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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.1119.85400" ID-GBIF-Dataset="418bd521-e4b9-4ee6-981f-a6f471eab287" ID-Pensoft-Pub="1313-2970-1119-1" ID-Pensoft-UUID="F158A7C5261D52B69288A62F3C777CAF" ID-ZooBank="A3BE91C6B79344E1A886A803BF104D8B" ModsDocID="1313-2970-1119-1" checkinTime="1661532673136" checkinUser="pensoft" docAuthor="Jirapatrasilp, Parin, Sutcharit, Chirasak &amp; Panha, Somsak" docDate="2022" docId="3ED453755B705B66888718AF13B99A51" docLanguage="en" docName="ZooKeys 1119: 1-115" docOrigin="ZooKeys 1119" docPubDate="2022-08-25" docSource="http://dx.doi.org/10.3897/zookeys.1119.85400" docTitle="Pupina peguensis Benson 1860" docType="treatment" docVersion="4" id="F158A7C5261D52B69288A62F3C777CAF" lastPageNumber="1" masterDocId="F158A7C5261D52B69288A62F3C777CAF" masterDocTitle="Annotated checklist of the operculated land snails from Thailand (Mollusca, Gastropoda, Caenogastropoda): the family Pupinidae, with descriptions of several new species and subspecies, and notes on classification of Pupina Vignard, 1829 and Pupinella Gray, 1850 from mainland Southeast Asia" masterLastPageNumber="115" masterPageNumber="1" pageNumber="1" updateTime="1661537468408" updateUser="ExternalLinkService">
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<mods:title>Annotated checklist of the operculated land snails from Thailand (Mollusca, Gastropoda, Caenogastropoda): the family Pupinidae, with descriptions of several new species and subspecies, and notes on classification of Pupina Vignard, 1829 and Pupinella Gray, 1850 from mainland Southeast Asia</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Jirapatrasilp, Parin</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-5591-6724</mods:nameIdentifier>
<mods:affiliation>Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand</mods:affiliation>
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<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Sutcharit, Chirasak</mods:namePart>
<mods:affiliation>Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand</mods:affiliation>
<mods:nameIdentifier type="email">jirasak4@yahoo.com</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Panha, Somsak</mods:namePart>
<mods:affiliation>Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Bangkok 10330, Thailand &amp; Academy of Science, The Royal Society of Thailand, Bangkok 10300, Thailand</mods:affiliation>
<mods:nameIdentifier type="email">somsak.pan@chula.ac.th</mods:nameIdentifier>
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<mods:title>ZooKeys</mods:title>
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<mods:date>2022</mods:date>
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<mods:number>2022-08-25</mods:number>
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<mods:number>1119</mods:number>
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<mods:url>http://dx.doi.org/10.3897/zookeys.1119.85400</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.1119.85400</mods:identifier>
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<treatment ID-GBIF-Taxon="199839009" LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51" httpUri="http://treatment.plazi.org/id/3ED453755B705B66888718AF13B99A51" lastPageNumber="1" pageId="0" pageNumber="1">
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<taxonomicName LSID="3ED45375-5B70-5B66-8887-18AF13B99A51" authority="Benson, 1860" authorityName="Benson" authorityYear="1860" class="Gastropoda" family="Pupinidae" genus="Pupina" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Pupina peguensis" order="Architaenioglossa" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="peguensis">Pupina peguensis Benson, 1860</taxonomicName>
</paragraph>
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<paragraph pageId="0" pageNumber="1">
<figureCitation captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. External shell morphology of three mainland Southeast Asian Pupina species groups A Pupina artata from the Pupina artata species group, CUMZ 12003 from Ban Ping Khong, Chiang Mai B Pupina peguensis from the Pupina arula species group, CUMZ 12094 from Khao Tham Phra Temple, Chiang Rai C Pupina siamensis from the Pupina arula species group, CUMZ 12052 from Sri Thong Cave, Sra Keo, and D Pupina tchehelensis from the Pupina aureola species group, CUMZ 12136 from limestone mountain, Phang Nga. Red frames focus on the parietal tooth and posterior canal; blue frames focus on the curvature of the apertural lip when observed from lateral view." figureDoi="10.3897/zookeys.1119.85400.figure18" httpUri="https://binary.pensoft.net/fig/735379" pageId="0" pageNumber="1">Figs 18B</figureCitation>
<figureCitation captionStart="Figure 25" captionStartId="F25" captionText="Figure 25. Live specimens of A-C Pupina artata: specimens A CUMZ 12020 from Buri Ratchawanaram Temple, Ratchaburi B CUMZ 12022 from Golden Dragon Cave, Ratchaburi, and C CUMZ 12037 from Wua Ta Lap Island, Surat Thani D, E Pupina bensoni sp. nov. D paratype CUMZ 12045 / 2 from Khao Wong Cave, Uthai Thani and E specimen CUMZ 12047 from Tham Namthip Bureau of Monks, Uthai Thani F, G Pupina peguensis: specimens F CUMZ 12050 from Chai Thong Wararam Temple, Nakhon Sawan and G CUMZ 12051 from Tham Saeng Wiset Bureau of Monks, Nakhon Sawan H Pupina siamensis, specimen CUMZ 12069 from Khao Chi Chan Buddha Image, Chonburi. All not to scale." figureDoi="10.3897/zookeys.1119.85400.figure25" httpUri="https://binary.pensoft.net/fig/735386" pageId="0" pageNumber="1">, 25F, G</figureCitation>
<figureCitation captionStart="Figure 26" captionStartId="F26" captionText="Figure 26. A Pupina bensoni sp. nov., paratype CUMZ 12046 / 1 from Khao Wong Cave, Uthai Thani B Pupina hungerfordiana, specimen NHMUK 91.3.14.686 - 7 from Hsaddan Koo C Pupina billeti, holotype MNHN-IM- 2000 - 35841 D Pupina verneaui, syntype MNHN-IM- 2000 - 35843 from Ha-Giang, Tonkin E, F Pupina peguensis E syntype of Pupina blanfordi NHMUK 1888.12.4.100 from Pegu and F specimen NHMUK ex. Cuming coll. from Lao Mountains, Camboja. Photo: P. Maestrati, MNHN (C, D)." figureDoi="10.3897/zookeys.1119.85400.figure26" httpUri="https://binary.pensoft.net/fig/735387" pageId="0" pageNumber="1">, 26E, F</figureCitation>
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Shells of Pupina arula species group from mainland Southeast Asia A-G Pupina peguensis A syntype of Pupina blanfordi NHMUK 1888.12.4.100 and specimens B CUMZ 12105 C CUMZ 12050 D CUMZ 12103 E NHMUK ex. Cuming coll. F CUMZ 12094, and G CUMZ 12108 H-J Pupina crosseana H lectotype MNHN-IM- 2000 - 35834 I paralectotype RBINS MT 966 / 10591, and J specimen CUMZ 12049 K, L Pupina siamensis K lectotype SMF 109948 and L specimen CUMZ 12052 M-P Pupina bilabiata sp. nov. M holotype CUMZ 12073 / 1 and specimens N CUMZ 12079 O CUMZ 12081, and P CUMZ 12086 Q, R Pupina godwinausteni sp. nov. Q holotype CUMZ 12090 / 1 and R paratype CUMZ 12090 / 2. Photo: P. Maestrati, MNHN (H), F. Trus, RBINS (I)." figureDoi="10.3897/zookeys.1119.85400.figure28" httpUri="https://binary.pensoft.net/fig/735389" pageId="0" pageNumber="1">, 28A-G</figureCitation>
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. A-D Pupina peguensis: specimens A CUMZ 12050 from Chai Thong Wararam Temple, Nakhon Sawan B CUMZ 12105 from Thep Phithak Punnaram Temple, Nakhon Ratchasima C CUMZ 12108 from Tham Thep Bandan Temple, Phetchabun, and D CUMZ 12094 from Khao Tham Phra Temple, Chiang Rai E, F Pupina crosseana E lectotype MNHN-IM- 2000 - 35834 from Cambodge and F paralectotype RBINS MT 966 / 10591 from Phnom-Rohan, Cambodge. Photo: P. Maestrati, MNHN (E), F. Trus, RBINS (F)." figureDoi="10.3897/zookeys.1119.85400.figure30" httpUri="https://binary.pensoft.net/fig/735391" pageId="0" pageNumber="1">, 30A-D</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="reference_group">
<paragraph pageId="0" pageNumber="1">
<taxonomicName authorityName="Benson" authorityYear="1860" class="Gastropoda" family="Pupinidae" genus="Pupina" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Pupina peguensis" order="Architaenioglossa" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="peguensis">Pupina peguensis</taxonomicName>
Benson, 1860: 192, 193. Type locality: Pegu [Bago Region, Myanmar].
<bibRefCitation DOI="https://doi.org/10.5962/bhl.title.11957" author="Nevill, G" journalOrPublisher="Indian Museum, Calcutta" pageId="0" pageNumber="1" refId="B104" refString="Nevill, G, 1878. Hand List of Mollusca in the Indian Museum, Calcutta. Part 1. Gastropoda. Pulmonata and Prosobranchia-Neurobranchia. Indian Museum, Calcutta, DOI: https://doi.org/10.5962/bhl.title.11957" title="Hand List of Mollusca in the Indian Museum, Calcutta. Part 1. Gastropoda. Pulmonata and Prosobranchia-Neurobranchia." url="https://doi.org/10.5962/bhl.title.11957" year="1878">Nevill 1878</bibRefCitation>
: 300, Shuay-Gheen, Burma [Shwegyin, Bago Region, Myanmar]; Zwagabin [Zwekabin Taung mountain, Hpa-An District, Kayin State, Myanmar].
<bibRefCitation author="Tripathy, B" journalOrPublisher="Records of the Zoological Survey of India" pageId="0" pageNumber="1" pagination="508 - 510" refId="B177" refString="Tripathy, B, Sajan, S, 2019. Pupina peguensis Benson, 1860 (Mollusca: Pupinidae): Tracing a long lost molluscan type. Records of the Zoological Survey of India 119: 508 - 510" title="Pupina peguensis Benson, 1860 (Mollusca: Pupinidae): Tracing a long lost molluscan type." volume="119" year="2019">Tripathy and Sajan 2019</bibRefCitation>
: 508, fig. 1.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<taxonomicName authorityName="Theobald" authorityYear="1864" class="Gastropoda" family="Pupinidae" genus="Pupina" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Pupina blanfordi" order="Architaenioglossa" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="blanfordi">Pupina blanfordi</taxonomicName>
Theobald, 1864: 247, 248. Type locality: Pegu. Hanley and Theobald 1870: 4, pl. 7, fig. 6.
<bibRefCitation author="Reeve, L" journalOrPublisher="Journal of Systematic Palaeontology" pageId="0" pageNumber="1" refId="B139" refString="Reeve, L, 1878. Conchologia Iconica: Illustrations of the shells of molluscous animals. Volume 20, Monograph of the genus Pupinidae, pls 1-10. Lovell Reeve &amp; Co, London, [without pagination]." title="Conchologia Iconica: Illustrations of the shells of molluscous animals. Volume 20, Monograph of the genus Pupinidae, pls 1 - 10. Lovell Reeve &amp; Co, London, [without pagination]." year="1878">Reeve 1878</bibRefCitation>
:
<taxonomicName authorityName="L. Pfeiffer" authorityYear="1853" family="Pupinidae" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="" order="Architaenioglossa" pageId="0" pageNumber="1" rank="family">Pupinidae</taxonomicName>
, pl. 1, sp. 6.
<bibRefCitation author="Godwin-Austen, HH" journalOrPublisher="Journal of Malacology" pageId="0" pageNumber="1" refId="B51" refString="Godwin-Austen, HH, 1897. Land and freshwater mollusca of India, including South Arabia, Baluchistan, Afghanistan, Kashmir, Nepal, Burma, Pegu, Tenasserim, Malaya Peninsula, Ceylon and other islands of the Indian Ocean. Vol. 2, Part 7: 1-46." title="Land and freshwater mollusca of India, including South Arabia, Baluchistan, Afghanistan, Kashmir, Nepal, Burma, Pegu, Tenasserim, Malaya Peninsula, Ceylon and other islands of the Indian Ocean. Vol. 2, Part 7: 1 - 46." year="1897">Godwin-Austen 1897</bibRefCitation>
: 41, pl. 69, fig. 2, 2a, b. Syn. nov.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<taxonomicName authorityName="Theobald" authorityYear="1864" class="Gastropoda" family="Pupinidae" genus="Pupina" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Pupina (Tylotoechus) blanfordi" order="Architaenioglossa" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="blanfordi" subGenus="Tylotoechus">Pupina (Tylotoechus) blanfordi</taxonomicName>
-
<bibRefCitation author="Kobelt, W" journalOrPublisher="Friedlaender und Sohn, Berlin" pageId="0" pageNumber="1" refId="B76" refString="Kobelt, W, 1902. Cyclophoridae. Das Tierreich. R. Friedlaender und Sohn, Berlin" title="Cyclophoridae. Das Tierreich. R." year="1902">Kobelt 1902</bibRefCitation>
: 309, 310.
<bibRefCitation author="Gude, GK" journalOrPublisher="Taylor and Francis, London" pageId="0" pageNumber="1" refId="B60" refString="Gude, GK, 1921. The Fauna of British India including Ceylon and Burma. Mollusca-- III. Land Operculates. Taylor and Francis, London" title="The Fauna of British India including Ceylon and Burma. Mollusca- - III. Land Operculates." year="1921">Gude 1921</bibRefCitation>
: 194.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<taxonomicName authorityName="Benson" authorityYear="1860" class="Gastropoda" family="Pupinidae" genus="Pupina" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Pupina (Tylotoechus) peguensis" order="Architaenioglossa" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="peguensis" subGenus="Tylotoechus">Pupina (Tylotoechus) peguensis</taxonomicName>
-
<bibRefCitation author="Kobelt, W" journalOrPublisher="Friedlaender und Sohn, Berlin" pageId="0" pageNumber="1" refId="B76" refString="Kobelt, W, 1902. Cyclophoridae. Das Tierreich. R. Friedlaender und Sohn, Berlin" title="Cyclophoridae. Das Tierreich. R." year="1902">Kobelt 1902</bibRefCitation>
: 319.
<bibRefCitation author="Gude, GK" journalOrPublisher="Taylor and Francis, London" pageId="0" pageNumber="1" refId="B60" refString="Gude, GK, 1921. The Fauna of British India including Ceylon and Burma. Mollusca-- III. Land Operculates. Taylor and Francis, London" title="The Fauna of British India including Ceylon and Burma. Mollusca- - III. Land Operculates." year="1921">Gude 1921</bibRefCitation>
: 197.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<taxonomicName authorityName="Pfeiffer" authorityYear="1861" class="Gastropoda" family="Pupinidae" genus="Pupina" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Pupina mouhoti" order="Architaenioglossa" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="mouhoti">Pupina mouhoti</taxonomicName>
[non Pfeiffer]-
<bibRefCitation author="Bavay, A" journalOrPublisher="Journal de Conchyliologie" pageId="0" pageNumber="1" refId="B8" refString="BEDO (2017) Land Snails: Checklist of Molluscan Biodiversity in Thailand.BEDO, Bangkok, 300 pp. [in Thai]" year="1904">BEDO 2017</bibRefCitation>
: 91.
<bibRefCitation author="Sutcharit, C" journalOrPublisher="Zino Publishing, Bangkok" pageId="0" pageNumber="1" refId="B156" refString="Sutcharit, C, Tongkerd, P, Panha, S, 2018. Land snails: The invaluable bio-resources for the Kingdom of Thailand. Zino Publishing, Bangkok" title="Land snails: The invaluable bio-resources for the Kingdom of Thailand." year="2018">Sutcharit et al. 2018</bibRefCitation>
: figs 4-2-7, 5-13h.
<bibRefCitation DOI="https://doi.org/10.3897/zookeys.834.28800" author="Inkhavilay, K" journalOrPublisher="ZooKeys" pageId="0" pageNumber="1" pagination="1 - 166" refId="B71" refString="Inkhavilay, K, Sutcharit, C, Bantaowong, U, Chanabun, R, Siriwut, W, Srisonchai, R, Pholyotha, A, Jirapatrasilp, P, Panha, S, 2019. Annotated checklist of the terrestrial molluscs from Laos (Mollusca, Gastropoda). ZooKeys 834: 1 - 166, DOI: https://doi.org/10.3897/zookeys.834.28800" title="Annotated checklist of the terrestrial molluscs from Laos (Mollusca, Gastropoda)." url="https://doi.org/10.3897/zookeys.834.28800" volume="834" year="2019">Inkhavilay et al. 2019</bibRefCitation>
: 46, fig. 15g, Ngoy Town, Ngoy District, Luang Phrabang Province, Laos.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="type material">
<paragraph pageId="0" pageNumber="1">Type material examined.</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Holotype</emphasis>
of
<taxonomicName authorityName="Benson" authorityYear="1860" class="Gastropoda" family="Pupinidae" genus="Pupina" kingdom="Animalia" lsidName="Pupina peguensis" order="Architaenioglossa" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="peguensis">
<emphasis italics="true" pageId="0" pageNumber="1">Pupina peguensis</emphasis>
</taxonomicName>
NZSI M.32940/9 from
<normalizedToken originalValue="Shuay-Gheen">'Shuay-Gheen'</normalizedToken>
, Burma figured in
<bibRefCitation author="Tripathy, B" journalOrPublisher="Records of the Zoological Survey of India" pageId="0" pageNumber="1" pagination="508 - 510" refId="B177" refString="Tripathy, B, Sajan, S, 2019. Pupina peguensis Benson, 1860 (Mollusca: Pupinidae): Tracing a long lost molluscan type. Records of the Zoological Survey of India 119: 508 - 510" title="Pupina peguensis Benson, 1860 (Mollusca: Pupinidae): Tracing a long lost molluscan type." volume="119" year="2019">Tripathy and Sajan (2019</bibRefCitation>
: fig. 1).
<emphasis bold="true" italics="true" pageId="0" pageNumber="1">Syntype</emphasis>
of
<taxonomicName authorityName="Theobald" authorityYear="1864" class="Gastropoda" family="Pupinidae" genus="Pupina" kingdom="Animalia" lsidName="Pupina blanfordi" order="Architaenioglossa" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="blanfordi">
<emphasis italics="true" pageId="0" pageNumber="1">Pupina blanfordi</emphasis>
</taxonomicName>
NHMUK 1888.12.4.100 (1 shell; Figs
<figureCitation captionStart="Figure 26" captionStartId="F26" captionText="Figure 26. A Pupina bensoni sp. nov., paratype CUMZ 12046 / 1 from Khao Wong Cave, Uthai Thani B Pupina hungerfordiana, specimen NHMUK 91.3.14.686 - 7 from Hsaddan Koo C Pupina billeti, holotype MNHN-IM- 2000 - 35841 D Pupina verneaui, syntype MNHN-IM- 2000 - 35843 from Ha-Giang, Tonkin E, F Pupina peguensis E syntype of Pupina blanfordi NHMUK 1888.12.4.100 from Pegu and F specimen NHMUK ex. Cuming coll. from Lao Mountains, Camboja. Photo: P. Maestrati, MNHN (C, D)." figureDoi="10.3897/zookeys.1119.85400.figure26" httpUri="https://binary.pensoft.net/fig/735387" pageId="0" pageNumber="1">26E</figureCitation>
,
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Shells of Pupina arula species group from mainland Southeast Asia A-G Pupina peguensis A syntype of Pupina blanfordi NHMUK 1888.12.4.100 and specimens B CUMZ 12105 C CUMZ 12050 D CUMZ 12103 E NHMUK ex. Cuming coll. F CUMZ 12094, and G CUMZ 12108 H-J Pupina crosseana H lectotype MNHN-IM- 2000 - 35834 I paralectotype RBINS MT 966 / 10591, and J specimen CUMZ 12049 K, L Pupina siamensis K lectotype SMF 109948 and L specimen CUMZ 12052 M-P Pupina bilabiata sp. nov. M holotype CUMZ 12073 / 1 and specimens N CUMZ 12079 O CUMZ 12081, and P CUMZ 12086 Q, R Pupina godwinausteni sp. nov. Q holotype CUMZ 12090 / 1 and R paratype CUMZ 12090 / 2. Photo: P. Maestrati, MNHN (H), F. Trus, RBINS (I)." figureDoi="10.3897/zookeys.1119.85400.figure28" httpUri="https://binary.pensoft.net/fig/735389" pageId="0" pageNumber="1">28A</figureCitation>
) from Pegu.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="materials_examined">
<paragraph pageId="0" pageNumber="1">Other material examined.</paragraph>
<paragraph pageId="0" pageNumber="1">
<materialsCitation collectingDate="2017-06-09" collectorName="Specimen, Cuming" country="Camboja" county="Pfeiffer" location="Lao Mountains" municipality="Figs" specimenCount="73" stateProvince="Tak Fa District, Nakhon Sawan Province">
<collectorName>Specimen</collectorName>
NHMUK ex.
<collectorName>Cuming</collectorName>
coll. (1 shell; labelled as
<taxonomicName authorityName="Pfeiffer" authorityYear="1861" class="Gastropoda" family="Pupinidae" genus="Pupina" kingdom="Animalia" lsidName="Pupina mouhoti" order="Architaenioglossa" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="mouhoti">
<emphasis italics="true" pageId="0" pageNumber="1">Pupina mouhoti</emphasis>
</taxonomicName>
,
<collectingCounty>Pfeiffer</collectingCounty>
;
<collectingMunicipality>Figs</collectingMunicipality>
<figureCitation captionStart="Figure 26" captionStartId="F26" captionText="Figure 26. A Pupina bensoni sp. nov., paratype CUMZ 12046 / 1 from Khao Wong Cave, Uthai Thani B Pupina hungerfordiana, specimen NHMUK 91.3.14.686 - 7 from Hsaddan Koo C Pupina billeti, holotype MNHN-IM- 2000 - 35841 D Pupina verneaui, syntype MNHN-IM- 2000 - 35843 from Ha-Giang, Tonkin E, F Pupina peguensis E syntype of Pupina blanfordi NHMUK 1888.12.4.100 from Pegu and F specimen NHMUK ex. Cuming coll. from Lao Mountains, Camboja. Photo: P. Maestrati, MNHN (C, D)." figureDoi="10.3897/zookeys.1119.85400.figure26" httpUri="https://binary.pensoft.net/fig/735387" pageId="0" pageNumber="1">26F</figureCitation>
,
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Shells of Pupina arula species group from mainland Southeast Asia A-G Pupina peguensis A syntype of Pupina blanfordi NHMUK 1888.12.4.100 and specimens B CUMZ 12105 C CUMZ 12050 D CUMZ 12103 E NHMUK ex. Cuming coll. F CUMZ 12094, and G CUMZ 12108 H-J Pupina crosseana H lectotype MNHN-IM- 2000 - 35834 I paralectotype RBINS MT 966 / 10591, and J specimen CUMZ 12049 K, L Pupina siamensis K lectotype SMF 109948 and L specimen CUMZ 12052 M-P Pupina bilabiata sp. nov. M holotype CUMZ 12073 / 1 and specimens N CUMZ 12079 O CUMZ 12081, and P CUMZ 12086 Q, R Pupina godwinausteni sp. nov. Q holotype CUMZ 12090 / 1 and R paratype CUMZ 12090 / 2. Photo: P. Maestrati, MNHN (H), F. Trus, RBINS (I)." figureDoi="10.3897/zookeys.1119.85400.figure28" httpUri="https://binary.pensoft.net/fig/735389" pageId="0" pageNumber="1">28E</figureCitation>
) from
<location LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51:004F0C76487474922D67EFD0AF9250F9" country="Camboja" county="Pfeiffer" municipality="Figs" name="Lao Mountains" stateProvince="Tak Fa District, Nakhon Sawan Province">Lao Mountains</location>
,
<collectingCountry name="Cambodia">Camboja</collectingCountry>
. CUMZ 12050 (78 shells and
<specimenCount type="generic">73 specimens</specimenCount>
in ethanol;
<location LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51:79CA69FD298B2B6382D28896A93A8464" country="Camboja" county="Pfeiffer" municipality="Figs" name="Figs" stateProvince="Tak Fa District, Nakhon Sawan Province">Figs</location>
<figureCitation captionStart="Figure 25" captionStartId="F25" captionText="Figure 25. Live specimens of A-C Pupina artata: specimens A CUMZ 12020 from Buri Ratchawanaram Temple, Ratchaburi B CUMZ 12022 from Golden Dragon Cave, Ratchaburi, and C CUMZ 12037 from Wua Ta Lap Island, Surat Thani D, E Pupina bensoni sp. nov. D paratype CUMZ 12045 / 2 from Khao Wong Cave, Uthai Thani and E specimen CUMZ 12047 from Tham Namthip Bureau of Monks, Uthai Thani F, G Pupina peguensis: specimens F CUMZ 12050 from Chai Thong Wararam Temple, Nakhon Sawan and G CUMZ 12051 from Tham Saeng Wiset Bureau of Monks, Nakhon Sawan H Pupina siamensis, specimen CUMZ 12069 from Khao Chi Chan Buddha Image, Chonburi. All not to scale." figureDoi="10.3897/zookeys.1119.85400.figure25" httpUri="https://binary.pensoft.net/fig/735386" pageId="0" pageNumber="1">25F</figureCitation>
,
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Shells of Pupina arula species group from mainland Southeast Asia A-G Pupina peguensis A syntype of Pupina blanfordi NHMUK 1888.12.4.100 and specimens B CUMZ 12105 C CUMZ 12050 D CUMZ 12103 E NHMUK ex. Cuming coll. F CUMZ 12094, and G CUMZ 12108 H-J Pupina crosseana H lectotype MNHN-IM- 2000 - 35834 I paralectotype RBINS MT 966 / 10591, and J specimen CUMZ 12049 K, L Pupina siamensis K lectotype SMF 109948 and L specimen CUMZ 12052 M-P Pupina bilabiata sp. nov. M holotype CUMZ 12073 / 1 and specimens N CUMZ 12079 O CUMZ 12081, and P CUMZ 12086 Q, R Pupina godwinausteni sp. nov. Q holotype CUMZ 12090 / 1 and R paratype CUMZ 12090 / 2. Photo: P. Maestrati, MNHN (H), F. Trus, RBINS (I)." figureDoi="10.3897/zookeys.1119.85400.figure28" httpUri="https://binary.pensoft.net/fig/735389" pageId="0" pageNumber="1">28C</figureCitation>
,
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. A-D Pupina peguensis: specimens A CUMZ 12050 from Chai Thong Wararam Temple, Nakhon Sawan B CUMZ 12105 from Thep Phithak Punnaram Temple, Nakhon Ratchasima C CUMZ 12108 from Tham Thep Bandan Temple, Phetchabun, and D CUMZ 12094 from Khao Tham Phra Temple, Chiang Rai E, F Pupina crosseana E lectotype MNHN-IM- 2000 - 35834 from Cambodge and F paralectotype RBINS MT 966 / 10591 from Phnom-Rohan, Cambodge. Photo: P. Maestrati, MNHN (E), F. Trus, RBINS (F)." figureDoi="10.3897/zookeys.1119.85400.figure30" httpUri="https://binary.pensoft.net/fig/735391" pageId="0" pageNumber="1">30A</figureCitation>
) from
<location LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51:D39818994C84FB7EBA1F8BE1BBDA5980" country="Camboja" county="Pfeiffer" municipality="Figs" name="Chai Thong Wararam Temple" stateProvince="Tak Fa District, Nakhon Sawan Province">Chai Thong Wararam Temple</location>
,
<collectingRegion isEnumeration="true">
Tak
<collectingRegion inferredBySuffix="true">Fa District</collectingRegion>
, Nakhon Sawan Province
</collectingRegion>
,
<collectingDate value="2017-06-09">9 June 2017</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2020-12-06" collectorName="Fig." country="Cambodia" location="Tham Saeng Wiset Bureau of Monks" specimenCount="125" stateProvince="Tak Fa District, Nakhon Sawan Province">
CUMZ 12051 (170 shells and
<specimenCount type="generic">125 specimens</specimenCount>
in ethanol;
<collectorName>Fig.</collectorName>
<figureCitation captionStart="Figure 25" captionStartId="F25" captionText="Figure 25. Live specimens of A-C Pupina artata: specimens A CUMZ 12020 from Buri Ratchawanaram Temple, Ratchaburi B CUMZ 12022 from Golden Dragon Cave, Ratchaburi, and C CUMZ 12037 from Wua Ta Lap Island, Surat Thani D, E Pupina bensoni sp. nov. D paratype CUMZ 12045 / 2 from Khao Wong Cave, Uthai Thani and E specimen CUMZ 12047 from Tham Namthip Bureau of Monks, Uthai Thani F, G Pupina peguensis: specimens F CUMZ 12050 from Chai Thong Wararam Temple, Nakhon Sawan and G CUMZ 12051 from Tham Saeng Wiset Bureau of Monks, Nakhon Sawan H Pupina siamensis, specimen CUMZ 12069 from Khao Chi Chan Buddha Image, Chonburi. All not to scale." figureDoi="10.3897/zookeys.1119.85400.figure25" httpUri="https://binary.pensoft.net/fig/735386" pageId="0" pageNumber="1">25G</figureCitation>
) from
<location LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51:1F79C7EC0F984FE87EE717F440D75CF5" country="Cambodia" name="Tham Saeng Wiset Bureau of Monks" stateProvince="Tak Fa District, Nakhon Sawan Province">Tham Saeng Wiset Bureau of Monks</location>
,
<collectingRegion isEnumeration="true">
Tak
<collectingRegion inferredBySuffix="true">Fa District</collectingRegion>
, Nakhon Sawan Province
</collectingRegion>
,
<collectingDate value="2020-12-06">6 Dec. 2020</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2017-09-18" collectorName="Figs" country="Cambodia" location="Pak Chong District, Nakhon Ratchasima Province" specimenCount="1" stateProvince="Pak Chong District, Nakhon Ratchasima Province">
CUMZ 12105 (2 shells;
<collectorName>Figs</collectorName>
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Shells of Pupina arula species group from mainland Southeast Asia A-G Pupina peguensis A syntype of Pupina blanfordi NHMUK 1888.12.4.100 and specimens B CUMZ 12105 C CUMZ 12050 D CUMZ 12103 E NHMUK ex. Cuming coll. F CUMZ 12094, and G CUMZ 12108 H-J Pupina crosseana H lectotype MNHN-IM- 2000 - 35834 I paralectotype RBINS MT 966 / 10591, and J specimen CUMZ 12049 K, L Pupina siamensis K lectotype SMF 109948 and L specimen CUMZ 12052 M-P Pupina bilabiata sp. nov. M holotype CUMZ 12073 / 1 and specimens N CUMZ 12079 O CUMZ 12081, and P CUMZ 12086 Q, R Pupina godwinausteni sp. nov. Q holotype CUMZ 12090 / 1 and R paratype CUMZ 12090 / 2. Photo: P. Maestrati, MNHN (H), F. Trus, RBINS (I)." figureDoi="10.3897/zookeys.1119.85400.figure28" httpUri="https://binary.pensoft.net/fig/735389" pageId="0" pageNumber="1">28B</figureCitation>
,
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. A-D Pupina peguensis: specimens A CUMZ 12050 from Chai Thong Wararam Temple, Nakhon Sawan B CUMZ 12105 from Thep Phithak Punnaram Temple, Nakhon Ratchasima C CUMZ 12108 from Tham Thep Bandan Temple, Phetchabun, and D CUMZ 12094 from Khao Tham Phra Temple, Chiang Rai E, F Pupina crosseana E lectotype MNHN-IM- 2000 - 35834 from Cambodge and F paralectotype RBINS MT 966 / 10591 from Phnom-Rohan, Cambodge. Photo: P. Maestrati, MNHN (E), F. Trus, RBINS (F)." figureDoi="10.3897/zookeys.1119.85400.figure30" httpUri="https://binary.pensoft.net/fig/735391" pageId="0" pageNumber="1">30B</figureCitation>
) from Thep Phithak Punnaram Temple,
<collectingRegion isEnumeration="true">Pak Chong District, Nakhon Ratchasima Province</collectingRegion>
,
<collectingDate value="2017-09-18">18 Sept. 2017</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2020-09-03" collectorName="Tham Wua Daeng Temple" country="Cambodia" location="Phakdi" specimenCount="1" stateProvince="Chumphon District, Chaiyaphum Province">
CUMZ 12107 (
<specimenCount type="generic">1 specimen</specimenCount>
in ethanol) from
<collectorName>Tham Wua Daeng Temple</collectorName>
,
<location LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51:460E827BB66F5CF3C7B63DF2FB6BEB3E" country="Cambodia" name="Phakdi" stateProvince="Chumphon District, Chaiyaphum Province">Phakdi</location>
<collectingRegion isEnumeration="true">Chumphon District, Chaiyaphum Province</collectingRegion>
,
<collectingDate value="2020-09-03">3 Sept. 2020</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2007-10-21" collectorName="Figs" country="Cambodia" location="Wichian Buri District, Phetchabun Province" specimenCount="1" stateProvince="Wichian Buri District, Phetchabun Province">
CUMZ 12108 (8 shells;
<collectorName>Figs</collectorName>
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Shells of Pupina arula species group from mainland Southeast Asia A-G Pupina peguensis A syntype of Pupina blanfordi NHMUK 1888.12.4.100 and specimens B CUMZ 12105 C CUMZ 12050 D CUMZ 12103 E NHMUK ex. Cuming coll. F CUMZ 12094, and G CUMZ 12108 H-J Pupina crosseana H lectotype MNHN-IM- 2000 - 35834 I paralectotype RBINS MT 966 / 10591, and J specimen CUMZ 12049 K, L Pupina siamensis K lectotype SMF 109948 and L specimen CUMZ 12052 M-P Pupina bilabiata sp. nov. M holotype CUMZ 12073 / 1 and specimens N CUMZ 12079 O CUMZ 12081, and P CUMZ 12086 Q, R Pupina godwinausteni sp. nov. Q holotype CUMZ 12090 / 1 and R paratype CUMZ 12090 / 2. Photo: P. Maestrati, MNHN (H), F. Trus, RBINS (I)." figureDoi="10.3897/zookeys.1119.85400.figure28" httpUri="https://binary.pensoft.net/fig/735389" pageId="0" pageNumber="1">28G</figureCitation>
,
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. A-D Pupina peguensis: specimens A CUMZ 12050 from Chai Thong Wararam Temple, Nakhon Sawan B CUMZ 12105 from Thep Phithak Punnaram Temple, Nakhon Ratchasima C CUMZ 12108 from Tham Thep Bandan Temple, Phetchabun, and D CUMZ 12094 from Khao Tham Phra Temple, Chiang Rai E, F Pupina crosseana E lectotype MNHN-IM- 2000 - 35834 from Cambodge and F paralectotype RBINS MT 966 / 10591 from Phnom-Rohan, Cambodge. Photo: P. Maestrati, MNHN (E), F. Trus, RBINS (F)." figureDoi="10.3897/zookeys.1119.85400.figure30" httpUri="https://binary.pensoft.net/fig/735391" pageId="0" pageNumber="1">30C</figureCitation>
) from Tham Thep Bandan Temple,
<collectingRegion isEnumeration="true">Wichian Buri District, Phetchabun Province</collectingRegion>
,
<collectingDate value="2007-10-21">21 Oct. 2007</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2020-08-03" collectorName="Tham Pha Ta Phon" country="Cambodia" location="Noen Maprang District, Phitsanulok Province" specimenCount="1" stateProvince="Noen Maprang District, Phitsanulok Province">
CUMZ 12109 (1 shell) from
<collectorName>Tham Pha Ta Phon</collectorName>
,
<collectingRegion isEnumeration="true">Noen Maprang District, Phitsanulok Province</collectingRegion>
,
<collectingDate value="2020-08-03">3 Aug. 2020</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2017-06-08" country="Cambodia" location="Tham Wang Na Bureau of Monks" specimenCount="1" stateProvince="Noen Maprang District, Phitsanulok Province">
CUMZ 12110 (1 shell) from
<location LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51:C8ADBEE3141215BF7060E49087BEE0C7" country="Cambodia" name="Tham Wang Na Bureau of Monks" stateProvince="Noen Maprang District, Phitsanulok Province">Tham Wang Na Bureau of Monks</location>
,
<collectingRegion isEnumeration="true">Noen Maprang District, Phitsanulok Province</collectingRegion>
,
<collectingDate value="2017-06-08">8 June 2017</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2009-12-01" country="Cambodia" location="Tham Pet Tham Thong Forest" specimenCount="1" stateProvince="Takhli District, Nakhon Sawan Province">
CUMZ 12172 (15 shells) from
<location LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51:E06A3341827395492585FF0F5D096E3D" country="Cambodia" name="Tham Pet Tham Thong Forest" stateProvince="Takhli District, Nakhon Sawan Province">Tham Pet Tham Thong Forest</location>
Park,
<collectingRegion isEnumeration="true">Takhli District, Nakhon Sawan Province</collectingRegion>
,
<collectingDate value="2009-12-01">1 Dec. 2009</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2008-01-09" collectorName="Figs" country="Cambodia" location="Mueang Chiang Rai District, Chiang Rai Province" specimenCount="1" stateProvince="Mueang Chiang Rai District, Chiang Rai Province">
CUMZ 12094 (12 shells;
<collectorName>Figs</collectorName>
<figureCitation captionStart="Figure 18" captionStartId="F18" captionText="Figure 18. External shell morphology of three mainland Southeast Asian Pupina species groups A Pupina artata from the Pupina artata species group, CUMZ 12003 from Ban Ping Khong, Chiang Mai B Pupina peguensis from the Pupina arula species group, CUMZ 12094 from Khao Tham Phra Temple, Chiang Rai C Pupina siamensis from the Pupina arula species group, CUMZ 12052 from Sri Thong Cave, Sra Keo, and D Pupina tchehelensis from the Pupina aureola species group, CUMZ 12136 from limestone mountain, Phang Nga. Red frames focus on the parietal tooth and posterior canal; blue frames focus on the curvature of the apertural lip when observed from lateral view." figureDoi="10.3897/zookeys.1119.85400.figure18" httpUri="https://binary.pensoft.net/fig/735379" pageId="0" pageNumber="1">18B</figureCitation>
,
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Shells of Pupina arula species group from mainland Southeast Asia A-G Pupina peguensis A syntype of Pupina blanfordi NHMUK 1888.12.4.100 and specimens B CUMZ 12105 C CUMZ 12050 D CUMZ 12103 E NHMUK ex. Cuming coll. F CUMZ 12094, and G CUMZ 12108 H-J Pupina crosseana H lectotype MNHN-IM- 2000 - 35834 I paralectotype RBINS MT 966 / 10591, and J specimen CUMZ 12049 K, L Pupina siamensis K lectotype SMF 109948 and L specimen CUMZ 12052 M-P Pupina bilabiata sp. nov. M holotype CUMZ 12073 / 1 and specimens N CUMZ 12079 O CUMZ 12081, and P CUMZ 12086 Q, R Pupina godwinausteni sp. nov. Q holotype CUMZ 12090 / 1 and R paratype CUMZ 12090 / 2. Photo: P. Maestrati, MNHN (H), F. Trus, RBINS (I)." figureDoi="10.3897/zookeys.1119.85400.figure28" httpUri="https://binary.pensoft.net/fig/735389" pageId="0" pageNumber="1">28F</figureCitation>
,
<figureCitation captionStart="Figure 30" captionStartId="F30" captionText="Figure 30. A-D Pupina peguensis: specimens A CUMZ 12050 from Chai Thong Wararam Temple, Nakhon Sawan B CUMZ 12105 from Thep Phithak Punnaram Temple, Nakhon Ratchasima C CUMZ 12108 from Tham Thep Bandan Temple, Phetchabun, and D CUMZ 12094 from Khao Tham Phra Temple, Chiang Rai E, F Pupina crosseana E lectotype MNHN-IM- 2000 - 35834 from Cambodge and F paralectotype RBINS MT 966 / 10591 from Phnom-Rohan, Cambodge. Photo: P. Maestrati, MNHN (E), F. Trus, RBINS (F)." figureDoi="10.3897/zookeys.1119.85400.figure30" httpUri="https://binary.pensoft.net/fig/735391" pageId="0" pageNumber="1">30D</figureCitation>
) from Khao Tham Phra Temple,
<collectingRegion isEnumeration="true">Mueang Chiang Rai District, Chiang Rai Province</collectingRegion>
,
<collectingDate value="2008-01-09">9 Jan. 2008</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2008-10-25" collectorName="Tham Phajarui Temple" country="Cambodia" location="Pa Daet District, Chiang Rai Province" specimenCount="1" stateProvince="Pa Daet District, Chiang Rai Province">
CUMZ 12095 (1 shell) from
<collectorName>Tham Phajarui Temple</collectorName>
,
<collectingRegion isEnumeration="true">Pa Daet District, Chiang Rai Province</collectingRegion>
,
<collectingDate value="2008-10-25">25 Oct. 2008</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2015-10-23" country="Cambodia" location="Luang Cave" specimenCount="1" stateProvince="Mae Sai District, Chiang Rai Province">
CUMZ 12096 (9 shells) from
<location LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51:2F785F0F20E420F9217AC8E067DA8C3C" country="Cambodia" name="Luang Cave" stateProvince="Mae Sai District, Chiang Rai Province">Luang Cave</location>
,
<collectingRegion isEnumeration="true">Mae Sai District, Chiang Rai Province</collectingRegion>
,
<collectingDate value="2015-10-23">23 Oct. 2015</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2009-11-29" collectorName="Tham Phra Bamphen Bun Temple" country="Cambodia" location="Phan District, Chiang Rai Province" specimenCount="1" stateProvince="Phan District, Chiang Rai Province">
CUMZ 12097 (3 shells) from
<collectorName>Tham Phra Bamphen Bun Temple</collectorName>
,
<collectingRegion isEnumeration="true">Phan District, Chiang Rai Province</collectingRegion>
,
<collectingDate value="2009-11-29">29 Nov. 2009</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2017-10-06" collectorName="Mae Lana" country="Cambodia" location="Pang Mapha District, Mae Hong Son Province" specimenCount="1" stateProvince="Pang Mapha District, Mae Hong Son Province">
CUMZ 12098 (1 shell) from
<collectorName>Mae Lana</collectorName>
checkpoint,
<collectingRegion isEnumeration="true">Pang Mapha District, Mae Hong Son Province</collectingRegion>
,
<collectingDate value="2017-10-06">6 Oct. 2017</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2015-01-18" collectorName="Mae Lana" country="Cambodia" location="Pang Mapha District, Mae Hong Son Province" specimenCount="1" stateProvince="Pang Mapha District, Mae Hong Son Province">
CUMZ 12099 (5 shells) from
<collectorName>Mae Lana</collectorName>
junction,
<collectingRegion isEnumeration="true">Pang Mapha District, Mae Hong Son Province</collectingRegion>
,
<collectingDate value="2015-01-18">18 Jan. 2015</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2015-01-18" collectorName="Mueang" country="Cambodia" location="Pha Daeng Cave" specimenCount="1" stateProvince="Mae Hong Son District, Mae Hong Son Province">
CUMZ 12100 (6 shells) from
<location LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51:F66A622972D6DB81EB566E687ED2835D" country="Cambodia" name="Pha Daeng Cave" stateProvince="Mae Hong Son District, Mae Hong Son Province">Pha Daeng Cave</location>
,
<collectorName>Mueang</collectorName>
<collectingRegion isEnumeration="true">Mae Hong Son District, Mae Hong Son Province</collectingRegion>
,
<collectingDate value="2015-01-18">18 Jan. 2015</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2017-10-05" country="Cambodia" location="Mueang" municipality="Pha Daeng Cave" specimenCount="1" stateProvince="Mae Hong Son District, Mae Hong Son Province">
CUMZ 12101 (
<specimenCount type="generic">1 specimen</specimenCount>
in ethanol) from
<collectingMunicipality>Pha Daeng Cave</collectingMunicipality>
,
<location LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51:1A6242D7AC4BC91D195ED58D7564C8B8" country="Cambodia" municipality="Pha Daeng Cave" name="Mueang" stateProvince="Mae Hong Son District, Mae Hong Son Province">Mueang</location>
<collectingRegion isEnumeration="true">Mae Hong Son District, Mae Hong Son Province</collectingRegion>
,
<collectingDate value="2017-10-05">5 Oct. 2017</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2020-12-03" collectorName="Mueang" country="Cambodia" location="Pha Daeng Cave" specimenCount="1" stateProvince="Mae Hong Son District, Mae Hong Son Province">
CUMZ 12102 (6 shells) from
<location LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51:2AA5C111405EC989FB00608775961BA4" country="Cambodia" name="Pha Daeng Cave" stateProvince="Mae Hong Son District, Mae Hong Son Province">Pha Daeng Cave</location>
,
<collectorName>Mueang</collectorName>
<collectingRegion isEnumeration="true">Mae Hong Son District, Mae Hong Son Province</collectingRegion>
,
<collectingDate value="2020-12-03">3 Dec. 2020</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2015-10-24" collectorName="Doi Ang Khang" country="Cambodia" location="Fang District, Chiang Mai Province" specimenCount="1" stateProvince="Fang District, Chiang Mai Province">
CUMZ 12187 (2 shells) from
<collectorName>Doi Ang Khang</collectorName>
,
<collectingRegion isEnumeration="true">Fang District, Chiang Mai Province</collectingRegion>
,
<collectingDate value="2015-10-24">24 Oct. 2015</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2009-10-11" collectorName="Fig." country="Cambodia" location="Mueang" municipality="Pha Tub Cave" specimenCount="3" stateProvince="Nan District, Nan Province">
CUMZ 12103 (
<specimenCount type="generic">3 specimens</specimenCount>
in ethanol;
<collectorName>Fig.</collectorName>
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Shells of Pupina arula species group from mainland Southeast Asia A-G Pupina peguensis A syntype of Pupina blanfordi NHMUK 1888.12.4.100 and specimens B CUMZ 12105 C CUMZ 12050 D CUMZ 12103 E NHMUK ex. Cuming coll. F CUMZ 12094, and G CUMZ 12108 H-J Pupina crosseana H lectotype MNHN-IM- 2000 - 35834 I paralectotype RBINS MT 966 / 10591, and J specimen CUMZ 12049 K, L Pupina siamensis K lectotype SMF 109948 and L specimen CUMZ 12052 M-P Pupina bilabiata sp. nov. M holotype CUMZ 12073 / 1 and specimens N CUMZ 12079 O CUMZ 12081, and P CUMZ 12086 Q, R Pupina godwinausteni sp. nov. Q holotype CUMZ 12090 / 1 and R paratype CUMZ 12090 / 2. Photo: P. Maestrati, MNHN (H), F. Trus, RBINS (I)." figureDoi="10.3897/zookeys.1119.85400.figure28" httpUri="https://binary.pensoft.net/fig/735389" pageId="0" pageNumber="1">28D</figureCitation>
) from
<collectingMunicipality>Pha Tub Cave</collectingMunicipality>
,
<location LSID="urn:lsid:plazi:treatment:3ED453755B705B66888718AF13B99A51:225C2E979AD1422606B5E21B4B56DB8E" country="Cambodia" municipality="Pha Tub Cave" name="Mueang" stateProvince="Nan District, Nan Province">Mueang</location>
<collectingRegion isEnumeration="true">Nan District, Nan Province</collectingRegion>
,
<collectingDate value="2009-10-11">11 Oct. 2009</collectingDate>
</materialsCitation>
.
<materialsCitation collectingDate="2007-10-09" collectorName="Tham Pha Nang Khoi Temple" country="Thailand" location="Phrae Province" specimenCount="1" stateProvince="Phrae Province">
CUMZ 12104 (24 shells) from
<collectorName>Tham Pha Nang Khoi Temple</collectorName>
, Rong Kwang District,
<collectingRegion country="Thailand" name="Phrae">Phrae Province</collectingRegion>
,
<collectingCountry name="Thailand">Thailand</collectingCountry>
,
<collectingDate value="2007-10-09">9 Oct. 2007</collectingDate>
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="diagnosis">
<paragraph pageId="0" pageNumber="1">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="1">Shell globose to ovate-fusiform; last whorl ca. 75-80% of shell height. Apertural lip thickened but not expanded; apertural lip curved when observed from lateral view. Columellar tooth fin-shaped or curved downward like an earlobe.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="differential diagnosis">
<paragraph pageId="0" pageNumber="1">Differential diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="1">
<taxonomicName authorityName="Benson" authorityYear="1860" class="Gastropoda" family="Pupinidae" genus="Pupina" kingdom="Animalia" lsidName="Pupina peguensis" order="Architaenioglossa" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="peguensis">
<emphasis italics="true" pageId="0" pageNumber="1">Pupina peguensis</emphasis>
</taxonomicName>
is similar to
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. arula" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="arula">
<emphasis italics="true" pageId="0" pageNumber="1">P. arula</emphasis>
</taxonomicName>
in shell shape and a curved apertural lip when observed from lateral view, but differs in having a glossy shell surface. This species is also similar to
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. exclamationis" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="exclamationis">
<emphasis italics="true" pageId="0" pageNumber="1">P. exclamationis</emphasis>
</taxonomicName>
in having a glossy surface and a curved apertural lip when observed from lateral view, but differs in having a more ovate shell shape and a more distinct parietal callus.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="distribution">
<paragraph pageId="0" pageNumber="1">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="1">
Myanmar (
<bibRefCitation DOI="https://doi.org/10.1080/00222936008697305" author="Benson, WH" journalOrPublisher="Journal de Conchyliologie" pageId="0" pageNumber="1" refId="B10" refString="Benson, WH, 1860. Characters of new land-shells from Burmah and the Andamans. Annals and Magazine of Natural History, Ser 3 6: 190-195. https://doi.org/10.1080/00222936008697305" title="Characters of new land-shells from Burmah and the Andamans. Annals and Magazine of Natural History, Ser 3 6: 190 - 195." url="https://doi.org/10.1080/00222936008697305" year="1860">Benson 1860</bibRefCitation>
;
<bibRefCitation author="Theobald, W" journalOrPublisher="Journal of the Asiatic Society of Bengal" pageId="0" pageNumber="1" pagination="238 - 250" refId="B168" refString="Theobald, W, 1864. Notes on the variation of some Indian and Burmese Helicidae, with an attempt at their re-arrangement, together with descriptions of new Burmese Gasteropoda. Journal of the Asiatic Society of Bengal 33 (3): 238 - 250" title="Notes on the variation of some Indian and Burmese Helicidae, with an attempt at their re-arrangement, together with descriptions of new Burmese Gasteropoda." volume="33" year="1864">Theobald 1864</bibRefCitation>
), Luang Phrabang Province, Laos (
<bibRefCitation DOI="https://doi.org/10.3897/zookeys.834.28800" author="Inkhavilay, K" journalOrPublisher="ZooKeys" pageId="0" pageNumber="1" pagination="1 - 166" refId="B71" refString="Inkhavilay, K, Sutcharit, C, Bantaowong, U, Chanabun, R, Siriwut, W, Srisonchai, R, Pholyotha, A, Jirapatrasilp, P, Panha, S, 2019. Annotated checklist of the terrestrial molluscs from Laos (Mollusca, Gastropoda). ZooKeys 834: 1 - 166, DOI: https://doi.org/10.3897/zookeys.834.28800" title="Annotated checklist of the terrestrial molluscs from Laos (Mollusca, Gastropoda)." url="https://doi.org/10.3897/zookeys.834.28800" volume="834" year="2019">Inkhavilay et al. 2019</bibRefCitation>
), northern, northeastern, and central Thailand.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="remarks">
<paragraph pageId="0" pageNumber="1">Remarks.</paragraph>
<paragraph pageId="0" pageNumber="1">
Given that the holotype of
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. peguensis" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="peguensis">
<emphasis italics="true" pageId="0" pageNumber="1">P. peguensis</emphasis>
</taxonomicName>
and the syntype of
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. blanfordi" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="blanfordi">
<emphasis italics="true" pageId="0" pageNumber="1">P. blanfordi</emphasis>
</taxonomicName>
are highly similar in shell shape and size, and their type localities belong to the same area,
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. blanfordi" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="blanfordi">
<emphasis italics="true" pageId="0" pageNumber="1">P. blanfordi</emphasis>
</taxonomicName>
is regarded herein as a junior subjective synonym of
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. peguensis" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="peguensis">
<emphasis italics="true" pageId="0" pageNumber="1">P. peguensis</emphasis>
</taxonomicName>
. This species was previously identified as
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. mouhoti" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="mouhoti">
<emphasis italics="true" pageId="0" pageNumber="1">P. mouhoti</emphasis>
</taxonomicName>
(
<bibRefCitation author="Bavay, A" journalOrPublisher="Journal de Conchyliologie" pageId="0" pageNumber="1" refId="B8" refString="BEDO (2017) Land Snails: Checklist of Molluscan Biodiversity in Thailand.BEDO, Bangkok, 300 pp. [in Thai]" year="1904">BEDO 2017</bibRefCitation>
;
<bibRefCitation author="Sutcharit, C" journalOrPublisher="Zino Publishing, Bangkok" pageId="0" pageNumber="1" refId="B156" refString="Sutcharit, C, Tongkerd, P, Panha, S, 2018. Land snails: The invaluable bio-resources for the Kingdom of Thailand. Zino Publishing, Bangkok" title="Land snails: The invaluable bio-resources for the Kingdom of Thailand." year="2018">Sutcharit et al. 2018</bibRefCitation>
). However, compared to the type specimens of
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. mouhoti" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="mouhoti">
<emphasis italics="true" pageId="0" pageNumber="1">P. mouhoti</emphasis>
</taxonomicName>
,
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. peguensis" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="peguensis">
<emphasis italics="true" pageId="0" pageNumber="1">P. peguensis</emphasis>
</taxonomicName>
has a longer and wider posterior canal. In addition, the apertural lip when observed from lateral view of
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. peguensis" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="peguensis">
<emphasis italics="true" pageId="0" pageNumber="1">P. peguensis</emphasis>
</taxonomicName>
is curved and its columellar tooth is curved downward like an earlobe.
</paragraph>
<paragraph pageId="0" pageNumber="1">
All specimens in the
<taxonomicName authorityName="Benson" authorityYear="1856" class="Gastropoda" family="Pupinidae" genus="Pupina" kingdom="Animalia" lsidName="Pupina arula" order="Architaenioglossa" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="arula">
<emphasis italics="true" pageId="0" pageNumber="1">Pupina arula</emphasis>
</taxonomicName>
species group from Thailand with an ovate shell shape and a curved apertural lip when observed from lateral view are herein identified as
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. peguensis" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="peguensis">
<emphasis italics="true" pageId="0" pageNumber="1">P. peguensis</emphasis>
</taxonomicName>
(Fig.
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Shells of Pupina arula species group from mainland Southeast Asia A-G Pupina peguensis A syntype of Pupina blanfordi NHMUK 1888.12.4.100 and specimens B CUMZ 12105 C CUMZ 12050 D CUMZ 12103 E NHMUK ex. Cuming coll. F CUMZ 12094, and G CUMZ 12108 H-J Pupina crosseana H lectotype MNHN-IM- 2000 - 35834 I paralectotype RBINS MT 966 / 10591, and J specimen CUMZ 12049 K, L Pupina siamensis K lectotype SMF 109948 and L specimen CUMZ 12052 M-P Pupina bilabiata sp. nov. M holotype CUMZ 12073 / 1 and specimens N CUMZ 12079 O CUMZ 12081, and P CUMZ 12086 Q, R Pupina godwinausteni sp. nov. Q holotype CUMZ 12090 / 1 and R paratype CUMZ 12090 / 2. Photo: P. Maestrati, MNHN (H), F. Trus, RBINS (I)." figureDoi="10.3897/zookeys.1119.85400.figure28" httpUri="https://binary.pensoft.net/fig/735389" pageId="0" pageNumber="1">28A-G</figureCitation>
). However, these specimens exhibit a variable shell size (smaller with shell height 6.1 mm, shell width 4.6 mm; Fig.
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Shells of Pupina arula species group from mainland Southeast Asia A-G Pupina peguensis A syntype of Pupina blanfordi NHMUK 1888.12.4.100 and specimens B CUMZ 12105 C CUMZ 12050 D CUMZ 12103 E NHMUK ex. Cuming coll. F CUMZ 12094, and G CUMZ 12108 H-J Pupina crosseana H lectotype MNHN-IM- 2000 - 35834 I paralectotype RBINS MT 966 / 10591, and J specimen CUMZ 12049 K, L Pupina siamensis K lectotype SMF 109948 and L specimen CUMZ 12052 M-P Pupina bilabiata sp. nov. M holotype CUMZ 12073 / 1 and specimens N CUMZ 12079 O CUMZ 12081, and P CUMZ 12086 Q, R Pupina godwinausteni sp. nov. Q holotype CUMZ 12090 / 1 and R paratype CUMZ 12090 / 2. Photo: P. Maestrati, MNHN (H), F. Trus, RBINS (I)." figureDoi="10.3897/zookeys.1119.85400.figure28" httpUri="https://binary.pensoft.net/fig/735389" pageId="0" pageNumber="1">28A</figureCitation>
, to larger with shell height 9.6 mm; shell width 7.1 mm; Fig.
<figureCitation captionStart="Figure 28" captionStartId="F28" captionText="Figure 28. Shells of Pupina arula species group from mainland Southeast Asia A-G Pupina peguensis A syntype of Pupina blanfordi NHMUK 1888.12.4.100 and specimens B CUMZ 12105 C CUMZ 12050 D CUMZ 12103 E NHMUK ex. Cuming coll. F CUMZ 12094, and G CUMZ 12108 H-J Pupina crosseana H lectotype MNHN-IM- 2000 - 35834 I paralectotype RBINS MT 966 / 10591, and J specimen CUMZ 12049 K, L Pupina siamensis K lectotype SMF 109948 and L specimen CUMZ 12052 M-P Pupina bilabiata sp. nov. M holotype CUMZ 12073 / 1 and specimens N CUMZ 12079 O CUMZ 12081, and P CUMZ 12086 Q, R Pupina godwinausteni sp. nov. Q holotype CUMZ 12090 / 1 and R paratype CUMZ 12090 / 2. Photo: P. Maestrati, MNHN (H), F. Trus, RBINS (I)." figureDoi="10.3897/zookeys.1119.85400.figure28" httpUri="https://binary.pensoft.net/fig/735389" pageId="0" pageNumber="1">28G</figureCitation>
). The shell shape is also variable from globose as in the type material (see
<bibRefCitation author="Tripathy, B" journalOrPublisher="Records of the Zoological Survey of India" pageId="0" pageNumber="1" pagination="508 - 510" refId="B177" refString="Tripathy, B, Sajan, S, 2019. Pupina peguensis Benson, 1860 (Mollusca: Pupinidae): Tracing a long lost molluscan type. Records of the Zoological Survey of India 119: 508 - 510" title="Pupina peguensis Benson, 1860 (Mollusca: Pupinidae): Tracing a long lost molluscan type." volume="119" year="2019">Tripathy and Sajan 2019</bibRefCitation>
: fig. 1), to more ovate and ovate-fusiform. As this species is also similar to
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. exclamationis" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="exclamationis">
<emphasis italics="true" pageId="0" pageNumber="1">P. exclamationis</emphasis>
</taxonomicName>
, more sampling, with both morphometric and molecular phylogenetic analyses, are needed to resolve the relationship between these two species and reveal the extent of genetic differentiation or cryptic diversity within the
<taxonomicName class="Gastropoda" family="Pupinidae" kingdom="Animalia" lsidName="P. peguensis" order="Architaenioglossa" pageId="0" pageNumber="1" rank="species" species="peguensis">
<emphasis italics="true" pageId="0" pageNumber="1">P. peguensis</emphasis>
</taxonomicName>
morphotype.
</paragraph>
</subSubSection>
</treatment>
</document>