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<document id="96DD35ECD7CDD0D53B988BF2056914D1" ID-CLB-Dataset="4210" ID-DOI="10.5852/ejt.2021.767.1495" ID-GBIF-Dataset="67bc6a73-4137-4de3-9dc4-ce87972f742d" ID-ISSN="2118-9773" ID-Zenodo-Dep="5514839" ID-ZooBank="9F019E02-95E7-4B6A-B5BC-01F8E26F3466" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1631927261920" checkinUser="felipe" docAuthor="Nahok, Benchawan, Tumpeesuwan, Sakboworn &amp; Tumpeesuwan, Chanidaporn" docDate="2021" docId="03AC87E7C325FFC29F7FFEE391E1BC45" docLanguage="en" docName="ejt-767_142-166.pdf" docOrigin="European Journal of Taxonomy 767 (1)" docSource="http://dx.doi.org/10.5852/ejt.2021.767.1495" docStyle="DocumentStyle:EF2B578F1D15862ADE45B0C07C620911.14:EJT.2018-.journal_article.type1" docStyleId="EF2B578F1D15862ADE45B0C07C620911" docStyleName="EJT.2018-.journal_article.type1" docStyleVersion="14" docTitle="Landouria tuberculata Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan 2021, sp. nov." docType="treatment" docUuid="84133944-D5C9-49A3-9997-8E1C7B335938" docUuidSource="ZooBank" docVersion="11" lastPageNumber="155" masterDocId="FF95FF9FC32EFFCF9D1FFF9B931BBE42" masterDocTitle="Five new species of the land snail genus Landouria Godwin-Austen, 1918 (Gastropoda, Camaenidae) from northeastern Thailand, with note on genitalia and radula morphology of Landouria diplogramma (Möllendorff, 1902)" masterLastPageNumber="166" masterPageNumber="142" pageNumber="153" updateTime="1699449512379" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0" zenodo-license-treatments="UNSPECIFIED">
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<mods:title id="B280BBCCFF684E705A814C920DF75A12">Five new species of the land snail genus Landouria Godwin-Austen, 1918 (Gastropoda, Camaenidae) from northeastern Thailand, with note on genitalia and radula morphology of Landouria diplogramma (Möllendorff, 1902)</mods:title>
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<mods:namePart id="CE93ECA69DB070A073658E496AEADFFA">Nahok, Benchawan</mods:namePart>
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<mods:affiliation id="21254FABE632290C2A134F0B80376079">Department of Biology, Faculty of Science, Mahasarakham University, Kantharawichai District, Maha Sarakham, 44150 Thailand. &amp; Educational Research Development and Demonstration Institute, Srinakharinwirot University, Ongkharak District, Nakhon Nayok, 26120 Thailand.</mods:affiliation>
<mods:nameIdentifier id="FB0919F73AF0FC8793BE1CA04F7BE47F" type="email">nahok.b@gmail.com</mods:nameIdentifier>
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<mods:namePart id="FB9F01ADBDC3B0CEF31E224549AADA3B">Tumpeesuwan, Sakboworn</mods:namePart>
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<mods:affiliation id="21325CFB8288D2CF26C457D983E8568B">Department of Biology, Faculty of Science, Mahasarakham University, Kantharawichai District, Maha Sarakham, 44150 Thailand. &amp; Palaeontological Research and Education Centre, Mahasarakham University, Kantharawichai District, Maha Sarakham, 44150 Thailand.</mods:affiliation>
<mods:nameIdentifier id="1EA988FB6440DEDD2031B234B7BD1081" type="email">stumpeesuwan@yahoo.com</mods:nameIdentifier>
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<mods:namePart id="E35C7E2CE68318F8F8761307185DC3E3">Tumpeesuwan, Chanidaporn</mods:namePart>
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<mods:affiliation id="4002AE34226EA99769B6445053A9C8AA">Department of Biology, Faculty of Science, Mahasarakham University, Kantharawichai District, Maha Sarakham, 44150 Thailand.</mods:affiliation>
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<mods:title id="C212A1B194B7BC213D8D9F5018048506">European Journal of Taxonomy</mods:title>
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<treatment id="03AC87E7C325FFC29F7FFEE391E1BC45" ID-DOI="http://doi.org/10.5281/zenodo.5514729" ID-GBIF-Taxon="188498238" ID-Zenodo-Dep="5514729" LSID="urn:lsid:zoobank.org:act:84133944-D5C9-49A3-9997-8E1C7B335938" httpUri="http://treatment.plazi.org/id/03AC87E7C325FFC29F7FFEE391E1BC45" lastPageId="13" lastPageNumber="155" pageId="11" pageNumber="153">
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<taxonomicName id="4C054D72C325FFC49F7FFEE39068BFD1" ID-CoL="8QGJ8" authority="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan, 2021" authorityName="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan" authorityYear="2021" box="[608,883,376,403]" class="Gastropoda" family="Camaenidae" genus="Landouria" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="153" phylum="Mollusca" rank="species" species="tuberculata" status="sp. nov.">
<emphasis id="B971EAE3C325FFC49F7FFEE39068BFD1" bold="true" box="[608,883,376,403]" italics="true" pageId="11" pageNumber="153">Landouria tuberculata</emphasis>
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<taxonomicNameLabel id="A2425798C325FFC49E65FEE290CFBFD1" box="[890,980,377,403]" pageId="11" pageNumber="153" rank="species">sp. nov.</taxonomicNameLabel>
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<paragraph id="8BBA36F1C325FFC49F54FE2490F3BF9B" blockId="11.[377,1210,376,474]" box="[587,1000,447,474]" pageId="11" pageNumber="153">
<figureCitation id="133E2A74C325FFC49F54FE2491B2BF9B" box="[587,681,447,473]" captionStart="Fig" captionStartId="9.[189,232,1849,1875]" captionTargetBox="[220,1368,265,1811]" captionTargetId="figure-16@9.[220,1368,265,1811]" captionTargetPageId="9" captionText="Fig. 3. External shell morphology of Landouria spp. from northeastern Thailand. A. Landouria circinata sp. nov., holotype (NHMSU-00023). B. L. tuberculata sp. nov., holotype (NHMSU-00025). C. L. trochomorphoides sp. nov., holotype (NHMSU-00027). D. L. chloritoides sp. nov., holotype (NHMSU-00029). E. L. elegans sp. nov., holotype (NHMSU-00032). F. L. diplogramma (Möllendorff, 1902) comb. nov. (NHMSU-00034). Scale bar = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5514845" httpUri="https://zenodo.org/record/5514845/files/figure.png" pageId="11" pageNumber="153">Figs 3B</figureCitation>
,
<figureCitation id="133E2A74C325FFC49FA9FE2491C1BF9B" box="[694,730,447,473]" captionStart="Fig" captionStartId="12.[189,232,1921,1947]" captionTargetBox="[193,1395,278,1870]" captionTargetId="figure-12@12.[193,1395,265,1883]" captionTargetPageId="12" captionText="Fig. 4. Living adults of Landouria spp. from northeastern Thailand. A. Landouria circinata sp. nov. B. L. tuberculata sp. nov. C. L. trochomorphoides sp. nov. D. L. chloritoides sp. nov. E. L. elegans sp. nov. F. L. diplogramma (Möllendorff, 1902) comb. nov. Scale bars = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5514847" httpUri="https://zenodo.org/record/5514847/files/figure.png" pageId="11" pageNumber="153">4B</figureCitation>
,
<figureCitation id="133E2A74C325FFC49FF9FE249011BF98" box="[742,778,447,474]" captionStart="Fig" captionStartId="15.[189,232,1816,1842]" captionTargetBox="[283,1352,278,1767]" captionTargetId="figure-15@15.[232,1356,265,1778]" captionTargetPageId="15" captionText="Fig. 5. Illustrations of the genital system of Landouria spp. from northeastern Thailand. A. Landouria circinata sp. nov., paratype (NHMSU-00024). B. L. tuberculata sp. nov., paratype (NHMSU-00026). C. L. trochomorphoides sp. nov., paratype (NHMSU-00028). D. L. chloritoides sp. nov., paratype (NHMSU-00030). E. L. elegans sp. nov., paratype (NHMSU-00033). F. L. diplogramma (Möllendorff, 1902) comb. nov. (NHMSU-00034). Internal sculpture of a = flagellum; b = penis; c = vagina. Scale bars = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5514849" httpUri="https://zenodo.org/record/5514849/files/figure.png" pageId="11" pageNumber="153">5B</figureCitation>
,
<figureCitation id="133E2A74C325FFC49E08FE249045BF9B" box="[791,862,447,473]" captionStart="Fig" captionStartId="18.[189,232,1783,1809]" captionTargetBox="[244,1343,265,1744]" captionTargetId="figure-12@18.[244,1343,265,1744]" captionTargetPageId="18" captionText="Fig. 6.Radula morphology ofLandouria spp. from northeasternThailand.AC, GR. Right side of central teeth. DF. Left side of central teeth. AC. Landouria circinata sp. nov., paratype (NHMSU-00024). DF. L. tuberculata sp. nov., paratype (NHMSU-00026). GI. L. trochomorphoides sp. nov., paratype (NHMSU-00028). JL. L. chloritoides sp. nov., paratype (NHMSU-00030). MO. L. elegans sp. nov., paratype (NHMSU-00033). PR. L. diplogramma (Möllendorff, 1902) comb. nov. (NHMSU-00034). Numbers indicate order of lateral and marginal teeth. Abbreviation: c = central tooth. Scale bars = 25 µm." figureDoi="http://doi.org/10.5281/zenodo.5514851" httpUri="https://zenodo.org/record/5514851/files/figure.png" pageId="11" pageNumber="153">6DF</figureCitation>
;
<tableCitation id="C687034AC325FFC49E76FE2490F3BF9B" box="[873,1000,447,473]" captionStart-0="Table 1" captionStart-1="Table 2" captionStart-2="Table 3" captionStart-3="Table 4" captionStartId-0="2.[185,255,857,885]" captionStartId-1="4.[186,256,600,628]" captionStartId-2="6.[185,255,695,723]" captionStartId-3="10.[185,255,544,572]" captionTargetBox-0="[195,1921,987,1444]" captionTargetBox-1="[197,1907,827,1685]" captionTargetBox-2="[212,1909,868,1619]" captionTargetBox-3="[196,1908,758,1739]" captionTargetPageId-0="2" captionTargetPageId-1="4" captionTargetPageId-2="6" captionTargetPageId-3="10" captionText-0="Table 1. Shell measurements (mm) of six new species of the genus Landouria Godwin-Austen, 1918 from northeastern Thailand." captionText-1="Table 2. Locality data and identification of specimens of Landouria Godwin-Austen, 1918 used for mtDNA sequencing, including collection catalogue numbers and GenBank accession numbers. Abbreviations: NHMSU = Natural History Museum, Mahasarakham University; HNHM = Hungarian Natural History Museum; AM = Australian Museum. References: 1 = Tumpeesuwan &amp; Tumpeesuwan (2019); 2 = Hirano et al. (2014); 3 = Köhler et al. (2018)." captionText-2="Table 3. Pairwise p-distance values of Landouria Godwin-Austen, 1918 based on mitochondrial COI sequences." captionText-3="Table 4. Comparison of shells, radulae, and genital systems of Landouria spp. from northeastern Thailand. References: 1 = Tumpeesuwan &amp; Tumpeesuwan (2019); 2 = Nahok (2020); 3 = this study. *In Tumpeesuwan &amp; Tumpeesuwan (2019) indicated the number of teeth per row in Landouria strobiloides varies from 3 to 53. This lowest row has only 3 teeth. It is the uppermost row, probably lacerated and peel off. Unfortunately, we could not re-checked the complete specimens in this time being, so, this point need further investigation and proof." httpUri-0="http://table.plazi.org/id/DF7A6679C32CFFCD9DA6FCC295B0BD37" httpUri-1="http://table.plazi.org/id/DF7A6679C32AFFCB9DA5FDC39249BC9F" httpUri-2="http://table.plazi.org/id/DF7A6679C328FFC99DA6FD2C96FFBC91" httpUri-3="http://table.plazi.org/id/DF7A6679C324FFC59DA6FDBB969CBCE7" pageId="11" pageNumber="153" tableUuid-0="DF7A6679C32CFFCD9DA6FCC295B0BD37" tableUuid-1="DF7A6679C32AFFCB9DA5FDC39249BC9F" tableUuid-2="DF7A6679C328FFC99DA6FD2C96FFBC91" tableUuid-3="DF7A6679C324FFC59DA6FDBB969CBCE7">Tables 14</tableCitation>
</paragraph>
</subSubSection>
<subSubSection id="C31F657AC325FFC49DA2FD9B92C8BCCD" pageId="11" pageNumber="153" type="diagnosis">
<paragraph id="8BBA36F1C325FFC49DA2FD9B922DBC58" blockId="11.[189,310,512,538]" box="[189,310,512,538]" pageId="11" pageNumber="153">
<heading id="D0F2819DC325FFC49DA2FD9B922DBC58" bold="true" box="[189,310,512,538]" fontSize="11" level="3" pageId="11" pageNumber="153" reason="3">
<emphasis id="B971EAE3C325FFC49DA2FD9B922DBC58" bold="true" box="[189,310,512,538]" pageId="11" pageNumber="153">Diagnosis</emphasis>
</heading>
</paragraph>
<paragraph id="8BBA36F1C325FFC49DA2FDB492C8BCCD" blockId="11.[189,1398,559,655]" pageId="11" pageNumber="153">
Shell moderately keeled, with numerous tiny tubercles on shell surface (
<figureCitation id="133E2A74C325FFC49929FDB4978DBC0B" box="[1078,1174,559,585]" captionStart="Fig" captionStartId="12.[189,232,1921,1947]" captionTargetBox="[193,1395,278,1870]" captionTargetId="figure-12@12.[193,1395,265,1883]" captionTargetPageId="12" captionText="Fig. 4. Living adults of Landouria spp. from northeastern Thailand. A. Landouria circinata sp. nov. B. L. tuberculata sp. nov. C. L. trochomorphoides sp. nov. D. L. chloritoides sp. nov. E. L. elegans sp. nov. F. L. diplogramma (Möllendorff, 1902) comb. nov. Scale bars = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5514847" httpUri="https://zenodo.org/record/5514847/files/figure.png" pageId="11" pageNumber="153">Fig. 4B</figureCitation>
). Flagellum like a short protrusion with a pointed tip, penis swollen, base of gametolytic sac narrow. Radula with slender, lanceolate central teeth.
</paragraph>
</subSubSection>
<subSubSection id="C31F657AC325FFC49DA2FD2C922BBD61" pageId="11" pageNumber="153" type="etymology">
<paragraph id="8BBA36F1C325FFC49DA2FD2C925FBC93" blockId="11.[189,324,695,721]" box="[189,324,695,721]" pageId="11" pageNumber="153">
<heading id="D0F2819DC325FFC49DA2FD2C925FBC93" bold="true" box="[189,324,695,721]" fontSize="11" level="3" pageId="11" pageNumber="153" reason="3">
<emphasis id="B971EAE3C325FFC49DA2FD2C925FBC93" bold="true" box="[189,324,695,721]" pageId="11" pageNumber="153">Etymology</emphasis>
</heading>
</paragraph>
<paragraph id="8BBA36F1C325FFC49DA2FD7E922BBD61" blockId="11.[189,1399,741,803]" pageId="11" pageNumber="153">
The specific epithet
<taxonomicName id="4C054D72C325FFC49CA0FD7E915EBCBD" authorityName="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan" authorityYear="2021" box="[447,581,741,767]" class="Gastropoda" family="Camaenidae" genus="Landouria" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="153" phylum="Mollusca" rank="species" species="tuberculata">
<emphasis id="B971EAE3C325FFC49CA0FD7E915EBCBD" box="[447,581,741,767]" italics="true" pageId="11" pageNumber="153">tuberculata</emphasis>
</taxonomicName>
refers to the shell sculpture consisting of numerous tiny tubercles (
<figureCitation id="133E2A74C325FFC49DD9FC939239BD61" box="[198,290,776,803]" captionStart="Fig" captionStartId="12.[189,232,1921,1947]" captionTargetBox="[193,1395,278,1870]" captionTargetId="figure-12@12.[193,1395,265,1883]" captionTargetPageId="12" captionText="Fig. 4. Living adults of Landouria spp. from northeastern Thailand. A. Landouria circinata sp. nov. B. L. tuberculata sp. nov. C. L. trochomorphoides sp. nov. D. L. chloritoides sp. nov. E. L. elegans sp. nov. F. L. diplogramma (Möllendorff, 1902) comb. nov. Scale bars = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5514847" httpUri="https://zenodo.org/record/5514847/files/figure.png" pageId="11" pageNumber="153">Fig. 4B</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C31F657AC325FFC49DA2FCD1915CBAC6" pageId="11" pageNumber="153" type="materials_examined">
<paragraph id="8BBA36F1C325FFC49DA2FCD19277BD26" blockId="11.[189,364,842,868]" box="[189,364,842,868]" pageId="11" pageNumber="153">
<heading id="D0F2819DC325FFC49DA2FCD19277BD26" bold="true" box="[189,364,842,868]" fontSize="11" level="3" pageId="11" pageNumber="153" reason="3">
<emphasis id="B971EAE3C325FFC49DA2FCD19277BD26" bold="true" box="[189,364,842,868]" pageId="11" pageNumber="153">Type material</emphasis>
</heading>
</paragraph>
<paragraph id="8BBA36F1C325FFC49DF5FCE3974FBDBE" blockId="11.[189,1399,888,1021]" pageId="11" pageNumber="153">
<materialsCitation id="3B6D3CACC325FFC49DF5FCE3974FBDBE" ID-GBIF-Occurrence="3357769305" collectingDate="2017-09-23" collectionCode="UW" collectorName="B. Nahok &amp; K. Tanmuangpak &amp; U. Chanlabut" country="Thailand" elevation="647" latitude="17.437525" location="Wat Thepnimit" longLatPrecision="1" longitude="101.27819" municipality="Phu Rua District" pageId="11" pageNumber="153" specimenCode="NHMSU-00025" specimenCount="1" stateProvince="Loei Province" typeStatus="holotype">
<emphasis id="B971EAE3C325FFC49DF5FCE39247BDD0" bold="true" box="[234,348,888,914]" pageId="11" pageNumber="153">
<typeStatus id="54BE8853C325FFC49DF5FCE39247BDD0" box="[234,348,888,914]" pageId="11" pageNumber="153" type="holotype">Holotype</typeStatus>
</emphasis>
<collectingCountry id="F3127661C325FFC49DA2FC07924EBDF4" box="[189,341,924,950]" name="Thailand" pageId="11" pageNumber="153">THAILAND</collectingCountry>
<specimenCount id="9D03FD78C325FFC49C72FC0092BFBDF7" box="[365,420,923,949]" pageId="11" pageNumber="153">shell</specimenCount>
(SH =
<quantity id="4CFD9B14C325FFC49CE7FC00914EBDF4" box="[504,597,923,950]" metricMagnitude="-3" metricUnit="m" metricValue="9.6" pageId="11" pageNumber="153" unit="mm" value="9.6">9.6 mm</quantity>
, SW =
<quantity id="4CFD9B14C325FFC49FB5FC07900EBDF4" box="[682,789,924,950]" metricMagnitude="-2" metricUnit="m" metricValue="1.44" pageId="11" pageNumber="153" unit="mm" value="14.4">14.4 mm</quantity>
, AH =
<quantity id="4CFD9B14C325FFC49E78FC0090D8BDF4" box="[871,963,923,950]" metricMagnitude="-3" metricUnit="m" metricValue="5.2" pageId="11" pageNumber="153" unit="mm" value="5.2">5.2 mm</quantity>
, AW =
<quantity id="4CFD9B14C325FFC49906FC00976EBDF4" box="[1049,1141,923,950]" metricMagnitude="-3" metricUnit="m" metricValue="5.5" pageId="11" pageNumber="153" unit="mm" value="5.5">5.5 mm</quantity>
, UW =
<quantity id="4CFD9B14C325FFC499D0FC009636BDF4" box="[1231,1325,923,950]" metricMagnitude="-3" metricUnit="m" metricValue="3.5" pageId="11" pageNumber="153" unit="mm" value="3.5">3.5 mm</quantity>
);
<collectingRegion id="49C1F813C325FFC4985EFC009233BD9B" country="Thailand" name="Loei" pageId="11" pageNumber="153">Loei Province</collectingRegion>
,
<collectingMunicipality id="6BDEAC8BC325FFC49C2CFC2492E3BD9B" box="[307,504,959,985]" pageId="11" pageNumber="153">Phu Rua District</collectingMunicipality>
,
<location id="8EDA602AC325FFC49F1DFC2491AFBD9B" LSID="urn:lsid:plazi:treatment:03AC87E7C325FFC29F7FFEE391E1BC45:8EDA602AC325FFC49F1DFC2491AFBD9B" box="[514,692,959,985]" country="Thailand" latitude="17.437525" longLatPrecision="1" longitude="101.27819" municipality="Phu Rua District" name="Wat Thepnimit" pageId="11" pageNumber="153" stateProvince="Loei Province">Wat Thepnimit</location>
;
<geoCoordinate id="EE315036C325FFC49FA0FC269063BD9B" box="[703,888,957,985]" degrees="17" direction="north" minutes="26" orientation="latitude" pageId="11" pageNumber="153" precision="1" seconds="15.09" value="17.437525">17°2615.09″ N</geoCoordinate>
,
<geoCoordinate id="EE315036C325FFC49E9CFC269753BD9B" box="[899,1096,957,985]" degrees="101" direction="east" minutes="16" orientation="longitude" pageId="11" pageNumber="153" precision="1" seconds="41.46" value="101.27819">101°1641.46″ E</geoCoordinate>
; alt.
<quantity id="4CFD9B14C325FFC49960FC2497D0BD9B" box="[1151,1227,959,985]" metricMagnitude="2" metricUnit="m" metricValue="6.47" pageId="11" pageNumber="153" unit="m" value="647.0">
<elevation id="0028D1C2C325FFC49960FC2497D0BD9B" box="[1151,1227,959,985]" metricMagnitude="2" metricUnit="m" metricValue="6.47" pageId="11" pageNumber="153" unit="m" value="647.0">647 m</elevation>
</quantity>
;
<date id="FFBB1031C325FFC499C9FC249669BD9B" box="[1238,1394,959,985]" pageId="11" pageNumber="153" value="2017-09-23">
<collectingDate id="EFFFE9D9C325FFC499C9FC249669BD9B" box="[1238,1394,959,985]" pageId="11" pageNumber="153" value="2017-09-23">23 Sep. 2017</collectingDate>
</date>
;
<collectorName id="26F05327C325FFC49DA2FC78922BBDBE" box="[189,304,994,1021]" pageId="11" pageNumber="153">B. Nahok</collectorName>
,
<collectorName id="26F05327C325FFC49C23FC78911EBDBE" box="[316,517,994,1021]" pageId="11" pageNumber="153">K. Tanmuangpak</collectorName>
and
<collectorName id="26F05327C325FFC49F21FC7891C0BDBE" box="[574,731,994,1021]" pageId="11" pageNumber="153">U. Chanlabut</collectorName>
leg.;
<specimenCode id="DBA39E8AC325FFC49E02FC7990C6BDBF" box="[797,989,994,1021]" pageId="11" pageNumber="153">NHMSU-00025</specimenCode>
. (
<figureCitation id="133E2A74C325FFC49EEEFC799750BDBE" box="[1009,1099,994,1020]" captionStart="Fig" captionStartId="9.[189,232,1849,1875]" captionTargetBox="[220,1368,265,1811]" captionTargetId="figure-16@9.[220,1368,265,1811]" captionTargetPageId="9" captionText="Fig. 3. External shell morphology of Landouria spp. from northeastern Thailand. A. Landouria circinata sp. nov., holotype (NHMSU-00023). B. L. tuberculata sp. nov., holotype (NHMSU-00025). C. L. trochomorphoides sp. nov., holotype (NHMSU-00027). D. L. chloritoides sp. nov., holotype (NHMSU-00029). E. L. elegans sp. nov., holotype (NHMSU-00032). F. L. diplogramma (Möllendorff, 1902) comb. nov. (NHMSU-00034). Scale bar = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5514845" httpUri="https://zenodo.org/record/5514845/files/figure.png" pageId="11" pageNumber="153">Fig. 3B</figureCitation>
)
</materialsCitation>
</paragraph>
<paragraph id="8BBA36F1C325FFC49DF5FBBF915CBAC6" blockId="11.[189,1398,1060,1157]" pageId="11" pageNumber="153">
<materialsCitation id="3B6D3CACC325FFC49DF5FBBF9158BAC6" ID-GBIF-Occurrence="3357769304" accessionNumber="MN449402 and MZ435747" collectingDate="2017-09-23" collectionCode="MN, MZ" collectorName="GenBank No." country="Thailand" elevation="647" latitude="17.437525" location="THAILAND" longLatPrecision="1" longitude="101.27819" municipality="Phu Rua District" pageId="11" pageNumber="153" specimenCode="NHMSU-00026" specimenCount="17" stateProvince="Loei" typeStatus="paratype">
<emphasis id="B971EAE3C325FFC49DF5FBBF927CBA7C" bold="true" box="[234,359,1060,1086]" pageId="11" pageNumber="153">
<typeStatus id="54BE8853C325FFC49DF5FBBF927CBA7C" box="[234,359,1060,1086]" pageId="11" pageNumber="153" type="paratype">Paratypes</typeStatus>
</emphasis>
<collectingCountry id="F3127661C325FFC49DA2FBDC924EBA23" box="[189,341,1095,1121]" name="Thailand" pageId="11" pageNumber="153">THAILAND</collectingCountry>
<specimenCount id="9D03FD78C325FFC49C72FBDC92CCBA23" box="[365,471,1095,1121]" pageId="11" pageNumber="153">10 shells</specimenCount>
,
<specimenCount id="9D03FD78C325FFC49CFDFBDC919CBA23" box="[482,647,1095,1121]" pageId="11" pageNumber="153">7 living specs</specimenCount>
; same collection data as for holotype; GenBank No.
<accessionNumber id="9456AB12C325FFC499F0FBDC966DBA23" box="[1263,1398,1095,1121]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/MN449402" pageId="11" pageNumber="153">MN449402</accessionNumber>
and
<accessionNumber id="9456AB12C325FFC49DF0FBF1926CBAC7" box="[239,375,1130,1157]" httpUri="https://www.ncbi.nlm.nih.gov/nuccore/MZ435747" pageId="11" pageNumber="153" type="GenBank">MZ435747</accessionNumber>
;
<specimenCode id="DBA39E8AC325FFC49C9DFBF19158BAC6" box="[386,579,1130,1156]" pageId="11" pageNumber="153">NHMSU-00026</specimenCode>
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C31F657AC325FFC49DA2FB3091FDB9CA" pageId="11" pageNumber="153" type="description">
<paragraph id="8BBA36F1C325FFC49DA2FB309255BA87" blockId="11.[189,334,1195,1221]" box="[189,334,1195,1221]" pageId="11" pageNumber="153">
<heading id="D0F2819DC325FFC49DA2FB309255BA87" bold="true" box="[189,334,1195,1221]" fontSize="11" level="3" pageId="11" pageNumber="153" reason="3">
<emphasis id="B971EAE3C325FFC49DA2FB309255BA87" bold="true" box="[189,334,1195,1221]" pageId="11" pageNumber="153">Description</emphasis>
</heading>
</paragraph>
<paragraph id="8BBA36F1C325FFC49DA2FB41922CBBC3" blockId="11.[188,1399,1240,1409]" pageId="11" pageNumber="153">
SHELL (
<figureCitation id="133E2A74C325FFC49C0CFB419275BAB6" box="[275,366,1242,1268]" captionStart="Fig" captionStartId="9.[189,232,1849,1875]" captionTargetBox="[220,1368,265,1811]" captionTargetId="figure-16@9.[220,1368,265,1811]" captionTargetPageId="9" captionText="Fig. 3. External shell morphology of Landouria spp. from northeastern Thailand. A. Landouria circinata sp. nov., holotype (NHMSU-00023). B. L. tuberculata sp. nov., holotype (NHMSU-00025). C. L. trochomorphoides sp. nov., holotype (NHMSU-00027). D. L. chloritoides sp. nov., holotype (NHMSU-00029). E. L. elegans sp. nov., holotype (NHMSU-00032). F. L. diplogramma (Möllendorff, 1902) comb. nov. (NHMSU-00034). Scale bar = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5514845" httpUri="https://zenodo.org/record/5514845/files/figure.png" pageId="11" pageNumber="153">Fig. 3B</figureCitation>
,
<tableCitation id="C687034AC325FFC49C66FB4192D4BAB6" box="[377,463,1242,1268]" captionStart="Table 1" captionStartId="2.[185,255,857,885]" captionTargetBox="[195,1921,987,1444]" captionTargetPageId="2" captionText="Table 1. Shell measurements (mm) of six new species of the genus Landouria Godwin-Austen, 1918 from northeastern Thailand." httpUri="http://table.plazi.org/id/DF7A6679C32CFFCD9DA6FCC295B0BD37" pageId="11" pageNumber="153" tableUuid="DF7A6679C32CFFCD9DA6FCC295B0BD37">Table 1</tableCitation>
). Dextral, medium-sized, tightly coiled, with 6½ slightly convex whorls. Suture rather shallow, apex obtuse with high conical spire. Body whorl angular, with tubercles all over its surface, without distinct incised spiral lines. Colour red brown in living specimens (
<figureCitation id="133E2A74C325FFC499A9FABB960EBB78" box="[1206,1301,1312,1338]" captionStart="Fig" captionStartId="12.[189,232,1921,1947]" captionTargetBox="[193,1395,278,1870]" captionTargetId="figure-12@12.[193,1395,265,1883]" captionTargetPageId="12" captionText="Fig. 4. Living adults of Landouria spp. from northeastern Thailand. A. Landouria circinata sp. nov. B. L. tuberculata sp. nov. C. L. trochomorphoides sp. nov. D. L. chloritoides sp. nov. E. L. elegans sp. nov. F. L. diplogramma (Möllendorff, 1902) comb. nov. Scale bars = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5514847" httpUri="https://zenodo.org/record/5514847/files/figure.png" pageId="11" pageNumber="153">Fig. 4B</figureCitation>
), empty shells are paler (
<figureCitation id="133E2A74C325FFC49C97FAD892FCBB1C" box="[392,487,1347,1374]" captionStart="Fig" captionStartId="9.[189,232,1849,1875]" captionTargetBox="[220,1368,265,1811]" captionTargetId="figure-16@9.[220,1368,265,1811]" captionTargetPageId="9" captionText="Fig. 3. External shell morphology of Landouria spp. from northeastern Thailand. A. Landouria circinata sp. nov., holotype (NHMSU-00023). B. L. tuberculata sp. nov., holotype (NHMSU-00025). C. L. trochomorphoides sp. nov., holotype (NHMSU-00027). D. L. chloritoides sp. nov., holotype (NHMSU-00029). E. L. elegans sp. nov., holotype (NHMSU-00032). F. L. diplogramma (Möllendorff, 1902) comb. nov. (NHMSU-00034). Scale bar = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5514845" httpUri="https://zenodo.org/record/5514845/files/figure.png" pageId="11" pageNumber="153">Fig. 3B</figureCitation>
). Umbilicus deep and wide. Aperture obliquely oval, reflexed, and slightly thickened.
</paragraph>
<paragraph id="8BBA36F1C325FFC49DA2FA339279B8FA" blockId="11.[186,1399,1448,1720]" pageId="11" pageNumber="153">
GENITAL SYSTEM (n = 3) (
<figureCitation id="133E2A74C325FFC49CC0FA339121BB81" box="[479,570,1448,1475]" captionStart="Fig" captionStartId="15.[189,232,1816,1842]" captionTargetBox="[283,1352,278,1767]" captionTargetId="figure-15@15.[232,1356,265,1778]" captionTargetPageId="15" captionText="Fig. 5. Illustrations of the genital system of Landouria spp. from northeastern Thailand. A. Landouria circinata sp. nov., paratype (NHMSU-00024). B. L. tuberculata sp. nov., paratype (NHMSU-00026). C. L. trochomorphoides sp. nov., paratype (NHMSU-00028). D. L. chloritoides sp. nov., paratype (NHMSU-00030). E. L. elegans sp. nov., paratype (NHMSU-00033). F. L. diplogramma (Möllendorff, 1902) comb. nov. (NHMSU-00034). Internal sculpture of a = flagellum; b = penis; c = vagina. Scale bars = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5514849" httpUri="https://zenodo.org/record/5514849/files/figure.png" pageId="11" pageNumber="153">Fig. 5B</figureCitation>
,
<tableCitation id="C687034AC325FFC49F59FA3391BABB80" box="[582,673,1448,1474]" captionStart="Table 4" captionStartId="10.[185,255,544,572]" captionTargetBox="[196,1908,758,1739]" captionTargetPageId="10" captionText="Table 4. Comparison of shells, radulae, and genital systems of Landouria spp. from northeastern Thailand. References: 1 = Tumpeesuwan &amp; Tumpeesuwan (2019); 2 = Nahok (2020); 3 = this study. *In Tumpeesuwan &amp; Tumpeesuwan (2019) indicated the number of teeth per row in Landouria strobiloides varies from 3 to 53. This lowest row has only 3 teeth. It is the uppermost row, probably lacerated and peel off. Unfortunately, we could not re-checked the complete specimens in this time being, so, this point need further investigation and proof." httpUri="http://table.plazi.org/id/DF7A6679C324FFC59DA6FDBB969CBCE7" pageId="11" pageNumber="153" tableUuid="DF7A6679C324FFC59DA6FDBB969CBCE7">Table 4</tableCitation>
). Penis cylindrical, swollen in the middle part, internally with rather wavy, corrugated longitudinal pilasters and a short, hollow verge with 23 lobes around its opening. Epiphallus evenly cylindrical, as long as penis. Flagellum like a short protrusion with a pointed tip, without nodes, internally with longitudinal slender pilasters. Vas deferens cylindrical, narrow, entering in the lower half of the flagellum. Vagina relatively short, internally with five corrugated longitudinal pilasters. Free oviduct very short. Gametolytic sac slightly thicker at base, with a long, narrow and thin cylindrical tube, and at its distal end a small, swollen spherical sac. Prostate gland very long. Uterus long and thin.
</paragraph>
<paragraph id="8BBA36F1C325FFC49DA2F97B91FDB9CA" blockId="11.[189,1400,1760,1928]" pageId="11" pageNumber="153">
RADULA (n = 3). Comprises 126130 transverse rows with 7987 teeth per row; radular formula: (28 30)-(1113)-1-(1113)-(2830). Central tooth symmetric, slender lanceolate, and unicuspid (
<figureCitation id="133E2A74C325FFC49816F8989672B95C" box="[1289,1385,1795,1822]" captionStart="Fig" captionStartId="18.[189,232,1783,1809]" captionTargetBox="[244,1343,265,1744]" captionTargetId="figure-12@18.[244,1343,265,1744]" captionTargetPageId="18" captionText="Fig. 6.Radula morphology ofLandouria spp. from northeasternThailand.AC, GR. Right side of central teeth. DF. Left side of central teeth. AC. Landouria circinata sp. nov., paratype (NHMSU-00024). DF. L. tuberculata sp. nov., paratype (NHMSU-00026). GI. L. trochomorphoides sp. nov., paratype (NHMSU-00028). JL. L. chloritoides sp. nov., paratype (NHMSU-00030). MO. L. elegans sp. nov., paratype (NHMSU-00033). PR. L. diplogramma (Möllendorff, 1902) comb. nov. (NHMSU-00034). Numbers indicate order of lateral and marginal teeth. Abbreviation: c = central tooth. Scale bars = 25 µm." figureDoi="http://doi.org/10.5281/zenodo.5514851" httpUri="https://zenodo.org/record/5514851/files/figure.png" pageId="11" pageNumber="153">Fig. 6D</figureCitation>
). Lateral teeth larger than the central tooth, lanceolate, bicuspid, with large endocone and sharp cusp (
<figureCitation id="133E2A74C325FFC49DD9F8D29233B926" box="[198,296,1865,1892]" captionStart="Fig" captionStartId="18.[189,232,1783,1809]" captionTargetBox="[244,1343,265,1744]" captionTargetId="figure-12@18.[244,1343,265,1744]" captionTargetPageId="18" captionText="Fig. 6.Radula morphology ofLandouria spp. from northeasternThailand.AC, GR. Right side of central teeth. DF. Left side of central teeth. AC. Landouria circinata sp. nov., paratype (NHMSU-00024). DF. L. tuberculata sp. nov., paratype (NHMSU-00026). GI. L. trochomorphoides sp. nov., paratype (NHMSU-00028). JL. L. chloritoides sp. nov., paratype (NHMSU-00030). MO. L. elegans sp. nov., paratype (NHMSU-00033). PR. L. diplogramma (Möllendorff, 1902) comb. nov. (NHMSU-00034). Numbers indicate order of lateral and marginal teeth. Abbreviation: c = central tooth. Scale bars = 25 µm." figureDoi="http://doi.org/10.5281/zenodo.5514851" httpUri="https://zenodo.org/record/5514851/files/figure.png" pageId="11" pageNumber="153">Fig. 6D</figureCitation>
). Marginal teeth tricuspid, with small sharp endocone; large, slender mesocone; and small v-shaped ectocone with two cusps (
<figureCitation id="133E2A74C325FFC49F41F8F691C2B9CA" box="[606,729,1901,1928]" captionStart="Fig" captionStartId="18.[189,232,1783,1809]" captionTargetBox="[244,1343,265,1744]" captionTargetId="figure-12@18.[244,1343,265,1744]" captionTargetPageId="18" captionText="Fig. 6.Radula morphology ofLandouria spp. from northeasternThailand.AC, GR. Right side of central teeth. DF. Left side of central teeth. AC. Landouria circinata sp. nov., paratype (NHMSU-00024). DF. L. tuberculata sp. nov., paratype (NHMSU-00026). GI. L. trochomorphoides sp. nov., paratype (NHMSU-00028). JL. L. chloritoides sp. nov., paratype (NHMSU-00030). MO. L. elegans sp. nov., paratype (NHMSU-00033). PR. L. diplogramma (Möllendorff, 1902) comb. nov. (NHMSU-00034). Numbers indicate order of lateral and marginal teeth. Abbreviation: c = central tooth. Scale bars = 25 µm." figureDoi="http://doi.org/10.5281/zenodo.5514851" httpUri="https://zenodo.org/record/5514851/files/figure.png" pageId="11" pageNumber="153">Fig. 6EF</figureCitation>
).
</paragraph>
</subSubSection>
<caption id="DF7A6679C322FFC39DA2F81A976BB9A0" ID-DOI="http://doi.org/10.5281/zenodo.5514847" ID-Zenodo-Dep="5514847" httpUri="https://zenodo.org/record/5514847/files/figure.png" pageId="12" pageNumber="154" startId="12.[189,232,1921,1947]" targetBox="[193,1395,278,1870]" targetPageId="12">
<paragraph id="8BBA36F1C322FFC39DA2F81A976BB9A0" blockId="12.[189,1399,1921,2018]" pageId="12" pageNumber="154">
<emphasis id="B971EAE3C322FFC39DA2F81A9215B9D9" bold="true" box="[189,270,1921,1947]" pageId="12" pageNumber="154">Fig. 4.</emphasis>
Living adults of
<taxonomicName id="4C054D72C322FFC39CFCF81A9182B9DE" box="[483,665,1921,1948]" class="Gastropoda" family="Camaenidae" genus="Landouria" kingdom="Animalia" order="Stylommatophora" pageId="12" pageNumber="154" phylum="Mollusca" rank="species" species="undetermined">
<emphasis id="B971EAE3C322FFC39CFCF81A9144B9D9" box="[483,607,1921,1947]" italics="true" pageId="12" pageNumber="154">Landouria</emphasis>
spp.
</taxonomicName>
from northeastern Thailand.
<emphasis id="B971EAE3C322FFC39EE5F81A9714B9D9" bold="true" box="[1018,1039,1921,1947]" pageId="12" pageNumber="154">A</emphasis>
.
<taxonomicName id="4C054D72C322FFC3993EF81A960BB9D9" authority="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan, 2021" authorityName="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan" authorityYear="2021" box="[1057,1296,1921,1947]" class="Gastropoda" family="Camaenidae" genus="Landouria" kingdom="Animalia" order="Stylommatophora" pageId="12" pageNumber="154" phylum="Mollusca" rank="species" species="circinata" status="sp. nov.">
<emphasis id="B971EAE3C322FFC3993EF81A960BB9D9" box="[1057,1296,1921,1947]" italics="true" pageId="12" pageNumber="154">Landouria circinata</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2425798C322FFC39805F819966CB9DE" box="[1306,1399,1922,1948]" pageId="12" pageNumber="154" rank="species">sp. nov.</taxonomicNameLabel>
<emphasis id="B971EAE3C322FFC39DA2F83F93CAB9FC" bold="true" box="[189,209,1956,1982]" pageId="12" pageNumber="154">B</emphasis>
.
<taxonomicName id="4C054D72C322FFC39DFDF83E9291B9FC" authority="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan, 2021" authorityName="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan" authorityYear="2021" box="[226,394,1956,1982]" class="Gastropoda" family="Camaenidae" genus="Landouria" kingdom="Animalia" order="Stylommatophora" pageId="12" pageNumber="154" phylum="Mollusca" rank="species" species="tuberculata" status="sp. nov.">
<emphasis id="B971EAE3C322FFC39DFDF83E9291B9FC" box="[226,394,1956,1982]" italics="true" pageId="12" pageNumber="154">L. tuberculata</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2425798C322FFC39C8AF83E92E8B9FD" box="[405,499,1957,1983]" pageId="12" pageNumber="154" rank="species">sp. nov.</taxonomicNameLabel>
<emphasis id="B971EAE3C322FFC39CE2F83F9109B9FC" bold="true" box="[509,530,1956,1982]" pageId="12" pageNumber="154">C</emphasis>
.
<taxonomicName id="4C054D72C322FFC39F3BF83E9007B9FC" authority="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan, 2021" authorityName="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan" authorityYear="2021" box="[548,796,1956,1982]" class="Gastropoda" family="Camaenidae" genus="Landouria" kingdom="Animalia" order="Stylommatophora" pageId="12" pageNumber="154" phylum="Mollusca" rank="species" species="trochomorphoides" status="sp. nov.">
<emphasis id="B971EAE3C322FFC39F3BF83E9007B9FC" box="[548,796,1956,1982]" italics="true" pageId="12" pageNumber="154">L. trochomorphoides</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2425798C322FFC39E39F83E909FB9FD" box="[806,900,1957,1983]" pageId="12" pageNumber="154" rank="species">sp. nov.</taxonomicNameLabel>
<emphasis id="B971EAE3C322FFC39E91F83F90B8B9FC" bold="true" box="[910,931,1956,1982]" pageId="12" pageNumber="154">D</emphasis>
.
<taxonomicName id="4C054D72C322FFC39EAAF83E9778B9FC" authority="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan, 2021" authorityName="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan" authorityYear="2021" box="[949,1123,1956,1982]" class="Gastropoda" family="Camaenidae" genus="Landouria" kingdom="Animalia" order="Stylommatophora" pageId="12" pageNumber="154" phylum="Mollusca" rank="species" species="chloritoides" status="sp. nov.">
<emphasis id="B971EAE3C322FFC39EAAF83E9778B9FC" box="[949,1123,1956,1982]" italics="true" pageId="12" pageNumber="154">L. chloritoides</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2425798C322FFC39971F83E97D0B9FD" box="[1134,1227,1957,1983]" pageId="12" pageNumber="154" rank="species">sp. nov.</taxonomicNameLabel>
<emphasis id="B971EAE3C322FFC399CAF83F97F2B9FC" bold="true" box="[1237,1257,1956,1982]" pageId="12" pageNumber="154">E</emphasis>
.
<taxonomicName id="4C054D72C322FFC399E4F83E966CB9FC" authority="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan, 2021" authorityName="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan" authorityYear="2021" box="[1275,1399,1956,1982]" class="Gastropoda" family="Camaenidae" genus="Landouria" kingdom="Animalia" order="Stylommatophora" pageId="12" pageNumber="154" phylum="Mollusca" rank="species" species="elegans" status="sp. nov.">
<emphasis id="B971EAE3C322FFC399E4F83E966CB9FC" box="[1275,1399,1956,1982]" italics="true" pageId="12" pageNumber="154">L. elegans</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2425798C322FFC39DA2F8539203B9A0" box="[189,280,1992,2018]" pageId="12" pageNumber="154" rank="species">sp. nov.</taxonomicNameLabel>
<emphasis id="B971EAE3C322FFC39C01F853922BB9A0" bold="true" box="[286,304,1992,2018]" pageId="12" pageNumber="154">F</emphasis>
.
<taxonomicName id="4C054D72C322FFC39C20F85391F4B9A0" authority="(Mollendorff, 1902)" authorityName="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan" authorityYear="2021" baseAuthorityName="Mollendorff" baseAuthorityYear="1902" box="[319,751,1991,2018]" class="Gastropoda" family="Camaenidae" genus="Landouria" kingdom="Animalia" order="Stylommatophora" pageId="12" pageNumber="154" phylum="Mollusca" rank="species" species="diplogramma" status="comb. nov.">
<emphasis id="B971EAE3C322FFC39C20F85392E7B9A3" box="[319,508,1991,2017]" italics="true" pageId="12" pageNumber="154">L. diplogramma</emphasis>
(
<bibRefCitation id="EF944B00C322FFC39F14F85C91FCB9A0" author="Mollendorff O. F." box="[523,743,1991,2018]" pageId="12" pageNumber="154" pagination="153 - 160" refId="ref10489" refString="Mollendorff O. F. 1902. Neue Arten und Unterarten von Fruhstorfer in Siam gesammelt. Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft 34 (9 - 10): 153 - 160." type="journal article" year="1902">Möllendorff, 1902</bibRefCitation>
)
</taxonomicName>
<taxonomicNameLabel id="A2425798C322FFC39FE8F85C9063B9A0" box="[759,888,1991,2018]" pageId="12" pageNumber="154" rank="species">comb. nov.</taxonomicNameLabel>
Scale bars = 10 mm.
</paragraph>
</caption>
<subSubSection id="C31F657AC323FFC29DA2FE92960EBF37" pageId="13" pageNumber="155" type="discussion">
<paragraph id="8BBA36F1C323FFC29DA2FE929234BF61" blockId="13.[189,303,265,291]" box="[189,303,265,291]" pageId="13" pageNumber="155">
<heading id="D0F2819DC323FFC29DA2FE929234BF61" bold="true" box="[189,303,265,291]" fontSize="11" level="3" pageId="13" pageNumber="155" reason="3">
<emphasis id="B971EAE3C323FFC29DA2FE929234BF61" bold="true" box="[189,303,265,291]" pageId="13" pageNumber="155">Remarks</emphasis>
</heading>
</paragraph>
<paragraph id="8BBA36F1C323FFC29DA2FEAC960EBF37" blockId="13.[189,1398,311,373]" pageId="13" pageNumber="155">
<taxonomicName id="4C054D72C323FFC29DA2FEAC92DCBF13" authority="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan, 2021" authorityName="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan" authorityYear="2021" box="[189,455,311,337]" class="Gastropoda" family="Camaenidae" genus="Landouria" kingdom="Animalia" order="Stylommatophora" pageId="13" pageNumber="155" phylum="Mollusca" rank="species" species="tuberculata" status="sp. nov.">
<emphasis id="B971EAE3C323FFC29DA2FEAC92DCBF13" box="[189,455,311,337]" italics="true" pageId="13" pageNumber="155">Landouria tuberculata</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2425798C323FFC29CCFFEA39137BF10" box="[464,556,312,338]" pageId="13" pageNumber="155" rank="species">sp. nov.</taxonomicNameLabel>
differs from other Thai species of
<taxonomicName id="4C054D72C323FFC29ED2FEAC9753BF13" authorityName="Godwin-Austen" authorityYear="1918" box="[973,1096,311,337]" class="Gastropoda" family="Camaenidae" genus="Landouria" kingdom="Animalia" order="Stylommatophora" pageId="13" pageNumber="155" phylum="Mollusca" rank="genus">
<emphasis id="B971EAE3C323FFC29ED2FEAC9753BF13" box="[973,1096,311,337]" italics="true" pageId="13" pageNumber="155">Landouria</emphasis>
</taxonomicName>
, by the shell sculpture of numerous tiny tubercles (
<figureCitation id="133E2A74C323FFC29CF7FEC1915EBF36" box="[488,581,346,372]" captionStart="Fig" captionStartId="12.[189,232,1921,1947]" captionTargetBox="[193,1395,278,1870]" captionTargetId="figure-12@12.[193,1395,265,1883]" captionTargetPageId="12" captionText="Fig. 4. Living adults of Landouria spp. from northeastern Thailand. A. Landouria circinata sp. nov. B. L. tuberculata sp. nov. C. L. trochomorphoides sp. nov. D. L. chloritoides sp. nov. E. L. elegans sp. nov. F. L. diplogramma (Möllendorff, 1902) comb. nov. Scale bars = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5514847" httpUri="https://zenodo.org/record/5514847/files/figure.png" pageId="13" pageNumber="155">Fig. 4B</figureCitation>
). Lateral teeth of radula bicuspid and lanceolate (
<figureCitation id="133E2A74C323FFC29997FEC19612BF37" box="[1160,1289,346,373]" captionStart="Fig" captionStartId="18.[189,232,1783,1809]" captionTargetBox="[244,1343,265,1744]" captionTargetId="figure-12@18.[244,1343,265,1744]" captionTargetPageId="18" captionText="Fig. 6.Radula morphology ofLandouria spp. from northeasternThailand.AC, GR. Right side of central teeth. DF. Left side of central teeth. AC. Landouria circinata sp. nov., paratype (NHMSU-00024). DF. L. tuberculata sp. nov., paratype (NHMSU-00026). GI. L. trochomorphoides sp. nov., paratype (NHMSU-00028). JL. L. chloritoides sp. nov., paratype (NHMSU-00030). MO. L. elegans sp. nov., paratype (NHMSU-00033). PR. L. diplogramma (Möllendorff, 1902) comb. nov. (NHMSU-00034). Numbers indicate order of lateral and marginal teeth. Abbreviation: c = central tooth. Scale bars = 25 µm." figureDoi="http://doi.org/10.5281/zenodo.5514851" httpUri="https://zenodo.org/record/5514851/files/figure.png" pageId="13" pageNumber="155">Fig. 6DE</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C31F657AC323FFC29DA2FE0091E1BC45" pageId="13" pageNumber="155" type="distribution">
<paragraph id="8BBA36F1C323FFC29DA2FE00924DBFF7" blockId="13.[189,342,411,437]" box="[189,342,411,437]" pageId="13" pageNumber="155">
<heading id="D0F2819DC323FFC29DA2FE00924DBFF7" bold="true" box="[189,342,411,437]" fontSize="11" level="3" pageId="13" pageNumber="155" reason="3">
<emphasis id="B971EAE3C323FFC29DA2FE00924DBFF7" bold="true" box="[189,342,411,437]" pageId="13" pageNumber="155">Distribution</emphasis>
</heading>
</paragraph>
<paragraph id="8BBA36F1C323FFC29DA2FE5191E1BC45" blockId="13.[189,1398,458,520]" pageId="13" pageNumber="155">
<taxonomicName id="4C054D72C323FFC29DA2FE5192D3BFA6" authority="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan, 2021" authorityName="Nahok &amp; Tumpeesuwan &amp; Tumpeesuwan" authorityYear="2021" box="[189,456,458,484]" class="Gastropoda" family="Camaenidae" genus="Landouria" kingdom="Animalia" order="Stylommatophora" pageId="13" pageNumber="155" phylum="Mollusca" rank="species" species="tuberculata" status="sp. nov.">
<emphasis id="B971EAE3C323FFC29DA2FE5192D3BFA6" box="[189,456,458,484]" italics="true" pageId="13" pageNumber="155">Landouria tuberculata</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2425798C323FFC29CCDFE509134BFA7" box="[466,559,459,485]" pageId="13" pageNumber="155" rank="species">sp. nov.</taxonomicNameLabel>
is currently only known from the area surrounding Phu Rua National Park, Phu Rua District,
<collectingRegion id="49C1F813C323FFC29CC9FE769165BC45" box="[470,638,493,519]" country="Thailand" name="Loei" pageId="13" pageNumber="155">Loei Province</collectingRegion>
,
<collectingCountry id="F3127661C323FFC29F96FE7691EEBC45" box="[649,757,493,519]" name="Thailand" pageId="13" pageNumber="155">Thailand</collectingCountry>
.
</paragraph>
</subSubSection>
</treatment>
</document>