis>
had been re-described and illustrated by Sutcharit &amp; Panha (2008). The unique characters are a depressed conic shell of medium size (width up to
<quantity id="0D3DE9DCA4614B5FFBD6867167EEB823" box="[1100,1183,1609,1633]" metricMagnitude="-2" metricUnit="m" metricValue="2.11" pageId="28" pageNumber="49" unit="mm" value="21.1">21.1 mm</quantity>
; height up to
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), deep and narrow impressed suture, and long radial grooves that appeared on the last 3764 grooves (n = 32; figs 5, 6A). The genitalia are bent at the epiphallic caecum, and the proximal part of dart apparatus is with a thick and swollen sac-like structure (figs 7A, 7B). Spermatophore has one spine on the tail filament close to the sperm sac (fig. 7C). Living snails have monochrome red to dark red on tail (
<figureCitation id="52FE58BCA4634B5DFD9884DA6147BAB8" box="[514,566,1250,1275]" captionStart="Figure 3" captionStartId="13.[98,166,1547,1569]" captionTargetBox="[103,1152,168,1525]" captionTargetId="figure-2@13.[99,1155,164,1528]" captionTargetPageId="13" captionText="Figure 3 A representative ML tree showing only the Taphrenalla clade.The synopsis of the radial groove on shell sculpture, genitalia showing location of dart apparatus, and live specimens of Taphrenalla spp. Results of species delimitation analyses based on the COI gene using ABGD and PTP.Black circles represent nodes with both ML (when Ē 70%) and BI (when Ē 0.95) support.White circles represent nodes with ML BS support but no BI PP support (when &lt;0.95). Each clade colour refers to the distribution range in fig. 1." figureDoi="http://doi.org/10.5281/zenodo.8343295" httpUri="https://zenodo.org/record/8343295/files/figure.png" pageId="30" pageNumber="51">fig. 3</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>