307 lines
45 KiB
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307 lines
45 KiB
XML
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<mods:title id="BF39564038738AFFA26146D60742D5E7">Systematic revision of the limestone karst-restricted land snail genus Aenigmatoconcha (Eupulmonata: Helicarionidae), with description of a new species</mods:title>
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<mods:namePart id="AD069479F2EC8880D99B13CD5122EB32">Pholyotha, Arthit</mods:namePart>
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<mods:namePart id="C1F97863E9155D6EFBFE57D176A4DE2D">Sutcharit, Chirasak</mods:namePart>
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<mods:affiliation id="B47D2B0324A2CD98A10453A76562438F">Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand.</mods:affiliation>
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<mods:namePart id="921B866409686C99767AFA27086D84D3">Tongkerd, Piyoros</mods:namePart>
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<mods:affiliation id="EB07FFFD2C28A7055DE24745F7D5DE1D">Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand.</mods:affiliation>
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<treatment id="03DE8788A43DFFB938CEFD6CFED643E4" ID-DOI="http://doi.org/10.5281/zenodo.5528128" ID-GBIF-Taxon="188498258" ID-Zenodo-Dep="5528128" LSID="urn:lsid:plazi:treatment:03DE8788A43DFFB938CEFD6CFED643E4" httpUri="http://treatment.plazi.org/id/03DE8788A43DFFB938CEFD6CFED643E4" lastPageId="11" lastPageNumber="66" pageId="9" pageNumber="64">
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<taxonomicName id="4C774D1DA43DFFBB38CEFD6CFB3E432E" ID-CoL="64Y97" authority="Tumpeesuwan & Tumpeesuwan, 2017" authorityName="Tumpeesuwan & Tumpeesuwan" authorityYear="2017" box="[401,1187,765,792]" class="Gastropoda" family="Helicarionidae" genus="Aenigmatoconcha" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="64" phylum="Mollusca" rank="species" species="clivicola">
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<emphasis id="B903EA8CA43DFFBB38CEFD6CFD44432E" bold="true" box="[401,729,765,792]" italics="true" pageId="9" pageNumber="64">Aenigmatoconcha clivicola</emphasis>
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<bibRefCitation id="EFE64B6FA43DFFBB3BBFFD6FFB3E432E" author="Tumpeesuwan C. & Tumpeesuwan S." box="[736,1187,766,792]" pageId="9" pageNumber="64" pagination="170 - 176" refId="ref13001" refString="Tumpeesuwan C. & Tumpeesuwan S. 2017. Discovery of an overlooked helicarionid land snail (Helicarionidae: Durgellinae) from northeastern Thailand, with description of a new genus and new species, and a note on radula morphology and genital system. Raffles Bulletin of Zoology 65: 174 - 181. Tumpeesuwan C. & Tumpeesuwan S. 2018. Aenigmatoconcha sumonthai, a new helicarionid land snail from Chumphon Province, Southern Thailand (Helicarionidae: Durgellinae). Raffles Bulletin of Zoology 66: 170 - 176. http: // doi. org / 10.5281 / zenodo. 4504598" type="journal article" year="2017">Tumpeesuwan & Tumpeesuwan, 2017</bibRefCitation>
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<paragraph id="8BC8369EA43DFFBB3BDCFCB0FC2C430D" blockId="9.[401,1187,765,827]" box="[643,945,800,827]" pageId="9" pageNumber="64">
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<figureCitation id="134C2A1BA43DFFBB3BDCFCB0FD51430D" box="[643,716,801,827]" captionStart="Fig" captionStartId="2.[189,232,1962,1988]" captionTargetBox="[189,1398,678,1922]" captionTargetId="figure-108@2.[189,1398,678,1922]" captionTargetPageId="2" captionText="Fig. 1. Geographic distribution of four species of Aenigmatoconcha Tumpeesuwan & Tumpeesuwan, 2017 based on specimens examined herein." figureDoi="http://doi.org/10.5281/zenodo.5528136" httpUri="https://zenodo.org/record/5528136/files/figure.png" pageId="9" pageNumber="64">Figs 1</figureCitation>
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,
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<figureCitation id="134C2A1BA43DFFBB3B84FCB0FC9C430D" box="[731,769,801,827]" captionStart="Fig" captionStartId="6.[189,232,1790,1816]" captionTargetBox="[189,1397,265,1744]" captionTargetId="figure-14@6.[189,1398,265,1744]" captionTargetPageId="6" captionText="Fig. 2. Maximum likelihood tree of Aenigmatoconcha Tumpeesuwan & Tumpeesuwan, 2017 based on the COI gene dataset. Numbers on nodes indicate the ML bootstrap values (left) and Bayesian posterior probabilities (right) values. Pictures of living snails are not to scale." figureDoi="http://doi.org/10.5281/zenodo.5528138" httpUri="https://zenodo.org/record/5528138/files/figure.png" pageId="9" pageNumber="64">2A</figureCitation>
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,
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<figureCitation id="134C2A1BA43DFFBB3A52FCB0FCCB430D" box="[781,854,801,827]" captionStart="Fig" captionStartId="10.[189,232,1676,1702]" captionTargetBox="[189,1398,264,1638]" captionTargetId="figure-14@10.[189,1398,264,1638]" captionTargetPageId="10" captionText="Fig. 4. Shells of Aenigmatoconcha spp. A–B. A. clivicola Tumpeesuwan & Tumpeesuwan, 2017, specimen CUMZ 7928 from the type locality (A) and specimen CUMZ 7930 from Loei Province (B). C–F. A. mitis (Pfeiffer, 1863) comb. nov., syntypes NHMUK ex. Cuming collection (C), specimen CUMZ 7921 from Phetchaburi Province (D), specimen CUMZ 7687 from Kanchanaburi Province (E), and specimen CUMZ 7885 from Nakhon Sawan Province (F)." figureDoi="http://doi.org/10.5281/zenodo.5528146" httpUri="https://zenodo.org/record/5528146/files/figure.png" pageId="9" pageNumber="64">4A–B</figureCitation>
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,
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<figureCitation id="134C2A1BA43DFFBB3A3DFCB1FCEF430D" box="[866,882,800,827]" captionStart="Fig" captionStartId="12.[189,232,1790,1816]" captionTargetBox="[189,1398,264,1743]" captionTargetId="figure-14@12.[189,1398,264,1743]" captionTargetPageId="12" captionText="Fig. 5. Genital system and mantle lobes of Aenigmatoconcha clivicola Tumpeesuwan & Tumpeesuwan, 2017, specimen CUMZ 7929 from the type locality. A. Mantle lobes. B. General view of genitalia; inset showing flagellum encircled with a loose tissue before clearing it out. C. Internal structure of penis. D. Internal structure of epiphallus and epiphallic caecum. White arrow indicates the junction between penis and epiphallus." figureDoi="http://doi.org/10.5281/zenodo.5528148" httpUri="https://zenodo.org/record/5528148/files/figure.png" pageId="9" pageNumber="64">5</figureCitation>
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,
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<figureCitation id="134C2A1BA43DFFBB3A20FCB0FC2C430D" box="[895,945,801,827]" captionStart="Fig" captionStartId="21.[189,232,1823,1849]" captionTargetBox="[194,1394,950,1780]" captionTargetId="figure-243@21.[189,1398,945,1784]" captionTargetPageId="21" captionText="Fig. 10. Radula morphology of Aenigmatoconcha spp. A. A. clivicola Tumpeesuwan & Tumpeesuwan, 2017, specimen CUMZ 7929 from Loei Province. B. A. mitis (Pfeiffer, 1863) comb. nov., specimen CUMZ 7687 from Kanchanaburi Province. C. A. sumonthai Tumpeesuwan & Tumpeesuwan, 2018, specimen CUMZ 7922 from Chumphon Province. D. A. eunetis Pholyotha & Panha sp. nov., paratype CUMZ 7933 from Uthai Thani Province. Central tooth in yellow. Scale bars: A–C = 100 μm; D = 50 μm." figureDoi="http://doi.org/10.5281/zenodo.5528158" httpUri="https://zenodo.org/record/5528158/files/figure.png" pageId="9" pageNumber="64">10A</figureCitation>
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<treatmentCitationGroup id="AB6711B0A43DFFBB39E2FCF3FB6F43A9" pageId="9" pageNumber="64">
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<taxonomicName id="4C774D1DA43DFFBB39E2FCF3FBCE434A" ID-CoL="64Y97" authority="Tumpeesuwan & Tumpeesuwan, 2017: 184 - 187" authorityName="Tumpeesuwan & Tumpeesuwan" authorityPageNumber="184 - 187" authorityYear="2017" box="[189,1107,866,893]" class="Gastropoda" family="Helicarionidae" genus="Aenigmatoconcha" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="64" phylum="Mollusca" rank="species" species="clivicola">
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<emphasis id="B903EA8CA43DFFBB39E2FCF3FD9E434A" box="[189,515,866,892]" italics="true" pageId="9" pageNumber="64">Aenigmatoconcha clivicola</emphasis>
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<bibRefCitation id="EFE64B6FA43DFFBB3B52FCF2FBCE434A" author="Tumpeesuwan C. & Tumpeesuwan S." box="[525,1107,866,893]" pageId="9" pageNumber="64" pagination="170 - 176" refId="ref13001" refString="Tumpeesuwan C. & Tumpeesuwan S. 2017. Discovery of an overlooked helicarionid land snail (Helicarionidae: Durgellinae) from northeastern Thailand, with description of a new genus and new species, and a note on radula morphology and genital system. Raffles Bulletin of Zoology 65: 174 - 181. Tumpeesuwan C. & Tumpeesuwan S. 2018. Aenigmatoconcha sumonthai, a new helicarionid land snail from Chumphon Province, Southern Thailand (Helicarionidae: Durgellinae). Raffles Bulletin of Zoology 66: 170 - 176. http: // doi. org / 10.5281 / zenodo. 4504598" type="journal article" year="2017">Tumpeesuwan & Tumpeesuwan, 2017: 184–187</bibRefCitation>
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, figs 2–5.
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<typeStatus id="54CC883CA43DFFBB3D8DFCF2FA91434B" box="[1234,1292,867,893]" pageId="9" pageNumber="64">Type</typeStatus>
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locality: Phu Pha Lom Limestone Hill in Mueang District,
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<collectingRegion id="49B3F87CA43DFFBB3A69FC14FC4343A9" box="[822,990,901,927]" country="Thailand" name="Loei" pageId="9" pageNumber="64">Loei Province</collectingRegion>
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, northeastern
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<collectingCountry id="F360760EA43DFFBB3DDDFC14FB7343A9" box="[1154,1262,901,927]" name="Thailand" pageId="9" pageNumber="64">Thailand</collectingCountry>
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.
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<paragraph id="8BC8369EA43DFFBB39E2FC56FB8143D7" blockId="9.[189,1052,967,993]" box="[189,1052,967,993]" pageId="9" pageNumber="64">
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<taxonomicName id="4C774D1DA43DFFBB39E2FC56FD9D43D7" ID-CoL="64Y97" authorityName="Tumpeesuwan & Tumpeesuwan" authorityYear="2017" box="[189,512,967,993]" class="Gastropoda" family="Helicarionidae" genus="Aenigmatoconcha" kingdom="Animalia" order="Stylommatophora" pageId="9" pageNumber="64" phylum="Mollusca" rank="species" species="clivicola">
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<emphasis id="B903EA8CA43DFFBB39E2FC56FD9D43D7" box="[189,512,967,993]" italics="true" pageId="9" pageNumber="64">Aenigmatoconcha clivicola</emphasis>
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</taxonomicName>
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– Tumpeesuwan & Tumpeesuwan 2018: 171.
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<subSubSection id="C36D6515A43DFFBB39E2FB99FD82448C" pageId="9" pageNumber="64" type="materials_examined">
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<paragraph id="8BC8369EA43DFFBB39E2FB99FE374414" blockId="9.[189,426,1032,1058]" box="[189,426,1032,1058]" pageId="9" pageNumber="64">
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<emphasis id="B903EA8CA43DFFBB39E2FB99FE374414" bold="true" box="[189,426,1032,1058]" pageId="9" pageNumber="64">Material examined</emphasis>
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</heading>
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</paragraph>
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<paragraph id="8BC8369EA43DFFBB39E2FBA6FD82448C" blockId="9.[189,1399,1077,1211]" pageId="9" pageNumber="64">
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<materialsCitation id="3B1F3CC3A43DFFBB39E2FBA6FDB74442" ID-GBIF-Occurrence="3357771308" collectionCode="CUMZ" country="Thailand" latitude="17.554611" location="Phu Pha Lom Limestone Hill" longLatPrecision="1" longitude="101.86817" municipality="Mueang District" pageId="9" pageNumber="64" specimenCode="CUMZ 7928" specimenCount="61" stateProvince="Loei Province">
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<collectingCountry id="F360760EA43DFFBB39E2FBA6FEC84467" box="[189,341,1079,1105]" name="Thailand" pageId="9" pageNumber="64">THAILAND</collectingCountry>
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•
|
||
<specimenCount id="9D71FD17A43DFFBB3837FBA6FE3A4467" box="[360,423,1079,1105]" pageId="9" pageNumber="64">39 sh</specimenCount>
|
||
,
|
||
<specimenCount id="9D71FD17A43DFFBB38EFFBA6FE724467" box="[432,495,1079,1105]" pageId="9" pageNumber="64">22 sp</specimenCount>
|
||
;
|
||
<collectingRegion id="49B3F87CA43DFFBB38A7FBA6FD034467" box="[504,670,1079,1105]" country="Thailand" name="Loei" pageId="9" pageNumber="64">Loei Province</collectingRegion>
|
||
,
|
||
<collectingMunicipality id="6BACACE4A43DFFBB3BF8FBA6FCFA4467" box="[679,871,1079,1105]" pageId="9" pageNumber="64">Mueang District</collectingMunicipality>
|
||
,
|
||
<location id="8EA86045A43DFFBB3A2FFBA6FB5D4467" LSID="urn:lsid:plazi:treatment:03DE8788A43DFFB938CEFD6CFED643E4:8EA86045A43DFFBB3A2FFBA6FB5D4467" box="[880,1216,1079,1105]" country="Thailand" latitude="17.554611" longLatPrecision="1" longitude="101.86817" municipality="Mueang District" name="Phu Pha Lom Limestone Hill" pageId="9" pageNumber="64" stateProvince="Loei Province">Phu Pha Lom Limestone Hill</location>
|
||
;
|
||
<geoCoordinate id="EE435059A43DFFBB3D96FBA4FAEF4467" box="[1225,1394,1077,1105]" degrees="17" direction="north" minutes="33" orientation="latitude" pageId="9" pageNumber="64" precision="1" seconds="16.6" value="17.554611">17°33′16.6″ N</geoCoordinate>
|
||
,
|
||
<geoCoordinate id="EE435059A43DFFBB39E2FBC9FEE74442" box="[189,378,1112,1140]" degrees="101" direction="east" minutes="52" orientation="longitude" pageId="9" pageNumber="64" precision="1" seconds="05.4" value="101.86817">101°52′05.4″ E</geoCoordinate>
|
||
;
|
||
<specimenCode id="DBD19EE5A43DFFBB38D2FBCBFDB74442" box="[397,554,1114,1140]" collectionCode="CUMZ" country="0" httpUri="http://grbio.org/cool/da58-p0xp" name="Cameroon University, Museum of Zoology" pageId="9" pageNumber="64">CUMZ 7928</specimenCode>
|
||
</materialsCitation>
|
||
•
|
||
<materialsCitation id="3B1F3CC3A43DFFBB3B11FBCBFAEA4442" ID-GBIF-Occurrence="3357771311" box="[590,1399,1114,1140]" collectionCode="CUMZ" country="Thailand" latitude="17.554611" location="Phu Pha Lom Limestone Hill" longLatPrecision="1" longitude="101.86817" municipality="Mueang District" pageId="9" pageNumber="64" specimenCode="CUMZ 7929" specimenCount="18" stateProvince="Loei">
|
||
<specimenCount id="9D71FD17A43DFFBB3B11FBCBFD1A4442" box="[590,647,1114,1140]" pageId="9" pageNumber="64">6 sh</specimenCount>
|
||
,
|
||
<specimenCount id="9D71FD17A43DFFBB3BC6FBCBFD7D4442" box="[665,736,1114,1140]" pageId="9" pageNumber="64">12 sp</specimenCount>
|
||
; same collection data as for preceding;
|
||
<specimenCode id="DBD19EE5A43DFFBB3D85FBCBFAEA4442" box="[1242,1399,1114,1140]" collectionCode="CUMZ" country="0" httpUri="http://grbio.org/cool/da58-p0xp" name="Cameroon University, Museum of Zoology" pageId="9" pageNumber="64">CUMZ 7929</specimenCode>
|
||
</materialsCitation>
|
||
•
|
||
<materialsCitation id="3B1F3CC3A43DFFBB398EFBECFD86448C" ID-GBIF-Occurrence="3357771317" collectionCode="CUMZ" country="Thailand" county="Na Duang District" latitude="17.577806" location="Na Duang District" longLatPrecision="1" longitude="101.89308" municipality="Limestone" pageId="9" pageNumber="64" specimenCode="CUMZ 7930" specimenCount="48" stateProvince="Loei Province">
|
||
<specimenCount id="9D71FD17A43DFFBB398EFBECFE8844A1" box="[209,277,1149,1175]" pageId="9" pageNumber="64">48 sh</specimenCount>
|
||
;
|
||
<collectingRegion id="49B3F87CA43DFFBB387CFBECFE5044A1" box="[291,461,1149,1175]" country="Thailand" name="Loei" pageId="9" pageNumber="64">Loei Province</collectingRegion>
|
||
,
|
||
<collectingCounty id="62A94E12A43DFFBB3883FBEFFD2744A1" box="[476,698,1149,1176]" pageId="9" pageNumber="64">Na Duang District</collectingCounty>
|
||
,
|
||
<locationDeviation id="9A7A2FBCA43DFFBB3B97FBECFB2B44AE" box="[712,1206,1149,1176]" pageId="9" pageNumber="64">Limestone outcrops at Wat Tham Pha Ya</locationDeviation>
|
||
;
|
||
<geoCoordinate id="EE435059A43DFFBB3D9BFBEAFAEF44AE" box="[1220,1394,1147,1176]" degrees="17" direction="north" minutes="34" orientation="latitude" pageId="9" pageNumber="64" precision="1" seconds="40.1" value="17.577806">17°34′40.1″ N</geoCoordinate>
|
||
,
|
||
<geoCoordinate id="EE435059A43DFFBB39E2FB0EFEE9448D" box="[189,372,1183,1211]" degrees="101" direction="east" minutes="53" orientation="longitude" pageId="9" pageNumber="64" precision="1" seconds="35.1" value="101.89308">101°53′35.1″ E</geoCoordinate>
|
||
;
|
||
<specimenCode id="DBD19EE5A43DFFBB38DEFB31FD86448C" box="[385,539,1184,1210]" collectionCode="CUMZ" country="0" httpUri="http://grbio.org/cool/da58-p0xp" name="Cameroon University, Museum of Zoology" pageId="9" pageNumber="64">CUMZ 7930</specimenCode>
|
||
</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C36D6515A43DFFBB39E2FB70FECF478F" pageId="9" pageNumber="64" type="description">
|
||
<paragraph id="8BC8369EA43DFFBB39E2FB70FED344CD" blockId="9.[189,334,1249,1275]" box="[189,334,1249,1275]" pageId="9" pageNumber="64">
|
||
<heading id="D08081F2A43DFFBB39E2FB70FED344CD" bold="true" box="[189,334,1249,1275]" fontSize="11" level="3" pageId="9" pageNumber="64" reason="3">
|
||
<emphasis id="B903EA8CA43DFFBB39E2FB70FED344CD" bold="true" box="[189,334,1249,1275]" pageId="9" pageNumber="64">Description</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8BC8369EA43DFFBB39E2FA81FEA445A2" blockId="9.[189,1401,1296,1428]" pageId="9" pageNumber="64">
|
||
SHELL (
|
||
<figureCitation id="134C2A1BA43DFFBB3849FA81FE05451C" box="[278,408,1296,1323]" captionStart="Fig" captionStartId="10.[189,232,1676,1702]" captionTargetBox="[189,1398,264,1638]" captionTargetId="figure-14@10.[189,1398,264,1638]" captionTargetPageId="10" captionText="Fig. 4. Shells of Aenigmatoconcha spp. A–B. A. clivicola Tumpeesuwan & Tumpeesuwan, 2017, specimen CUMZ 7928 from the type locality (A) and specimen CUMZ 7930 from Loei Province (B). C–F. A. mitis (Pfeiffer, 1863) comb. nov., syntypes NHMUK ex. Cuming collection (C), specimen CUMZ 7921 from Phetchaburi Province (D), specimen CUMZ 7687 from Kanchanaburi Province (E), and specimen CUMZ 7885 from Nakhon Sawan Province (F)." figureDoi="http://doi.org/10.5281/zenodo.5528146" httpUri="https://zenodo.org/record/5528146/files/figure.png" pageId="9" pageNumber="64">Fig. 4A–B</figureCitation>
|
||
). Shell strongly depressed to depressed, medium-sized (shell width
|
||
<quantity id="4C8F9B7BA43DFFBB3D9CFA81FAEF451D" box="[1219,1394,1296,1323]" metricMagnitude="-2" metricUnit="m" metricValue="1.8649999999999998" metricValueMax="2.01" metricValueMin="1.72" pageId="9" pageNumber="64" unit="mm" value="18.65" valueMax="20.1" valueMin="17.2">17.2–20.1 mm</quantity>
|
||
, shell height 8.6–10.0 mm), rather thin to slightly solid and translucent, milky to pale whitish-horny colour, well-rounded body whorl, little elevated spire, impressed suture, obvious varix, and open umbilicus.
|
||
</paragraph>
|
||
<paragraph id="8BC8369EA43DFFBB39E2FA2DFD50462A" blockId="9.[189,1399,1467,1564]" pageId="9" pageNumber="64">
|
||
EXTERNAL FEATURES (
|
||
<figureCitation id="134C2A1BA43DFFBB38EDFA2AFD8C45E3" box="[434,529,1467,1493]" captionStart="Fig" captionStartId="6.[189,232,1790,1816]" captionTargetBox="[189,1397,265,1744]" captionTargetId="figure-14@6.[189,1398,265,1744]" captionTargetPageId="6" captionText="Fig. 2. Maximum likelihood tree of Aenigmatoconcha Tumpeesuwan & Tumpeesuwan, 2017 based on the COI gene dataset. Numbers on nodes indicate the ML bootstrap values (left) and Bayesian posterior probabilities (right) values. Pictures of living snails are not to scale." figureDoi="http://doi.org/10.5281/zenodo.5528138" httpUri="https://zenodo.org/record/5528138/files/figure.png" pageId="9" pageNumber="64">Figs 2A</figureCitation>
|
||
,
|
||
<figureCitation id="134C2A1BA43DFFBB3B7DFA2AFDDB45E3" box="[546,582,1467,1494]" captionStart="Fig" captionStartId="12.[189,232,1790,1816]" captionTargetBox="[189,1398,264,1743]" captionTargetId="figure-14@12.[189,1398,264,1743]" captionTargetPageId="12" captionText="Fig. 5. Genital system and mantle lobes of Aenigmatoconcha clivicola Tumpeesuwan & Tumpeesuwan, 2017, specimen CUMZ 7929 from the type locality. A. Mantle lobes. B. General view of genitalia; inset showing flagellum encircled with a loose tissue before clearing it out. C. Internal structure of penis. D. Internal structure of epiphallus and epiphallic caecum. White arrow indicates the junction between penis and epiphallus." figureDoi="http://doi.org/10.5281/zenodo.5528148" httpUri="https://zenodo.org/record/5528148/files/figure.png" pageId="9" pageNumber="64">5A</figureCitation>
|
||
). Animal with five well-developed mantle lobes. Left and right shell lobes thin, pale yellowish to fleshy-grey colour and spread with small whitish dots. Three dorsal lobes crescent-shaped and smaller than shell lobes.
|
||
</paragraph>
|
||
<paragraph id="8BC8369EA43DFFBB39E2F9D2FDBC4706" blockId="9.[187,1400,1603,1841]" pageId="9" pageNumber="64">
|
||
GENITALIA (
|
||
<figureCitation id="134C2A1BA43DFFBB3819F9D2FE544668" box="[326,457,1603,1630]" captionStart="Fig" captionStartId="12.[189,232,1790,1816]" captionTargetBox="[189,1398,264,1743]" captionTargetId="figure-14@12.[189,1398,264,1743]" captionTargetPageId="12" captionText="Fig. 5. Genital system and mantle lobes of Aenigmatoconcha clivicola Tumpeesuwan & Tumpeesuwan, 2017, specimen CUMZ 7929 from the type locality. A. Mantle lobes. B. General view of genitalia; inset showing flagellum encircled with a loose tissue before clearing it out. C. Internal structure of penis. D. Internal structure of epiphallus and epiphallic caecum. White arrow indicates the junction between penis and epiphallus." figureDoi="http://doi.org/10.5281/zenodo.5528148" httpUri="https://zenodo.org/record/5528148/files/figure.png" pageId="9" pageNumber="64">Fig. 5B–D</figureCitation>
|
||
). Atrium (at) enlarged and very short. Penis (p) rather long cylindrical, penial sheath (ps) very thick and covering entire penis, and penial caecum (pc) rather small protruding. Inner wall of penis covered with trapezoid-shaped penial pilasters (pp) arranged in oblique rows. Epiphallus (e1 + e2) as long as penis: e1 long and slender, and e2 bulbous and about half e1 length. Inner sculpture of e1 with very small and thin longitudinal folds to nearly smooth surface. Inner sculpture of e2 trapezoid to conical pilasters. Epiphallic caecum (ec) long with thick penial retractor muscle (prm) attached at tip. Flagellum (fl) small and short.
|
||
</paragraph>
|
||
<paragraph id="8BC8369EA43DFFBB39E2F8C9FECF478F" blockId="9.[189,1398,1880,1977]" pageId="9" pageNumber="64">
|
||
RADULA (
|
||
<figureCitation id="134C2A1BA43DFFBB3873F8C9FE074744" box="[300,410,1880,1906]" captionStart="Fig" captionStartId="21.[189,232,1823,1849]" captionTargetBox="[194,1394,950,1780]" captionTargetId="figure-243@21.[189,1398,945,1784]" captionTargetPageId="21" captionText="Fig. 10. Radula morphology of Aenigmatoconcha spp. A. A. clivicola Tumpeesuwan & Tumpeesuwan, 2017, specimen CUMZ 7929 from Loei Province. B. A. mitis (Pfeiffer, 1863) comb. nov., specimen CUMZ 7687 from Kanchanaburi Province. C. A. sumonthai Tumpeesuwan & Tumpeesuwan, 2018, specimen CUMZ 7922 from Chumphon Province. D. A. eunetis Pholyotha & Panha sp. nov., paratype CUMZ 7933 from Uthai Thani Province. Central tooth in yellow. Scale bars: A–C = 100 μm; D = 50 μm." figureDoi="http://doi.org/10.5281/zenodo.5528158" httpUri="https://zenodo.org/record/5528158/files/figure.png" pageId="9" pageNumber="64">Fig. 10A</figureCitation>
|
||
). Teeth arranged in anteriorly V-shaped rows with half row consisting of about 70– 76 teeth at middle plate. Central, lateral, and marginal teeth monocuspid and spatulate-shaped with curved cusp.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<caption id="DF086616A43EFFB839E2F91DFC0A4702" ID-DOI="http://doi.org/10.5281/zenodo.5528146" ID-Zenodo-Dep="5528146" httpUri="https://zenodo.org/record/5528146/files/figure.png" pageId="10" pageNumber="65" startId="10.[189,232,1676,1702]" targetBox="[189,1398,264,1638]" targetPageId="10">
|
||
<paragraph id="8BC8369EA43EFFB839E2F91DFC0A4702" blockId="10.[189,1399,1676,1844]" pageId="10" pageNumber="65">
|
||
<emphasis id="B903EA8CA43EFFB839E2F91DFE8C4690" bold="true" box="[189,273,1676,1702]" pageId="10" pageNumber="65">Fig. 4.</emphasis>
|
||
Shells of
|
||
<taxonomicName id="4C774D1DA43EFFB838C8F91CFD514690" box="[407,716,1676,1703]" class="Gastropoda" family="Helicarionidae" genus="Aenigmatoconcha" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="65" phylum="Mollusca" rank="species" species="undefined-a">
|
||
<emphasis id="B903EA8CA43EFFB838C8F91CFDF04691" box="[407,621,1677,1703]" italics="true" pageId="10" pageNumber="65">Aenigmatoconcha</emphasis>
|
||
spp.
|
||
<emphasis id="B903EA8CA43EFFB83BE8F91DFD514690" bold="true" box="[695,716,1676,1702]" pageId="10" pageNumber="65">A</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="B903EA8CA43EFFB83B93F91CFD724691" bold="true" box="[716,751,1677,1703]" pageId="10" pageNumber="65">–B</emphasis>
|
||
.
|
||
<taxonomicName id="4C774D1DA43EFFB83A5CF91CFAEF4691" authority="Tumpeesuwan & Tumpeesuwan, 2017" authorityName="Tumpeesuwan & Tumpeesuwan" authorityYear="2017" box="[771,1394,1677,1703]" class="Gastropoda" family="Helicarionidae" genus="Aenigmatoconcha" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="65" phylum="Mollusca" rank="species" species="clivicola">
|
||
<emphasis id="B903EA8CA43EFFB83A5CF91CFC124691" box="[771,911,1677,1703]" italics="true" pageId="10" pageNumber="65">A. clivicola</emphasis>
|
||
<bibRefCitation id="EFE64B6FA43EFFB83AC3F91CFAEF4691" author="Tumpeesuwan C. & Tumpeesuwan S." box="[924,1394,1677,1703]" pageId="10" pageNumber="65" pagination="170 - 176" refId="ref13001" refString="Tumpeesuwan C. & Tumpeesuwan S. 2017. Discovery of an overlooked helicarionid land snail (Helicarionidae: Durgellinae) from northeastern Thailand, with description of a new genus and new species, and a note on radula morphology and genital system. Raffles Bulletin of Zoology 65: 174 - 181. Tumpeesuwan C. & Tumpeesuwan S. 2018. Aenigmatoconcha sumonthai, a new helicarionid land snail from Chumphon Province, Southern Thailand (Helicarionidae: Durgellinae). Raffles Bulletin of Zoology 66: 170 - 176. http: // doi. org / 10.5281 / zenodo. 4504598" type="journal article" year="2017">Tumpeesuwan & Tumpeesuwan, 2017</bibRefCitation>
|
||
</taxonomicName>
|
||
, specimen CUMZ 7928 from the type locality (A) and specimen CUMZ 7930 from Loei Province (B).
|
||
<emphasis id="B903EA8CA43EFFB839E2F942FF6E46DB" bold="true" box="[189,243,1747,1774]" pageId="10" pageNumber="65">C–F</emphasis>
|
||
.
|
||
<taxonomicName id="4C774D1DA43EFFB8385AF942FDB546D8" authority="(Pfeiffer, 1863)" baseAuthorityName="Pfeiffer" baseAuthorityYear="1863" box="[261,552,1747,1774]" class="Gastropoda" family="Helicarionidae" genus="Aenigmatoconcha" kingdom="Animalia" order="Stylommatophora" pageId="10" pageNumber="65" phylum="Mollusca" rank="species" species="mitis" status="comb. nov.">
|
||
<emphasis id="B903EA8CA43EFFB8385AF942FEF946DB" box="[261,356,1747,1773]" italics="true" pageId="10" pageNumber="65">A. mitis</emphasis>
|
||
(
|
||
<bibRefCitation id="EFE64B6FA43EFFB83828F942FDBD46D8" author="Pfeiffer L." box="[375,544,1747,1774]" pageId="10" pageNumber="65" pagination="268 - 278" refId="ref11649" refString="Pfeiffer L. 1863. Descriptions of thirty-six new land shells from the collection of H. Cuming, Esq. Proceedings of the Zoological Society of London 30: 268 - 278. [Published in parts, dates follow Duncan (1937)]." type="journal article" year="1863">Pfeiffer, 1863</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A23057F7A43EFFB83B6CF942FD2646D8" box="[563,699,1747,1774]" pageId="10" pageNumber="65" rank="species">comb. nov.</taxonomicNameLabel>
|
||
, syntypes NHMUK ex. Cuming collection (C), specimen CUMZ 7921 from Phetchaburi Province (D), specimen CUMZ 7687 from Kanchanaburi Province (E), and specimen CUMZ 7885 from Nakhon Sawan Province (F).
|
||
</paragraph>
|
||
</caption>
|
||
<subSubSection id="C36D6515A43FFFB939E2FE99FDBF4142" pageId="11" pageNumber="66" type="distribution">
|
||
<paragraph id="8BC8369EA43FFFB939E2FE99FECB4114" blockId="11.[189,342,264,290]" box="[189,342,264,290]" pageId="11" pageNumber="66">
|
||
<heading id="D08081F2A43FFFB939E2FE99FECB4114" bold="true" box="[189,342,264,290]" fontSize="11" level="3" pageId="11" pageNumber="66" reason="3">
|
||
<emphasis id="B903EA8CA43FFFB939E2FE99FECB4114" bold="true" box="[189,342,264,290]" pageId="11" pageNumber="66">Distribution</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8BC8369EA43FFFB939E2FEA6FDBF4142" blockId="11.[189,1398,311,372]" pageId="11" pageNumber="66">
|
||
This species is currently known only from 2 localities: the
|
||
<typeStatus id="54CC883CA43FFFB93A26FEA9FC314164" box="[889,940,312,338]" pageId="11" pageNumber="66">type</typeStatus>
|
||
locality (Phu Pha Lom) and limestone hills in
|
||
<collectingRegion id="49B3F87CA43FFFB93849FECBFE264142" box="[278,443,346,372]" country="Thailand" name="Loei" pageId="11" pageNumber="66">Loei Province</collectingRegion>
|
||
(
|
||
<figureCitation id="134C2A1BA43FFFB93894FECBFD8D4142" box="[459,528,346,372]" captionStart="Fig" captionStartId="2.[189,232,1962,1988]" captionTargetBox="[189,1398,678,1922]" captionTargetId="figure-108@2.[189,1398,678,1922]" captionTargetPageId="2" captionText="Fig. 1. Geographic distribution of four species of Aenigmatoconcha Tumpeesuwan & Tumpeesuwan, 2017 based on specimens examined herein." figureDoi="http://doi.org/10.5281/zenodo.5528136" httpUri="https://zenodo.org/record/5528136/files/figure.png" pageId="11" pageNumber="66">Fig. 1</figureCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C36D6515A43FFFB939E2FE0DFED643E4" pageId="11" pageNumber="66" type="discussion">
|
||
<paragraph id="8BC8369EA43FFFB939E2FE0DFEB24180" blockId="11.[189,303,412,438]" box="[189,303,412,438]" pageId="11" pageNumber="66">
|
||
<heading id="D08081F2A43FFFB939E2FE0DFEB24180" bold="true" box="[189,303,412,438]" fontSize="11" level="3" pageId="11" pageNumber="66" reason="3">
|
||
<emphasis id="B903EA8CA43FFFB939E2FE0DFEB24180" bold="true" box="[189,303,412,438]" pageId="11" pageNumber="66">Remarks</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8BC8369EA43FFFB939E2FE5BFED643E4" blockId="11.[188,1399,458,978]" pageId="11" pageNumber="66">
|
||
The genitalia were originally described but without examination of the internal sculpture of penis. In this study, we examined
|
||
<specimenCount id="9D71FD17A43FFFB938EEFE7FFD8E4231" box="[433,531,493,520]" pageId="11" pageNumber="66" type="adult">20 adult</specimenCount>
|
||
topotypic specimens to provide descriptions of the penial sheath, penial caecum, and flagellum that were not included in the original description. The penial sheath is very large and covers the entire penis (
|
||
<figureCitation id="134C2A1BA43FFFB93B5CFDA5FDC34279" box="[515,606,564,591]" captionStart="Fig" captionStartId="12.[189,232,1790,1816]" captionTargetBox="[189,1398,264,1743]" captionTargetId="figure-14@12.[189,1398,264,1743]" captionTargetPageId="12" captionText="Fig. 5. Genital system and mantle lobes of Aenigmatoconcha clivicola Tumpeesuwan & Tumpeesuwan, 2017, specimen CUMZ 7929 from the type locality. A. Mantle lobes. B. General view of genitalia; inset showing flagellum encircled with a loose tissue before clearing it out. C. Internal structure of penis. D. Internal structure of epiphallus and epiphallic caecum. White arrow indicates the junction between penis and epiphallus." figureDoi="http://doi.org/10.5281/zenodo.5528148" httpUri="https://zenodo.org/record/5528148/files/figure.png" pageId="11" pageNumber="66">Fig. 5B</figureCitation>
|
||
). The short penial caecum, an extension of the penis, is located near the penis and epiphallus junction (
|
||
<figureCitation id="134C2A1BA43FFFB93B04FDC6FD274244" box="[603,698,599,626]" captionStart="Fig" captionStartId="12.[189,232,1790,1816]" captionTargetBox="[189,1398,264,1743]" captionTargetId="figure-14@12.[189,1398,264,1743]" captionTargetPageId="12" captionText="Fig. 5. Genital system and mantle lobes of Aenigmatoconcha clivicola Tumpeesuwan & Tumpeesuwan, 2017, specimen CUMZ 7929 from the type locality. A. Mantle lobes. B. General view of genitalia; inset showing flagellum encircled with a loose tissue before clearing it out. C. Internal structure of penis. D. Internal structure of epiphallus and epiphallic caecum. White arrow indicates the junction between penis and epiphallus." figureDoi="http://doi.org/10.5281/zenodo.5528148" httpUri="https://zenodo.org/record/5528148/files/figure.png" pageId="11" pageNumber="66">Fig. 5B</figureCitation>
|
||
). This character is visible when penial sheath is removed. Its internal sculpture is rather smooth, unlike the penial sculpture, with its trapezoid-shaped pilasters (
|
||
<figureCitation id="134C2A1BA43FFFB93999FD0CFEBF428E" box="[198,290,669,696]" captionStart="Fig" captionStartId="12.[189,232,1790,1816]" captionTargetBox="[189,1398,264,1743]" captionTargetId="figure-14@12.[189,1398,264,1743]" captionTargetPageId="12" captionText="Fig. 5. Genital system and mantle lobes of Aenigmatoconcha clivicola Tumpeesuwan & Tumpeesuwan, 2017, specimen CUMZ 7929 from the type locality. A. Mantle lobes. B. General view of genitalia; inset showing flagellum encircled with a loose tissue before clearing it out. C. Internal structure of penis. D. Internal structure of epiphallus and epiphallic caecum. White arrow indicates the junction between penis and epiphallus." figureDoi="http://doi.org/10.5281/zenodo.5528148" httpUri="https://zenodo.org/record/5528148/files/figure.png" pageId="11" pageNumber="66">Fig. 5C</figureCitation>
|
||
). The Australian helicarionids,
|
||
<taxonomicName id="4C774D1DA43FFFB93BD0FD0FFC66428E" authority="Stanisic, 2010" authorityName="Stanisic" authorityYear="2010" box="[655,1019,669,696]" class="Gastropoda" family="Helicarionidae" genus="Nitor" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="66" phylum="Mollusca" rank="species" species="whitneyae">
|
||
<emphasis id="B903EA8CA43FFFB93BD0FD0FFCD04281" box="[655,845,669,695]" italics="true" pageId="11" pageNumber="66">Nitor whitneyae</emphasis>
|
||
Stanisic, 2010
|
||
</taxonomicName>
|
||
has a penial caecum while other
|
||
<taxonomicName id="4C774D1DA43FFFB939E2FD50FF6142EC" authorityName="Gude" authorityYear="1911" box="[189,252,705,730]" class="Gastropoda" family="Helicarionidae" genus="Nitor" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="66" phylum="Mollusca" rank="genus">
|
||
<emphasis id="B903EA8CA43FFFB939E2FD50FF6142EC" box="[189,252,705,730]" italics="true" pageId="11" pageNumber="66">Nitor</emphasis>
|
||
</taxonomicName>
|
||
taxa do not have a penial caecum (Hyman & Köhler 2018). In Southeast Asian ariophantids, the presence or absence of a penial caecum is a discriminating character among species in genera such as
|
||
<taxonomicName id="4C774D1DA43FFFB939E1FC96FE644317" authority="Gray, 1847" authorityName="Gray" authorityYear="1847" box="[190,505,775,801]" class="Gastropoda" family="Ariophantidae" genus="Macrochlamys" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="66" phylum="Mollusca" rank="genus">
|
||
<emphasis id="B903EA8CA43FFFB939E1FC96FEF04317" box="[190,365,775,801]" italics="true" pageId="11" pageNumber="66">Macrochlamys</emphasis>
|
||
Gray, 1847
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C774D1DA43FFFB93B69FC96FB9F4317" authority="Pholyotha & Panha, 2020" authorityName="Pholyotha & Panha" authorityYear="2020" box="[566,1026,775,801]" class="Gastropoda" family="Ariophantidae" genus="Taphrenalla" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="66" phylum="Mollusca" rank="genus">
|
||
<emphasis id="B903EA8CA43FFFB93B69FC96FD584317" box="[566,709,775,801]" italics="true" pageId="11" pageNumber="66">Taphrenalla</emphasis>
|
||
<bibRefCitation id="EFE64B6FA43FFFB93B92FC96FB9F4317" author="Pholyotha A. & Sutcharit C. & Tongkerd P. & Panha S." box="[717,1026,775,801]" pageId="11" pageNumber="66" pagination="1 - 100" refId="ref11850" refString="Pholyotha A., Sutcharit C., Tongkerd P. & Panha S. 2020 b. Integrative taxonomic revision of the land snail genus Sarika Godwin-Austen, 1907 in Thailand, with descriptions of nine new species (Eupulmonata, Ariophantidae). ZooKeys 976: 1 - 100. https: // doi. org / 10.3897 / zookeys. 976.53859" type="journal article" year="2020">Pholyotha & Panha, 2020</bibRefCitation>
|
||
</taxonomicName>
|
||
, and is supported by molecular studies (
|
||
<bibRefCitation id="EFE64B6FA43FFFB93840FCBBFD844372" author="Pholyotha A. & Sutcharit C. & Panha S." box="[287,537,810,836]" pageId="11" pageNumber="66" pagination="763 - 781" refId="ref11732" refString="Pholyotha A., Sutcharit C. & Panha S. 2018. The land snail genus Macrochlamys Gray, 1847 from Thailand, with descriptions of five new species (Pulmonata: Ariophantidae). Raffles Bulletin of Zoology 66: 763 - 781." type="journal article" year="2018">
|
||
Pholyotha
|
||
<emphasis id="B903EA8CA43FFFB938C2FCBAFE4A4372" box="[413,471,810,836]" italics="true" pageId="11" pageNumber="66">et al.</emphasis>
|
||
2018
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFE64B6FA43FFFB93B79FCBBFDFC4372" author="Pholyotha A. & Sutcharit C. & Tongkerd P. & Jeratthitikul E. & Panha S." box="[550,609,810,836]" pageId="11" pageNumber="66" pagination="21 - 69" refId="ref11914" refString="Pholyotha A., Sutcharit C., Tongkerd P., Jeratthitikul E. & Panha S. 2021. Integrative systematics reveals the new land-snail genus Taphrenalla (Eupulmonata: Ariophantidae) with a description of nine new species from Thailand. Contributions to Zoology 90 (1): 21 - 69. https: // doi. org / 10.1163 / 18759866 - BJA 10013" type="journal article" year="2021">2021</bibRefCitation>
|
||
). The flagellum of
|
||
<taxonomicName id="4C774D1DA43FFFB93A1EFCBAFC554372" authorityName="Tumpeesuwan & Tumpeesuwan" authorityYear="2017" box="[833,968,810,837]" class="Gastropoda" family="Helicarionidae" genus="Aenigmatoconcha" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="66" phylum="Mollusca" rank="species" species="clivicola">
|
||
<emphasis id="B903EA8CA43FFFB93A1EFCBAFC554372" box="[833,968,810,837]" italics="true" pageId="11" pageNumber="66">A. clivicola</emphasis>
|
||
</taxonomicName>
|
||
is an extension of the epiphallus and is located near the insertion point of the vas deferens. It is somewhat small and short, and bound to the vas deferens by thin connective tissue. This feature is important for spermatophore formation before copulation (
|
||
<bibRefCitation id="EFE64B6FA43FFFB93815FC05FE434398" author="Tompa A. S." box="[330,478,916,942]" pageId="11" pageNumber="66" pagination="47 - 140" refId="ref12846" refString="Tompa A. S. 1984. Land snails (Stylommatophora). In: Tompa A. S. & Verdonk N. H. (eds) The Mollusca, Vol. 7: Reproduction: 47 - 140. Academic Press, London." type="book chapter" year="1984">Tompa 1984</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFE64B6FA43FFFB938B2FC05FDF44398" author="Baur B." box="[493,617,916,942]" pageId="11" pageNumber="66" pagination="197 - 217" refId="ref9873" refString="Baur B. 2010. Stylommatophoran Gastropods. In: Leonard J. L. & Cordoba-Aguilar A. (eds) The Evolution of Primary Sexual Characters in Animals: 197 - 217. Oxford University Press, USA." type="book chapter" year="2010">Baur 2010</bibRefCitation>
|
||
). However, during this study no spermatophores were observed in
|
||
<taxonomicName id="4C774D1DA43FFFB939E2FC26FED943E7" authorityName="Tumpeesuwan & Tumpeesuwan" authorityYear="2017" box="[189,324,951,977]" class="Gastropoda" family="Helicarionidae" genus="Aenigmatoconcha" kingdom="Animalia" order="Stylommatophora" pageId="11" pageNumber="66" phylum="Mollusca" rank="species" species="clivicola">
|
||
<emphasis id="B903EA8CA43FFFB939E2FC26FED943E7" box="[189,324,951,977]" italics="true" pageId="11" pageNumber="66">A. clivicola</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |