A cybertaxonomic revision of the micro-landsnail genus Plectostoma Adam (Mollusca, Caenogastropoda, Diplommatinidae), from Peninsular Malaysia, Sumatra and Indochina
Identifieur interne : 000445 ( Pmc/Corpus ); précédent : 000444; suivant : 000446A cybertaxonomic revision of the micro-landsnail genus Plectostoma Adam (Mollusca, Caenogastropoda, Diplommatinidae), from Peninsular Malaysia, Sumatra and Indochina
Auteurs : Thor-Seng Liew ; Jaap Jan Vermeulen ; Mohammad Effendi Bin Marzuki ; Menno SchilthuizenSource :
- ZooKeys [ 1313-2989 ] ; 2014.
Abstract
Url:
DOI: 10.3897/zookeys.393.6717
PubMed: 24715783
PubMed Central: 3974427
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PMC:3974427Le document en format XML
<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">A cybertaxonomic revision of the micro-landsnail genus <italic>Plectostoma</italic> Adam (Mollusca, Caenogastropoda, Diplommatinidae), from Peninsular Malaysia, Sumatra and Indochina</title><author><name sortKey="Liew, Thor Seng" sort="Liew, Thor Seng" uniqKey="Liew T" first="Thor-Seng" last="Liew">Thor-Seng Liew</name><affiliation><nlm:aff id="A1">Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Institute Biology Leiden, Leiden University, P.O. Box 9516, 2300 RA Leiden, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Jalan UMS, 88400, Kota Kinabalu, Sabah, Malaysia</nlm:aff></affiliation><affiliation><nlm:aff id="A4">Rimba, 4 Jalan 1/9D, 43650, Bandar Baru Bangi, Selangor, Malaysia</nlm:aff></affiliation></author><author><name sortKey="Vermeulen, Jaap Jan" sort="Vermeulen, Jaap Jan" uniqKey="Vermeulen J" first="Jaap Jan" last="Vermeulen">Jaap Jan Vermeulen</name><affiliation><nlm:aff id="A1">Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A5">jk.artandscience, Lauwerbes 8, 2318 AT, Leiden, The Netherlandss</nlm:aff></affiliation></author><author><name sortKey="Marzuki, Mohammad Effendi Bin" sort="Marzuki, Mohammad Effendi Bin" uniqKey="Marzuki M" first="Mohammad Effendi Bin" last="Marzuki">Mohammad Effendi Bin Marzuki</name><affiliation><nlm:aff id="A6">102, Jalan Muut, Kampung Sekaan Besar, 96250 Matu, Sarawak, Malaysia</nlm:aff></affiliation></author><author><name sortKey="Schilthuizen, Menno" sort="Schilthuizen, Menno" uniqKey="Schilthuizen M" first="Menno" last="Schilthuizen">Menno Schilthuizen</name><affiliation><nlm:aff id="A1">Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Institute Biology Leiden, Leiden University, P.O. Box 9516, 2300 RA Leiden, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Jalan UMS, 88400, Kota Kinabalu, Sabah, Malaysia</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">24715783</idno><idno type="pmc">3974427</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3974427</idno><idno type="RBID">PMC:3974427</idno><idno type="doi">10.3897/zookeys.393.6717</idno><date when="2014">2014</date><idno type="wicri:Area/Pmc/Corpus">000445</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">A cybertaxonomic revision of the micro-landsnail genus <italic>Plectostoma</italic> Adam (Mollusca, Caenogastropoda, Diplommatinidae), from Peninsular Malaysia, Sumatra and Indochina</title><author><name sortKey="Liew, Thor Seng" sort="Liew, Thor Seng" uniqKey="Liew T" first="Thor-Seng" last="Liew">Thor-Seng Liew</name><affiliation><nlm:aff id="A1">Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Institute Biology Leiden, Leiden University, P.O. Box 9516, 2300 RA Leiden, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Jalan UMS, 88400, Kota Kinabalu, Sabah, Malaysia</nlm:aff></affiliation><affiliation><nlm:aff id="A4">Rimba, 4 Jalan 1/9D, 43650, Bandar Baru Bangi, Selangor, Malaysia</nlm:aff></affiliation></author><author><name sortKey="Vermeulen, Jaap Jan" sort="Vermeulen, Jaap Jan" uniqKey="Vermeulen J" first="Jaap Jan" last="Vermeulen">Jaap Jan Vermeulen</name><affiliation><nlm:aff id="A1">Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A5">jk.artandscience, Lauwerbes 8, 2318 AT, Leiden, The Netherlandss</nlm:aff></affiliation></author><author><name sortKey="Marzuki, Mohammad Effendi Bin" sort="Marzuki, Mohammad Effendi Bin" uniqKey="Marzuki M" first="Mohammad Effendi Bin" last="Marzuki">Mohammad Effendi Bin Marzuki</name><affiliation><nlm:aff id="A6">102, Jalan Muut, Kampung Sekaan Besar, 96250 Matu, Sarawak, Malaysia</nlm:aff></affiliation></author><author><name sortKey="Schilthuizen, Menno" sort="Schilthuizen, Menno" uniqKey="Schilthuizen M" first="Menno" last="Schilthuizen">Menno Schilthuizen</name><affiliation><nlm:aff id="A1">Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Institute Biology Leiden, Leiden University, P.O. Box 9516, 2300 RA Leiden, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Jalan UMS, 88400, Kota Kinabalu, Sabah, Malaysia</nlm:aff></affiliation></author></analytic><series><title level="j">ZooKeys</title><idno type="ISSN">1313-2989</idno><idno type="eISSN">1313-2970</idno><imprint><date when="2014">2014</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><label>Abstract</label><pmc-comment>PageBreak</pmc-comment>
<p><italic>Plectostoma</italic> is a micro land snail restricted to limestone outcrops in Southeast Asia. <italic>Plectostoma</italic> was previously classified as a subgenus of <italic>Opisthostoma</italic> because of the deviation from regular coiling in many species in both taxa. This paper is the first of a two-part revision of the genus <italic>Plectostoma</italic>, and includes all non-Borneo species. In the present paper, we examined 214 collection samples of 31 species, and obtained 62 references, 290 pictures, and 155 3D-models of 29 <italic>Plectostoma</italic> species and 51 COI sequences of 19 species. To work with such a variety of taxonomic data, and then to represent it in an integrated, scaleable and accessible manner, we adopted up-to-date cybertaxonomic tools. All the taxonomic information, such as references, classification, species descriptions, specimen images, genetic data, and distribution data, were tagged and linked with cyber tools and web servers (e.g. Lifedesks, Google Earth, and Barcoding of Life Database). We elevated <italic>Plectostoma</italic> from subgenus to genus level based on morphological, ecological and genetic evidence. We revised the existing 21 <italic>Plectostoma</italic> species and described 10 new species, namely, <italic>P. dindingensis</italic><bold>sp. n.</bold>, <italic>P. mengaburensis</italic><bold>sp. n.</bold>, <italic>P. whitteni</italic><bold>sp. n.</bold>, <italic>P. kayiani</italic><bold>sp. n.</bold>, <italic>P. davisoni</italic><bold>sp. n.</bold>, <italic>P. relauensis</italic><bold>sp. n.</bold>, <italic>P. kubuensis</italic><bold>sp. n.</bold>, <italic>P. tohchinyawi</italic><bold>sp. n.</bold>, <italic>P. tenggekensis</italic><bold>sp. n.</bold>, and <italic>P. ikanensis</italic><bold>sp. n.</bold> All the synthesised, semantic-tagged, and linked taxonomic information is made freely and publicly available online.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Adam, H" uniqKey="Adam H">H Adam</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Adam, H" uniqKey="Adam H">H 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uniqKey="Sutton T">T Sutton</name></author><author><name sortKey="Caithness, N" uniqKey="Caithness N">N Caithness</name></author><author><name sortKey="Pahwa, Js" uniqKey="Pahwa J">JS Pahwa</name></author><author><name sortKey="Burgess, M" uniqKey="Burgess M">M Burgess</name></author><author><name sortKey="Gray, Wa" uniqKey="Gray W">WA Gray</name></author><author><name sortKey="White, Rj" uniqKey="White R">RJ White</name></author><author><name sortKey="Jones, Ac" uniqKey="Jones A">AC Jones</name></author><author><name sortKey="Bisby, Fa" uniqKey="Bisby F">FA Bisby</name></author><author><name sortKey="Culham, A" uniqKey="Culham A">A Culham</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Zhang, Zq" uniqKey="Zhang Z">ZQ Zhang</name></author></analytic></biblStruct></listBibl></div1></back></TEI><pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Zookeys</journal-id><journal-id journal-id-type="iso-abbrev">Zookeys</journal-id><journal-id journal-id-type="publisher-id">ZooKeys</journal-id><journal-title-group><journal-title>ZooKeys</journal-title></journal-title-group><issn pub-type="ppub">1313-2989</issn><issn pub-type="epub">1313-2970</issn><publisher><publisher-name>Pensoft Publishers</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">24715783</article-id><article-id pub-id-type="pmc">3974427</article-id><article-id pub-id-type="doi">10.3897/zookeys.393.6717</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group></article-categories><title-group><article-title>A cybertaxonomic revision of the micro-landsnail genus <italic>Plectostoma</italic> Adam (Mollusca, Caenogastropoda, Diplommatinidae), from Peninsular Malaysia, Sumatra and Indochina</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Liew</surname><given-names>Thor-Seng</given-names></name><xref ref-type="aff" rid="A1">1</xref><xref ref-type="aff" rid="A2">2</xref><xref ref-type="aff" rid="A3">3</xref><xref ref-type="aff" rid="A4">4</xref><uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/7D912940-C78C-4E43-9BBA-F0A49E360028">http://zoobank.org/7D912940-C78C-4E43-9BBA-F0A49E360028</uri></contrib><contrib contrib-type="author"><name><surname>Vermeulen</surname><given-names>Jaap Jan</given-names></name><xref ref-type="aff" rid="A1">1</xref><xref ref-type="aff" rid="A5">5</xref><uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/F3BCEB3D-0214-45BB-9A37-FCDC9AD1B36B">http://zoobank.org/F3BCEB3D-0214-45BB-9A37-FCDC9AD1B36B</uri></contrib><contrib contrib-type="author"><name><surname>Marzuki</surname><given-names>Mohammad Effendi bin</given-names></name><xref ref-type="aff" rid="A6">6</xref><uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/F93444BE-BC25-46E1-85CB-EE1A7231AFA8">http://zoobank.org/F93444BE-BC25-46E1-85CB-EE1A7231AFA8</uri></contrib><contrib contrib-type="author"><name><surname>Schilthuizen</surname><given-names>Menno</given-names></name><xref ref-type="aff" rid="A1">1</xref><xref ref-type="aff" rid="A2">2</xref><xref ref-type="aff" rid="A3">3</xref><uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/683D0AB7-CDD8-4FAA-94B6-436F3BFB8873">http://zoobank.org/683D0AB7-CDD8-4FAA-94B6-436F3BFB8873</uri></contrib></contrib-group><aff id="A1"><label>1</label>Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Netherlands</aff><aff id="A2"><label>2</label>Institute Biology Leiden, Leiden University, P.O. Box 9516, 2300 RA Leiden, The Netherlands</aff><aff id="A3"><label>3</label>Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Jalan UMS, 88400, Kota Kinabalu, Sabah, Malaysia</aff><aff id="A4"><label>4</label>Rimba, 4 Jalan 1/9D, 43650, Bandar Baru Bangi, Selangor, Malaysia</aff><aff id="A5"><label>5</label>jk.artandscience, Lauwerbes 8, 2318 AT, Leiden, The Netherlandss</aff><aff id="A6"><label>6</label>102, Jalan Muut, Kampung Sekaan Besar, 96250 Matu, Sarawak, Malaysia</aff><author-notes><corresp>Corresponding author: Thor-Seng Liew (<email xlink:type="simple">thorsengliew@gmail.com</email>)</corresp><fn fn-type="edited-by"><p>Academic editor: F. Köhler</p></fn></author-notes><pub-date pub-type="collection"><year>2014</year></pub-date><pub-date pub-type="epub"><day>25</day><month>3</month><year>2014</year></pub-date><issue>393</issue><fpage>1</fpage><lpage>107</lpage><history><date date-type="received"><day>29</day><month>11</month><year>2013</year></date><date date-type="accepted"><day>27</day><month>2</month><year>2014</year></date></history><permissions><copyright-statement>Thor-Seng Liew, Jaap Jan Vermeulen, Mohammad Effendi bin Marzuki, Menno Schilthuizen</copyright-statement><license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0"><license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p></license></permissions><self-uri content-type="zoobank" xlink:type="simple" xlink:href="http://zoobank.org/5B035E76-BBD1-4A44-A5D8-C2140E6168F1">http://zoobank.org/5B035E76-BBD1-4A44-A5D8-C2140E6168F1</self-uri><abstract><label>Abstract</label><pmc-comment>PageBreak</pmc-comment>
<p><italic>Plectostoma</italic> is a micro land snail restricted to limestone outcrops in Southeast Asia. <italic>Plectostoma</italic> was previously classified as a subgenus of <italic>Opisthostoma</italic> because of the deviation from regular coiling in many species in both taxa. This paper is the first of a two-part revision of the genus <italic>Plectostoma</italic>, and includes all non-Borneo species. In the present paper, we examined 214 collection samples of 31 species, and obtained 62 references, 290 pictures, and 155 3D-models of 29 <italic>Plectostoma</italic> species and 51 COI sequences of 19 species. To work with such a variety of taxonomic data, and then to represent it in an integrated, scaleable and accessible manner, we adopted up-to-date cybertaxonomic tools. All the taxonomic information, such as references, classification, species descriptions, specimen images, genetic data, and distribution data, were tagged and linked with cyber tools and web servers (e.g. Lifedesks, Google Earth, and Barcoding of Life Database). We elevated <italic>Plectostoma</italic> from subgenus to genus level based on morphological, ecological and genetic evidence. We revised the existing 21 <italic>Plectostoma</italic> species and described 10 new species, namely, <italic>P. dindingensis</italic><bold>sp. n.</bold>, <italic>P. mengaburensis</italic><bold>sp. n.</bold>, <italic>P. whitteni</italic><bold>sp. n.</bold>, <italic>P. kayiani</italic><bold>sp. n.</bold>, <italic>P. davisoni</italic><bold>sp. n.</bold>, <italic>P. relauensis</italic><bold>sp. n.</bold>, <italic>P. kubuensis</italic><bold>sp. n.</bold>, <italic>P. tohchinyawi</italic><bold>sp. n.</bold>, <italic>P. tenggekensis</italic><bold>sp. n.</bold>, and <italic>P. ikanensis</italic><bold>sp. n.</bold> All the synthesised, semantic-tagged, and linked taxonomic information is made freely and publicly available online.</p></abstract><kwd-group><label>Keywords</label><kwd>Sundaland</kwd><kwd>karst habitat</kwd><kwd>shell</kwd><kwd>e-taxonomy</kwd><kwd>tropical</kwd><kwd>semantic tag</kwd><kwd>taxonomic impediment</kwd></kwd-group></article-meta><notes><sec sec-type="Citation"><title>Citation</title><p>Liew T-S, Vermeulen JJ, bin Marzuki ME, Schilthuizen M (2014) A cybertaxonomic revision of the micro-landsnail genus <italic>Plectostoma</italic> Adam (Mollusca, Caenogastropoda, Diplommatinidae), from Peninsular Malaysia, Sumatra and Indochina. ZooKeys 393: 1–107. doi: <ext-link ext-link-type="doi" xlink:href="10.3897/zookeys.393.6717">10.3897/zookeys.393.6717</ext-link></p></sec></notes></front><body><sec><title>Introduction</title><p>The purpose of this paper is twofold. Firstly, we demonstrate an updated workflow that uses several free tools to semantically tag and link different types of information during taxonomic revision. This approach allows the taxonomist to manage information in a more effective manner, making good quality data accessible and scaleable, which is essential for the taxonomist himself, future taxonomists and other users. Secondly, we revise the taxonomy of the genus <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> based on the materials that have been accumulated since the last revision on the <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species of this region about five decades ago (<xref rid="B6" ref-type="bibr">van Benthem Jutting 1952</xref>, <xref rid="B8" ref-type="bibr">1961</xref>). We revised the non-Bornean <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> by using the redefined description of shell characters, which are better suited for the representation of the shell ontogeny and shell form. These shell character descriptions constitute a species hypothesis for each species, which is then discussed in terms of its genetic variation and biogeography.</p><p>Therefore, in this introduction section, we briefly review current issues in taxonomy, especially on the importance of taxonomic data management. After that, we introduce the taxonomy history of the genus <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> and the taxonomic problems of the genus. In the methodology section, we describe in detail the procedure to incorporate cybertaxonomy tools in the taxonomic revision workflow. Finally, we discuss the taxonomy of the genus <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> in the results and discussion sections.</p><sec sec-type="Current issues in taxonomy"><title>Current issues in taxonomy</title><p>Taxonomy is arguably man’s oldest profession and an important fundamental discipline for many other biological fields (<xref rid="B47" ref-type="bibr">Hedgpeth 1961</xref>, <xref rid="B130" ref-type="bibr">Wilson 2004</xref>). It helps us to inventory the biodiversity on earth by naming each classified group of organisms that shares certain attributes based on, for example, genetic and morphological evidence. However, the taxa names are not necessarily maintained in perpetuity, as classifications can be changed when more specimens and novel attributes are examined and compared. In fact, the continuous improvement of the existing classification scheme of any taxon is a fundamental characteristic in taxonomic science. As a result, a taxon name is a summary of hundreds of years’ of taxonomists’ attempts to classify biodiversity.</p><p>Despite this long history, only a small fraction of biodiversity has been named (<xref rid="B74" ref-type="bibr">Mora et al. 2011</xref>). Certainly, more resources (money, time and taxonomists) are needed to describe the remaining unknown biodiversity. For example, <xref rid="B21" ref-type="bibr">Carbayo and Marques (2011)</xref> estimated that over two hundred billion US dollars are needed to describe the remaining ca. 5.5 million undescribed species. They indicate that half of the cost is to be invested on training taxonomists and project budgets. Although it is not being discussed in <xref rid="B21" ref-type="bibr">Carbayo and Marques (2011)</xref>, it is not hard to imaging that a significant proportion of the amount were spent on travelling to various museums for studying type specimens and other samples, compiling literature and maintaining all <pmc-comment>PageBreak</pmc-comment>this information. Even if there were no financial constraint, given the current pace in describing species (ca. 18000 species per year, see <xref rid="B133" ref-type="bibr">Zhang 2008</xref>), about 400 years are still needed to describe all the species, which translates into 10 generations of taxonomists (<xref rid="B79" ref-type="bibr">Padial et al. 2010</xref>).</p><p>All organisms could eventually be scientifically described, but there are several pressing contemporary issues that could affect the advancement of taxonomy as a science. Firstly, the population of taxonomists is declining (<xref rid="B124" ref-type="bibr">Wägele et al. 2011</xref>), which will directly reduce the pace of species discovery and capacity building for future taxonomists. Secondly, the majority of taxonomic resources (e.g. type materials, literature, expertise) are centralised in developed countries, whereas the remaining undescribed species are concentrated in developing countries (<xref rid="B91" ref-type="bibr">Rodrigues et al. 2010</xref>). Neither taxonomists from developed countries nor taxonomists from developing countries can work effectively without having both biodiversity and taxonomic resources readily available to be integrated. Thirdly, many unnamed species are likely to go extinct under the current rates of habitat destruction in biodiversity hotspots (<xref rid="B42" ref-type="bibr">Giam et al. 2011</xref>). Thus, taxonomists have to find ways to be able to work more effectively and accelerate the pace of species discovery.</p><p>Several proposals to improve the practice of taxonomy have been published (e.g. <xref rid="B126" ref-type="bibr">Webb et al. 2010</xref>, <xref rid="B124" ref-type="bibr">Wägele et al. 2011</xref>, <xref rid="B71" ref-type="bibr">Miller et al. 2012</xref>). One of these suggests that taxonomists embrace the information and internet technologies in their routine workflow. This is particularly relevant because taxonomy is a data-rich science and taxonomists have to deal with all past taxonomic works, in terms of literature, collection data, genetic data, and ecology, that have been accumulated over hundreds of years. Thus, computer and internet technologies have become important tools to manage all these taxonomic data, especially regarding data storage, dissemination, and retrieval.</p><p>Over the past few years, many web-based databases have been developed to accommodate almost all kinds of the data that taxonomists deal with. For example, museum specimen data are held in the Global Biodiversity Information Facility (<ext-link ext-link-type="uri" xlink:href="http://www.gbif.org">http://www.gbif.org</ext-link>, <xref rid="B37" ref-type="bibr">Edwards 2004</xref>), morphology data (images) in Morphbank (<ext-link ext-link-type="uri" xlink:href="http://www.morphbank.net">http://www.morphbank.net</ext-link>), genetic data in the Barcoding of Life Database (<ext-link ext-link-type="uri" xlink:href="http://www.barcodinglife.com">http://www.barcodinglife.com</ext-link>, <xref rid="B90" ref-type="bibr">Ratnasingham and Hebert 2007</xref>) and GenBank (<ext-link ext-link-type="uri" xlink:href="http://www.ncbi.nlm.nih.gov/genbank">http://www.ncbi.nlm.nih.gov/genbank</ext-link>, <xref rid="B15" ref-type="bibr">Benson et al. 1997</xref>), nomenclature data in ZooBank (<ext-link ext-link-type="uri" xlink:href="http://zoobank.org">http://zoobank.org</ext-link>, <xref rid="B89" ref-type="bibr">Pyle and Michel 2008</xref>), and literature in the Biodiversity Heritage Library (<ext-link ext-link-type="uri" xlink:href="http://www.biodiversitylibrary.org">http://www.biodiversitylibrary.org</ext-link>, <xref rid="B45" ref-type="bibr">Gwinn and Rinaldo 2009</xref>). Although the need to use these facilities to improve taxonomy has been emphasised (e.g. <xref rid="B65" ref-type="bibr">Maddison et al. 2012</xref>, <xref rid="B128" ref-type="bibr">Wheeler et al. 2012</xref>, Miller et al. 2013), the majority of taxonomists has yet to join this movement. This hesitance may be caused by taxonomists’ fear for investing extra work to supply the data, and their doubt of the usefulness of these poorly integrated databases.</p><p>Recently, tools have become available for managing and integrating various types of taxonomic data by using either top-down approaches (e.g. Encyclopedia of Life <ext-link ext-link-type="uri" xlink:href="http://eol.org">http://eol.org</ext-link>, <xref rid="B129" ref-type="bibr">Wilson 2003</xref>) or bottom-up approaches (e.g. Lifedesks <pmc-comment>PageBreak</pmc-comment><ext-link ext-link-type="uri" xlink:href="http://www.lifedesks.org">http://www.lifedesks.org</ext-link>, offered by EOL; Scratchpads <ext-link ext-link-type="uri" xlink:href="http://scratchpads.eu">http://scratchpads.eu</ext-link>, <xref rid="B103" ref-type="bibr">Smith et al. 2009</xref>). The top-down mechanism acts as an automatic data aggregator that harvests and pools semantic-tagged information from different databases. The bottom-up mechanism, on the other hand, acts as quality controller that checks, links and tags the different types of taxonomic data. Obviously, the key factor that determines the success of cybertaxonomy and the main challenge of these integration processes is the quality of the underlying taxonomic data (<xref rid="B82" ref-type="bibr">Parr et al. 2012</xref>).</p><p>Many data in the databases are outdated (e.g. nomenclature change), incorrect due to the limitation of technology (e.g. automation text extraction, <xref rid="B80" ref-type="bibr">Page 2011</xref>) or human error (e.g. misidentification, <xref rid="B132" ref-type="bibr">Yesson et al. 2007</xref>), and not linked or semantic-tagged. In fact, taxonomists spend much of the time in their careers to validate, link and tag all the existing and new taxonomic information. Thus, their contribution to the quality control of the data is essential. However, each database has its unique needs, standards, and format for the data and thus it might require taxonomists to do redundant work, such as uploading and key in the same data in different formats as required by the database.</p><p>In addition to the possible redundant efforts that need to be spent, it is not very clear how taxonomists as data suppliers would benefit most from such databases. In view of this, we demonstrate a working example of how these existing databases and platforms can be incorporated into revisionary taxonomic study processes, which begin with managing new specimen information, then establishing identity (literature study and examination of specimens in museum collections), and then writing taxonomic treatises (<xref rid="B131" ref-type="bibr">Winston 1999</xref>). Our working example consists of a revision of the taxonomy of the genus <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> of Peninsular Malaysia, Indochina and Sumatra (i.e. non-Bornean) by using cybertaxonomy tools during the entire revisionary taxonomy process. With these cybertaxonomy tools, we show that various kinds of taxonomic information can be semantically tagged, linked and integrated in a more user-friendly interface (<xref rid="B87" ref-type="bibr">Penev et al. 2011</xref>). In addition, we demonstrate that using this technology and these databases could facilitate the hypothesis-testing nature of taxonomic research that deals with vast amounts of different kinds of information (e.g. <xref rid="B102" ref-type="bibr">Sluys 2013</xref>). The validated, linked and tagged data that are generated during the taxonomic revision in turn facilitates forthcoming taxonomy studies that will be conducted by future taxonomists. Finally, all this species information is made readily available and accessible for other users such as ecologists and conservationists, especially for those who have limited budget and are from developing countries.</p></sec><sec sec-type="An overview of genus Plectostoma’s taxonomy history"><title>An overview of genus <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>’s taxonomy history</title><p>The study of <pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> sensu stricto and <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> started during the English colonisation in India and North Borneo in the 1860s (<xref rid="B16" ref-type="bibr">Blanford and Blanford 1860</xref>; <xref rid="B2" ref-type="bibr">Adam 1865b</xref>). All early collections were made by English officers, who later described the species themselves or sent their collections to other malacologists for species identification and publication. Until 1880, there was only one <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species known–<italic><named-content content-type="taxon-name">Plectostoma decrespignyi</named-content></italic> from Labuan, North Borneo. Between the 1880s and 1900s, A. H. Everett, C. Hose and S. Beddome collected more <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> specimens from North Borneo and Sarawak, where they were working as either British colonial administrators or naturalists. Based on these specimens, a total of 20 <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species were eventually described from Borneo (<xref rid="B43" ref-type="bibr">Godwin-Austen 1889</xref>, <xref rid="B44" ref-type="bibr">1890</xref>, <xref rid="B20" ref-type="bibr">Boettger 1893</xref>, <xref rid="B105" ref-type="bibr">Smith 1893b</xref>, <xref rid="B106" ref-type="bibr">1894</xref>, <xref rid="B107" ref-type="bibr">1904</xref>, <xref rid="B108" ref-type="bibr">1905a</xref>, <xref rid="B41" ref-type="bibr">Fulton 1901</xref>). Many of these were originally described under <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>, which was considered a subgenus of <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>.</p><p>After the intensive malacological documentation in Borneo in the late 19th and early 20<sup>th</sup> century, more material of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> was collected in Sarawak and Sabah during explorations which principally had a different purpose, by the geologist G.E. Wilford and his associates in 1960s, by soil scientists K. Auffenberg and D.K. Dorman in the late 1980s, and by the botanist J.J. Vermeulen in the 1980s and early 1990s. All of this material was revised by <xref rid="B122" ref-type="bibr">Vermeulen (1994)</xref>, under the genus <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>, which resulted in the description of 27 new <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species from Borneo. Since the 2000s, <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> has attracted more interest, also regarding its phylogeny and evolution (<xref rid="B96" ref-type="bibr">Schilthuizen 2003</xref>, <xref rid="B100" ref-type="bibr">Schilthuizen et al. 2006</xref>, <xref rid="B127" ref-type="bibr">Webster et al. 2012</xref>) and ecology (<xref rid="B97" ref-type="bibr">Schilthuizen et al. 2003a</xref>, <xref rid="B99" ref-type="bibr">2005</xref>).</p><p>In addition to the Bornean <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>, two other <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species were described from Southern Thailand and Southern Vietnam in the 1900s (<xref rid="B117" ref-type="bibr">Sykes 1903</xref>, <xref rid="B35" ref-type="bibr">Dautzenberg and Fischer 1905</xref>). Between the 1930s and 1960s, M.W.F. Tweedie (then director of the Raffles Museum of Singapore) collected many <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> specimens during explorations of Peninsular Malaysia. These specimens were later described as 13 new <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species (<xref rid="B119" ref-type="bibr">Tomlin 1938</xref>, <xref rid="B120" ref-type="bibr">1948</xref>, <xref rid="B6" ref-type="bibr">van Benthem Jutting 1952</xref>, <xref rid="B8" ref-type="bibr">1961</xref>). The research on the taxonomy of non-Bornean <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> ceased in the 1960s with the retirement of Tweedie and the death of van Benthem Jutting. However, ecological investigations of one <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species–<italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic>–had started in 1960s by A.J. Berry, a professor at University Malaya (<xref rid="B10" ref-type="bibr">Berry 1961</xref>, <xref rid="B11" ref-type="bibr">1962</xref>, <xref rid="B12" ref-type="bibr">1963</xref>, <xref rid="B13" ref-type="bibr">1964</xref>, <xref rid="B14" ref-type="bibr">1966</xref>).</p><p>No publications on <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> from Peninsula Malaysia appeared between 1966 and 1996, but many <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> specimens were collected by Geoffrey W. H. Davison, mainly from the state of Kelantan, and other localities in Malaysia (e.g. <xref rid="B34" ref-type="bibr">Davison and Kiew 1990</xref>). In 1996, <italic><named-content content-type="taxon-name">Plectostoma klongsangensis</named-content></italic> was described from Southern Thailand (<xref rid="B81" ref-type="bibr">Panha 1996</xref>). After that, four new species were described from Southern Thailand and Peninsula Malaysia (<xref rid="B63" ref-type="bibr">Maassen 2001</xref>). One year later, the first <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> from Sumatra–<italic><named-content content-type="taxon-name">Plectostoma kitteli</named-content></italic>, was decribed by <xref rid="B64" ref-type="bibr">Maassen (2002)</xref>.</p><p>To sum up, a total of 69 <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> are currently known. The hotspots of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> diversity are the Malay Peninsula (including the southern part of Thailand) and Borneo, harbouring 19 and 48 species, respectively. In addition, one species is known from Sumatra and another from Southern Vietnam.</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="Current taxonomic status of Plectostoma from Indochina, Sumatra and Peninsular Malaysia"><title>Current taxonomic status of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> from Indochina, Sumatra and Peninsular Malaysia</title><p>The last revision of the non-Bornean <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species was done by <xref rid="B6" ref-type="bibr">van Benthem Jutting (1952</xref>, <xref rid="B8" ref-type="bibr">1961</xref>). It is important to note that she described 10 new species on the basis of shell characters from only 21 samples from 16 locations. However, it is problematic to use conventional shell descriptions in this genus. The striking shell form of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> has attracted the attention of malacologists, but it also poses a challenge to describe the shell accurately. As mentioned by <xref rid="B6" ref-type="bibr">Benthem-Jutting (1952)</xref>, “… it is evident that in such irregular shells … the measurements can only be given approximately, and never indicate the real proportion of the shell…” and “…after comparing over and over again did I succeed in checking the points of difference [between species], but even then it remained difficult to bring the true nature of these minor details into adequate words”. To date, <xref rid="B122" ref-type="bibr">Vermeulen’s (1994)</xref> approach in describing the <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> shell is the most comprehensive, but it is still difficult to recognise the species from the written description alone.</p><p>As mentioned in the taxonomic history section above, additional <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> specimens were collected by Geoffrey W. H. Davison in Peninsular Malaysia in the 1990s. Furthermore, we collected more specimens, including living ones for genetic study, during field trips to Peninsular Malaysian limestone hills between 2010 and 2011. These <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> specimens are valuable to improve the taxonomy of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> in Peninsular Malaysia. Thus, it is timely to revise the non-Bornean <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> based on these recently collected specimens by re-examining the species hypotheses formed on the basis of shell morphology. In addition, we also update the knowledge of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> regarding conservation status, distribution, and genetics.</p></sec></sec><sec sec-type="materials|methods"><title>Materials and methods</title><sec sec-type="Taxonomic data mining, storing and tagging (Appendix 1)"><title>Taxonomic data mining, storing and tagging (<xref ref-type="app" rid="APP1">Appendix 1</xref>)</title><p><bold>Literature.</bold> In addition to traditional searching of, for example, the Zoological Record, we also searched for the terms “Geothauma, Plectostoma and Opisthostoma” in Google Scholar (on 21<sup>st</sup> November 2012) and the Biodiversity Heritage Library (on 19<sup>th</sup> November 2012). A URL link to the full-text article was provided as listed in bibliography whenever possible (e.g. from <ext-link ext-link-type="uri" xlink:href="http://www.biodiversitylibrary.org">http://www.biodiversitylibrary.org</ext-link>). Each of the articles was tagged with the relevant <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> taxon names. All the relevant references were catalogued as individual contents in bibliography of the <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> Lifedesks pages (<ext-link ext-link-type="uri" xlink:href="http://opisthostoma.lifedesks.org/biblio">http://opisthostoma.lifedesks.org/biblio</ext-link>). All the relevant bibliography metadata of each article were entered and stored according to the standard data entries of BibTeX (<ext-link ext-link-type="uri" xlink:href="http://www.bibtex.org/">http://www.bibtex.org/</ext-link>) as implemented in Lifedesks. Furthermore, the bibliography metadata can be exported as BibTeX formatted file (.bib), which all the tagged metadata can be retrieved and reused. For video tutorials, see <xref ref-type="app" rid="APP1">Appendix 1(1 and 2)</xref>.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Nomenclature information and classification.</bold> After the relevant literature was identified, the relevant taxon names were extracted and organised by using the classification tool (tree editor) of Lifedesks (for detailed descriptions of the methodology, see: <ext-link ext-link-type="uri" xlink:href="http://help.lifedesks.org/classification/edit">http://help.lifedesks.org/classification/edit</ext-link>). The classification and nomenclature information can be downloaded and saved according to the DarwinCore standard as xml file. For video tutorials, see <xref ref-type="app" rid="APP1">Appendix 1(3)</xref>.</p><p><bold>Species information.</bold> In addition, the original species descriptions and important notes from the literature were imported to individual species pages as quotations (for detailed descriptions of the methodology, see: <ext-link ext-link-type="uri" xlink:href="http://help.lifedesks.org/quickstart">http://help.lifedesks.org/quickstart</ext-link>). New and unpublished data, together with the information extracted from literature were managed and stored in the relevant chapters under the headings of ‘Overview’, ‘Conservation’, ‘Description’, ‘Ecology and Distribution’, ‘Evolution and Systematics’, and ‘Relevance’ of each species webpage. As all the information was tagged accordingly in the form of xhtml format, the data of each species can be retrieved and reused. For video tutorials, see <xref ref-type="app" rid="APP1">Appendix 1(4)</xref>.</p><p><bold>Managing museum collections data.</bold> As common curation practice, each collection (i.e. one museum lot) consists of a specimen or multiple specimens, which are kept either dry (empty shells) or wet (shells with animals preserved in ethanol) that were collected at a single sampling occasion, for example from a particular location at a particular day/time. In this study, each collection is regarded as a sample. For each sample, there are two categories of information that can be extracted, namely the physical properties and metadata of the samples.</p><p>For the physical properties of the samples, we recorded the exact number of shells or (for samples >10 shells) categorised the samples into four categories of sample size: 1) > 10 (10–24 shells); 2) > 25 (25–49 shells); 3) > 50 (50–100 shells); and 4) > 100 (>100 shells). We also obtained shell morphology data as follows. Whenever possible, each registered collection was photographed. The images for each unique collection were imported into Lifedesks as an individual content. Each of the images was then linked with the species name. For video tutorials, see <xref ref-type="app" rid="APP1">Appendix 1(5)</xref>.</p><p>Metadata consisted of collection reference number, collector information, collecting date, and locality of each sample. This information was published in Lifedesks as image description, except for the collection reference number, which was published as image caption. In addition to presenting collection data in a tabular format (<xref ref-type="app" rid="APP1">Appendix 2</xref>), we also published the collection data in a more interactive manner, which can be used in Google Earth.</p><p>Whenever possible, location data of the collections were georeferenced. When the location description in the specimen label or in the publication was not clear, the itinerary of the collectors or expedition which had been published in other types of publication (e.g. maps and reports) was consulted. After the exact limestone hill where the collection had been made was identified, it was verified in Google Earth, after which the latitude and longitude were obtained. Although the coordinates as obtained from Google Earth are of high accuracy, they might be too accurate and lack an uncertainty estimate (<xref rid="B70" ref-type="bibr">Mesibov 2012</xref>). Thus, the coordinates that we report here should be inter<pmc-comment>PageBreak</pmc-comment>preted as the location of the limestone hill and not the exact spot where the specimens were collected.</p><p>A Keyhole Markup Language (KML) file, which comprises the location, images, and collection reference number of each of the museum collections, was created by using Google Earth Spreadsheet Mapper v3.0 (<ext-link ext-link-type="uri" xlink:href="http://www.google.com/earth/outreach/tutorials/spreadsheet3.html">http://www.google.com/earth/outreach/tutorials/spreadsheet3.html</ext-link>). For the data input for the spreadsheet (Template4), the species name was used for “Folder name”, and collection reference number for “Placemark Name”; the concatenation of species name and collection reference number for “Title”; URL of each collection’s original image in Lifedesks was named “Image URL”; detailed collection data as “Paragraph Text”. After that, the data in Spreadsheet Mapper were converted into a KML file, which allows semantic-tagged collection information to be retrieved and reused. When the KML file is opened in Google Earth, each of the museum collection (specimens) is shown as a single landmark on the virtual earth. For video tutorials, see <xref ref-type="app" rid="APP1">Appendix 1(6)</xref>.</p></sec><sec sec-type="Specimen repositories"><title>Specimen repositories</title><p><abbrev>BMNH<def><p> Natural History Museum (previously known as British Museum (Natural History), London, United Kingdom.</p></def></abbrev></p><p><abbrev>BOR<def><p> BORNEENSIS collection, Institute for Tropical Biology and Conservation, Universiti Malaysia Sabah, Kota Kinabalu, Malaysia.</p></def></abbrev></p><p><abbrev>RMNH<def><p> Naturalis Biodiversity Center (previously known as Rijksmuseum van Natuurlijke Historie), Leiden, the Netherlands</p></def></abbrev></p><p><abbrev>V<def><p> Jaap Vermeulen’s private collection, Leiden, the Netherlands.</p></def></abbrev></p><p><abbrev>YSC<def><p> Chen Yansen’s private collection, Medan, Sumatra, Indonesia.</p></def></abbrev></p><p><abbrev>ZMA<def><p> Naturalis Biodiversity Center (previously known as Zoological Museum of Amsterdam), Leiden, the Netherlands.</p></def></abbrev></p></sec><sec sec-type="Establishing identity and revising the collection data"><title>Establishing identity and revising the collection data</title><p>In the routine of conventional taxonomic revision practice, samples are sorted into groups (e.g. morphospecies) based on their morphology and distribution. Then, each group is assigned to an existing species name–when the morphology, distribution and/or other important characteristics fit with the species’ morphological description and distributional range as mentioned in the literature; or a newly designed species name–when the characteristics do not fit to any of the named species. In some cases, different species identities may have been assigned to the same specimen by multiple taxonomists. Thus, for each specimen, the collection data (morphology, distribution, genetics and others) are immutable, but the species name is mutable.</p><pmc-comment>PageBreak</pmc-comment>
<p>The key of this process is the taxonomists, who gather, integrate, sort and analyse, not only the biological specimens, but diverse and vast amounts of information from hundreds of specimens (<xref rid="B102" ref-type="bibr">Sluys 2013</xref>). This task has become more challenging for taxonomists and their successors because of the accumulative nature of taxonomic information. Thus, taxonomists have been using information technology to assist their routine work since the 1980s (<xref rid="B51" ref-type="bibr">Heywood 1974</xref>, <xref rid="B68" ref-type="bibr">Maxted 1992</xref>). However, the potential of information technology to be used by taxonomists remains underexploited, except for data storage. In fact, in additional to data storage, this technology can improve the efficiency and effectiveness of taxonomists in integrating, sorting, analysing and disseminating the information from the specimens.</p><p>As mentioned above, all key information from specimens and literature was digitised and tagged. Here, we integrated different types of information for different processes in taxonomy revision. For the specimen sorting process, we used a KML file to link and present the unique collection number, images and location data for each specimen. Each museum collection was shown on Google Earth as a landmark and these landmarks were sorted into respective species folders. When each landmark was selected (by clicking it), the information of the morphology (as shown in images), location (text description and map), and other relevant information was visible to the user (taxonomist). Species identification can be done in a single platform (i.e. Google Earth), where the morphological variation within a species or between the specimens across the genus’ geographical range can be examined. Based on this information, the species identity of the landmark (specimen) was determined by either keeping the landmark in the same species folder or moving to the other species folder. Likewise, whenever the coordinate of a specimen location was wrong, the landmark was edited by moving it to the correct location. Whenever necessary, the specimen itself was examined. For video tutorials, see <xref ref-type="app" rid="APP1">Appendix 1(7)</xref>.</p><p>After all the collection species identifications and location data in the KML file were verified and corrected, the data in the KML file were extracted with a customised Python script to update the data in Spreadsheet mapper and image species link in Lifedesks for all specimens in an automatic manner. This saved much time compared to the traditional laborious method of manually updating the collection database specimen by specimen after sorting and identification. Lastly, the taxonomic content was written in Lifedesks and then exported to the appropriate format and layout for publication. For video tutorials, see <xref ref-type="app" rid="APP1">Appendix 1(8)</xref>.</p><p>Since the information was stored and tagged digitally, a simple program can be customised to retrieve, integrate and process the data from many different online/offline databases and files. In our case, we used scripting language Python 2.73 (<ext-link ext-link-type="uri" xlink:href="http://www.python.org">http://www.python.org</ext-link>). Its “urllib” and “re” modules were used to retrieve information from Internet resources and searching for patterns in text. In the same way, additional specimen information from Lifedesks (i.e. image pages) can be integrated into the KML file. Simultaneously, the literature and species information compiled in Lifedesks can be retrieved easily when necessary.</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="Species delimitation"><title>Species delimitation</title><p>The application of a species concept in <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> has been particular problematic. Nowadays, the most widely accepted species concepts (e.g., the biological species concept; <xref rid="B69" ref-type="bibr">Mayr 1942</xref>) include some aspect of genetic and/or reproductive cohesion. Such species concepts are, however, difficult to apply in taxa like <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>, where all species and populations are restricted to isolated limestone outcrops. This island-like (allopatric) distribution pattern suggests very limited gene flow between populations. Furthermore, it would be impractical to verify experimentally “potential interbreeding” for each population, as <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> populations occur in hundreds of different limestone outcrops.</p><p>Previous species circumscriptions in this genus have been mainly based on a morphological species concept. In gastropods, this is common practice. However, shell shape or even some shell structures, such as rib intensity, can be genetically variable and/or phenotypically plastic under different environmental conditions (e.g. <xref rid="B11" ref-type="bibr">Berry 1962</xref>, <xref rid="B58" ref-type="bibr">Kemp and Bertness 1984</xref>). Nevertheless, when the intra- and inter-specific variations in shells are understood (see below “Morphological analysis”), a morphological species concept may be used as one of the guidelines in species delimitation.</p><p>Finally, the phylogenetic species concept (sensu <xref rid="B30" ref-type="bibr">Cracraft 1983</xref>–“A species is the smallest diagnosable cluster of individual organism within which there is a parental pattern of ancestry and descent”) also cannot be used in the case of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species, again because of their populations’ allopatric distribution on isolated limestone outcrops. The deposition age of Peninsular Malaysian limestone ranges from the Ordovician to the late Triassic (<italic>ca.</italic> 480 mya–200 mya). Though the exact time at which the limestone outcrop was exposed is unknown, <xref rid="B7" ref-type="bibr">van Benthem Jutting (1960)</xref> believed this began to happen after the Cretaceous - <italic>ca.</italic> 140 mya. Given the fact that <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species have been found on relatively young, Miocene age, limestone (<italic>ca.</italic> 24 mya–5 mya) in Borneo, it is likely that <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species will have colonised limestone hills in Peninsular Malaysia soon after the limestone hills were exposed. Hence, populations of the same <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species on each isolated limestone hill could have been separated for a long time and thus these isolated populations would appear as a several deeply diverged lineages in the phylogenetic tree (Liew TS, unpublished data). In view of this, the blind application of a phylogenetic species concept could inflate the number of species in <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>.</p><p>Because of these problems, in this study, we have used elements of different species concepts for the delimitation of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species. First we used a set of shell characters for initial morphological species delimitation. These groups were then checked for their distribution ranges. Whenever more than one of these morphological species was found at the same hill, we examined the genetic dissimilarity (in DNA barcode, COI) between these sympatric morphological species. In addition, the intra-specific genetic divergence was examined among several geographically separated populations of the same species (see below “COI Barcoding”). By reciprocal illumination from the morphological, genetic and distribution data, we determined for each species a set of characters that is stable within a species and diagnosable between <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species. We applied the same approach to the Bornean species (Liew et al. in prep).</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="Morphological analysis"><title>Morphological analysis</title><p>In conventional conchology, shell descriptions and measurements are mainly made based on the standard apertural view of the shell. In this view, the shell is positioned so that the columella is vertical and the shell is rotated around this columella axis until the aperture faces the user. After the apertural view of the shell is set, the shell linear measurements are taken and descriptions of other shell characters are made (see, e.g., <xref rid="B122" ref-type="bibr">Vermeulen 1994</xref>). However, the irregularity in the orientation of the aperture of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> shells, caused by the distortion in the shell coiling hinder this traditional conchological approach.</p><p>We feel that in <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> the usefulness of this traditional approach is limited because of the presence of the tuba that deviate from the coiling axis of the spire. The varying length and coiling mode of the tuba prevent any standardisation of the ‘frontal’ view of the spire (<xref ref-type="fig" rid="F1">Figure 1</xref>; see also <xref rid="B94" ref-type="bibr">Sasaki 2010</xref>, who refers to “different direction in shell-mouth opening”). Nevertheless, all previous authors use this method to describe and illustrate <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> shells. Here, we proposed a better approach to describe the shells of this, conchologically unusual, genus.</p><fig id="F1" orientation="portrait" position="float"><label>Figures 1.</label><caption><p>Six shell views. The shaded circle is the operculum. The frontal view (perpendicular on the operculum) was set as reference perspective for the other views: back, right lateral, left lateral, top, and bottom.</p></caption><graphic xlink:href="zookeys-393-001-g001"></graphic></fig><p>In <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>, the only part of the shell that can serve as a landmark to determine a frontal view, while at the same time fixing the position of both the spire as well as the tuba, is not the aperture, but the constriction, the point where the spire ends and the tuba begins, and operculum rests. Therefore, we determined the frontal view of the shell as follows: the shell is held with the coiling axis of the spire vertical and with the operculum perpendicular to the line of view, on the right side (because all <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> shells are dextral). Left lateral view, back view and right lateral view are obtained by turning the shell 90°, 180°, and 270° from this starting point (<xref ref-type="fig" rid="F1">Figure 1</xref>). Accordingly, all the shell characters were described based on this positioning scheme.</p><p><bold>Specimen images and 3D models.</bold> To examine the shell morphology, at least one digital 3D model of each species was obtained. We used microcomputed X-ray tomography (CT) to obtain 3D models of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> shells. For each species, several empty shells and ethanol-preserved shells with the soft body inside, spaning the morphological variation breadth, were selected. Microcomputed tomography was carried in a high-resolution micro-CT scanner (SkyScan, model 1172, Aartselaar, Belgium). 3D models were created from the reconstructed images with the manufacturer’s software CT Analyser v1.12.0.0 (Skyscan©) and saved as digital polygon mesh objects (*.ply). The 3D models were then simplified by quadric edge collapse decimation (to ca. 200,000 faces) as implemented in the program MeshLab v1.3.2 (<xref rid="B22" ref-type="bibr">Cignoni et al. 2008</xref>).</p><p>The position and orientation of the 3D digital shell was manipulated so that the shell columella was in parallel with the z-axis and the operculum outer side was visible from a user perspective (<xref ref-type="fig" rid="F1">Figure 1</xref>, <xref ref-type="app" rid="APP1">Appendix 1(9)</xref>). Then, the outer operculum view of the shell was regarded as frontal view. After that, the field of view of the 3D model was set to orthographic, and an image was taken for each of the six perspectives: frontal view (A), left lateral view (B), back view (C), right lateral view (D), top view (E), and bottom view (F). In addition, two images were made of the constriction teeth of <pmc-comment>PageBreak</pmc-comment>the parietal (G) and basal (H) inner shell whorl after clipping of the 3D model. All manipulation and imaging was done with MeshLab v1.3.2 (<xref rid="B22" ref-type="bibr">Cignoni et al. 2008</xref>). Thus, a total of eight images were made for each species (A–H). For video tutorials, see <xref ref-type="app" rid="APP1">Appendix 1(10)</xref>.</p><p><bold>Shell characters and descriptions.</bold> The shell is an accretionary exoskeleton of the snail. The overall shape of the shell, which resembles a 3D spiral, results from changes in the curvature and torsion during the shell accretionary process, and form changes in aperture form during shell growth (<xref rid="B78" ref-type="bibr">Okamoto 1988</xref>). However, the exact quantifications of these changes might exceed the requirements of the practical purpose of this taxonomic paper. Therefore, we used traditional linear measurements to quantify the shell form, in a way that these measurements abstract the shell ontogeny and its 3D spiral properties.</p><p>After the six views of a shell were determined as described (<xref ref-type="fig" rid="F1">Figure 1</xref>), the shell whorls were described for each of six major parts, according to the shell ontogeny order: (1) apex–protoconch and the first teleoconch (<xref ref-type="fig" rid="F2">Figure 2A</xref>, and <xref ref-type="fig" rid="F3">3</xref>); (2) apical <pmc-comment>PageBreak</pmc-comment>spire–the whole teleoconch except the last 1 1/2 whorls before the constriction (<xref ref-type="fig" rid="F2">Figure 2B</xref>, and <xref ref-type="fig" rid="F4">4</xref>); (3) basal spire–the last 1 1/2 whorls before the constriction (<xref ref-type="fig" rid="F2">Figure 2B</xref>, and <xref ref-type="fig" rid="F5">5</xref>); (4) constriction–the narrowest transitional part of the whorl between spire and tuba (<xref ref-type="fig" rid="F2">Figure 2A</xref>, <xref ref-type="fig" rid="F6">6</xref>, and <xref ref-type="fig" rid="F7">7</xref>); (5) tuba whorl–teleoconch after the constriction (<xref ref-type="fig" rid="F2">Figure 2A</xref>, and <xref ref-type="fig" rid="F8">8</xref>); and (6) aperture and peristome (<xref ref-type="fig" rid="F2">Figure 2A</xref>, <xref ref-type="fig" rid="F9">9</xref>, and <xref ref-type="fig" rid="F10">10</xref>). The first three parts <pmc-comment>PageBreak</pmc-comment>constitute the shell spire, for which size and shape were quantified from the left lateral view. The height, width and number of whorls of the shell were measured and counted from the spire (<xref ref-type="fig" rid="F11">Figure 11A</xref>, and <xref ref-type="fig" rid="F11">B</xref>). In addition to the description of the general shell form, we recorded the shell surface ornamentations, namely, (7) fine spiral striation (<xref ref-type="fig" rid="F12">Figure 12</xref>), and (8) distinct radial ribs (<xref ref-type="fig" rid="F2">Figures 2A</xref> and <xref ref-type="fig" rid="F13">13</xref>).</p><fig id="F2" orientation="portrait" position="float"><label>Figures 2.</label><caption><p>Morphological terminology and shell measurements. <bold>A</bold> shell part terminology as used in the species descriptions is shown in an example shell <bold>B</bold> Two shell examples show the basal spire that consist of the last two whorls of the spire, and the apical spire that consist of the remaining spire whorls.</p></caption><graphic xlink:href="zookeys-393-001-g002"></graphic></fig><fig id="F3" orientation="portrait" position="float"><label>Figures 3.</label><caption><p>Apex forms in shell left lateral view. The degree of shell apex depression results from the growth regime of the teleoconch after the protoconch.</p></caption><graphic xlink:href="zookeys-393-001-g003"></graphic></fig><fig id="F4" orientation="portrait" position="float"><label>Figures 4.</label><caption><p>Two basic forms of apical spire whorls: <bold>A</bold> oblong conical <bold>B</bold> depressed conical.</p></caption><graphic xlink:href="zookeys-393-001-g004"></graphic></fig><fig id="F5" orientation="portrait" position="float"><label>Figures 5.</label><caption><p>Three basic forms of shell spire basal whorls in left lateral view. Panels from left to right of each of the shell forms: quantifications of the shell form in top view; example of the shell form with a low number of whorls in left lateral view; example of the shell form with a higher number of whorls in left lateral view. <bold>A</bold> conical <bold>B</bold> ovoid <bold>C</bold> ellipsoid.</p></caption><graphic xlink:href="zookeys-393-001-g005"></graphic></fig><fig id="F6" orientation="portrait" position="float"><label>Figures 6.</label><caption><p>Different types of parietal constriction teeth before the operculum resting site, in bottom view. <bold>A</bold> two long lamellae run parallel to the whorl growing direction <bold>B</bold> two short ridges run parallel to the whorl growing direction, each knob-shaped at one end <bold>C</bold> a single ridge runs parallel to the whorl growing direction, knob-shaped at one end <bold>D</bold> no tooth.</p></caption><graphic xlink:href="zookeys-393-001-g006"></graphic></fig><fig id="F7" orientation="portrait" position="float"><label>Figures 7.</label><caption><p>Different types of basal constriction teeth in top view. <bold>A</bold> two teeth after operculum resting site: one ridge runs parallel to the whorl growing direction; the other ridge has a knob at one end and runs perpendicular to the whorl growing direction <bold>B</bold> no tooth.</p></caption><graphic xlink:href="zookeys-393-001-g007"></graphic></fig><fig id="F8" orientation="portrait" position="float"><label>Figures 8.</label><caption><p>Different tuba coiling regimes in bottom view. <bold>A</bold> tuba type 1–tuba coiling as regularly as the spire last whorl <bold>B</bold> tuba type 2–tuba gradually coiling downward and then in a different direction <bold>C</bold> tuba type 3–tuba bent abruptly. Arrows point to the constriction.</p></caption><graphic xlink:href="zookeys-393-001-g008"></graphic></fig><fig id="F9" orientation="portrait" position="float"><label>Figures 9.</label><caption><p>Positioning scheme for the description of aperture and peristomes. <bold>A</bold> anterior, posterior, right and left lateral sides of aperture and peristome are defined according to the orientation of the shell relative to the active animal <bold>B</bold> Three shell examples showing the defined positions of aperture and peristome. Red arrows point to the aperture area with the densest ribs.</p></caption><graphic xlink:href="zookeys-393-001-g009"></graphic></fig><fig id="F10" orientation="portrait" position="float"><label>Figures 10.</label><caption><p>Aperture and peristome forms. <bold>A</bold> outer and inner peristome <bold>B</bold> Simple aperture without outer peristome <bold>C</bold> aperture with double peristomes in which the outer peristome is prominent <bold>D</bold> aperture with double peristomes in which the outer peristome is equally prominent as the inner.</p></caption><graphic xlink:href="zookeys-393-001-g010"></graphic></fig><fig id="F11" orientation="portrait" position="float"><label>Figures 11.</label><caption><p>Shell measurements. <bold>A</bold> height and width of spire <bold>B</bold> number of whorls <bold>C</bold> spire last whorl length, tuba whorl length and umbilicus openness.</p></caption><graphic xlink:href="zookeys-393-001-g011"></graphic></fig><fig id="F12" orientation="portrait" position="float"><label>Figures 12.</label><caption><p>Spiral lines on the shell surface are shown in, from top to bottom, 100 × magnification under the dissecting microscope, 500 × and 2000 × magnification under scanning electron microscope. <bold>A</bold> shell with both thick and thin spiral lines <bold>B</bold> shell with only thin spiral lines. Each corresponding image in <bold>A</bold> and <bold>B</bold> is at the same scale.</p></caption><graphic xlink:href="zookeys-393-001-g012"></graphic></fig><fig id="F13" orientation="portrait" position="float"><label>Figures 13.</label><caption><p>Radial ribs. <bold>A</bold> measurement of number of ribs per mm at the whorl above the constriction in left lateral view <bold>B</bold> rib intensity, from left to right: thick ribs, thin ribs, and abraded ribs <bold>C</bold> three different rib shapes, from left to right: single-humped, slightly curved and straight.</p></caption><graphic xlink:href="zookeys-393-001-g013"></graphic></fig><p><bold>(1) Apex</bold> – Ranges from “distinctly convex”, via “moderately convex”, to “slightly convex” (<xref ref-type="fig" rid="F3">Figure 3</xref>). The slightly convex apex has a teleoconch that grows with less torsion and greater curvature than the protoconch, and vice versa in the distinctly convex apex.</p><p><bold>(2) Apical spire</bold> – Similar to the apex form, less torsion and greater curvature in shell growth produce a “depressed conical” apical spire, and the reverse produces an “oblong conical” apical spire. The oblong and depressed conical shape of the spire can be estimated by measuring the ratio between the apical spire height and width (<xref ref-type="fig" rid="F4">Figure 4</xref>).</p><p><bold>(3) Basal spire</bold> – The curvature of the basal spire determines the final form of the spire. The width of the spire base is related to how tightly the basal spire whorl coils towards the shell columella, and to the whorl width. The basal spire shape categories, namely, conical, ovoid, and ellipsoid, can be estimated by comparing the difference <pmc-comment>PageBreak</pmc-comment>between two whorl width measurements from three consecutive whorl peripheries (<xref ref-type="fig" rid="F5">Figure 5</xref>). This measurement is made at both the left and the right side of the shell, in left lateral view. The spire umbilicus may be open, partially closed or totally closed by the tuba.</p><p><bold>(4) Constriction</bold> – The constriction is a short transitional and narrow part of the whorl between the spire and tuba (<xref ref-type="fig" rid="F2">Figure 2A</xref>). This is the furthest point to where the snail can retract into the shell and where the operculum rests (<xref ref-type="fig" rid="F1">Figure 1</xref>, and <xref ref-type="fig" rid="F2">2A</xref>). Inside the constriction of some species, there are calcareous structures that protrude from the inner shell wall–constriction teeth (<xref ref-type="fig" rid="F6">Figure 6</xref> and <xref ref-type="fig" rid="F7">7</xref>). The number, shape, and location of the constriction teeth are taxonomically informative characters.</p><p><bold>(5) Tuba</bold> – It is difficult to describe the variable forms of tuba in words. For the sake of convenience, we categorised the tuba into three coiling regimes, which represent <pmc-comment>PageBreak</pmc-comment>how many times torsion changes drastically. These are: type 1–no drastic changes in torsion at the beginning of the tuba as compared to the spire; type 2–drastic changes in torsion at the beginning and midway of the tuba; and type 3–drastic changes in torsion <pmc-comment>PageBreak</pmc-comment>at the beginning of tuba (<xref ref-type="fig" rid="F8">Figure 8</xref>). In addition to the coiling regime, the overall shell form is also determined by the final tuba length. However, estimation of tuba length is difficult. Hence, we quantified the tuba length by estimating the ratio of the tuba periphery length on the one hand, and the spire last whorl periphery length on the other. In addition, we estimated the proportion of the tuba that attaches to the spire. Finally, <pmc-comment>PageBreak</pmc-comment>the difference between tuba forms can also be determined by comparing the shell perspective where the aperture opening is best visible.</p><p><bold>(6) Aperture and peristome</bold> – Before we characterise the aperture form, we define the anatomical position of the aperture according to the orientation of the animal inside (<xref ref-type="fig" rid="F9">Figure 9</xref>). Our definitions for the four areas of aperture side correspond to the conventional terminology used for the aperture of a regularly coiling shell: Anterior = palatal side, posterior = columellar side, right lateral = parietal side and left lateral = basal side. The convenient way to recognise these aperture areas is by identifying the posterior area where it has the densest ribs (<xref ref-type="fig" rid="F9">Figure 9B</xref>). After that we describe and compare the shape of the aperture and the outer peristome (<xref ref-type="fig" rid="F10">Figure 10</xref>). The shell has either a simple or a double peristome, and this character is species specific in <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>. The prominence and shape of the outer peristome is described by how much the outer peristome is projected at the anterior, posterior, and both lateral sides, as compared to the inner peristome (<xref ref-type="fig" rid="F10">Figure 10A</xref>).</p><p><bold>(7) Spiral lines</bold> – Spiral line sculpture on the shell is composed of a row of granulated micro-structures. The intensity of the spiral lines depends on the size of these micro-structures. In general, Thick spiral lines should be visible under the dissecting microscope at 50× magnification (<xref ref-type="fig" rid="F12">Figure 12A</xref>), whereas thin spiral lines are hardly visible at 50× magnification, but are visible at 100× magnification (<xref ref-type="fig" rid="F12">Figures 12A and B</xref>, <xref ref-type="app" rid="APP1">Appendix 3</xref>). Thick spiral lines are more widely spaced (< 7 lines per 100 μm) than thin lines (> 10 lines per 100 μm). Thin lines may be present in between sparse thick lines (<xref ref-type="fig" rid="F12">Figure 12A</xref>), but on living snails these may fade away when the snail ages and may be hardly visible in old empty shells.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>(8) Radial ribs</bold> – Radial ribs are produced by a change of shell ontogeny in both shell accretion direction and aperture dimensions. During rib formation, the shell material accretion direction around the aperture changes from longitudinal to orthogonal; meanwhile, aperture size increases and probably aperture shape changes. Thus, the formation of each rib represents a discontinuity in shell growth in the longitudinal direction and a change of the aperture shape and size. In view of these, three characters can be observed in the radial ribs.</p><pmc-comment>PageBreak</pmc-comment>
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<p>The first character is the total number of ribs and the spacing between them on the shell. The number of ribs for a species can be highly variable between different individuals. The ribs are not evenly distributed on the shell surface; for example, the spacing between ribs consistently increases from the apex to the last whorl of the spire <pmc-comment>PageBreak</pmc-comment>(<xref ref-type="app" rid="APP1">Appendix 4</xref>). In view of this, we describe the rib density as the number of ribs within 1 mm on the whorl above the shell constriction in left lateral view (<xref ref-type="fig" rid="F13">Figure 13A</xref>).</p><p>The second character is the intensity of the ribs in terms of length and thickness. Generally, rib length is related to the spacing between the ribs; for example, the greater the spacing (whorl length) before the rib, the longer the rib projects from the whorl periphery (Liew T.S. unpublished data). Thus, the lengths of the ribs, from the apex to the last whorl of the shell, change in a trend similar to the rib spacing. Besides, the thickness of ribs can vary between species, but less so within species. The thickness of the ribs depends on the number of shell layers and the thicker the ribs, the more likely it is that ribs persist in old shells (<xref ref-type="fig" rid="F13">Figure 13B</xref>).</p><pmc-comment>PageBreak</pmc-comment>
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<p>The third character is the form of the ribs. As mentioned above, each radial rib on a <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> shell was actually a deformed aperture during shell ontogeny. Thus, a more biologically meaningful way to describe the radial ribs is to compare the rib edge form (deformed aperture) to the whorl before the rib (regular aperture). The rib form can be either the same or different in the spire and tuba parts of the shell. As is the case with the aperture and peristome, the rib edge is either slightly or distinctly projected at its anterior side (i.e. at whorl periphery) as compared to the lateral sides (hereafter “rib plate”). In addition, the shape of the rib plate can be straight, slightly curved, or single-humped (<xref ref-type="fig" rid="F13">Figure 13C</xref>). Although thinner ribs are easily abraded in old shells, at least the rib plate form can be inferred from the abraded scar at the whorl periphery (<xref ref-type="fig" rid="F13">Figure 13B</xref>).</p><p>Finally, we noticed variation among species in the rib inclination, which can be estimated with respect to the coiling axis. However, it is difficult to quantify this inclination accurately. Nevertheless, we add this character into the description of the shell in a qualitative manner in the terms of orthoclin (i.e. ribs are almost straight with respect to the collumella, as in <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic>), prosoclin (i.e. ribs are distinctly tilted with respect to the collumella, as in <italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic>).</p><p>In brief, shells of all species were described following a template consisting of three major elements (<xref ref-type="table" rid="T1">Table 1</xref>). First, the shell was described in several parts which represent the chronology of the shell ontogeny. Second, we defined a set of characters that can be determined in each shell part. Lastly, we described each character in either a quantitative or quantitative manner. We regard this template as a morphological model for the taxonomy of the genus <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>, and each element in this template may be updated by future taxonomists when necessary. We present the final description of each species in a uniform telegraphic format (e.g. semantic-tagged by bold text, and colon “:”) so that these morphological data can be mined effectively. By doing this, we hope to reduce the redundant process where species descriptions are done de novo each time a taxonomist revises the same taxa (<xref rid="B36" ref-type="bibr">Deans et al. 2012</xref>).</p><table-wrap id="T1" orientation="portrait" position="float"><label>Table 1.</label><caption><p>Format for species shell description.</p></caption><table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Shell parts (<xref ref-type="fig" rid="F2">Figure 2</xref>)</th><th rowspan="1" colspan="1">Characters for shell part<xref ref-type="table-fn" rid="TN1.1">†</xref></th><th rowspan="1" colspan="1">Descriptions (see section “Morphological analysis” for details).</th></tr><tr><td rowspan="1" colspan="1">Apex.</td><td rowspan="1" colspan="1">Shape: #.</td><td rowspan="1" colspan="1"><xref ref-type="fig" rid="F3">Figure 3</xref>.</td></tr><tr><td rowspan="1" colspan="1">Spire.</td><td rowspan="1" colspan="1">Height: #.<break></break>Width: #.<break></break>Number of whorls: #.<break></break>Apical spire shape: #.<break></break>Basal spire shape: #.<break></break>Whorl periphery: #.<break></break>Umbilicus: #.</td><td rowspan="1" colspan="1"><xref ref-type="fig" rid="F2">Figure 2</xref>.<break></break><xref ref-type="fig" rid="F4">Figure 4</xref>.<break></break><xref ref-type="fig" rid="F5">Figure 5</xref>.<break></break><xref ref-type="fig" rid="F11">Figure 11</xref>.</td></tr><tr><td rowspan="1" colspan="1">Constriction.</td><td rowspan="1" colspan="1">Parietal teeth: #.<break></break>Basal teeth: #.</td><td rowspan="1" colspan="1"><xref ref-type="fig" rid="F6">Figure 6</xref>.<break></break><xref ref-type="fig" rid="F7">Figure 7</xref>.</td></tr><tr><td rowspan="1" colspan="1">Tuba.</td><td rowspan="1" colspan="1">Coiling direction: # and aperture visible in # view.<break></break>Tuba whorl length in proportion to spire last whorl: #.<break></break>Proportion of tuba that attaches to spire: #.</td><td rowspan="1" colspan="1"><xref ref-type="fig" rid="F1">Figure 1</xref>.<break></break><xref ref-type="fig" rid="F8">Figure 8</xref>.<break></break><xref ref-type="fig" rid="F11">Figure 11</xref>.</td></tr><tr><td rowspan="1" colspan="1">Aperture and peristome.</td><td rowspan="1" colspan="1">Peristome: #.<break></break>Outer peristome shape: #.</td><td rowspan="1" colspan="1"><xref ref-type="fig" rid="F9">Figure 9</xref>.<break></break><xref ref-type="fig" rid="F10">Figure 10</xref>.</td></tr><tr><td rowspan="1" colspan="1">Spiral lines.</td><td rowspan="1" colspan="1">Thick lines: #.<break></break>Thin lines: #.</td><td rowspan="1" colspan="1"><xref ref-type="fig" rid="F12">Figure 12</xref>.</td></tr><tr><td rowspan="1" colspan="1">Radial ribs.</td><td rowspan="1" colspan="1">Rib density: # ribs per mm.<break></break>Rib intensity: #.<break></break>Shape: #.<break></break>Inclination: #.</td><td rowspan="1" colspan="1"><xref ref-type="fig" rid="F13">Figure 13</xref>.</td></tr></tbody></table><table-wrap-foot><fn id="TN1.1"><p>† “#” is the description for each shell character for <xref ref-type="fig" rid="F1">Figure 1</xref>–<xref ref-type="fig" rid="F13">13</xref>.</p></fn></table-wrap-foot></table-wrap><p><bold>Digital model.</bold> Pictures are more effective than verbal descriptions for shell morphology. However, it is not feasible to have hundreds of pictures taken for each perspective of a shell. Many non-linear characteristics of a shell cannot effectively be representedby 2D images. Thus, an interactive 3D model shell improves the dissemination of morphological information. Presenting 3D models in digital publication has started five year ago (<xref rid="B92" ref-type="bibr">Ruthensteiner and Heß 2008</xref>), and since then more taxonomists have taken the initiative to embed 3D models in e-papers. However, in this paper we have refrained from embed 3D models in the paper itself, since this limits further analysis by readers. Instead, we provide sets of 3D data in *.blend files, which consists of all 3D models and which can be opened in Blender v2.63 (<ext-link ext-link-type="uri" xlink:href="http://www.blender.org">www.blender.org</ext-link>). The 3D models in the blend file can be exported to *.ply format, which can be opened in MeshLab v1.3.2 (<xref rid="B22" ref-type="bibr">Cignoni et al. 2008</xref>). Both Blender and MeshLab are freeware and can be used to analyse the 3D model further (e.g. measurements, modification, etc). For video tutorials, see <xref ref-type="app" rid="APP1">Appendix 1(11 and 12)</xref>.</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="Conservation status assessment"><title>Conservation status assessment</title><p>We propose the conservation status for each species by following IUCN Red List Criteria and guidelines (<xref rid="B56" ref-type="bibr">IUCN Standards and Petitions Subcommittee 2013</xref>). We assess the conservation status based on our fieldwork in Malaysia between the year 2010 and year 2013 and the information obtained from the museum collections.</p></sec><sec sec-type="Molecular phylogeny and COI barcoding"><title>Molecular phylogeny and COI barcoding</title><p><bold>Taxon sampling.</bold> A total of 27 ingroup taxa of the genus <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> (n=11) and <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> (n=16) were included in this study: six <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> species from Borneo, five <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> from Peninsular Malaysia, nine <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species from Borneo, and seven <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> from Peninsular Malaysia. All of these ingroup taxa were selected on the basis of their distribution and shell forms which are representative for about 150 species in both genera. In addition to the ingroup taxa, eight outgroup taxa were included in the phylogenetic analysis. Sequence data for these outgroup taxa, which include three genera of the <named-content content-type="taxon-name">Diplommatinidae</named-content> and a species of the <named-content content-type="taxon-name">Cochlostomatidae</named-content>, were obtained from <xref rid="B127" ref-type="bibr">Webster et al. (2012)</xref>. The details of these specimens and the Genbank accession numbers are listed in <xref ref-type="table" rid="T2">Table 2</xref>.</p><table-wrap id="T2" orientation="portrait" position="float"><label>Table 2.</label><caption><p>Details of the taxa used for phylogenetic analysis and DNA barcoding.</p></caption><table frame="hsides" rules="groups"><tbody><tr><th rowspan="2" colspan="1">no.</th><th rowspan="2" colspan="1">Species</th><th rowspan="2" colspan="1">Voucher codes</th><th rowspan="2" colspan="1">Locality (Country, state, location, Lat, Long)</th><th rowspan="1" colspan="4">Genbank accession no.</th></tr><tr><th rowspan="1" colspan="1">COI</th><th rowspan="1" colspan="1">16S</th><th rowspan="1" colspan="1">18S</th><th rowspan="1" colspan="1">28S</th></tr><tr><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Cochlostoma septemspirale</named-content></italic> (Wagner, 1897)</td><td rowspan="1" colspan="1">RMNH.MOL.119825</td><td rowspan="1" colspan="1">Switzerland.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753326">HM753326</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753497">HM753497</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753423">HM753423</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753367">HM753367</ext-link></td></tr><tr><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Arinia paricostata</named-content></italic> (Vermeulen, 1996)</td><td rowspan="1" colspan="1">RMNH.MOL.119779</td><td rowspan="1" colspan="1">Malaysia, Sabah.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753500">HM753500</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753441">HM753441</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753384">HM753384</ext-link></td></tr><tr><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Diplommatina canaliculata</named-content></italic> (Moellendorff, 1886)</td><td rowspan="1" colspan="1">RMNH.MOL.119783</td><td rowspan="1" colspan="1">Malaysia, Pahang.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753338">HM753338</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753504">HM753504</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753445">HM753445</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753388">HM753388</ext-link></td></tr><tr><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Diplommatina gomantongensis</named-content></italic> (Smith, 1894)</td><td rowspan="1" colspan="1">RMNH.MOL.119800</td><td rowspan="1" colspan="1">Malaysia, Sabah.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753342">HM753342</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753509">HM753509</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753451">HM753451</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753394">HM753394</ext-link></td></tr><tr><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Diplommatina rubicunda</named-content></italic> (Von Martens, 1864)</td><td rowspan="1" colspan="1">RMNH.MOL.119819</td><td rowspan="1" colspan="1">Malaysia, Sabah.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753363">HM753363</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753520">HM753520</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753477">HM753477</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753417">HM753417</ext-link></td></tr><tr><td rowspan="1" colspan="1">6</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Diplommatina rubra</named-content></italic> (Godwin Austen, 1889)</td><td rowspan="1" colspan="1">RMNH.MOL.119797</td><td rowspan="1" colspan="1">Malaysia, Sabah.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753346">HM753346</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753514">HM753514</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753456">HM753456</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753399">HM753399</ext-link></td></tr><tr><td rowspan="1" colspan="1">7</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Palaina albata</named-content></italic> (Beddome, 1889)</td><td rowspan="1" colspan="1">RMNH.MOL.119810</td><td rowspan="1" colspan="1">Belau, Peleliu.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753354">HM753354</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753531">HM753531</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753469">HM753469</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753409">HM753409</ext-link></td></tr><tr><td rowspan="1" colspan="1">8</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Palaina striolata</named-content></italic> (Crosse, 1866)</td><td rowspan="1" colspan="1">RMNH.MOL.119812</td><td rowspan="1" colspan="1">Belau, Ngerekebesang.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753356">HM753356</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753533">HM753533</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753470">HM753470</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="HM753410">HM753410</ext-link></td></tr><tr><td rowspan="1" colspan="1">9</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Opisthostoma hailei</named-content></italic> Solem, 1964</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">Malaysia, Sabah, <named-content content-type="dwc:verbatimCoordinates">5.536, 118.213</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250856">KC250856</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250897">KC250897</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250923">KC250923</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250948">KC250948</ext-link></td></tr><tr><td rowspan="1" colspan="1">10</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Opisthostoma holzmarkii</named-content></italic> Thompson, 1978</td><td rowspan="1" colspan="1">BOR 5580</td><td rowspan="1" colspan="1">Malaysia, Sarawak, <named-content content-type="dwc:verbatimCoordinates">3.804, 113.78</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250871">KC250871</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250912">KC250912</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250937">KC250937</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250962">KC250962</ext-link></td></tr><tr><td rowspan="1" colspan="1">11</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> sp.</td><td rowspan="1" colspan="1">BOR 5561</td><td rowspan="1" colspan="1">Malaysia, Sarawak, <named-content content-type="dwc:verbatimCoordinates">4.044, 114.815</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250876">KC250876</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250917">KC250917</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250942">KC250942</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250967">KC250967</ext-link></td></tr><tr><td rowspan="1" colspan="1">12</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Opisthostoma brachyacrum brachyacrum</named-content></italic> (Thompson, 1978)</td><td rowspan="1" colspan="1">BOR 5548</td><td rowspan="1" colspan="1">Malaysia, Sarawak, <named-content content-type="dwc:verbatimCoordinates">1.319, 110.291</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250874">KC250874</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250915">KC250915</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250940">KC250940</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250965">KC250965</ext-link></td></tr><tr><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Opisthostoma brachyacrum brachyacrum</named-content></italic> (Thompson, 1978)</td><td rowspan="1" colspan="1">BOR 5593</td><td rowspan="1" colspan="1">Malaysia, Sarawak, <named-content content-type="dwc:verbatimCoordinates">3.804, 113.78</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250873">KC250873</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250914">KC250914</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250939">KC250939</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250964">KC250964</ext-link></td></tr><tr><td rowspan="1" colspan="1">14</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> sp.</td><td rowspan="1" colspan="1">BOR 5561</td><td rowspan="1" colspan="1">Malaysia, Sarawak, <named-content content-type="dwc:verbatimCoordinates">4.044, 114.815</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250878">KC250878</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250920">KC250920</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250945">KC250945</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250970">KC250970</ext-link></td></tr><tr><td rowspan="1" colspan="1">15</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Opisthostoma vermiculum</named-content></italic> Clements & Vermeulen, 2008</td><td rowspan="1" colspan="1">BOR 5526</td><td rowspan="1" colspan="1">Malaysia, Perak, <named-content content-type="dwc:verbatimCoordinates">4.55, 101.131</named-content>.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250910">KC250910</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">16</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Opisthostoma paulucciae</named-content></italic> Crosse & Nevill, 1879</td><td rowspan="1" colspan="1">BOR 5558</td><td rowspan="1" colspan="1">Malaysia, Pahang, <named-content content-type="dwc:verbatimCoordinates">4.487, 101.976</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250866">KC250866</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250907">KC250907</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">17</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Opisthostoma tenuicostatum</named-content></italic> van Benthem Jutting, 1952</td><td rowspan="1" colspan="1">BOR 5525</td><td rowspan="1" colspan="1">Malaysia, Kelantan, <named-content content-type="dwc:verbatimCoordinates">5.09, 102.22</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250869">KC250869</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250935">KC250935</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250960">KC250960</ext-link></td></tr><tr><td rowspan="1" colspan="1">18</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Opisthostoma platycephalum</named-content></italic> van Benthem Jutting, 1952</td><td rowspan="1" colspan="1">BOR 5576</td><td rowspan="1" colspan="1">Malaysia, Pahang, <named-content content-type="dwc:verbatimCoordinates">3.908, 103.147</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250864">KC250864</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250905">KC250905</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250931">KC250931</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250956">KC250956</ext-link></td></tr><tr><td rowspan="1" colspan="1">19</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Opisthostoma tenerum</named-content></italic> van Benthem Jutting, 1952</td><td rowspan="1" colspan="1">BOR 5577</td><td rowspan="1" colspan="1">Malaysia, Pahang, <named-content content-type="dwc:verbatimCoordinates">3.984, 103.144</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250861">KC250861</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250902">KC250902</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250928">KC250928</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250953">KC250953</ext-link></td></tr><tr><td rowspan="1" colspan="1">20</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma mirabile</named-content></italic> (Smith, 1893)</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">Malaysia, Sabah, <named-content content-type="dwc:verbatimCoordinates">5.531, 118.071</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250867">KC250867</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250908">KC250908</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250933">KC250933</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250958">KC250958</ext-link></td></tr><tr><td rowspan="1" colspan="1">21</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma obliquedentatum</named-content></italic> (Vermeulen, 1994)</td><td rowspan="1" colspan="1">BOR 5607</td><td rowspan="1" colspan="1">Malaysia, Sabah, <named-content content-type="dwc:verbatimCoordinates">4.726, 116.615</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250880">KC250880</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250922">KC250922</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250947">KC250947</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250972">KC250972</ext-link></td></tr><tr><td rowspan="1" colspan="1">22</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma hosei</named-content></italic> (Godwin-Austen, 1890)</td><td rowspan="1" colspan="1">BOR 5592</td><td rowspan="1" colspan="1">Malaysia, Sarawak, <named-content content-type="dwc:verbatimCoordinates">3.804, 113.78</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250860">KC250860</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250901">KC250901</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250927">KC250927</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250952">KC250952</ext-link></td></tr><tr><td rowspan="1" colspan="1">23</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma austeni</named-content></italic> (Smith, 1894)</td><td rowspan="1" colspan="1">BOR 5589</td><td rowspan="1" colspan="1">Malaysia, Sarawak, <named-content content-type="dwc:verbatimCoordinates">1.311, 110.293</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250863">KC250863</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250904">KC250904</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250930">KC250930</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250955">KC250955</ext-link></td></tr><tr><td rowspan="1" colspan="1">24</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma wallacei busauense</named-content></italic> (Smith, 1893)</td><td rowspan="1" colspan="1">BOR 5545</td><td rowspan="1" colspan="1">Malaysia, Sarawak, <named-content content-type="dwc:verbatimCoordinates">1.323, 110.3</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250875">KC250875</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250916">KC250916</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250941">KC250941</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250966">KC250966</ext-link></td></tr><tr><td rowspan="1" colspan="1">25</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma stellasubis</named-content></italic> (Vermeulen, 1994)</td><td rowspan="1" colspan="1">BOR 5588</td><td rowspan="1" colspan="1">Malaysia, Sarawak, <named-content content-type="dwc:verbatimCoordinates">3.804, 113.78</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250858">KC250858</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250899">KC250899</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250925">KC250925</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250950">KC250950</ext-link></td></tr><tr><td rowspan="1" colspan="1">26</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma pulchellum</named-content></italic> (Godwin-Austen, 1890)</td><td rowspan="1" colspan="1">BOR 5563</td><td rowspan="1" colspan="1">Malaysia, Sarawak, <named-content content-type="dwc:verbatimCoordinates">4.044, 114.815</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250857">KC250857</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250898">KC250898</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250924">KC250924</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250949">KC250949</ext-link></td></tr><tr><td rowspan="1" colspan="1">27</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma cookei</named-content></italic> (Smith, 1894)</td><td rowspan="1" colspan="1">BOR 5591</td><td rowspan="1" colspan="1">Malaysia, Sarawak, <named-content content-type="dwc:verbatimCoordinates">3.804, 113.78</named-content>.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250918">KC250918</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250943">KC250943</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250968">KC250968</ext-link></td></tr><tr><td rowspan="1" colspan="1">28</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma grandispinosum</named-content></italic> (Godwin-Austen, 1889)</td><td rowspan="1" colspan="1">BOR 5590</td><td rowspan="1" colspan="1">Malaysia, Sarawak, <named-content content-type="dwc:verbatimCoordinates">3.804, 113.78</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250879">KC250879</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250921">KC250921</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250946">KC250946</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250971">KC250971</ext-link></td></tr><tr><td rowspan="1" colspan="1">29</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">BOR 5509</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone hill on the right hand side of the road no. 8 direction to Gua Musang, opposite to Quarry Damai, <named-content content-type="dwc:verbatimCoordinates">4.896, 102.137</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420288">KC420288</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">30</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">BOR 5505</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Bukit Sejuk near Quarry Damai, along the road no. 8 from Gua Musang to Kuala Krai, <named-content content-type="dwc:verbatimCoordinates">4.905, 102.121</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420287">KC420287</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">31</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">BOR 5505</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Bukit Sejuk near Quarry Damai, along the road no. 8 from Gua Musang to Kuala Krai, <named-content content-type="dwc:verbatimCoordinates">4.905, 102.121</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420294">KC420294</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">32</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">BOR 5506</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone hills `Ciku 2`. In the FELDA plantation Ciku 2, <named-content content-type="dwc:verbatimCoordinates">4.924, 102.177</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420305">KC420305</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">33</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">BOR 5506</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone hills `Ciku 2`. In the FELDA plantation Ciku 2, <named-content content-type="dwc:verbatimCoordinates">4.924, 102.177</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420303">KC420303</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">34</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">BOR 5572</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone in FELDA Ciku 5, <named-content content-type="dwc:verbatimCoordinates">5.004, 102.2</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420271">KC420271</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">35</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">BOR 5509</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone hill on the right hand side of the road no. 8 direction to Gua Musang, opposite to Quarry Damai, <named-content content-type="dwc:verbatimCoordinates">4.896, 102.137</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420298">KC420298</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">36</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">BOR 5505</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Bukit Sejuk near Quarry Damai, along the road no. 8 from Gua Musang to Kuala Krai, <named-content content-type="dwc:verbatimCoordinates">4.905, 102.121</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420269">KC420269</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">37</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5512</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone hill near Kampung Paloh, on the right hand side of the road no 8 to Gua Musang, <named-content content-type="dwc:verbatimCoordinates">4.992, 102.228</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420304">KC420304</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">38</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5512</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone hill near Kampung Paloh, on the right hand side of the road no 8 to Gua Musang, <named-content content-type="dwc:verbatimCoordinates">4.992, 102.228</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420273">KC420273</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">39</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5515</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone hill in rubber estate (Ulu Kumbang FELCRA), <named-content content-type="dwc:verbatimCoordinates">4.69, 101.989</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420295">KC420295</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">40</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma davisoni</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5508</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone hill on the right hand side of the road D29, km 17 from Jelawang to Gua Musang, <named-content content-type="dwc:verbatimCoordinates">4.985, 101.965</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250872">KC250872</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250913">KC250913</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250938">KC250938</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250963">KC250963</ext-link></td></tr><tr><td rowspan="1" colspan="1">41</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma davisoni</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5508</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone hill on the right hand side of the road D29, km 17 from Jelawang to Gua Musang, <named-content content-type="dwc:verbatimCoordinates">4.985, 101.965</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420264">KC420264</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">42</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5612</td><td rowspan="1" colspan="1">Malaysia, Pahang, Dinding, <named-content content-type="dwc:verbatimCoordinates">3.846, 102.378</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420285">KC420285</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">43</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5504</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Gua Ikan. Loc 1, <named-content content-type="dwc:verbatimCoordinates">5.353, 102.026</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420286">KC420286</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">44</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5507</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Gua Ikan. Loc 1, <named-content content-type="dwc:verbatimCoordinates">5.353, 102.026</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420302">KC420302</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">45</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5504</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Gua Ikan. Loc 1, <named-content content-type="dwc:verbatimCoordinates">5.353, 102.026</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420301">KC420301</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">46</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5504</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Gua Ikan. Loc 1, <named-content content-type="dwc:verbatimCoordinates">5.353, 102.026</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250862">KC250862</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250903">KC250903</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250929">KC250929</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250954">KC250954</ext-link></td></tr><tr><td rowspan="1" colspan="1">47</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5507</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Gua Ikan. Loc 1, <named-content content-type="dwc:verbatimCoordinates">5.353, 102.026</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420278">KC420278</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">48</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5504</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Gua Ikan. Loc 1, <named-content content-type="dwc:verbatimCoordinates">5.353, 102.026</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420268">KC420268</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">49</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kakiense</named-content></italic> (Tomlin, 1948)</td><td rowspan="1" colspan="1">BOR 5516</td><td rowspan="1" colspan="1">Malaysia, Perlis, Kaki Bukit, <named-content content-type="dwc:verbatimCoordinates">6.645, 100.202</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420265">KC420265</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">50</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5533</td><td rowspan="1" colspan="1">Malaysia, Terengganu, Tasik Kenyir. Gua Bewah, <named-content content-type="dwc:verbatimCoordinates">4.851, 102.723</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420289">KC420289</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">51</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5533</td><td rowspan="1" colspan="1">Malaysia, Terengganu, Tasik Kenyir. Gua Bewah, <named-content content-type="dwc:verbatimCoordinates">4.851, 102.723</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420307">KC420307</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">52</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5519</td><td rowspan="1" colspan="1">Malaysia, Perlis, Bukit Kubu. Loc 3, <named-content content-type="dwc:verbatimCoordinates">6.404, 100.144</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420272">KC420272</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">53</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5519</td><td rowspan="1" colspan="1">Malaysia, Perlis, Bukit Kubu. Loc 3, <named-content content-type="dwc:verbatimCoordinates">6.404, 100.144</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420270">KC420270</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">54</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic> (Sykes, 1902)</td><td rowspan="1" colspan="1">BOR 5510</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone hill in Kampung Bayu. About 337 km from Kuala Lumpur by road no. 8, <named-content content-type="dwc:verbatimCoordinates">5.09, 102.22</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420279">KC420279</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">55</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic> (Sykes, 1902)</td><td rowspan="1" colspan="1">BOR 5510</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone hill in Kampung Bayu. About 337 km from Kuala Lumpur by road no. 8, <named-content content-type="dwc:verbatimCoordinates">5.09, 102.22</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420300">KC420300</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">56</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic> (Sykes, 1902)</td><td rowspan="1" colspan="1">BOR 5571</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Limestone in FELDA Ciku 4, <named-content content-type="dwc:verbatimCoordinates">5.043, 102.196</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420275">KC420275</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">57</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5574</td><td rowspan="1" colspan="1">Malaysia, Pahang, Sri Jaya, Gunung Mengapur, <named-content content-type="dwc:verbatimCoordinates">3.731, 102.821</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420299">KC420299</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">58</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5574</td><td rowspan="1" colspan="1">Malaysia, Pahang, Sri Jaya, Gunung Mengapur, <named-content content-type="dwc:verbatimCoordinates">3.731, 102.821</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420267">KC420267</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">59</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5520</td><td rowspan="1" colspan="1">Malaysia, Pahang, Bukit Serdam, <named-content content-type="dwc:verbatimCoordinates">3.83, 101.927</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250877">KC250877</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250919">KC250919</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250944">KC250944</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250969">KC250969</ext-link></td></tr><tr><td rowspan="1" colspan="1">60</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5520</td><td rowspan="1" colspan="1">Malaysia, Pahang, Bukit Serdam, <named-content content-type="dwc:verbatimCoordinates">3.83, 101.927</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420283">KC420283</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">61</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma relauensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5511</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Taman Negara, Sungai Relau Station. Gua Gajah, <named-content content-type="dwc:verbatimCoordinates">4.642, 102.063</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420277">KC420277</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">62</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic> (Tomlin, 1938)</td><td rowspan="1" colspan="1">BOR 5559</td><td rowspan="1" colspan="1">Malaysia, Pahang, Bukit Chintamanis, <named-content content-type="dwc:verbatimCoordinates">3.446, 102.014</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420297">KC420297</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">63</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5539</td><td rowspan="1" colspan="1">Malaysia, Pahang, Kenong Rimba Park. Gunung Kesong, <named-content content-type="dwc:verbatimCoordinates">4.194, 102.177</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420276">KC420276</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">64</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5542</td><td rowspan="1" colspan="1">Malaysia, Pahang, Kenong Rimba Park. Gunung Tangkup, <named-content content-type="dwc:verbatimCoordinates">4.194, 102.177</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420296">KC420296</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">65</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5541</td><td rowspan="1" colspan="1">Malaysia, Pahang, Kenong Rimba Park. Gunung Telahup, <named-content content-type="dwc:verbatimCoordinates">4.194, 102.177</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420266">KC420266</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">66</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5569</td><td rowspan="1" colspan="1">Malaysia, Pahang, Gua Bama, <named-content content-type="dwc:verbatimCoordinates">4.194, 101.967</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420280">KC420280</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">67</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5569</td><td rowspan="1" colspan="1">Malaysia, Pahang, Gua Bama, <named-content content-type="dwc:verbatimCoordinates">4.194, 101.967</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250868">KC250868</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250909">KC250909</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250934">KC250934</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250959">KC250959</ext-link></td></tr><tr><td rowspan="1" colspan="1">68</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5575</td><td rowspan="1" colspan="1">Malaysia, Pahang, Gua Charas, <named-content content-type="dwc:verbatimCoordinates">3.908, 103.147</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250859">KC250859</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250900">KC250900</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250926">KC250926</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250951">KC250951</ext-link></td></tr><tr><td rowspan="1" colspan="1">69</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">BOR 5537</td><td rowspan="1" colspan="1">Malaysia, Pahang, Gunung Jebak Puyuh, near Gunung Senyum, <named-content content-type="dwc:verbatimCoordinates">3.7, 102.453</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420291">KC420291</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">70</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">BOR 5537</td><td rowspan="1" colspan="1">Malaysia, Pahang, Gunung Jebak Puyuh, near Gunung Senyum, <named-content content-type="dwc:verbatimCoordinates">3.7, 102.453</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250870">KC250870</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250911">KC250911</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250936">KC250936</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250961">KC250961</ext-link></td></tr><tr><td rowspan="1" colspan="1">71</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5513</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Taman Negara, Sungai Relau Station. Gua Gajah, <named-content content-type="dwc:verbatimCoordinates">4.642, 102.063</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420292">KC420292</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">72</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5513</td><td rowspan="1" colspan="1">Malaysia, Kelantan, Taman Negara, Sungai Relau Station. Gua Gajah, <named-content content-type="dwc:verbatimCoordinates">4.642, 102.063</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420274">KC420274</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">73</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5557</td><td rowspan="1" colspan="1">Malaysia, Pahang, Limestone hill on the left hand site of the road no. 8 toward Kuala Lipis. Near Kampung Chegar Perah I and II FELDA, <named-content content-type="dwc:verbatimCoordinates">4.487, 101.976</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420306">KC420306</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">74</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5557</td><td rowspan="1" colspan="1">Malaysia, Pahang, Limestone hill on the left hand site of the road no. 8 toward Kuala Lipis. Near Kampung Chegar Perah I and II FELDA, <named-content content-type="dwc:verbatimCoordinates">4.487, 101.976</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420293">KC420293</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">75</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5557</td><td rowspan="1" colspan="1">Malaysia, Pahang, Limestone hill on the left hand site of the road no. 8 toward Kuala Lipis. Near Kampung Chegar Perah I and II FELDA, <named-content content-type="dwc:verbatimCoordinates">4.487, 101.976</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250865">KC250865</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250906">KC250906</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250932">KC250932</ext-link></td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC250957">KC250957</ext-link></td></tr><tr><td rowspan="1" colspan="1">76</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5536</td><td rowspan="1" colspan="1">Malaysia, Terengganu, Tasik Kenyir. Gua Taat, <named-content content-type="dwc:verbatimCoordinates">4.842, 102.721</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420284">KC420284</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">77</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">BOR 5536</td><td rowspan="1" colspan="1">Malaysia, Terengganu, Tasik Kenyir. Gua Taat, <named-content content-type="dwc:verbatimCoordinates">4.842, 102.721</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420282">KC420282</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">78</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma umbilicatum</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5503</td><td rowspan="1" colspan="1">Malaysia, Pahang, Gua Tongkat, <named-content content-type="dwc:verbatimCoordinates">3.891, 102.473</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420281">KC420281</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr><tr><td rowspan="1" colspan="1">79</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma umbilicatum</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">BOR 5503</td><td rowspan="1" colspan="1">Malaysia, Pahang, Gua Tongkat, <named-content content-type="dwc:verbatimCoordinates">3.891, 102.473</named-content>.</td><td rowspan="1" colspan="1"><ext-link ext-link-type="gen" xlink:href="KC420290">KC420290</ext-link></td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1">n.a.</td></tr></tbody></table></table-wrap><pmc-comment>PageBreak</pmc-comment>
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<p><bold>DNA extraction, PCR and sequencing.</bold> DNA extraction was done for each specimen (entire animal and its shell), with the E.Z.N.A. Mollusc DNA kit (OMEGA bio-tek). We followed the manufacturer’s extraction protocol. After extraction, PCR was carried out to amplify four regions, namely, 16S (mitochondrial, <xref rid="B83" ref-type="bibr">Palumbi 1996</xref>), COI (mitochondrial, <xref rid="B40" ref-type="bibr">Folmer et al. 1994</xref>), 28S (nuclear, <xref rid="B84" ref-type="bibr">Park and Foighil 2000</xref>), and 18S (nuclear, <xref rid="B113" ref-type="bibr">Stothard et al. 2000</xref>). We used the PCR reactions and programs of Webster et al. (2000). Positive PCR products were sequenced by Macrogen sequencing service (Macrogen Inc., Europe).</p><p><bold>Phylogenetic inferences.</bold> Alignment of sequences was done with Bioedit v7.1.3 (<xref rid="B46" ref-type="bibr">Hall 1999</xref>) and adjusted manually. The final aligned data matrix consists of 2241 positions, of which 2092 can be aligned unambiguously (<xref ref-type="app" rid="APP1">Appendix 17</xref>). The remaining 149 characters (91 from 16S and 58 from 28S) were excluded from further analysis. Mr.Modeltest v2.3 (<xref rid="B77" ref-type="bibr">Nylander 2004</xref>) was used to select the most appropriate model, based on the Akaike Information Criterion (AIC) for 16S, 18S and 28S, and as well as each of the three codon positions for COI. The best fits were: the GTR+I+Γ model for 16S, 28S, COI(2nd codon) and COI (3rd codon); the GTR+Γ for COI(1st codon); and the SYM+I+Γ for 18S.</p><p>Three phylogenetic analyses were done, namely, Bayesian inference (BI), Maximum Likelihood analysis (ML), and Parsimony analyses (PA). Bayesian inference was run in MrBayes v3.2.1 (<xref rid="B53" ref-type="bibr">Huelsenbeck and Ronquist 2001</xref>) with the following setting: mcmc ngen=5000000; nchains=4; samplefreq=100; average deviation of split frequencies < 0.01; and a burn-in value of 25%. A Maximum Likelihood analysis was run using RaxML v7.2.6 (<xref rid="B112" ref-type="bibr">Stamatakis 2006</xref>) as implemented on CIPRES portal v2.2 (<xref rid="B72" ref-type="bibr">Miller et al. 2010</xref>) on all the genes together, with 1000 rapid bootstraps using GTR + U. The data was divided into six partitions, all analyzed with a GTR substitution model. The Parsimony analyses were run using PAUP v4.0b (<xref rid="B114" ref-type="bibr">Swofford 1998</xref>). Gaps were set as fifth character state. A bootstrapped heuristic search with 1000 bootstrap replicates, and 10 random addition sequence heuristic search replicates, with no rearrangement limit per replicate was carried out, with 50% as the minimum bootstrap support included.</p><p><bold>COI Barcoding.</bold> As mentioned above, the shell morphology alone may not be sufficient to delimit many species that have greater morphological variation within species. Thus, in addition to shell morphology, we sought another way to make the decision about the species delimitation. DNA barcoding is used to provide more insight into the species delimitation for gastropods (<xref rid="B19" ref-type="bibr">Boeters and Knebelsberger 2012</xref>, <xref rid="B88" ref-type="bibr">Puillandre et al. 2012</xref>). Therefore, we sequenced COI (mitochondrial, <xref rid="B40" ref-type="bibr">Folmer et al. 1994</xref>) for standard DNA barcoding analysis for a greater number of individuals than we used for the phylogenetic analysis. In total, we obtained COI sequences from 51 individuals, which comprise 19 species (including 8 new species) (<xref ref-type="table" rid="T2">Table 2</xref>). Then, the pairwise genetic distances for all the 51 sequences were computed by using Kimura 2-parameters in MEGA5 (<xref rid="B118" ref-type="bibr">Tamura et al. 2011</xref>).</p><p><bold>Genetic data repository.</bold> The collection information for the specimens used were uploaded, stored and managed in Barcoding of Life Database (BOLD, <ext-link ext-link-type="uri" xlink:href="http://www.boldsystems.org">http://www.boldsystems.org</ext-link>, <xref rid="B90" ref-type="bibr">Ratnasingham and Hebert 2007</xref>). After that, all DNA sequences were <pmc-comment>PageBreak</pmc-comment>uploaded to NCBI GenBank (<ext-link ext-link-type="uri" xlink:href="http://www.ncbi.nlm.nih.gov/genbank">http://www.ncbi.nlm.nih.gov/genbank</ext-link>, <xref rid="B15" ref-type="bibr">Benson et al. 1997</xref>) via BOLD. Genetic data deposited in GenBank can be retrieved easily with the Python tool Biopython (e.g. Module “Entrez”) (<ext-link ext-link-type="uri" xlink:href="http://biopython.org">http://biopython.org</ext-link>, <xref rid="B29" ref-type="bibr">Cock et al. 2009</xref>).</p></sec></sec><sec><title>Data resources</title><p>The data underpinning the analysis reported in this paper are deposited in the Figshare Repository at <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.6084/m9.figshare.830412">http://dx.doi.org/10.6084/m9.figshare.830412</ext-link>.</p></sec><sec><title>Results and discussion</title><sec sec-type="Taxonomic data depository and cybertaxonomy"><title>Taxonomic data depository and cybertaxonomy</title><p>While uploading the data of 31 species, 62 references, 214 collections, 290 pictures, and 31 species pages to Lifedesks, they were simultaneously tagged and linked to each other (<xref ref-type="fig" rid="F14">Figure 14</xref>). The textual information can be downloaded and accessed offline in the xhtml format (source code) (<xref ref-type="fig" rid="F15">Figures 15</xref>). Similarly, the taxonomic information that was used in creating the KML file can be retrieved in KML format (source code) (<xref ref-type="fig" rid="F15">Figures 15</xref>, <xref ref-type="app" rid="APP1">Appendix 5</xref>). In addition to the taxonomic data, a total of 155 3D models of 29 <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species were constructed, which belong to 86 samples. For the ease of specimen and species comparison, all 3D models were saved in ten separate. blend files, each containing 20 layers (<xref ref-type="app" rid="APP1">Appendix 6–15</xref>). The 3D models (i.e. specimens) that belong to the same species were saved in the same layer.</p><fig id="F14" orientation="portrait" position="float"><label>Figures 14.</label><caption><p>User interface of the Lifedesks online database and Google Earth. Lifedesks was used for data storage, management, and tagging; Google Earth was used for data sorting and exploration.</p></caption><graphic xlink:href="zookeys-393-001-g014"></graphic></fig><fig id="F15" orientation="portrait" position="float"><label>Figures 15.</label><caption><p>The raw tagged and linked data (in XML format) that underlay the Lifedesks online database and Google Earth can be accessed and used without the need of a specific platform.</p></caption><graphic xlink:href="zookeys-393-001-g015"></graphic></fig><p>Most taxonomists will have established their own workflow when working with taxonomic data. However, often only fractions of such data are published in a taxonomic revision paper. Thus, extra work would be needed to upload the data to online data platforms whenever a taxonomic revision is done in a traditional manner (i.e. not using cybertaxonomic tools). Here, we demonstrated that, in fact, no extra works is required if a cybertaxonomic workflow is adopted. Taxonomists themselves, who maintain, tag, link and create the data will benefit most by using these cybertaxonomy tools. For example, with some existing tools, such as Google Earth, the information can be integrated, explored and analysed in an interactive way, which will increase the efficiency of the taxonomic methodology (sensu Figure 1 in <xref rid="B102" ref-type="bibr">Sluys 2013</xref>; <xref ref-type="app" rid="APP1">Appendix 5</xref>).</p><p>Furthermore, all textual information is preserved, and can be simply accessed in raw format. In other words, this information is not locked and can be retrieved and integrated with very basic programming skills (such as Python) even when the desirable platform (internet) and software are not available. More importantly, adopting this workflow in taxonomy practice will help realise the vision of <xref rid="B128" ref-type="bibr">Wheeler et al. (2012)</xref>, that “The resultant cyber-enabled taxonomy, or cybertaxonomy, would open access to biodiversity data to developing nations, assure access to reliable data about species, and change how scientists and citizens alike access, use and think about biological diversity information”.</p><pmc-comment>PageBreak</pmc-comment>
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</sec><sec sec-type="Morphological analysis"><title>Morphological analysis</title><p><xref ref-type="table" rid="T3">Table 3</xref> shows the morphological data matrix of 31 <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species and 11 general qualitative shell characters. Twenty-seven out of 31 <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species have a unique set of shell character states. The remaining four species share two unique sets of general characters states, namely, 1) <italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> (van Benthem Jutting, 1952), and <italic><named-content content-type="taxon-name">Plectostoma laemodes</named-content></italic> (van Benthem Jutting, 1961); and 2) <italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic> sp. n., and <italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic> sp. n. However, each of these four species is distinguishable by other shell characters (see diagnosis and species description in the taxonomy section).</p><table-wrap id="T3" orientation="portrait" position="float"><label>Table 3.</label><caption><p>Characters matrix for 31 <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species.</p></caption><table frame="hsides" rules="groups"><tbody><tr><th rowspan="1" colspan="1">Species that share the same general character state</th><th rowspan="1" colspan="1">Species</th><th rowspan="1" colspan="1">Apex form</th><th rowspan="1" colspan="1">Apical spire form</th><th rowspan="1" colspan="1">Basal spire form</th><th rowspan="1" colspan="1">Parietal constriction teeth</th><th rowspan="1" colspan="1">Basal constriction teeth</th><th rowspan="1" colspan="1">Tuba form</th><th rowspan="1" colspan="1">Aperture view</th><th rowspan="1" colspan="1">Peristome types</th><th rowspan="1" colspan="1">Spiral lines</th><th rowspan="1" colspan="1">Rib shape</th><th rowspan="1" colspan="1">Rib thickness</th></tr><tr><td rowspan="1" colspan="1">‡</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1">‡</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma laemodes</named-content></italic> (van Benthem Jutting, 1961)</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1">|</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1">|</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1">§</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n. Form BOR 5504</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1">§</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n. Form BOR 5507</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma annandalei</named-content></italic> (Sykes, 1903)</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">?</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kayiani</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">2 or 3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma charasense</named-content></italic> (Tomlin, 1948)</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">2 or 3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma davisoni</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">2 or 3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kakiense</named-content></italic> (Tomlin, 1948)</td><td rowspan="1" colspan="1">2 or 3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kitteli</named-content></italic> (Maassen, 2002)</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma klongsangensis</named-content></italic> (Panha, 1997)</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">?</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">2 or 3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic> (Sykes, 1902)</td><td rowspan="1" colspan="1">1 or 2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic> (van Benthem Jutting, 1961)</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma relauensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">2 or 3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic> (Tomlin, 1938)</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma sciaphilum</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic> (Maassen, 2001)</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma tenggekensis</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma tonkinianum</named-content></italic> (Dautzenberg & Fischer, 1905)</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma umbilicatum</named-content></italic> (van Benthem Jutting, 1952)</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic> sp. n.</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">2</td></tr></tbody></table><table-wrap-foot><fn><p><bold>Characters Character states:</bold> Apex form (<xref ref-type="fig" rid="F4">Figure 4</xref>) 1 = distinctly convex; 2 = moderately convex; 3 = slightly convex.</p><p>Apical spire form (<xref ref-type="fig" rid="F5">Figure 5</xref>) 1 = oblong; 2 = depressed.</p><p>Basal spire form (<xref ref-type="fig" rid="F6">Figure 6</xref>) 1 = conical; 2 = ovoid; 3 = ellipsoid.</p><p>Parietal constriction teeth (<xref ref-type="fig" rid="F7">Figure 7</xref>) 1 = two long lamella; 2 = two knob-shaped teeth; 3 = single knob-shaped tooth; 4 = no tooth.</p><p>Basal constriction teeth (<xref ref-type="fig" rid="F8">Figure 8</xref>) 1 = two knob-shaped teeth; 2 = no tooth.</p><p>Tuba form (<xref ref-type="fig" rid="F9">Figure 9</xref>) 1 = tuba type 1; 2 = tuba type 2; 3 = tuba type 3.</p><p>Aperture view 1 = frontal view; 2 = right lateral view; 3 = back view; and 4 = left lateral view.</p><p>Peristome types (<xref ref-type="fig" rid="F11">Figure 11</xref>) 1 = simple peristome; 2 = double peristome.</p><p>Spiral lines (<xref ref-type="fig" rid="F12">Figure 12</xref>) 1 = thick and thin; 2 = thin only.</p><p>Rib shape (<xref ref-type="fig" rid="F13">Figure 13C</xref>) 1 = single-humped; 2 = slightly curved; 3 = straight.</p><p>Rib thickness (<xref ref-type="fig" rid="F13">Figure 13B</xref>) 1 = thick; 2 = thin.</p></fn></table-wrap-foot></table-wrap><p>Our shell shape characterisation approach that views the shell as a petrified ontogeny provides a set of distinguishable general characters for species delimitation (<xref ref-type="table" rid="T3">Table 3</xref>). Most of the species in this study can be identified just by using these general shape characters. Furthermore, those species that are not distinguishable with these general shape characters, are distinguishable by using more specific shell shape and size characters (see Taxonomy Key).</p><p>However, it is important to note that the intra- and inter-specific variation in shell shape is more difficult to characterise than the variation in shell size. This is reflected in our species description, where the variation in shell size and countable shell characters, such as ribs, but not the variation in general shell shape are explicitly given. Nevertheless, the general qualitative shell shape characters do implicitly reflect the variation because many of these, such as spire shape, tuba, and spiral lines, are obtained by categorising the quantitative variation of these characters (e.g. <xref ref-type="fig" rid="F4">Figures 4</xref> and <xref ref-type="fig" rid="F5">5</xref>).</p></sec><sec sec-type="Conservation status assessment"><title>Conservation status assessment</title><p>Overall, we suggest that 10 of the non-Bornean <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species are threatened and <italic><named-content content-type="taxon-name">Plectostoma sciaphilum</named-content></italic> is extinct. Specifically, <italic><named-content content-type="taxon-name">Plectostoma umbilicatum</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma charasense</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma tenggekensis</named-content></italic> are in the Critically Endangered category; <italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic> is in the Endangered category; and <italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic> are in the Vulnerable category. All of these species, except <italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic>, occur in limestone hills in the State of Pahang, Malaysia, where many of these hills are being quarried or are at risk of being quarried. Our assessments would eventually be submitted to IUCN.</p></sec><sec sec-type="Molecular phylogeny and COI barcoding"><title>Molecular phylogeny and COI barcoding</title><p>As revealed by the Bayesian posterior probability (PP) and maximum likelihood analysis bootstrap (BS) values of the phylogenetic tree in <xref ref-type="fig" rid="F16">Figure 16</xref>, <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> is monophyletic (BI/ML/PA; 1.0/99/100) and is the sister taxon of the less well-supported clade of <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> + <italic><named-content content-type="taxon-name">Arinia</named-content></italic> (0.5/<70/<70). Within the <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> + <italic><named-content content-type="taxon-name">Arinia</named-content></italic> clade, all <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> except <italic><named-content content-type="taxon-name">Opisthostoma vermiculum</named-content></italic> form a well-supported clade (0.98/88/71). The divergence between these clades is similar to the divergence between other genera in <named-content content-type="taxon-name">Diplommatinidae</named-content>. Our phylogenetic analysis suggested that <italic><named-content content-type="taxon-name">Opisthostoma vermiculum</named-content></italic> Clements & Vermeulen, 2008 in Clements et al. (2008) has been incorrectly assigned to the genus <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>.</p><fig id="F16" orientation="portrait" position="float"><label>Figures 16.</label><caption><p>The phylogeny of <named-content content-type="taxon-name">Diplommatinidae</named-content> genera with <italic><named-content content-type="taxon-name">Cochlostoma septemspirale</named-content></italic> (Wagner, 1897) as outgroup. Bayesian inference 50% majority-rule consensus trees based on the concatenated dataset consisting of parts of 28S, 18S, COI, and 16S. Bayesian posterior probabilities, percent bootstrap support after 1000 maximum likelihood replicates, and percent bootstrap support after 1000 maximum parsimony replicates are shown for the major clades. Clade A consists of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species, Clade B consists of <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> species. The shell forms are shown for representative taxa. Details of the taxa can be found in <xref ref-type="table" rid="T1">Table 1 </xref>(no. 1–28, 42, 48, 61, 69, 71, 73, and 78). Scale bar = 1 mm.</p></caption><graphic xlink:href="zookeys-393-001-g016"></graphic></fig><p><xref ref-type="table" rid="T4">Table 4</xref> shows the Kimura 2-parameter distances for all sequence pairs within each group and the net average between groups of sequences (for sequence alignment see <xref ref-type="app" rid="APP1">Appendix 16</xref>). This reveals that all species pairs exceed a divergence of 10% (n = 169), with the exception of <italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic> vs. <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic> vs. <italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic>. Genetic divergence within each species is below 9% (n = 14, mean = 2.6%, SD = 0.1%), with the exception of <italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic>.</p><table-wrap id="T4" orientation="portrait" position="float"><label>Table 4.</label><caption><p>COI sequence divergence <xref ref-type="table-fn" rid="TN4.1">†</xref> within and between 19 <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species.</p></caption><table frame="hsides" rules="groups"><tbody><tr><th rowspan="2" colspan="1">Number of specimens</th><th rowspan="2" colspan="1">Divergence within groups <xref ref-type="table-fn" rid="TN4.2">‡</xref></th><th rowspan="1" colspan="19">Divergence between groups of sequences <xref ref-type="table-fn" rid="TN4.3">§</xref></th></tr><tr><th rowspan="1" colspan="1"></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma davisoni</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kakiense</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma relauensis</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic></th><th rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma umbilicatum</named-content></italic></th></tr><tr><td rowspan="1" colspan="1">8</td><td rowspan="1" colspan="1">0.06</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0.13</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic></td><td rowspan="1" colspan="1"><bold>0.09</bold>*</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0.00</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma davisoni</named-content></italic></td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic></td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">6</td><td rowspan="1" colspan="1">0.01</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic></td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.11</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kakiense</named-content></italic></td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.11</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.21</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0.00</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic></td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">0.08</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic></td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1"><bold>0.09</bold>*</td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0.01</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic></td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.10</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1">0.21</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0.00</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic></td><td rowspan="1" colspan="1">0.13</td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma relauensis</named-content></italic></td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1">0.13</td><td rowspan="1" colspan="1">0.21</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic></td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1">0.13</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.11</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">0.04</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic></td><td rowspan="1" colspan="1">0.11</td><td rowspan="1" colspan="1">0.11</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">n.a.</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic></td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.21</td><td rowspan="1" colspan="1">0.23</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.22</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0.00</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic></td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.21</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.21</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">5</td><td rowspan="1" colspan="1">0.03</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic></td><td rowspan="1" colspan="1">0.10</td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0.00</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic></td><td rowspan="1" colspan="1">0.13</td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.12</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.13</td><td rowspan="1" colspan="1">0.06</td><td rowspan="1" colspan="1">0.10</td><td rowspan="1" colspan="1">0.21</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1"></td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0.00</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma umbilicatum</named-content></italic></td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1">0.13</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.21</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.17</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.22</td><td rowspan="1" colspan="1">0.13</td><td rowspan="1" colspan="1">0.19</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1"></td></tr><tr><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">0.01</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic></td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.16</td><td rowspan="1" colspan="1">0.21</td><td rowspan="1" colspan="1">0.21</td><td rowspan="1" colspan="1">0.23</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.20</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.18</td><td rowspan="1" colspan="1">0.14</td><td rowspan="1" colspan="1">0.11</td><td rowspan="1" colspan="1">0.11</td><td rowspan="1" colspan="1">0.21</td><td rowspan="1" colspan="1">0.15</td><td rowspan="1" colspan="1">0.10</td><td rowspan="1" colspan="1">0.21</td></tr></tbody></table><table-wrap-foot><fn id="TN4.1"><p>† Analyses were conducted using the Kimura 2-parameter model (<xref rid="B59" ref-type="bibr">Kimura 1980</xref>). The analysis involved 51 nucleotide sequences. Codon positions included were 1st+2nd+3rd. All positions with less than 95% site coverage were eliminated. There were a total of 615 positions in the final dataset. These analyses were conducted in MEGA5 (<xref rid="B118" ref-type="bibr">Tamura et al. 2011</xref>).</p></fn><fn id="TN4.2"><p>‡ The number of base substitutions per site from averaging over all sequence pairs within each group.</p></fn><fn id="TN4.3"><p>§ The number of base substitutions per site from estimation of net average between groups of sequences. Genetic distances between two species smaller than 10% are shown in bold.</p></fn></table-wrap-foot></table-wrap><p>One key determinant for the success of DNA barcoding is prior knowledge of intra- and interspecific genetic distances for the barcoding marker in question. The optimum intra- and interspecific threshold in gastropods is higher than the conventional value (<xref rid="B49" ref-type="bibr">Hebert et al. 2003</xref>) of 3% (e.g. 4% in <xref rid="B33" ref-type="bibr">Davison et al. 2009</xref>, 6% in <xref rid="B60" ref-type="bibr">Köhler and Johnson 2012</xref>, 9.8%–25% in <xref rid="B85" ref-type="bibr">Parmakelis et al. 2013</xref>). In our study, we also found higher values of intraspecific variation and interspecific divergence’ of COI for three well-defined species, namely, <italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> are larger and smaller than 10%, respectively.</p><p>A study on a pulmonate limestone-dwelling micro-landsnail in the same region also suggests intraspecific COI divergence not exceeding 10% based on the Kimura 2-parameter model (<xref rid="B52" ref-type="bibr">Hoekstra and Schilthuizen 2011</xref>), and similar values were obtained for <italic><named-content content-type="taxon-name">Everettia</named-content></italic>, a Malaysian pulmonate not restricted to limestone (<xref rid="B62" ref-type="bibr">Liew et al. 2009</xref>). Hence, based on our results, together with the only other two studies on the COI variation of land snails in Sundaland, we advise caution in using a conventional threshold value in COI genetic variability for species delimitation, when the background genetic variability of COI is unknown for a particular taxon in a particular geographic region.</p></sec><sec sec-type="Key to Peninsular Malaysia Plectostoma"><title>Key to Peninsular Malaysia <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic></title><table-wrap content-type="key" orientation="portrait" id="d36e5994" position="anchor"><table frame="hsides" rules="groups"><tbody><tr><td rowspan="1" colspan="1">1</td><td rowspan="1" colspan="1">Oblong apical spire</td><td rowspan="1" colspan="1">2</td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Depressed apical spire</td><td rowspan="1" colspan="1">10</td></tr><tr><td rowspan="1" colspan="1">2</td><td rowspan="1" colspan="1">Tuba with type 1 coiling</td><td rowspan="1" colspan="1">3</td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Tuba with type 2 or 3 coiling</td><td rowspan="1" colspan="1">4</td></tr><tr><td rowspan="1" colspan="1">3</td><td rowspan="1" colspan="1">Constriction with 2 parietal teeth. Aperture visible when shell observed in left lateral view</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Constriction without parietal tooth. Aperture visible when shell observed in back view</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma umbilicatum</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">4</td><td rowspan="1" colspan="1">Tuba with type 2 coiling. Aperture visible when shell observed in right lateral view</td><td rowspan="1" colspan="1">5</td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Tuba with type 3 coiling. Aperture visible when shell observed in left lateral view</td><td rowspan="1" colspan="1">8</td></tr><tr><td rowspan="1" colspan="1"><pmc-comment>PageBreak</pmc-comment>5</td><td rowspan="1" colspan="1">Spire with both thick and thin spiral lines</td><td rowspan="1" colspan="1">6</td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Spire with only thin spiral lines</td><td rowspan="1" colspan="1">7</td></tr><tr><td rowspan="1" colspan="1">6</td><td rowspan="1" colspan="1">Left lateral side of outer peristome projected not more than two times the width of the right lateral side</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Left lateral side of outer peristome projected more than three times the width of the right lateral side</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma klongsangensis</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">7</td><td rowspan="1" colspan="1">Constriction with 2 basal teeth</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma charasense</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Constriction without basal tooth</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kitteli</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">8</td><td rowspan="1" colspan="1">Constriction without basal tooth</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma sciaphilum</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Constriction with 2 basal teeth</td><td rowspan="1" colspan="1">9</td></tr><tr><td rowspan="1" colspan="1">9</td><td rowspan="1" colspan="1">Constriction with 2 long lamella-shaped parietal teeth</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Constriction with knob-shaped parietal teeth</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">10</td><td rowspan="1" colspan="1">Tuba with type 1 coiling</td><td rowspan="1" colspan="1">11</td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Tuba with type 2 or 3 coiling</td><td rowspan="1" colspan="1">15</td></tr><tr><td rowspan="1" colspan="1">11</td><td rowspan="1" colspan="1">Constriction with 1 or 2 parietal teeth</td><td rowspan="1" colspan="1">12</td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Constriction without parietal teeth</td><td rowspan="1" colspan="1">13</td></tr><tr><td rowspan="1" colspan="1">12</td><td rowspan="1" colspan="1">Constriction with 1 parietal tooth</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Constriction with 2 parietal teeth</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">13</td><td rowspan="1" colspan="1">Ellipsoid basal spire</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Ovoid basal spire</td><td rowspan="1" colspan="1">14</td></tr><tr><td rowspan="1" colspan="1">14</td><td rowspan="1" colspan="1">More than 1/4 of the tuba visible in top view</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Less than 1/4 of the tuba visible in top view</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">15</td><td rowspan="1" colspan="1">Tuba with type 3 coiling</td><td rowspan="1" colspan="1">16</td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Tuba with type 2 coiling</td><td rowspan="1" colspan="1">18</td></tr><tr><td rowspan="1" colspan="1">16</td><td rowspan="1" colspan="1">Simple peristome</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma tonkinianum</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Double peristome</td><td rowspan="1" colspan="1">17</td></tr><tr><td rowspan="1" colspan="1">17</td><td rowspan="1" colspan="1">Aperture visible when shell observed in frontal view</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Aperture visible when shell observed in left lateral view</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">18</td><td rowspan="1" colspan="1">Aperture visible when shell observed in frontal or back views</td><td rowspan="1" colspan="1">19</td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Aperture visible when shell observed in right lateral view</td><td rowspan="1" colspan="1">24</td></tr><tr><td rowspan="1" colspan="1">19</td><td rowspan="1" colspan="1">Aperture visible when shell observed in back views</td><td rowspan="1" colspan="1">20</td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Aperture visible when shell observed in frontal views</td><td rowspan="1" colspan="1">21</td></tr><tr><td rowspan="1" colspan="1">20</td><td rowspan="1" colspan="1">Spire 1.6–1.9 mm high</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Spire 2.3–2.6 mm high</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">21</td><td rowspan="1" colspan="1">Ovoid basal spire</td><td rowspan="1" colspan="1">22</td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Conical basal spire</td><td rowspan="1" colspan="1">23</td></tr><tr><td rowspan="1" colspan="1">22</td><td rowspan="1" colspan="1">Constriction with 1 parietal tooth</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma relauensis</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Constriction with 2 parietal teeth</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma davisoni</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">23</td><td rowspan="1" colspan="1">Spire with both thick and thin spiral lines</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Spire with only thin spiral lines</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kayiani</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">24</td><td rowspan="1" colspan="1">Spire with straight radial ribs</td><td rowspan="1" colspan="1">25</td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Spire with slightly curved or single-humped radial ribs</td><td rowspan="1" colspan="1">28</td></tr><tr><td rowspan="1" colspan="1">25</td><td rowspan="1" colspan="1">Conical basal spire</td><td rowspan="1" colspan="1">26</td></tr><tr><td rowspan="1" colspan="1"><pmc-comment>PageBreak</pmc-comment>–</td><td rowspan="1" colspan="1">Ovoid basal spire</td><td rowspan="1" colspan="1">27</td></tr><tr><td rowspan="1" colspan="1">26</td><td rowspan="1" colspan="1">Whorl periphery distinctly convex</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Whorl periphery slightly convex</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma annandalei</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">27</td><td rowspan="1" colspan="1">Constriction without parietal teeth</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Constriction with 2 parietal teeth</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma kakiense</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">28</td><td rowspan="1" colspan="1">Conical basal spire</td><td rowspan="1" colspan="1">29</td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Ovoid basal spire</td><td rowspan="1" colspan="1">30</td></tr><tr><td rowspan="1" colspan="1">29</td><td rowspan="1" colspan="1">Spire 1.6–1.7 mm high</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma tenggekensis</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Spire 2.0–2.2 mm high</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">30</td><td rowspan="1" colspan="1">Whole tuba attaches to spire</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma laemodes</named-content></italic></td></tr><tr><td rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1">Less than 2/3 of the tuba attaches to spire</td><td rowspan="1" colspan="1"><italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic></td></tr></tbody></table></table-wrap></sec></sec><sec><title>Taxonomy</title><sec sec-type="taxon-treatment"><label>Genus</label><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma"><named-content content-type="genus">Plectostoma</named-content></named-content></title><p>http://species-id.net/wiki/Plectostoma</p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Plectostoma</named-content><named-content content-type="comment"> Adam 1865: 177. Type species: <italic><named-content content-type="taxon-name">Plectostoma DeCrespignii</named-content></italic> (by original designation)</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Geothauma</named-content><named-content content-type="comment"><xref rid="B32" ref-type="bibr">Crosse 1892</xref>: 282. Type species (by original designation): <italic><named-content content-type="taxon-name">Plectostoma grandispinosum</named-content></italic> (Godwin-Austen, 1889).</named-content></p></list-item></list><sec sec-type="treatment-Generic classification dispute"><title>Generic classification dispute.</title><p>The genus <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> was described by <xref rid="B16" ref-type="bibr">Blanford and Blanford (1860)</xref> based on one species–<italic><named-content content-type="taxon-name">Opisthostoma nilgiricum</named-content></italic> from India. <xref rid="B2" ref-type="bibr">Adam (1865b)</xref> described a second species of <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>, namely, <italic><named-content content-type="taxon-name">Opisthostoma decrespignyi</named-content></italic>, which he previously described under the new genus <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> (<xref rid="B1" ref-type="bibr">Adam 1865a</xref>). Nevertheless, <xref rid="B18" ref-type="bibr">Blanford (1867)</xref> concluded that the conchological differences between these two taxa were not enough to create a different genus. Instead, he suggested these could be two different subgenera. Next, another two subgenera–<italic><named-content content-type="taxon-name">Gyrostropha</named-content></italic> Ancey, 1887 and <italic><named-content content-type="taxon-name">Geothauma</named-content></italic> Crosse, 1892, were proposed for different forms of <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>. However, <xref rid="B104" ref-type="bibr">Smith (1893a)</xref> suggested that this subgeneric classification was not necessary until more data other than shell morphology were available. Since then, a classification into three subgenera within the genus <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>, namely, <italic><named-content content-type="taxon-name">Geothauma</named-content></italic>, <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> has generally been accepted (e.g. <xref rid="B67" ref-type="bibr">von Martens and Thiele 1908</xref>, <xref rid="B5" ref-type="bibr">van Benthem Jutting 1932</xref>, <xref rid="B6" ref-type="bibr">van Benthem Jutting 1952</xref>), until, in a recent review of the genus <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>, <xref rid="B121" ref-type="bibr">Vermeulen (1991</xref>, <xref rid="B122" ref-type="bibr">1994</xref>) followed a classification into only two subgenera, namely, <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>.</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Despite the distinct ecological niche differences (see below–Distribution and habitat) between <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>, it is not feasible to use this criterion in the genus identification, because information about the ecology is usually not available as most collections are made by soil sampling. After 150 years of work on <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>, it is still difficult to identify reliable apomorphic character states that can be used to distinguish between <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> (<xref rid="B121" ref-type="bibr">Vermeulen 1991</xref>, <xref rid="B122" ref-type="bibr">1994</xref>). Both share the character state of the constriction, which is a slight shrinkage in the whorl towards the end of the spire. When the animal retracts into its shell, its operculum rests at the constriction (<xref rid="B121" ref-type="bibr">Vermeulen 1991</xref>). It is, however, possible to make a morphological distinction between <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> on the basic of the shell colouration in a fully grown adult, which is orange or pinkish in <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> and white or pale yellowish in <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>. The colour differences between these two genera are very clear when comparing the living snails or freshly dead shell material (<xref ref-type="fig" rid="F17">Figure 17</xref>, and <xref ref-type="app" rid="APP1">Appendix 18</xref>). Some <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species have a regularly coiled tuba, and a shell form that is similar to <italic><named-content content-type="taxon-name">Arinia</named-content></italic>. However, <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> and <italic><named-content content-type="taxon-name">Arinia</named-content></italic> can be easily distinguished by shell colour differences. The shell colour in <italic><named-content content-type="taxon-name">Arinia</named-content></italic> is similar to that in <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>.</p><fig id="F17" orientation="portrait" position="float"><label>Figures 17.</label><caption><p>Photographs of 17 living <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species. <bold>A</bold><italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> (van Benthem Jutting, 1952), BOR 5569 <bold>B</bold><italic><named-content content-type="taxon-name">Plectostoma umbilicatum</named-content></italic> (van Benthem Jutting, 1952), BOR 5503 <bold>C</bold><italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic> (van Benthem Jutting, 1952), BOR 5520 <bold>D</bold><italic><named-content content-type="taxon-name">Plectostoma kakiense</named-content></italic> (Tomlin, 1948), BOR 5516 <bold>E</bold><italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic> sp. n., BOR 5574 <bold>F</bold><italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic> sp. n., BOR 5518 <bold>G</bold><italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic> sp. n., BOR 5536 <bold>H</bold><italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic> (van Benthem Jutting, 1952), BOR 5631 <bold>I</bold><italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic> (van Benthem Jutting, 1952), BOR 5515 <bold>J</bold><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n. Form BOR 5507, BOR 5507 <bold>K</bold><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n. Form BOR 5504, BOR 5504 <bold>L</bold><italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic> (Maassen, 2001), BOR 5537 <bold>M</bold><italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic> (van Benthem Jutting, 1952), BOR 5512 <bold>N</bold><italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> (van Benthem Jutting, 1952), BOR 5557 <bold>O</bold><italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic> (Sykes, 1902) Form BOR 5510, BOR 5510 <bold>P</bold><italic><named-content content-type="taxon-name">Plectostoma relauensis</named-content></italic> sp. n., BOR 5511 <bold>Q</bold><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> (Maassen, 2001), BOR 5505 <bold>R</bold><italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic> (Tomlin, 1938), BOR 5559 <bold>S</bold><italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> sp. n., BOR 5533.</p></caption><graphic xlink:href="zookeys-393-001-g017"></graphic></fig></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Protoconch is either slightly, moderately or distinctly convex (<xref ref-type="fig" rid="F3">Figure 3</xref>).</p><p><bold>Spire.</bold> Height: 1.0 mm–3.7 mm. Width: 0.85 mm–2.60 mm (<xref ref-type="fig" rid="F11">Figure 11</xref>). Number of whorls between 2 3/4–71/4. Apical spire shape: oblong or depressed conical (<xref ref-type="fig" rid="F4">Figure 4</xref>). Basal spire shape: conical, ovoid or ellipsoid (<xref ref-type="fig" rid="F5">Figure 5</xref>). Whorl periphery: flat, slightly, moderately or distinctly convex. Umbilicus: open, partially closed, or totally closed.</p><p><bold>Constriction.</bold> Parietal teeth: parietal side of inner constriction whorl (<xref ref-type="fig" rid="F2">Figure 2</xref>) with two long lamellae (<xref ref-type="fig" rid="F6">Figure 6A</xref>), two ridges with a knob at each end (<xref ref-type="fig" rid="F6">Figure 6B</xref>), one ridge with a knob at one end (<xref ref-type="fig" rid="F6">Figure 6C</xref>), or no teeth (<xref ref-type="fig" rid="F6">Figure 6D</xref>). Basal teeth: basal side of inner constriction whorl (<xref ref-type="fig" rid="F2">Figure 2</xref>) with no teeth (<xref ref-type="fig" rid="F7">Figure 7B</xref>), one ridge running parallel with the whorl growing direction, one ridge with a knob at one end running perpendicular to the whorl growing direction, or a combination of the latter two types (<xref ref-type="fig" rid="F7">Figure 7A</xref>).</p><p><bold>Tuba.</bold> Coiling direction: regular coiling (type 1, <xref ref-type="fig" rid="F8">Figure 8A</xref>) or distorted (Type 2, or 3) (<xref ref-type="fig" rid="F8">Figure 8B and C</xref>). Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 3/8–1 1/2. Proportion of tuba that attaches to spire: whole to none.</p><p><bold>Aperture and peristome.</bold> Peristome: simple aperture without outer peristome (<xref ref-type="fig" rid="F10">Figure 10B</xref>), or double peristome (<xref ref-type="fig" rid="F10">Figure 10C</xref> and <xref ref-type="fig" rid="F10">10D</xref>). Shape of outer peristome (<xref ref-type="fig" rid="F10">Figure 10A</xref>): same as inner peristome and uniformly round, or highly projected or slightly projected at either a particular side or at a several sides of anterior, poteriorior, left and right laterial (<xref ref-type="fig" rid="F9">Figure 9</xref> and <xref ref-type="fig" rid="F10">10A</xref>).</p><p><bold>Spiral lines.</bold> Either thick or thin, or only thin lines present (<xref ref-type="fig" rid="F12">Figure 12</xref>).</p><p><bold>Radial ribs.</bold> Rib density: 4–32 per mm on the spire’s last whorl in right lateral view (<xref ref-type="fig" rid="F13">Figure 13A</xref>). Intensity: thick or thin (<xref ref-type="fig" rid="F13">Figure 13B</xref>). Shape: straight, slightly curved, distinctly curved, single humped, single looped or double looped and the shape remaining the same or changing between between the spire and the tuba (<xref ref-type="fig" rid="F13">Figure 13C</xref>, but single-looped, and double-humped not shown). Inclination: from orthoclin to prosoclin.</p></sec><sec sec-type="treatment-Distribution and habitat"><title>Distribution and habitat.</title><p>The distribution range of <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> is about 4.6 million square kilometres within the extent limited by <named-content content-type="dwc:verbatimCoordinates">11°N, 97°E</named-content> and <named-content content-type="dwc:verbatimCoordinates">5°S, 120°E</named-content>. However probably less than 5% of this large area is covered by limestone outcrops where suitable habitat may exist for obligate karst taxa like <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>. The genus counts 79 species and occurs in Vietnam (1 species), Thailand (1), Peninsular Malaysia (28), Sumatra (1), and Borneo (48). Peninsular Malaysia, Sumatra and Borneo are part of the biogeographical region called Sundaland (<xref rid="B57" ref-type="bibr">Johnson 1964</xref>). <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> is found on most limestone hills. However, the genus is conspicuously absent on the limestone hills to the west of the central mountain ranges, such as the hills in the States of Perak and Kedah in Peninsular Malaysia, and in the northwestern half of Sumatra (<xref ref-type="fig" rid="F18">Figure 18</xref>). No species have been recorded from the east coast of Sumatra, where hardly any limestone outcrops exist.</p><fig id="F18" orientation="portrait" position="float"><label>Figures 18.</label><caption><p>Distributional records of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species. <bold>A</bold>–<bold>D</bold> distribution records for each of 31 <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species.</p></caption><graphic xlink:href="zookeys-393-001-g018"></graphic></fig><p>Based on collection data and our field experience, there is a distinct ecological divergence between <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> and <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>. This was already observed in the 19<sup>th</sup> century (<xref rid="B16" ref-type="bibr">Blanford and Blanford 1860</xref>, <xref rid="B31" ref-type="bibr">de Crespigny 1865</xref>, <xref rid="B17" ref-type="bibr">Blanford 1866</xref>), and also by <xref rid="B10" ref-type="bibr">Berry (1961)</xref>. <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> can only be found in limestone outcrops, where the rock face is its major habitat, although a few individuals can occasionally be found on vegetation debris below the limestone rock face. <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>, on the other hand, is a soil dweller, living in leaf litter on the forest floor. They are mostly but not exclusively found in forest over limestone bedrock (<xref rid="B98" ref-type="bibr">Schilthuizen et al. 2003b</xref>).</p></sec><sec sec-type="treatment-Phylogenetic relationships"><title>Phylogenetic relationships.</title><p>Our molecular phylogenetic analysis reveals that <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>, <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic>, and <italic><named-content content-type="taxon-name">Arinia</named-content></italic> are phylogenetically closely related (<xref ref-type="fig" rid="F16">Figure 16</xref>). It is important to point out that the phylogenetic relationships among <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>, <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> (except <italic><named-content content-type="taxon-name">Opisthostoma vermiculum</named-content></italic>), <italic><named-content content-type="taxon-name">Opisthostoma vermiculum</named-content></italic>, and <italic><named-content content-type="taxon-name">Arinia</named-content></italic> are unresolved. <xref ref-type="fig" rid="F16">Figure 16</xref> shows representative shell morphologies of the taxa that were included in the phylogenetic analysis, and it is clear that it is rather difficult to find shared derived characteristics (synapomorphies) in size, spire shape, or tuba coiling regime, for either <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> or <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>.</p><p>Nonetheless, we treat <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> and <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> as two separate genera based on their ecological divergence and differences in adult shell colouration. Similarly, we propose that <italic><named-content content-type="taxon-name">Opisthostoma vermiculum</named-content></italic> and <italic><named-content content-type="taxon-name">Arinia</named-content></italic> should be considered as two separate genera. However, this hypothesis needs further testing with more genetic data from <italic><named-content content-type="taxon-name">Opisthostoma vermiculum</named-content></italic> Clements & Vermeulen, 2008 (in Clements et al. 2008), the conchologically similar <italic><named-content content-type="taxon-name">Opisthostoma gittenbergeri</named-content></italic> Vermeulen & Clements, 2008 and further <italic><named-content content-type="taxon-name">Arinia</named-content></italic> species.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">dindingensis</named-content></named-content><named-content content-type="taxon-status">sp. n.</named-content></title><p>http://zoobank.org/529D652A-B50B-4B19-BD5D-0D769F9EBC4F</p><p>http://species-id.net/wiki/Plectostoma_dindingensis</p><p><xref ref-type="fig" rid="F19">Figure 19</xref><xref ref-type="app" rid="APP1">, Appendix 7</xref></p><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: BOR 5642(1). Paratypes: BOR 5612(3)</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic> the general shell form, in terms of apex, spire and tuba, but differs by lacking constriction teeth and having a more tightly coiled tuba (less than 1/4 of the tuba visible in top view).</p></sec><sec sec-type="treatment-Etymology"><title>Etymology.</title><p>This species is named after its type locality–Dinding.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: moderately convex.</p><p><bold>Spire.</bold> Height: 1.8–1.9 mm. Width: 1.4–1.5 mm. Number of whorls: 3 7/8–4 5/8. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: distinctly convex. Umbilicus: partially closed by tuba.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Constriction.</bold> Parietal teeth: none. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 1 and aperture visible from back view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 5/8–3/4. Proportion of tuba that attaches to the spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 6–8 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: orthoclin.</p><fig id="F19" orientation="portrait" position="float"><label>Figures 19.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic> sp. n. BOR 5642, Holotype. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g019"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. The exact location is unknown. The specimens are labeled as collected from “dinding”. It could be near Kampung Bukit Dinding, Pahang (<named-content content-type="dwc:verbatimCoordinates">3°49'41"N, 102°22'3"E</named-content>).</p><p>Distribution range. The species has only been recorded from the type locality (<xref ref-type="fig" rid="F18">Figure 18D</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Vulnerable (D2 ver. 10.1). The samples was collected from a living population in 1997. The population status remains unclear. The area around Kampung Bukit Dinding has been converted to plantation and no significant undisturbed forest coverage remains.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>See discussion under <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>.</p><pmc-comment>PageBreak</pmc-comment>
</sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">mengaburensis</named-content></named-content><named-content content-type="taxon-status">sp. n.</named-content></title><p>http://zoobank.org/3C81401C-9C4C-44C2-9A44-9F2F115D8D37</p><p>http://species-id.net/wiki/Plectostoma_mengaburensis</p><p><xref ref-type="fig" rid="F17">Figures 17E</xref><xref ref-type="fig" rid="F20">, 20</xref><xref ref-type="app" rid="APP1">, Appendix 7</xref></p><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: BOR 5643(1). Paratypes: BOR 5574(>25), V8822 (6)</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic> the general shell form, in the terms of apex, spire and tuba, but differs by lacking constriction teeth and having a less tightly coiled tuba (more than 1/4 of the tuba visible in top view).</p></sec><sec sec-type="treatment-Etymology"><title>Etymology.</title><p>This species is named after its type locality–Mengabur.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: moderately convex.</p><p><bold>Spire.</bold> Height: 1.9–2.2 mm. Width: 1.4–1.6 mm. Number of whorls: 3 7/8–4 1/2. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: distinctly convex. Umbilicus: open or half of the umbilicus closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: none. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 1 and aperture visible from back view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 5/8. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 7 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: orthoclin.</p><fig id="F20" orientation="portrait" position="float"><label>Figures 20.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic> sp. n. BOR 5643. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> left lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g020"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. An unnamed small limestone hill in the plantation near the large Bukit Mengabur quarry (<named-content content-type="dwc:verbatimCoordinates">3°43'50"N, 102°49'40"E</named-content>).</p><p>Distribution range. This species only occurs in the Mengabur limestone cluster (<xref ref-type="fig" rid="F18">Figure 18D</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Near Threatened. This species only occurs in the Mengabur limestone cluster, which is quite large (<italic>ca.</italic> 10 km<sup>2</sup>, estimated from Google Earth), and its vegetation cover is largely undisturbed. However, quarrying activities have started at the eastern part of the cluster and the whole limestone cluster is surrounded by plantation.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>See discussion under <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_sinyumensis"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">sinyumensis</named-content></named-content></title><p><named-content content-type="taxon-authority">(Maassen, 2001)</named-content></p><p>http://species-id.net/wiki/Plectostoma_sinyumensis</p><p><xref ref-type="fig" rid="F17">Figures 17L</xref><xref ref-type="fig" rid="F21">, 21</xref><xref ref-type="app" rid="APP1">, Appendix 7</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma sinyumensis</named-content><named-content content-type="comment"> Maassen, 2001: 52, figures 1, 6 & 7 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma sinyumensis</named-content><named-content content-type="comment"> Maassen, <xref rid="B23" ref-type="bibr">Clements (2007</xref>: 74).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma sinyumensis</named-content><named-content content-type="comment"> Maassen, Clements et al. (2008: 2760).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma sinyumensis</named-content><named-content content-type="comment"> Maassen, <xref rid="B127" ref-type="bibr">Webster et al. (2012</xref>: 628).</named-content></p></list-item><pmc-comment>PageBreak</pmc-comment>
</list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: ZMA 138439(1) (Seen). Paratypes: ZMA 138440(>25) (Seen), RMNH 81804(2) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>BOR 462(5), BOR 5537(>10), BOR 5623(>50).</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic> the general shell form, in terms of apex, apical spire and tuba, but differs by having an ellipsoid basal spire.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: moderately convex.</p><p><bold>Spire.</bold> Height: 1.6–1.9 mm. Width: 1.2–1.3 mm. Number of whorls: 3–3 3/4. Apical spire shape: depressed conical. Basal spire shape: ellipsoid. Whorl periphery: distinctly convex. Umbilicus: almost completely closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: none. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 1 and aperture visible between right lateral and back view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 5/8–3/4. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 9–10 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: orthoclin.</p><fig id="F21" orientation="portrait" position="float"><label>Figures 21.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic> (Maassen, 2001) BOR5537. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g021"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Gunung Senyum, Pahang (<named-content content-type="dwc:verbatimCoordinates">3°42'35"N, 102°26'3"E</named-content>).</p><p>Distribution range. In addition to the type locality, this species also was found at the Jebak Puyuh limestone outcrop, which lies about 1 km east of Gunung Senyum (<xref ref-type="fig" rid="F18">Figure 18D</xref>). BOR 462 was collected in Pulau Singa Besar, which is located about 300 km from type locality. The reliability of the collection data is dubious (the same was found for BOR 463 of <italic><named-content content-type="taxon-name">Plectostoma relauensis</named-content></italic>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Near Threatened. There are four limestone hills in this cluster. Gunung Senyum and Jebak Puyuh are the larger among these four hills. The former is gazetted as recreation forest but the latter has been at risk of destruction. Jebak Puyuh had been earmarked for quarrying several years ago, but the plan has been abandoned. <italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic> has been recorded from these two hills, but its status at the two smaller hills remains unknown. In a survey in July 2010, a living population was recorded at Jebak Puyuh, in an enclosed humid sinkhole. Intensive surveying on the limestone rock faces of Gunung Senyum and other parts of Jebak Puyuh has failed to retrieve any additional living individuals, probably because most of the rock faces were very dry.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>See discussion under <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_umbilicatum"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">umbilicatum</named-content></named-content></title><p><named-content content-type="taxon-authority">(van Benthem Jutting, 1952)</named-content></p><p>http://species-id.net/wiki/Plectostoma_umbilicatum</p><p><xref ref-type="fig" rid="F17">Figure 17B</xref><xref ref-type="fig" rid="F22">, 22</xref><xref ref-type="app" rid="APP1">, Appendix 8</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma umbilicatum</named-content><named-content content-type="comment"> van Benthem Jutting, 1952: 49, figure 25 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma umbilicatum</named-content><named-content content-type="comment"> van Benthem Jutting, <xref rid="B8" ref-type="bibr">van Benthem Jutting (1961</xref>: 39).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma umbilicatum</named-content><named-content content-type="comment"> van Benthem Jutting, <xref rid="B23" ref-type="bibr">Clements (2007</xref>: 74).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma umbilicatum</named-content><named-content content-type="comment"> van Benthem Jutting, Clements et al. (2008: 2760).</named-content></p></list-item><pmc-comment>PageBreak</pmc-comment>
</list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: ZMA 136070(1) (Seen). Paratypes: ZMA 136071(8) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>BOR 5503(>10), BOR 5625(>25).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic> the tuba form, but differs by having an oblong conical apical spire.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: moderately convex.</p><p><bold>Spire.</bold> Height: 2.0–2.3 mm. Width: 1.3–1.5 mm. Number of whorls: 4 5/8–4 3/8. Apical spire shape: oblong conical. Basal spire shape: ovoid. Whorl periphery: distinctly convex. Umbilicus: completely open or partially closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: none. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 1 and aperture visible from back view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 1/2–3/4. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 6–9 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: orthoclin.</p><fig id="F22" orientation="portrait" position="float"><label>Figures 22.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma umbilicatum</named-content></italic> (van Benthem Jutting, 1952) BOR5503. <bold>A</bold> frontal view <bold>B</bold> leftlateral view; <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g022"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Limestone hill cluster named Kota Tongkat (<named-content content-type="dwc:verbatimCoordinates">3°53'28"N, 102°28'23"E</named-content>).</p><p>Distribution range. It is only known from the type locality. All other adjacent limestone outcrops have been sampled, but only other <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species were found (<xref ref-type="fig" rid="F18">Figure 18D</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Critically Endangered (B2ab(iii)+C2a(i) ver. 10.1). The Kota Tongkat limestone cluster is surrounded by oil palm plantation and heavily degraded <pmc-comment>PageBreak</pmc-comment>forest. This species is only known from this limestone cluster. Recent soil samplings have not revealed any recent dead shells (Clements et al. 2008). Neverthelss, we found a living population with fewer than 100 individuals at a wet stalagmite of the entrance of one the caves during an intensive survey in May 2011. During that survey, we noticed that all other rock surfaces of the limestone outcrops in Kota Tongkat were very dry. Thus, the recorded and other unknown living populations are at risk of extinction because a long drought might wipe them out.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>See discussion under <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_siphonostomum"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">siphonostomum</named-content></named-content></title><p><named-content content-type="taxon-authority">(van Benthem Jutting, 1952)</named-content></p><p>http://species-id.net/wiki/Plectostoma_siphonostomum</p><p><xref ref-type="fig" rid="F17">Figure 17N</xref><xref ref-type="fig" rid="F23">, 23</xref><xref ref-type="app" rid="APP1">, Appendix 8</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma siphonostomum</named-content><named-content content-type="comment"> van Benthem Jutting, 1952: 52, figure 27 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma siphonostomum</named-content><named-content content-type="comment"> van Benthem Jutting, <xref rid="B8" ref-type="bibr">van Benthem Jutting (1961</xref>: 39).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma siphonostomum</named-content><named-content content-type="comment"> van Benthem Jutting, <xref rid="B13" ref-type="bibr">Berry (1964</xref>: 203).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: ZMA 136054(1) (Seen). Paratype: ZMA 136055(8) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>ZMA 162138(>100), BOR 5513 (>25), BOR 5521(>10), BOR 5557(>10), V 8199(>50), V 8223(>25).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> the tuba form, but differs by having an oblong conical apical and conical basal spire.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 1.9–2.1 mm. Width: 1.2–1.5 mm. Number of whorls: 4 3/4–5. Apical spire shape: oblong conical. Basal spire shape: conical. Whorl periphery: distinctly convex. Umbilicus: Open.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 1 and aperture visible between right lateral and back view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 3/8–1/2. Proportion of tuba that attaches to spire: at least 1/2.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 7–8 ribs per mm. Rib intensity: thin. Shape: slightly curved. Inclination: orthoclin.</p><fig id="F23" orientation="portrait" position="float"><label>Figures 23.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> (van Benthem Jutting, 1952) BOR5539. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g023"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Gua Siput, Taman Negara, Pahang (<named-content content-type="dwc:verbatimCoordinates">4°26'47"N, 102°14'44"E</named-content>).</p><p>Distribution range. <italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> has a similar distribution pattern as <italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> and often occurs sympatrically with that species. It can be found in many limestone outcrops in the valley between the Titingwangsa Range, Tahan Range and Benom Range (<xref ref-type="fig" rid="F18">Figure 18D</xref>).</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Least concern. Living populations of <italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> were recorded at several limestone hills during surveys between 2010 and 2011. Several of these are located within the National Park.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>See discussion under <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_panhai"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">panhai</named-content></named-content></title><p><named-content content-type="taxon-authority">(Maassen, 2001)</named-content></p><p>http://species-id.net/wiki/Plectostoma_panhai</p><p><xref ref-type="fig" rid="F24">Figure 24</xref><xref ref-type="app" rid="APP1">, Appendix 7</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Arinia panhai</named-content><named-content content-type="comment"> Maassen, 2001: 55, figure 4, 12 & 13 (original description).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: RMNH 81809(1) (Seen). Paratypes: RMNH 81810(2) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>ZMA 138438(1).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic> the general shell form, in the terms of apex, spire and tuba, but differs by having a single parietal constriction tooth.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: moderately convex.</p><p><bold>Spire.</bold> Height: 2.1–2.2 mm. Width: 1.6–1.7 mm. Number of whorls: 4 1/2–4 5/8. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: moderately convex. Umbilicus: open.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Constriction.</bold> Parietal teeth: one. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 1 and aperture visible from back view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 3/4. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 8–9 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: orthoclin.</p><fig id="F24" orientation="portrait" position="float"><label>Figures 24.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic> (Maassen, 2001) ZMA 138438. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g024"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Tham Krachaeng, Thailand (<named-content content-type="dwc:verbatimCoordinates">6°12'50"N, 101°12'9"E</named-content>). The location description in the original publication is not completely correct “(<named-content content-type="dwc:verbatimCoordinates">06°55'022"N, 101°12'160"E</named-content>)” (<xref rid="B63" ref-type="bibr">Maassen 2001</xref>).</p><p>Distribution range. This species is only known from the type locality (<xref ref-type="fig" rid="F18">Figure 18D</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Data Deficient.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>See discussion under <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_christae"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">christae</named-content></named-content></title><p><named-content content-type="taxon-authority">(Maassen, 2001)</named-content></p><p>http://species-id.net/wiki/Plectostoma_christae</p><p><xref ref-type="fig" rid="F17">Figures 17Q</xref><xref ref-type="fig" rid="F25">, 25</xref><xref ref-type="app" rid="APP1">, Appendix 6</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma christae</named-content><named-content content-type="comment"> Maassen, 2001: 52, figures 3, 10 & 11 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma jensi</named-content><named-content content-type="comment"> Maassen, 2001: 56, figures 5, 14 & 15 (original description), syn. n.</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: RMNH 81805(1) (Seen). Paratypes: RMNH 81806(1) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>RMNH 81807(1), RMNH 81808(1), ZMA 138436(1), ZMA 138437(2), BOR 3496(1), BOR 5505(>25), BOR 5506(>50), BOR 5509(>25), BOR 5572(2), V 12702(1), V 8314(>25), V 8406(2), V 9153(2), V 9207(>100), V 9285(3).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic> the general shell spire form but differs by having two parietal constriction teeth and aperture visible when shell observed in left lateral view.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: moderately convex.</p><p><bold>Spire.</bold> Height: 1.9–2.6 mm. Width: 1.4–1.8 mm. Number of whorls: 3 5/8–4 1/2. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: distinctly convex. Umbilicus: open.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 1 and aperture visible from left lateral view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 3/8–5/8. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Radial ribs.</bold> Rib density: 5–6 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: orthoclin.</p><fig id="F25" orientation="portrait" position="float"><label>Figures 25.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> (Maassen, 2001) BOR 5572. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g025"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Limestone hills, 16 km west of Gua Musang (<named-content content-type="dwc:verbatimCoordinates">4°54'46"N, 102°6'22"E</named-content>).</p><pmc-comment>PageBreak</pmc-comment>
<p>Distribution range. Limestone hills along the 50 km Northeast transect between <named-content content-type="dwc:verbatimCoordinates">4°38'51"N, 101°58'58"E</named-content> and <named-content content-type="dwc:verbatimCoordinates">5°0'13"N, 102°11'59"E</named-content> (<xref ref-type="fig" rid="F18">Figure 18D</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Near Threatened. Until today, this species has been recorded from at least six limestone hills. In a field survey in 2011 and 2012, living populations of <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> could be found on four of these hills. All of these are located near the road and are surrounded by oil palm plantation, although there is no immediate threat.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>, together with <italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma umbilicatum</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic> represent a group of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species that have a regulary coiled tuba (type 1 tuba). The species of this group occur only in Peninsular Malaysia and are genetically highly divergent (> 10% differences in COI) from the others (<xref ref-type="table" rid="T4">Table 4</xref>). All of the seven species are distributed allopatrically (<xref ref-type="fig" rid="F18">Figure 18D</xref>).</p><p>We synonymised <italic><named-content content-type="taxon-name">Plectostoma jensi</named-content></italic> with <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>, both of which were described from the same locality. <xref rid="B63" ref-type="bibr">Maassen (2001)</xref> distinguished between them by the slight difference in umbilicus opening and aperture tilting. In the material at our disposal, we recognised that these differences are intrapopulational variation. All individuals share the same diagnostic shell characters as mentioned above. In addition to the morphological evidence, the genetic variation between individuals with different shell forms is smaller than our species delimitation threshold of 10%.</p><p>Two species of this group, namely <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic>, have a wider distribution range than other species in this group. The two species occur parapatrically on the limestone hills in the centre of Peninsular Malaysia (<xref ref-type="fig" rid="F18">Figure 18D</xref>). On the other hand, very little is known of the distribution range of <italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic>. Although this species was reported only once and only from the type locality, it might also occur at other limestone sites near the type locality. <italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic> is very similar to <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>, but the two are separated by more than 150 km, and the limestone hills in between are occupied by three other <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species. The disjunct distribution and its single constriction tooth support the decision that <italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic> is a distinct species from <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>.</p><p>The remaining four species of this group, namely, <italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma umbilicatum</named-content></italic>, are site endemics, occurring at each of the four small limestone clusters in the centre of Peninsular Malaysia (<xref ref-type="fig" rid="F18">Figure 18D</xref>). These clusters are each quite isolated, with no other limestone hills within a 20 km radius.</p><p>Although these four species occur in adjacent limestone limestone clusters, and they have similar shell shapes, their taxonomic status are clear. The COI sequence divergence between these species is larger than 13% and each of them has a set of diagnostic shell characters (<xref ref-type="table" rid="T3">Tables 3</xref> and <xref ref-type="table" rid="T4">4</xref>). This may raise the question how each species evolved in each limestone cluster and how long these four species have been isolated. For example, a neighbouring species, <italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic>, has a similar distribution range as the former four species, but the morphological and genetic divergence is much smaller than in these four species. Presumably, the answer lies in the details of the geomorphological evolution of the limestone outcrops, which, however, remains largely unknown.</p><pmc-comment>PageBreak</pmc-comment>
</sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_crassipupa"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">crassipupa</named-content></named-content></title><p><named-content content-type="taxon-authority">(van Benthem Jutting, 1952)</named-content></p><p>http://species-id.net/wiki/Plectostoma_crassipupa</p><p><xref ref-type="fig" rid="F17">Figures 17I, M</xref><xref ref-type="fig" rid="F26">, 26</xref><xref ref-type="app" rid="APP1">, Appendix 6</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma crassipupa</named-content><named-content content-type="comment"> van Benthem Jutting, 1952: 51, figure 26 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma crassipupa</named-content><named-content content-type="comment"> van Benthem Jutting, <xref rid="B8" ref-type="bibr">van Benthem Jutting (1961</xref>: 39).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma crassipupa</named-content><named-content content-type="comment"> van Benthem Jutting, <xref rid="B23" ref-type="bibr">Clements (2007</xref>: 74).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma crassipupa</named-content><named-content content-type="comment"> van Benthem Jutting, Clements et al. (2008: 2760).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: ZMA 135994(1) (Seen). Paratype: ZMA 135993(>50) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>BOR 5512(>50), BOR 5515(2), BOR 5624(>10), BOR 5629(3), V 8392(>10), V 8407(10), V 8408(4), V 8437(>10), V 8898(7). V 8912(1), V 8956(1), V 9097(9), V 9157(>10), V 9326(>10), V 9353(3), V 9366(10).</p><p>Genetic distance between BOR 5512 and BOR 5515 is 14%.</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma sciaphilum</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic> the tuba form, but differs by having a slightly or moderately convex apex and depressed apical spire.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: slightly or moderately convex.</p><p><bold>Spire.</bold> Height: 1.3–3.0 mm. Width: 1.2–1.4 mm. Number of whorls: 3 1/2–4 1/2. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: moderately or distinctly convex. Umbilicus: closed by tuba (common) or partially open (rare).</p><p><bold>Constriction.</bold> Parietal teeth: none. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 3 and aperture visible from left lateral view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 1/2–3/4 Proportion of tuba that attaches to spire: whole.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 6–8 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: moderately prosoclin.</p><fig id="F26" orientation="portrait" position="float"><label>Figures 26.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic> (van Benthem Jutting, 1952) BOR 5512. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> left lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g026"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Gua Musang (<named-content content-type="dwc:verbatimCoordinates">4°52'59"N, 101°58'12"E</named-content>).</p><p>Distribution range. This species mainly occurs in the limestome hills that are located between Gua Musang and as far as 30 km radius of Gua Musang. One populations exists at a limestone hill that is located about 90 km north of Gua Musang (<xref ref-type="fig" rid="F18">Figure 18A</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Least Concern. Almost all the limestone hills are located along main roads and/or surrounded by oil palm plantation or cleared for urban development. However, several large limestone hills that hold the species are located in the well-protected National Park (Taman Negara), State of Pahang, Malaysia.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>The species is well-characterised, although it displays considerable variability in the shell shape (<xref ref-type="app" rid="APP1">Appendix 6</xref>).</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_tonkinianum"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">tonkinianum</named-content></named-content></title><p><named-content content-type="taxon-authority">(Dautzenberg & Fischer, 1905)</named-content></p><p>http://species-id.net/wiki/Plectostoma_tonkinianum</p><p><xref ref-type="fig" rid="F27">Figure 27</xref><xref ref-type="app" rid="APP1">, Appendix 8</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma tonkinianum</named-content><named-content content-type="comment"> Dautzenberg & Fischer, 1905: 444, plate 10 - figure 5, 6 and 7 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma tonkinianum</named-content><named-content content-type="comment"> Dautzenberg & Fischer, <xref rid="B95" ref-type="bibr">Saurin (1953</xref>: 134).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma tonkinianum</named-content><named-content content-type="comment"> Dautzenberg & Fischer, <xref rid="B9" ref-type="bibr">van Benthem Jutting (1962</xref>: 12).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma tonkinianum</named-content><named-content content-type="comment"> Dautzenberg & Fischer, <xref rid="B39" ref-type="bibr">Fischer (1963</xref>: 34).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Not seen.</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>ZMA 162136(1), V 10000(>25), V 10010(>25), V 10022(>25), V 11243(>25), V 11270(>25), V 11432(3), V 11502(>50), V 7937(5), V 9940(>10), V 9957(>25).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic> the general shell form, in terms of apex, spire and tuba, but differs by having two lamella-shaped constriction teeth and a higher spire (> 3 mm).</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 3.1–3.8 mm. Width: 2.4–2.6 mm. Number of whorls: 4 3/4–4 7/8. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: distinctly convex. Umbilicus: completely closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: two lamellae. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 3 and aperture visible in left lateral view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 5/8. Proportion of tuba that attaches to spire: whole.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Aperture and peristome.</bold> Peristome: simple peristome.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 6–7 ribs per mm. Rib intensity: thin. Shape: slightly curved to single-humped. Inclination: orthoclin.</p><fig id="F27" orientation="portrait" position="float"><label>Figures 27.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma tonkinianum</named-content></italic> (Dautzenberg & Fischer, 1905) V 9940. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view; <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g027"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. “Tonkin” (<xref rid="B35" ref-type="bibr">Dautzenberg and Fischer 1905</xref>). The exact locality should be the limestone hill near Chau Dao.</p><p>Distribution range. Limestone hills in the Province of Kien Giang (<xref ref-type="fig" rid="F18">Figure 18A</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Data deficient. Some recent dead shells have been collected but no information is available on the habitat and population status.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>In recent years, this species has been recorded at several limestone hills in the vicinity of Ha Tien, which is a popular tourism site. Although the type specimen cannot be located and the exact type locality cannot be determined, the shell morphology of the recent material fits well with the description in the original publication.</p><p>In the original publication, <xref rid="B35" ref-type="bibr">Dautzenberg and Fischer (1905)</xref> mentioned that this species was collected by M. Mansuy from Tonkin (French protectorate), which was a large area that included part of Southern China, and the Northern parts of Laos and Vietnam. <xref rid="B35" ref-type="bibr">Dautzenberg and Fischer (1905)</xref> further provided a list of locations where most of the snails were collected. One of these was, Chau Doc, which is located about 70 km from Ha Tien. Furthermore, a recent intensive land snail survey in the Northern Provinces of Laos, namely, Hua Phan and Luang Prabang, did not discover any <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species (Muratov and Abdou 2006). Hence, <italic><named-content content-type="taxon-name">Plectostoma tonkinianum</named-content></italic> probably occurs in the small limestone hill cluster in the coastal area of the Southern part of Vietnam and neighboring Cambodia.</p><pmc-comment>PageBreak</pmc-comment>
</sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">whitteni</named-content></named-content><named-content content-type="taxon-status">sp. n.</named-content></title><p>http://zoobank.org/37B4DE69-9255-4632-AF7F-8EB75D1D4CD8</p><p>http://species-id.net/wiki/Plectostoma_whitteni</p><p><xref ref-type="fig" rid="F17">Figures 17G</xref><xref ref-type="fig" rid="F28">, 28</xref><xref ref-type="app" rid="APP1">, Appendix 8</xref></p><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: BOR 5644(1). Paratypes: BOR 5536(>10), V 8802(1), V 8885(>10).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma tonkinianum</named-content></italic> the general shell form, in the terms of apex, spire and tuba, but differs by lacking constriction teeth and lower spire (< 2 mm).</p></sec><sec sec-type="treatment-Etymology"><title>Etymology.</title><p>This species is named after Dr. Tony Whitten, who was senior biodiversity specialist of the World Bank between 1995 and 2010, and is currently the Regional Director for Asia-Pacific Fauna & Flora International. Dr. Whitten has been actively promoting the protection of the biodiversity that is associated with limestone and has been involved in conservation action to protect limestone habitats.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 1.7–1.9 mm. Width: 1.4–1.6 mm. Number of whorls: 3 7/8–4 1/4. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: distinctly convex. Umbilicus: completely closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: none. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 3 and aperture visible in left lateral view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 1/2–5/8. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Radial ribs.</bold> Rib density: 6–7 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: orthoclin.</p><fig id="F28" orientation="portrait" position="float"><label>Figures 28.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic> sp. n. BOR 5644. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g028"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Gua Taat, Tasik Kenyir (<named-content content-type="dwc:verbatimCoordinates">4°51'3"N, 102°43'21"E</named-content>).</p><p>Distribution range. This species only occurs in Gua Taat, Tasik Kenyir (<xref ref-type="fig" rid="F18">Figure 18A</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Least Concern. Gua Taat is located in a prioritised protected National Park with good forest cover.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>The general shell form of <italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic> is similar to <italic><named-content content-type="taxon-name">Plectostoma tonkinianum</named-content></italic>. There is no genetic information for <italic><named-content content-type="taxon-name">Plectostoma tonkinianum</named-content></italic>, but <italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic> is genetically closer to <italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> than to any of the other 18 species. Nevertheless, <italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic> is considered a distinct species as compared to <italic><named-content content-type="taxon-name">Plectostoma tonkinianum</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> because of the lack of constriction teeth.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_sciaphilum"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">sciaphilum</named-content></named-content></title><p><named-content content-type="taxon-authority">(van Benthem Jutting, 1952)</named-content></p><p>http://species-id.net/wiki/Plectostoma_sciaphilum</p><p><xref ref-type="fig" rid="F29">Figure 29</xref><xref ref-type="app" rid="APP1">, Appendix 7</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma sciaphilum</named-content><named-content content-type="comment"> van Benthem Jutting, 1952: 45, figure 23 (original description).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: ZMA 136049(1) (Seen). Paratypes: ZMA 136050(>10) (Seen).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic> the tuba form, but differs by lacking basal constriction teeth.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 2.6–2.9 mm. Width: 1.5–1.6 mm. Number of whorls: 4 1/4–5 1/2. Apical spire shape: oblong conical. Basal spire shape: ovoid. Whorl periphery: distinctly convex. Umbilicus: completely closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 3 and aperture visible in left lateral view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 3/8. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 6–8 ribs per mm. Rib intensity: thin. Shape: slightly curved.</p><fig id="F29" orientation="portrait" position="float"><label>Figures 29.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma sciaphilum</named-content></italic> (van Benthem Jutting, 1952) ZMA 136050. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g029"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Bukit Panching, Pahang (<named-content content-type="dwc:verbatimCoordinates">3°53'28"N, 103°8'26"E</named-content>).</p><p>Distribution range. Endemic to Bukit Panching (not seen in <xref ref-type="fig" rid="F18">Figure 18A</xref> because its symbol overlaps with <italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Extinct. Its only habitat–Bukit Panching, has been completed quarried away (see also <xref rid="B101" ref-type="bibr">Schilthuizen and Clements 2008</xref>). The ruin is now inundated. The status of this species in a previous assessment (IUCN redlist) was: Critically Endangered B2ab(ii,iii) ver. 3.1 (<xref rid="B26" ref-type="bibr">Clements 2009a</xref>).</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>See under <italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic>.</p><pmc-comment>PageBreak</pmc-comment>
</sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_senex"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">senex</named-content></named-content></title><p><named-content content-type="taxon-authority">(van Benthem Jutting, 1952)</named-content></p><p>http://species-id.net/wiki/Plectostoma_senex</p><p><xref ref-type="fig" rid="F17">Figures 17H</xref><xref ref-type="fig" rid="F30">, 30</xref><xref ref-type="app" rid="APP1">, Appendix 7</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma senex</named-content><named-content content-type="comment"> van Benthem Jutting, 1952: 47, figure 24 (original description).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: ZMA 136052(1) (Seen). Paratypes: ZMA 136053(>10) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>RMNH 44723(5), BOR 460(1), BOR 5575(3), BOR 5603(2), BOR 5628(>100), BOR 5631(>10), V 5117(9).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma sciaphilum</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic> the tuba form, but differs by having two knob-shaped constriction teeth and fewer than 6 whorls.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 2.6–3.1 mm. Width: 1.6–1.7 mm. Number of whorls: 5 1/8–5 5/8. Apical spire shape: oblong conical. Basal spire shape: ovoid. Whorl periphery: moderately to distinctly convex. Umbilicus: completely closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: one transverse tooth.</p><p><bold>Tuba.</bold> Coiling direction: type 3 and aperture visible in left lateral view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 3/8. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Radial ribs.</bold> Rib density: 4 ribs per mm. Rib intensity: thin. Shape: slightly curved. Inclination: orthoclin.</p><fig id="F30" orientation="portrait" position="float"><label>Figures 30.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic> (van Benthem Jutting, 1952) BOR 5603. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g030"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Gua Charas, limestone hill near Sungai Lembing in the state of Pahang, Malaysia (<named-content content-type="dwc:verbatimCoordinates">3°54'27"N, 103°8'47"E</named-content>).</p><p>Distribution range. In addition to the type locality, this species had been recorded from Bukit Panching (<xref ref-type="fig" rid="F18">Figure 18A</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Critically Endangered (B2ab(iii)+C2a(i) ver. 10.1). This species is known from two locations. One of these, Bukit Panching, does not exist anymore. Two intensive surveys at the other location, Bukit Charas, were conducted in Feb. 2010 and May 2011. Two living populations with fewer than 50 individuals were found at wet staglamites near the cave. No other living population was found elsewhere on Bukit Charas. The status of this species in a previous assessment (IUCN redlist) was: vulnerable D2 ver. 3.1 (<xref rid="B27" ref-type="bibr">Clements 2009b</xref>).</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>See discussion under <italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_turriforme"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">turriforme</named-content></named-content></title><p><named-content content-type="taxon-authority">(van Benthem Jutting, 1952)</named-content></p><p>http://species-id.net/wiki/Plectostoma_turriforme</p><p><xref ref-type="fig" rid="F31">Figure 31</xref><xref ref-type="app" rid="APP1">, Appendix 8</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma turriforme</named-content><named-content content-type="comment"> van Benthem Jutting, 1952: 43, figure 22 (original description).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: ZMA 136068(1) (Seen). Paratypes: ZMA 136069(>10) (Seen).</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>BOR 461(2), BOR 5609(2).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma sciaphilum</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic> the tuba form, but differs by having two lamella-shaped constriction teeth and having more than 6 whorls.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 3.2 mm. Width: 1.6 mm. Number of whorls: 7. Apical spire shape: oblong conical. Basal spire shape: conical. Whorl periphery: distinctly convex. Umbilicus: completely closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: two.</p><p><bold>Tuba.</bold> Coiling direction: type 3 and aperture visible in left lateral view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 3/8. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 5 ribs per mm. Rib intensity: thin. Shape: slightly curved. Inclination: orthoclin.</p><fig id="F31" orientation="portrait" position="float"><label>Figures 31.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic> (van Benthem Jutting, 1952) BOR 5609. <bold>A</bold> frontal view <bold>B</bold> leftlateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g031"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Bukit Tenggek, Pahang (<named-content content-type="dwc:verbatimCoordinates">4°0'51"N, 103°9'34"E</named-content>).</p><p>Distribution range. In addition to the type locality, this species can be found at nearby Bukit Sagu (<xref ref-type="fig" rid="F18">Figure 18A</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Critically Endangered (B2ab(iii)+C2a(i) ver. 10.1). The whole Bukit Tenggek will disappear by 2014 because of quarrying activity. Moreover, more than half of Bukit Sagu has already been quarried away (see also <xref rid="B101" ref-type="bibr">Schilthuizen <pmc-comment>PageBreak</pmc-comment>and Clements 2008</xref>). Although living individuals had been recorded from Bukit Sagu in 1997 (BOR 5609), neither living nor recently dead individuals were found in a recent survey conducted in 2010 and 2011.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p><italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma sciaphilum</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic> are three very similar species than occur in the four lenticular easternmost limestone hills in Peninsular Malaysia (<xref ref-type="fig" rid="F18">Figure 18A</xref>). These four hills, namely, Bukit Panching, Bukit Charas, Bukit Sagu, and Bukit Tenggek are located along a 15 km longitudinal transect. <italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic> occurs at the two former sites and <italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic> at the two latter sites. <italic><named-content content-type="taxon-name">Plectostoma sciaphilum</named-content></italic> occurs sympatrically with <italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic> in Bukit Panching. These four hills (and thus the three species) are among the most isolated limestone outcrops in Peninsular Malaysia.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_palinhelix"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">palinhelix</named-content></named-content></title><p><named-content content-type="taxon-authority">(van Benthem Jutting, 1952)</named-content></p><p>http://species-id.net/wiki/Plectostoma_palinhelix</p><p><xref ref-type="fig" rid="F17">Figures 17C</xref><xref ref-type="fig" rid="F32">, 32</xref><xref ref-type="app" rid="APP1">, Appendix 12</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma palinhelix</named-content><named-content content-type="comment"> van Benthem Jutting, 1952: 40, figure 20 (original description).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: ZMA 136030(1) (Seen). Paratype: ZMA 136031(>10) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>BOR 466(1), BOR 5520(5), V 5104(5).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic> the spire and tuba form, but differs by having spire height between 1.6–1.9 mm.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: slightly convex.</p><p><bold>Spire.</bold> Height: 1.6–1.9 mm. Width: 1.1–1.3 mm. Number of whorls: 3 3/4–4 1/4. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: moderately to distinctly convex. Umbilicus: partially or completely closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible between right lateral and back view; the tuba coils upward until the first teleconch whorl of the spire. Tuba whorl length similar to that of the last whorl of the spire. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similer to inner peristome, projected all around, except the posterior part, where the two lateral sides are slightly more projected than the anterior side.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 7–8 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: orthoclin.</p><fig id="F32" orientation="portrait" position="float"><label>Figures 32.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic> (van Benthem Jutting, 1952) BOR 5520. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g032"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Bukit Serdam, Pahang (<named-content content-type="dwc:verbatimCoordinates">3°49'47"N, 101°55'36"E</named-content>).</p><p>Distribution range. To date, this species has been recorded from three limestone hills, namely Bukit Serdam, Gua Kechil and a small hill nearby (<xref ref-type="fig" rid="F18">Figure 18C</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Vulnerable (B2ab(iii) ver. 10.1). There are four limestone hills in this area, of which Bukit Serdam and its next unnamed hill support populations (assessment done in 2010 and 2011). Two of the hills, namely, Bukit Serdam <pmc-comment>PageBreak</pmc-comment>and Gunung Panas, are now being quarried. The smallest unnamed hill is highly degraded and Gua Kechil is surrounded by oil palm plantation.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>See discussion under <italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_retrovertens"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">retrovertens</named-content></named-content></title><p><named-content content-type="taxon-authority">(Tomlin, 1938)</named-content></p><p>http://species-id.net/wiki/Plectostoma_retrovertens</p><p><xref ref-type="fig" rid="F17">Figures 17R</xref><xref ref-type="fig" rid="F33">, 33</xref><xref ref-type="app" rid="APP1">, Appendix 13</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, 1938: 73, Plate 2 - figure 3 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, <xref rid="B6" ref-type="bibr">van Benthem Jutting (1952</xref>: 39).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, <xref rid="B10" ref-type="bibr">Berry (1961)</xref>.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, <xref rid="B11" ref-type="bibr">Berry (1962)</xref>.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, <xref rid="B12" ref-type="bibr">Berry (1963)</xref>.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, <xref rid="B13" ref-type="bibr">Berry (1964)</xref>.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, <xref rid="B14" ref-type="bibr">Berry (1966)</xref>.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, <xref rid="B110" ref-type="bibr">Solem (1966)</xref>.</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, <xref rid="B54" ref-type="bibr">Illert (1987</xref>: 800, figure 2a).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, <xref rid="B111" ref-type="bibr">Solem and Solem (1976</xref>: 31).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, <xref rid="B48" ref-type="bibr">Heller (2001</xref>: 426).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, <xref rid="B23" ref-type="bibr">Clements (2007</xref>: 74).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma retrovertens</named-content><named-content content-type="comment"> Tomlin, Clements et al. (2008: 2760).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: BMNH 1938.10.25.2(1) (Seen). Paratype: ZMA 136044(>10) (Seen).</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>RMNH 244699(1), RMNH 44725(8), ZMA 162133(>10), ZMA 162148(8), BOR 5559(3), BOR 5621(9), V 5124(9).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic> the spire and tuba form, but differs by having spire height between 2.3–2.6 mm.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: moderately convex.</p><p><bold>Spire.</bold> Height: 2.3–2.6 mm. Width: 1.5–1.7 mm. Number of whorls: 4 5/8–5. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: distinctly convex. Umbilicus: partially or completely closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible between right lateral and back view; the tuba coils upward until the first teleconch whorl of the spire. Tuba whorl length similar to the length of the last whorl of the spire. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similer to inner peristome, projected all around, except the posterior part, where two lateral sides are slightly more projected than the anterior side.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 7–9 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: orthoclin.</p><fig id="F33" orientation="portrait" position="float"><label>Figures 33.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic> (Tomlin, 1938) BOR 5559. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g033"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Bukit Chintamanis, Pahang (<named-content content-type="dwc:verbatimCoordinates">3°26'45"N, 102°0'51"E</named-content>).</p><p>Distribution range. Endemic to the type locality (<xref ref-type="fig" rid="F18">Figure 18C</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Critically Endangered (B2ab(iii)+C2a(i) ver. 10.1). This species is endemic to a single limestone hill–Bukit Chintamanis. Most of the western part of this <pmc-comment>PageBreak</pmc-comment>hill is gone due to the quarry activity in the past and this part is now covered by secondary vegetation. The rest of the outcrop is surrounded by plantation, which is subjected to periodic clearing and replanting. According to <xref rid="B10" ref-type="bibr">Berry (1961)</xref>, live individuals can only be found at the rock surface (Site A in <xref rid="B10" ref-type="bibr">Berry 1961</xref>). <xref rid="B11" ref-type="bibr">Berry (1962)</xref> reports finding several thousand individuals at Site A.</p><p>A recent intensive survey for <italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic> was done at the type locality in August 2010 and May 2011. Site A is a rock surface smaller than 20 m<sup>2</sup>. It appears that the vegetation cover and habitat structure of Site A has not changed as compared to the assessment done by <xref rid="B11" ref-type="bibr">Berry (1962</xref>, <xref rid="B13" ref-type="bibr">1964</xref>). However, only three live individuals were found during the survey in Aug. 2010. Furthermore, the surrounding of Bukit Chintamanis has been cleared in 2011. Judging by the population trends and habitat conditions together, this species is at the brink of extinction. The status of this species in a previous assessment (IUCN redlist) was: vulnerable D2 ver. 3.1 (<xref rid="B28" ref-type="bibr">Clements 2009c</xref>).</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p><italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic> are very distinct from other <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> by having a very long tuba in relation to their spire. In fact, a preliminary phylogenetic analysis suggests that these two are basal species for all Peninsular Malaysia’s <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>. These two species are located more 40 km apart on outcrops that belong to the same limestone facies (<xref ref-type="fig" rid="F18">Figure 18C</xref>). The two species are very similar in their shell shape, but <italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic> is about one-third larger than <italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic>. Furthermore, the genetic divergence between the two species is 10%.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">ikanensis</named-content></named-content><named-content content-type="taxon-status">sp. n.</named-content></title><p>http://zoobank.org/D450E51F-D3A1-49B7-ACA2-E70A5C42ED2C</p><p>http://species-id.net/wiki/Plectostoma_ikanensis</p><p><xref ref-type="fig" rid="F17">Figures 17J, K</xref><xref ref-type="fig" rid="F34">, 34</xref><xref ref-type="fig" rid="F35">, 35</xref><xref ref-type="app" rid="APP1">, Appendix 11</xref><xref ref-type="app" rid="APP1">, 14</xref></p><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: BOR 5645(1)</p><p>Paratypes: BOR 5507(6), BOR 5622(>50), BOR 5504(>10), V 9446(>100), V 9320(6).</p></sec><sec sec-type="treatment-Etymology"><title>Etymology.</title><p>This species is named after its type locality–Gua Ikan.</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma kayiani</named-content></italic> the general shell form, in terms of spire and tuba shape, but differs by having both thick and thin spiral lines.</p></sec><sec sec-type="treatment-Description for shell form 5504"><title>Description for shell form 5504</title><p>(<xref ref-type="fig" rid="F17">Figs 17K</xref> and <xref ref-type="fig" rid="F34">34</xref>, and <xref ref-type="app" rid="APP1">Appendix 14</xref>). <bold>Apex.</bold> Shape: moderately convex.</p><p><bold>Spire.</bold> Height: 1.7 mm. Width: 1.3 mm. Number of whorls: 4 1/8–4 1/4. Apical spire shape: depressed conical. Basal spire shape: conical. Whorl periphery: distinctly convex. Umbilicus: open.</p><p><bold>Constriction.</bold> Parietal teeth: none. Basal teeth: none</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in right lateral view. Tuba whorl length in proportion to spire last whorl: ca.3/4–7/8. Proportion of tuba that attaches to spire: whole.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similar to inner peristome, projected all around, except the posterior part, where left lateral sides are slightly more projected than the anterior and right lateral side.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Radial ribs.</bold> Rib density: 5 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: orthoclin.</p><fig id="F34" orientation="portrait" position="float"><label>Figures 34.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n. (Form BOR 5504) BOR 5504. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g034"></graphic></fig></sec><sec sec-type="treatment-Description for shell form 5507"><title>Description for shell form 5507</title><p>(<xref ref-type="fig" rid="F17">Figs 17J</xref> and <xref ref-type="fig" rid="F35">35</xref>, and <xref ref-type="app" rid="APP1">Appendix 11</xref>). <bold>Apex.</bold> Shape: moderately convex.</p><p><bold>Spire.</bold> Height: 1.8–1.9 mm. Width: 1.3 mm. Number of whorls: 3 7/8–4. Apical spire shape: depressed conical. Basal spire shape: conical. Whorl periphery: distinctly convex. Umbilicus: open.</p><p><bold>Constriction.</bold> Parietal teeth: none. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in left lateral view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 5/8. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Aperture with double peristome. Peristome: double peristomes. Outer peristome shape: similar to inner peristome, projected all around, except the posterior part, where left lateral sides are slightly more projected than the anterior and right lateral side.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 7 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: orthoclin.</p><fig id="F35" orientation="portrait" position="float"><label>Figures 35.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n. (Form BOR 5507) BOR 5645. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g035"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Gua Ikan in the State of Kelantan (<named-content content-type="dwc:verbatimCoordinates">5°21'9"N, 102°1'34"E</named-content>).</p><p>Distribution range. In addition to the type location, this species also occurs at nearby limestone hills as far as 30 km away (<xref ref-type="fig" rid="F18">Figure 18C</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Least concern. Living populations of <italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> were recorded at the type locality in 2011. The type locality is gazetted as recreation forest.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>The two forms of <italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> were found in the type locality at two different parts of the hill that within a distance of 100 m. The overall shell appearances of these two forms are very different, especially in terms of tuba coiling direction and spire shape. Interestingly, one of the <italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> forms (i.e. BOR 5507) is very similar to <italic><named-content content-type="taxon-name">Plectostoma davisoni</named-content></italic> on the basis of these two shell characters. In most of the other <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species examined in this study, tuba coiling direction and spire shape are rather stable characters within a species. Nevertheless, both <italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> forms lack a constriction which unite them and distinguish them from other similar species (see Diagnosis). In addition to the morphological evidence, we found that the genetic divergence of these two forms is smaller than 1% (<xref ref-type="table" rid="T4">Table 4</xref>), and our preliminary phylogenetic analysis shows that the two forms are reciprocally monophyletic. We therefore conclude that they be classified as the same species.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">kayiani</named-content></named-content><named-content content-type="taxon-status">sp. n.</named-content></title><p>http://zoobank.org/30F310AB-7709-4FF3-B5DD-5477AEBAA8BD</p><p>http://species-id.net/wiki/Plectostoma_kayiani</p><p><xref ref-type="fig" rid="F36">Figure 36</xref><xref ref-type="app" rid="APP1">, Appendix 11</xref></p><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: RMNH 330803 (1). Paratypes: V 8883(6), V 14243(1).</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Etymology"><title>Etymology.</title><p>This species is named after Kay Arnold and Ian Mellsop from New Zealand, who have generously supported wildlife conservation work in many parts of Peninsular Malaysia, including the forests around Lake Kenyir where this species was discovered.</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> the general shell form, in terms of spire and tuba shape, but differs by lacking thick spiral lines.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: slightly to moderately convex.</p><p><bold>Spire.</bold> Height: 1.4–1.6 mm. Width: 1.2–1.4 mm. Number of whorls: 3 1/2–3 3/4. Apical spire shape: depressed conical. Basal spire shape: conical. Whorl periphery: distinctly convex. Umbilicus: open.</p><p><bold>Constriction.</bold> Parietal teeth: none. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible from front view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 5/8–3/4. Proportion of tuba that attaches to spire: almost whole except the part near the aperture.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similer to inner peristome, projected all around, except the posterior part, where left lateral sides are slightly more projected than the anterior and right lateral side.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 8–10 ribs per mm. Rib intensity: thin. Shape: slightly curved. Inclination: orthoclin.</p><fig id="F36" orientation="portrait" position="float"><label>Figures 36.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma kayiani</named-content></italic> sp. n. RMNH 330803. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g036"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Gua Bewah, Tasik Kenyir (<named-content content-type="dwc:verbatimCoordinates">4°51'3"N, 102°43'21"E</named-content>).</p><p>Distribution range. To date, this species is only known to occur at two neighbouring limestone outcrops, namely, Gua Bewah and Gua Taat at the southern part of Tasik Kenyir (<xref ref-type="fig" rid="F18">Figure 18C</xref>).</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Least concern. The outcrops where this species is found, are partially submerged in Southeast-Asia’s largest man-made lake, Tasik Kenyir. Despite this, there is a good forest cover around and on the limestone outcrops. Furthermore, these two hills are located in a protected National Park.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>This species occurs sympatrically with <italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">davisoni</named-content></named-content><named-content content-type="taxon-status">sp. n.</named-content></title><p>http://zoobank.org/D54FF7DC-7DD6-44D3-8A53-C45DEAF5243E</p><p>http://species-id.net/wiki/Plectostoma_davisoni</p><p><xref ref-type="fig" rid="F37">Figure 37</xref><xref ref-type="app" rid="APP1">, Appendix 9</xref><xref ref-type="app" rid="APP1">, 10</xref></p><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: BOR 5646(1)</p><p>Paratypes: ZMA 162069(>25), ZMA 162070(3), ZMA 162071(>50), ZMA 162146(7), ZMA 162147(8), BOR 5508(>25), BOR 5626(>25), BOR 5641(9), V 8652(6), V 8929(>25), V 9206(6), V 8265(>10), V 8301(>25), V 9243(>50), V 9340(7), V 9417(7), V 14242(5).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma relauensis</named-content></italic> the general shell form, in terms of apex, spire, and tuba shape, but differs by having two parietal constriction teeth.</p></sec><sec sec-type="treatment-Etymology"><title>Etymology.</title><p>This species is named after Dr. Geoffrey Davison, who has been involved in the conservation of limestone hills in Malaysia, and has collected a lot of snail specimens, many of which are included in this revision of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> from Malaysia.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: slightly to moderately convex.</p><p><bold>Spire.</bold> Height: 1.6–2.0 mm. Width: 1.4–1.5 mm. Number of whorls: 3 5/8–4 7/8. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: moderately to distinctly convex. Umbilicus: open.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in frontal view. Tuba whorl length in proportion to spire last whorl: ca. 5/8–3/4. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similer to inner peristome, projected all around, except the posterior part, where the two lateral sides are slightly more projected than the anterior side.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 6–7 ribs per mm. Rib intensity: thin. Shape: straight. Inclination: orthoclin.</p><fig id="F37" orientation="portrait" position="float"><label>Figures 37.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma davisoni</named-content></italic> sp. n. BOR 5646. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g037"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Limestone hill on the right hand side of the road D29, at km 17 when travelling from Jelawang to Gua Musang (<named-content content-type="dwc:verbatimCoordinates">4°59'4"N, 101°57'53"E</named-content>).</p><p>Distribution range. This species has a very large distribution range, <italic>ca.</italic> 80 km diameter. It can be found in many limestone outcrops in the central part of Peninsular Malaysia, mainly in the State of Kelantan (<xref ref-type="fig" rid="F17">Figure 17D</xref>). In addition, it can also be found in a cluster of limestone hills located at upper Sungai Keniyam Kecil in Taman Negara (ca. 60 km from Gua Musang).</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Least concern. This is a widespread species. Although many hills in Kelantan are being degraded and surrounded by oil palm plantations, there are a few well protected hills in Taman Negara in Pahang, where this species occurs.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>This species is highly variable in the shell form (<xref ref-type="app" rid="APP1">Appendix 9</xref> and <xref ref-type="app" rid="APP1">Appendix 10</xref>), and has a very wide distribution range which partly overlaps with many other <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species (<xref ref-type="fig" rid="F18">Figure 18</xref>). It is very densely distributed in the State of Kelantan, parapatric with <italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic>. The species becomes more sparse toward the limestone hills in Taman Negara, Pahang (<xref ref-type="fig" rid="F18">Figure 18C</xref>). In view of this, it is possible that the species actually consists of two or more cryptic species, and thus more genetic data are needed.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">relauensis</named-content></named-content><named-content content-type="taxon-status">sp. n.</named-content></title><p>http://zoobank.org/7BC76202-2661-4F7C-A922-4DBA32622400</p><p>http://species-id.net/wiki/Plectostoma_relauensis</p><p><xref ref-type="fig" rid="F17">Figures 17P</xref><xref ref-type="fig" rid="F38">, 38</xref><xref ref-type="app" rid="APP1">, Appendix 13</xref></p><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: BOR 5647(1). Paratypes: BOR 463(2), BOR 5511(>25), V 8169(9).</p></sec><sec sec-type="treatment-Etymology"><title>Etymology.</title><p>This species is named after its type locality–Relau substation of Taman Negara, where Gua Gajah is located.</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma davisoni</named-content></italic> the general shell form, in terms of apex, spire, and tuba shape, but differs by having a single parietal constriction tooth.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: slightly to moderately convex.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Spire.</bold> Height: 1.5–1.9 mm. Width: 1.5–1.6 mm. Number of whorls: 3 3/8–3 5/8. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: moderately to distinctly convex. Umbilicus: open.</p><p><bold>Constriction.</bold> Parietal teeth: one. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in frontal view. Tuba whorl length in proportion to spire last whorl: ca. 1/2–5/8. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 4–7 ribs per mm. Rib intensity: thin. Shape: straight to slightly curved. Inclination: orthoclin.</p><fig id="F38" orientation="portrait" position="float"><label>Figures 38.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma relauensis</named-content></italic> sp. n. BOR 5511. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g038"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Gua Gajah in the Relau substation, Taman Negara. (<named-content content-type="dwc:verbatimCoordinates">4°38'15"N, 102°3'50"E</named-content>).</p><p>Distribution range. This species also occurs at a limestone hill located about 20 km north of the type locality. BOR 463 was collection in Pulau Singa Besar, which is located about 300 km from type locality. The reliability of the collection data is dubious (see also BOR 462 of <italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Least Concern. A large living population of <italic><named-content content-type="taxon-name">Plectostoma relauensis</named-content></italic> was found at Gua Gajah, which is located in the National Park.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>This species occurs sympatrically with <italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> on the same limestone hills. Despite the high density of limestone hills in the area, where many other <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species occur, <italic><named-content content-type="taxon-name">Plectostoma relauensis</named-content></italic> is only found in two of these (<xref ref-type="fig" rid="F18">Figure 18</xref>).</p><pmc-comment>PageBreak</pmc-comment>
</sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_kakiense"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">kakiense</named-content></named-content></title><p><named-content content-type="taxon-authority">(Tomlin, 1948)</named-content></p><p>http://species-id.net/wiki/Plectostoma_kakiense</p><p><xref ref-type="fig" rid="F17">Figures 17D</xref><xref ref-type="fig" rid="F39">, 39</xref><xref ref-type="app" rid="APP1">, Appendix 11</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma kakiense</named-content><named-content content-type="comment"> Tomlin, 1948: 225, Plate 2 - figure 5 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma kakiense</named-content><named-content content-type="comment"> Tomlin, <xref rid="B6" ref-type="bibr">van Benthem Jutting (1952</xref>: 39).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma kakiense</named-content><named-content content-type="comment"> Tomlin, <xref rid="B3" ref-type="bibr">Ali and Yaakob (2001</xref>: 145).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma</named-content><named-content content-type="comment"> sp., Razalli et al. (2010: figure 2C).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: BMNH 1948.10.2.3(1) (seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>ZMA 136009(1), ZMA 162094(5), BOR 445(1), BOR 5516(>25), BOR 5517(>10), BOR 5598(3), V 8789(2).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic> the general shell form, in terms of apex, spire, and tuba shape, but differs by having two parietal constriction teeth.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: slightly to moderately convex.</p><p><bold>Spire.</bold> Height: 1.7–2.1 mm. Width: 1.5–1.6 mm. Number of whorls: 3 3/4–4 1/4. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: moderately to distinctly convex. Umbilicus: partially closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in right lateral view. Tuba whorl length in proportion to spire last whorl: ca. 5/8–3/4. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similar to inner peristome, projected all around, except the posterior part, where the two lateral sides are slightly more projected than the anterior side.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 8–10 ribs per mm. Rib intensity: thin. Shape: straight to slightly curved. Inclination: moderately prosoclin.</p><fig id="F39" orientation="portrait" position="float"><label>Figures 39.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma kakiense</named-content></italic> (Tomlin, 1948) BOR 5516. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g039"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Kaki Bukit (<named-content content-type="dwc:verbatimCoordinates">6°38'42"N, 100°12'6"E</named-content>) (<xref ref-type="fig" rid="F18">Figure 18C</xref>). Distribution range. Wang Kelian limestone outcrops.</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Near threatened. To date, only two populations are known for two large limestone outcrops in the vast limestone formation in Perlis. These two locations are Kaki Bukit and Wang Kelian, both located in the protected Wang Kelian State Park.</p><p>Although Kaki Bukit is a large limestone outcrop with good forest cover, the population density of <italic><named-content content-type="taxon-name">Plectostoma kakiense</named-content></italic> is very low. During a survey in May 2011, this species was only found near the top of Kaki Bukit where several hundred individuals were found at one location (limestone wall) within an area of ca. 10 m<sup>2</sup>. There were several small populations (fewer than 10 individuals) in small pockets of suitable habitat. The population in Wang Kelian was recorded by <xref rid="B3" ref-type="bibr">Ali and Yaakob (2001)</xref>, but its status is not known. No live or dead individuals were collected at the dozen isolated limestone hills located within 5 km around Wang Kelian State Park.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>See discussion under <italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic>.</p><pmc-comment>PageBreak</pmc-comment>
</sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">kubuensis</named-content></named-content><named-content content-type="taxon-status">sp. n.</named-content></title><p>http://zoobank.org/7316ED06-4AF5-4E23-89E3-0BCE01E8B1C5</p><p>http://species-id.net/wiki/Plectostoma_kubuensis</p><p><xref ref-type="fig" rid="F17">Figures 17F</xref><xref ref-type="fig" rid="F40">, 40</xref><xref ref-type="app" rid="APP1">, Appendix 12</xref></p><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: BOR 5648(1). Paratypes: BOR 5518(>25), BOR 5519(>10).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma kakiense</named-content></italic> the general shell form, in terms of apex, spire, and tuba shape, but differs by lacking constriction teeth.</p></sec><sec sec-type="treatment-Etymology"><title>Etymology.</title><p>This species is named after its type locality–Bukit Kubu, Perlis.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: slightly to moderately convex.</p><p><bold>Spire.</bold> Height: 1.6–2.0 mm. Width: 1.3–1.5 mm. Number of whorls: 3 5/8–4 1/4. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: moderately to distinctly convex. Umbilicus: open or partially closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: none. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 3 and aperture visible in right lateral view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 5/8–3/4. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similar to inner peristome, projected all around, except the posterior part, where the left lateral sides are slightly more projected than the anterior and right lateral side.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 10–11 ribs per mm. Rib intensity: thin. Shape: straight to slightly curved. Inclination: moderately prosoclin.</p><fig id="F40" orientation="portrait" position="float"><label>Figures 40.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic> sp. n. BOR 5648. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g040"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Bukit Kubu (<named-content content-type="dwc:verbatimCoordinates">6°24'15"N, 100°8'37"E</named-content>) (<xref ref-type="fig" rid="F18">Figure 18C</xref>).</p><pmc-comment>PageBreak</pmc-comment>
<p>Distribution range. Endemic to the type locality.</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Endangered (B2ab(iii)+C2a(i) ver. 10.1). During a survey of a dozen isolated limestone hills of the State of Perlis in May 2011, only Bukit Kubu was found to support a living population of <italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic>, consisting of several hundred individuals at the top of Bukit Kubu. Several very small populations (< 50 individuals) live at the other part of Bukit Kubu where the habitat is relatively more exposed and dry. Bukit Kubu is gazetted by Perlis State government as a recreation forest, and its surroundings consist of urban development and paddy fields.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>From the conchological point of view, the overall shape and size of <italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma kakiense</named-content></italic> is almost the same. Nevertheless, two inconspicuous but significant shell characters, namely, basal and parietal constriction teeth, mark the difference between these two species. Despite the similarity in shell form, there is a great genetic distance between the two species (16%). The species occur at two limestone hills that lie about 30 km apart. During our survey, we could not find either species at the six limestone hills that are located between these two hills. A single <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> shell was recorded by <xref rid="B76" ref-type="bibr">Norhanis et al. (2010)</xref> from Pulau Dayang Bunting, Langkawi, and might belong to this species.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_laemodes"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">laemodes</named-content></named-content></title><p><named-content content-type="taxon-authority">(van Benthem Jutting, 1961)</named-content></p><p>http://species-id.net/wiki/Plectostoma_laemodes</p><p><xref ref-type="fig" rid="F41">Figure 41</xref><xref ref-type="app" rid="APP1">, Appendix 12</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma laemodes</named-content><named-content content-type="comment"> van Benthem Jutting, 1961: 40, plate 11 - figure 6 (original description).</named-content></p></list-item><pmc-comment>PageBreak</pmc-comment>
</list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: ZMA 136010(1) (Seen). Paratypes: ZMA 136011(3) (Seen), ZMA 136012(3) (Seen), ZMA 136013(4) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>RMNH 156267(7), V 8490(>25).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> the general shell form, in terms of apex, spire, and tuba shape, but differs by having the whole tuba attached to the spire.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 2.3–2.7 mm. Width: 1.6–1.8 mm. Number of whorls: 3 1/8–4 5/8. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: moderately to distinctly convex. Umbilicus: open or partially closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 3 and aperture visible in right lateral view. Tuba whorl length in proportion to spire last whorl: ca. 1/2–5/8. Proportion of tuba that attaches to spire: whole.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similar to inner peristome, projected all around, except the posterior part, where the left lateral sides are slightly more projected than the anterior and right lateral side.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 6–7 ribs per mm. Rib intensity: thin. Shape: straight to slightly curved. Inclination: prosoclin.</p><fig id="F41" orientation="portrait" position="float"><label>Figures 41.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma laemodes</named-content></italic> (van Benthem Jutting, 1961) ZMA 136011. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g041"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Batu Tai Gadjah, Ulu Keniyam Kechil, Taman Negara, Pahang Malaysia (<named-content content-type="dwc:verbatimCoordinates">4°37'0"N, 102°25'14"E</named-content>) (<xref ref-type="fig" rid="F18">Figure 18B</xref>).</p><p>Distribution range. This species mainly occurs in a cluster of limestone hills that are located at upper Sungai Keniyam Kecil in Taman Negara (<xref ref-type="fig" rid="F18">Figure 18B</xref>). In addition, <pmc-comment>PageBreak</pmc-comment>this species has been recorded from Bukit Jereng, Blau, Kelantan, which is about 80 km west from Sungai Keniyam Kecil.</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Least concern. Although no living population has been recorded so far, several large and well protected limestone hills in the Taman Negara probably support a viable population.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>From a conchological point of view, this species is similar to <italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic>; the two may be closely related. In view of biogeography, this species occurs parapatrically with <italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic>, which occurs in the limestone hills in between Bukit Jereng and the limestone cluster at Sungai Keniyam Kecil (<xref ref-type="fig" rid="F18">Figure 18B</xref>). Further genetic data are needed to verify the taxonomic status of <italic><named-content content-type="taxon-name">Plectostoma laemodes</named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_salpidomon"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">salpidomon</named-content></named-content></title><p><named-content content-type="taxon-authority">(van Benthem Jutting, 1952)</named-content></p><p>http://species-id.net/wiki/Plectostoma_salpidomon</p><p><xref ref-type="fig" rid="F17">Figures 17A</xref><xref ref-type="fig" rid="F42">, 42</xref><xref ref-type="app" rid="APP1">, Appendix 13</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma salpidomon</named-content><named-content content-type="comment"> van Benthem Jutting, 1952: 42, figure 21 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma salpidomon</named-content><named-content content-type="comment"> van Benthem Jutting, <xref rid="B8" ref-type="bibr">van Benthem Jutting (1961</xref>: 39).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: ZMA 136045(1) (Seen). Paratypes: ZMA 136046(>25) (Seen), ZMA 136047(2) (Seen), ZMA 136048(>10) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>ZMA 162137(>25), BOR 459(1), BOR 5539(>10), BOR 5540(>10), BOR 5541(5), BOR 5542(>25), BOR 5569(>25), BOR 5611(1), V 8171(4), V 8658(>25), V 8706(6), V 9116(7), V 9292(3).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma laemodes</named-content></italic> the general shell form, in terms of apex, spire, and tuba shape, but differs by having a tuba that attaches partly to the spire.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 2.5–2.8 mm. Width: 1.5–1.7 mm. Number of whorls: 4 5/8–5 1/4. Apical spire shape: depressed conical. Basal spire shape: ovoid. Whorl periphery: moderately to distinctly convex. Umbilicus: completely or partially closed by tuba.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in right lateral view. Tuba whorl length in proportion to spire last whorl: ca. 3/4–7/8. Proportion of tuba that attaches to spire: less than 2/3.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similar to inner peristome, projected all around, except the posterior part, where the two lateral sides are slightly more projected than the anterior side.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 5–6 ribs per mm. Rib intensity: thin. Shape: slightly curved. Inclination: prosoclin.</p><fig id="F42" orientation="portrait" position="float"><label>Figures 42.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> (van Benthem Jutting, 1952) BOR 5539. <bold>A</bold> frontal view <bold>B</bold> leftlateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g042"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Gua Bama, Pahang (<named-content content-type="dwc:verbatimCoordinates">4°11'37"N, 101°58'2"E</named-content>) (<xref ref-type="fig" rid="F18">Figure 18B</xref>).</p><p>Distribution range. <italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> can be found in many limestone hills in the valley in between the Titiwangsa, Tahan and Benom mountain ranges (<xref ref-type="fig" rid="F18">Figure 18B</xref>).</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Least concern. <italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> has a wide distribution range. Living populations have been recorded in many limestone hills and some of these are in protected areas.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>See discussion under <italic><named-content content-type="taxon-name">Plectostoma laemodes</named-content></italic>.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_kitteli"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">kitteli</named-content></named-content></title><p><named-content content-type="taxon-authority">(Maassen, 2002)</named-content></p><p>http://species-id.net/wiki/Plectostoma_kitteli</p><p><xref ref-type="fig" rid="F43">Figure 43</xref><xref ref-type="app" rid="APP1">, Appendix 11</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma kitteli</named-content><named-content content-type="comment"> Maassen, 2002: 176, figures 35 & 36 (original description).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: RMNH 92942(1) (seen). Paratypes: RMNH 92956(>25) (Seen), V 12697(9) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>BOR 1697(>10). YSC.</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma charasense</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma klongsangensis</named-content></italic> the general shell form, in terms of apex, spire, and tuba shape, but differs by lacking thick spiral lines and basal constriction teeth.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 2.9–3.0 mm. Width: 1.7–1.8 mm. Number of whorls: 5 5/8–5 3/4. Apical spire shape: oblong conical. Basal spire shape: conical. Whorl periphery: distinctly convex. Umbilicus: partially closed by tuba.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in right lateral view. Tuba whorl length in proportion to spire last whorl: ca. 3/4–7/8. Proportion of tuba that attaches to spire: less than 2/3.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similar to inner peristome, projected all around, except the posterior part, where the two lateral sides are slightly more projected than the anterior side.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 5–6 ribs per mm. Rib intensity: thin. Shape: slightly curved. Inclination: orthoclin.</p><fig id="F43" orientation="portrait" position="float"><label>Figures 43.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma kitteli</named-content></italic> (Maassen, 2002) V 12697. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g043"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Cave Pangian (local name Ngalau Pangian - <named-content content-type="dwc:verbatimCoordinates">0°27'45"S, 100°45'8"E</named-content>)</p><p>Distribution range. This species was also found at a limestone hill near Kampung Desa Gadut, West Sumatra (<named-content content-type="dwc:verbatimCoordinates">0°15'36"S, 100°44'16"E</named-content>). This hill is located about 20 km from the type locality. In addition, a private collector (Yansen Chen, pers. comm.) also collected the same species at Tiangko cave, about 230 km from the type locality (<xref ref-type="fig" rid="F18">Figure 18B</xref>). It therefore appears to have a wide distribution range in Sumatra. It may thus be expected that the limestone outcrops around the three hills mentioned, might support the same species.</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Data deficient. This species has only been recorded from three limestone hills in West Sumatra. The population status and actual distribution <pmc-comment>PageBreak</pmc-comment>range of this species remain unknown. Nevertheless, concern is warrented, as the surroundings of the type locality and several other limestone outcrops in the same area are highly degraded.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>There are large areas in the Northern and Western provinces of Sumatra that are covered by limestone hills (Verstappen and Genootschap 1973). Despite intensive malacological surveys in some of these areas (van Benthem Jutting 1959), no <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species was found until 1997. To date, <italic><named-content content-type="taxon-name">Plectostoma kitteli</named-content></italic> is the only <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species found in Sumatra (<xref rid="B64" ref-type="bibr">Maassen 2002</xref>).</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_klongsangensis"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">klongsangensis</named-content></named-content></title><p><named-content content-type="taxon-authority">(Panha, 1997)</named-content></p><p>http://species-id.net/wiki/Plectostoma_klongsangensis</p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma klongsangensis</named-content><named-content content-type="comment"> Panha, 1997: 133, figure 1 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma klongsangensis</named-content><named-content content-type="comment"> Panha, <xref rid="B50" ref-type="bibr">Hemmen and Hemmen (2001</xref>: 39).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma klongsangensis</named-content><named-content content-type="comment"> Panha, <xref rid="B75" ref-type="bibr">Nabhitabhata (2009</xref>: 50).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: CUIZMD 0001 (not seen). Paratypes: CUIZMDM 0002 (Not seen), CUIZMDM 0003 in ZMA (Not seen, specimen could not be located in ZMA).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma charasense</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma kitteli</named-content></italic> the general shell form, in terms of apex, spire, and tuba, but differs by having both thin and thick spiral lines, and the left lateral side of outer peristome projected more than three times the distance of the right lateral side of outer peristome.</p><p><bold>Description</bold> (estimated from figure in <xref rid="B81" ref-type="bibr">Panha 1996</xref>). <bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 2.3 mm. Width: 1.7 mm. Number of whorls: 5 1/2. Apical spire shape: oblong conical. Basal spire shape: conical. Whorl periphery: distinctly convex. Umbilicus: open.</p><p><bold>Constriction.</bold> Unknown.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in right lateral view. Tuba whorl length in proportion to spire last whorl: ca. 7/8. Proportion of tuba that attaches to spire: less than 2/3.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: different from inner peristome, the left lateral side of outer peristome is projected from inner peristome about 0.7 mm, and the right side of outer peristome about 0.1 mm, but narrowed toward the anterior part of outer peristome.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 3 ribs per mm. Rib intensity: thick. Shape: single-looped. Inclination: prosoclin.</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Khlong Saeng Wildlife Sanctuary Surat Thani Province, Thailand (<named-content content-type="dwc:verbatimCoordinates">8°31'13"N, 98°25'17"E</named-content>) (<xref ref-type="fig" rid="F18">Figure 18B</xref>).</p><p>Distribution range. Unknown.</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Data deficient. This species has been collected once in Khlong Saeng Wildlife Sanctuary.</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>From a conchological point of view, this species appears not to be related to <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species from the Malay Peninsula and Sumatra. Instead, it is almost identical to <italic><named-content content-type="taxon-name">Plectostoma mirabile</named-content></italic>, an endemic species from Gomantong Cave, Sabah, Borneo. These two species are separated by the South China Sea and located more than 2000 km apart. It remains to be determined whether this similarity is due to a disjunct distribution of closely related forms, or rather convergent shell evolution.</p><p>The taxonomic status of the species remains doubtful. In the original publication, <xref rid="B81" ref-type="bibr">Panha (1996)</xref> compared it with <italic><named-content content-type="taxon-name">Plectostoma heteropleuron</named-content></italic> (<xref rid="B122" ref-type="bibr">Vermeulen 1994</xref>) and <italic><named-content content-type="taxon-name">Plectostoma perspectivum</named-content></italic> (Vermeulen, 1994) from Northern Borneo, but not with <italic><named-content content-type="taxon-name">Plectostoma mirabile</named-content></italic> (Smith, 1893), which has an almost identical shell as <italic><named-content content-type="taxon-name">Plectostoma klongsangensis</named-content></italic>. This is all the more remarkable, as <italic><named-content content-type="taxon-name">Plectostoma mirabile</named-content></italic> (Smith, 1893) was treated in <xref rid="B122" ref-type="bibr">Vermeulen (1994)</xref>, where both <italic><named-content content-type="taxon-name">Plectostoma heteropleuron</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma perspectivum</named-content></italic> were originally described.</p><p>In conclusion, more data are needed to verify the taxonomic status and the interesting biogeography of this species.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_charasense"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">charasense</named-content></named-content></title><p><named-content content-type="taxon-authority">(Tomlin, 1948)</named-content></p><p>http://species-id.net/wiki/Plectostoma_charasense</p><p><xref ref-type="fig" rid="F44">Figure 44</xref><xref ref-type="app" rid="APP1">, Appendix 14</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma charasense</named-content><named-content content-type="comment"> Tomlin, 1948: 225, Plate 2 - figure 4 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma charasense</named-content><named-content content-type="comment"> Tomlin, <xref rid="B6" ref-type="bibr">van Benthem Jutting (1952</xref>: 42).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: BNHM 1948.10.2.2(1) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>ZMA 162063(>10), ZMA 162064(>10).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma kitteli</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma klongsangensis</named-content></italic> the general shell form, in terms of apex, spire, and tuba shape, but differs by lacking thick spiral lines and by having two basal constriction teeth.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 1.9–2.0 mm. Width: 1.3–1.4 mm. Number of whorls: 4 1/8. Apical spire shape: oblong conical. Basal spire shape: conical. Whorl periphery: distinctly convex. Umbilicus: open.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: two.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible from right lateral view. Tuba whorl length in proportion to spire last whorl: ca. 7/8. Proportion of tuba that attaches to spire: less than 1/2.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 4 ribs per mm. Rib intensity: thick. Shape: slightly curved to single-humped. Inclination: moderately prosoclin.</p><fig id="F44" orientation="portrait" position="float"><label>Figures 44.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma charasense</named-content></italic> (Tomlin, 1948) ZMA 162063. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g044"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Gua Charas, limestone hill near Sungai Lembing in the state of Pahang, Malaysia (<named-content content-type="dwc:verbatimCoordinates">3°54'27"N, 103°8'47"E</named-content>).</p><pmc-comment>PageBreak</pmc-comment>
<p>Distribution range. Until now, this species has been recorded from two neighbouring limestone hills, namely, Gua Charas and Bukit Panching. However, the whole hill of Bukit Panching has been lost to quarrying (<xref ref-type="fig" rid="F18">Figure 18B</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Critically Endangered (B2ab(iii)+C2a(i) ver. 10.1). Gua Charas might support a viable population. However, the hill is currently surrounded by oil palm plantation with a very narrow forested buffer zone. No living individuals or fresh dead shells were found at Gua Charas after exhaustive search during several sampling trips in 2010 and 2011.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>The taxonomic status of this species is stable. Although the two neighbouring species, <italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic>, are similar to <italic><named-content content-type="taxon-name">Plectostoma charasense</named-content></italic>, there are a few key shell characters that separate <italic><named-content content-type="taxon-name">Plectostoma charasense</named-content></italic> from the others. See discussion under <italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> for more discussion about the biogeography of this species.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">tohchinyawi</named-content></named-content><named-content content-type="taxon-status">sp. n.</named-content></title><p>http://zoobank.org/1D7B3A14-7D4D-42A1-B4BB-E12A71C560B5</p><p>http://species-id.net/wiki/Plectostoma_tohchinyawi</p><p><xref ref-type="fig" rid="F17">Figure 17S</xref><xref ref-type="fig" rid="F45">, 45</xref><xref ref-type="app" rid="APP1">, Appendix 14</xref></p><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: BOR 5649(1). Paratypes: BOR 5533(>10), BOR 5534(>10), BOR 5535(>10), V 8811(>25).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <pmc-comment>PageBreak</pmc-comment><italic><named-content content-type="taxon-name">Plectostoma charasense</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma kitteli</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma klongsangensis</named-content></italic> the general shell form, in terms of apex, spire, and tuba shape, but differs by having both thin and thick spiral lines, and the left lateral side of outer peristome projected not more than twice the width of the right lateral side of outer peristome.</p></sec><sec sec-type="treatment-Etymology"><title>Etymology.</title><p>This species is named after Dato’ Toh Chin Yaw, who was the Chairman of Industry, Trade and Environment Committee in the Terengganu State Government between 2008 and 2013. It is rare to find politicians working closely with the public for wildlife conservation. During his time in office, Dato’ Toh was tireless in his efforts to promote and preserve Terengganu’s natural heritage. One of his most influential decisions was getting the State to ban the hunting of threatened flying foxes (<italic><named-content content-type="taxon-name">Pteropus vampyrus</named-content></italic>). Before he left office, he was working with scientists to gazette the Kenyir Wildlife Corridor as a protected area and was helping to secure funds to improve anti-poaching efforts in that area.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 2.1–2.4 mm. Width: 1.3–1.5 mm. Number of whorls: 4 7/8–5 3/8. Apical spire shape: oblong conical. Basal spire shape: conical. Whorl periphery: distinctly convex. Umbilicus: open.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in right lateral view. Tuba whorl length in proportion to spire last whorl: ca. 7/8–8/8. Proportion of tuba that attaches to spire: less than 1/2.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similar to inner peristome, projected all around, except the posterior part, where the two lateral sides are distinctly more projected than the anterior side.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Radial ribs.</bold> Rib density: 4–6 ribs per mm. Rib intensity: thick. Shape: slightly curved to single-humped. Inclination: prosoclin.</p><fig id="F45" orientation="portrait" position="float"><label>Figures 45.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> sp. n. BOR 5649. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g045"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Gua Bewah, Tasik Kenyir (<named-content content-type="dwc:verbatimCoordinates">4°51'3"N, 102°43'21"E</named-content>).</p><p>Distribution range. To date, this species is only known to occur at two neighbouring limestone outcrops, namely, Gua Bewah and Gua Taat at the Southern part of Tasik Kenyir (<xref ref-type="fig" rid="F18">Figure 18B</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Least concern. The only two limestone outcrops where this species was found are partially submerged in Southeast Asia’s largest man-made lake, Tasik Kenyir. Nonetheless, there is good forest cover around and on the limestone outcrops. Furthermore, these two hills are located in the Taman Negara Pahang, which is a prioritised protected area.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>From a conchological point of view, this species is related to <italic><named-content content-type="taxon-name">Plectostoma charasense</named-content></italic>. Both are thought to be affiliated with the <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species from North Borneo. <italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> has a high conical spire, thick and projected radial ribs, and a long detached tuba. These are the typical characteristics for dozens of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species in North Borneo. In addition, <italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma charasense</named-content></italic> live on the five lenticular limestone outcrops that are the easternmost outcrops of the Malay Peninsula, thus closest geographically to Borneo (<xref ref-type="fig" rid="F18">Figure 18B</xref>).</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_annandalei"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">annandalei</named-content></named-content></title><p><named-content content-type="taxon-authority">(Sykes, 1903)</named-content></p><p>http://species-id.net/wiki/Plectostoma_annandalei</p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma annandalei</named-content><named-content content-type="comment"> Sykes, 1903: 198, Plate 20 - figures 4 & 5 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma annandalei</named-content><named-content content-type="comment"> Sykes, <xref rid="B61" ref-type="bibr">Laidlaw (1928</xref>: 36).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma annandalei</named-content><named-content content-type="comment"> Sykes, <xref rid="B6" ref-type="bibr">van Benthem Jutting (1952</xref>: 41).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Not seen.</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma tenggekensis</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic> the general shell form, in terms of spire and tuba shape, but differs by having slightly convex whorl periphery and straight ribs.</p></sec><sec sec-type="treatment-Description"><title>Description</title><p>(estimated from figure in <xref rid="B117" ref-type="bibr">Sykes 1903</xref>). <bold>Apex.</bold> Shape: moderately convex.</p><p><bold>Spire.</bold> Height: 2 mm. Width: 1.3 mm. Number of whorls: 4 1/2. Apical spire shape: depressed conical. Basal spire shape: conical. Whorl periphery: moderately convex. Umbilicus: Open.</p><p><bold>Constriction.</bold> Unknown.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in right lateral view. Tuba whorl length in proportion to spire last whorl: approximately the same as the spire’s last whorl length. Proportion of tuba that attaches to spire: less than 1/2.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: same as inner peristome and uniformly projected all around, except the posterior part.</p><p><bold>Spiral lines.</bold> Unknown</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Radial ribs.</bold> Rib density: 5 ribs per mm. Rib intensity: unknown. Shape: straight. Inclination: orthoclin.</p></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. This species is only known from its type locality, Jalor (Biserat). The exact location was not described in the original publication of this species. From the collectors’ report (Annandale and Robinson 1913), we estimated the location and name of the limestone hill from a map in the report. This hill was named Bukit Bayu. Later, we estimated its coordinates from Google Earth (<named-content content-type="dwc:verbatimCoordinates">6°16'48"N, 101°13'35"E</named-content>) (<xref ref-type="fig" rid="F18">Figure 18B</xref>).</p><p>Distribution range. Unknown.</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Data Deficient.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>This species has not been seen or collected after the original description. The type specimens cannot be located. <xref rid="B117" ref-type="bibr">Sykes (1903)</xref> mentioned that it is similar to <italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic> but he did not explain explicitly in what way. This is not unexpected, as <italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic> was the only one <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> known from Peninsular Malaysia at the time, although more than 20 <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species had already been described from Borneo. Thus, one can assume that the statement made by Sykes was based on the geographical proximity.</p><p>Interestingly, <italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic> was described from a limestone hill located just 8 km from the location where <italic><named-content content-type="taxon-name">Plectostoma annandalei</named-content></italic> was found. In addition to the geographical proximity, the two neighbouring species are similar in several shell characteristics. The shell spires are very similar in terms of number of whorls, overall shape and size. On the other hand, the main difference between these two species is the tuba coiling direction (type 1 vs. type 2).</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_praeco"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">praeco</named-content></named-content></title><p><named-content content-type="taxon-authority">(van Benthem Jutting, 1961)</named-content></p><p>http://species-id.net/wiki/Plectostoma_praeco</p><p><xref ref-type="fig" rid="F46">Figure 46</xref><xref ref-type="app" rid="APP1">, Appendix 14</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma praeco</named-content><named-content content-type="comment"> van Benthem Jutting, 1961: 39, Plate 2 - figure 5 (original description).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: ZMA 136034(1) (Seen). Paratypes: ZMA 136041(>50) (Seen), ZMA 136042(>25) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>ZMA 162150(>100), ZMA 162151(>100), ZMA 162152(>100), ZMA 162153(>10).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma tenggekensis</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma annandalei</named-content></italic> the general shell form, in terms of spire and tuba shape, but differs by having slightly curved ribs.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: distinctly convex.</p><p><bold>Spire.</bold> Height: 2.0–2.2 mm. Width: 1.8–2.0 mm. Number of whorls: 4 1/4–4 5/8. Apical spire shape: depressed conical. Basal spire shape: conical. Whorl periphery: moderately to distinctly convex. Umbilicus: open.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in right lateral view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 5/8–7/8. Proportion of tuba that attaches to spire: more than 1/3 but less than 1/2.</p><pmc-comment>PageBreak</pmc-comment>
<p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similar to inner peristome, projected all around, except the posterior part, where the left lateral sides are slightly more projected than the anterior and right lateral side.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 6–7 ribs per mm. Rib intensity: thin. Shape: slightly curved. Inclination: moderately prosoclin.</p><fig id="F46" orientation="portrait" position="float"><label>Figures 46.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic> (van Benthem Jutting, 1961) ZMA 162150. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g046"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Batu Che Derani, Taman Negara Pahang (<named-content content-type="dwc:verbatimCoordinates">4°35'57"N, 102°25'57"E</named-content>).</p><p>Distribution range. <italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic> has been recorded from four other hills at upper Sungai Keniyam Kecil, which is in the vicinity of the type locality (<xref ref-type="fig" rid="F18">Figure 18B</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Least concern. All the limestone hills where <italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic> occurs, are located in the National Park.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>There is no genetic information for <italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic>. Morphological similarity suggests that <italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic> and <italic><named-content content-type="taxon-name">Plectostoma tenggekensis</named-content></italic> are closely related. Each of the two species has a narrow distribution range and they are separated from each other by at least 50 km.</p></sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">tenggekensis</named-content></named-content><named-content content-type="taxon-status">sp. n.</named-content></title><p>http://zoobank.org/38A095DD-552F-4F77-8774-1E834531352C</p><p>http://species-id.net/wiki/Plectostoma_tenggekensis</p><p><xref ref-type="fig" rid="F47">Figure 47</xref><xref ref-type="app" rid="APP1">, Appendix 14</xref></p><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: BOR 5650(1). Paratypes: V 13554(5), BOR 444(4), BOR 5596(>10).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma annandalei</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic> the general shell from, in terms of spire and tuba shape, but differs by having single-humped shaped ribs.</p><pmc-comment>PageBreak</pmc-comment>
</sec><sec sec-type="treatment-Etymology"><title>Etymology.</title><p>This species is named after its type locality–Bukit Tenggek, Kuantan, Pahang.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: moderately convex.</p><p><bold>Spire.</bold> Height: 1.6–1.7 mm. Width: 1.3–1.4 mm. Number of whorls: 4–4 1/4. Apical spire shape: depressed. Basal spire shape: conical. Whorl periphery: moderately convex. Umbilicus: open.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in right lateral view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 5/8 of the spire last whorl length. Proportion of tuba that attaches to spire: less than 1/3.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similar to inner peristome, projected all around, except the posterior part, where the left lateral sides are slightly more projected than the anterior and right lateral side.</p><p><bold>Spiral lines.</bold> Thick lines: absent. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 6–7 ribs per mm. Rib intensity: thin. Shape: slightly curved. Inclination: moderately prosoclin.</p><fig id="F47" orientation="portrait" position="float"><label>Figures 47.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma tenggekensis</named-content></italic> sp. n. BOR 5650. <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g047"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. Bukit Tenggek, Pahang (<named-content content-type="dwc:verbatimCoordinates">4°0'51"N, 103°9'34"E</named-content>).</p><p>Distribution range. Endemic to type locality (<xref ref-type="fig" rid="F18">Figure 18B</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Critically Endangered (B2ab(iii)+C2a(i) ver. 10.1). The entire Bukit Tenggek will be gone by 2014 because of quarrying activity.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>See discussion under <italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic>.</p><pmc-comment>PageBreak</pmc-comment>
</sec></sec><sec sec-type="taxon-treatment"><title><named-content content-type="taxon-name" xlink:href="http://species-id.net/wiki/Plectostoma_laidlawi"><named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">laidlawi</named-content></named-content></title><p><named-content content-type="taxon-authority">(Sykes, 1902)</named-content></p><p>http://species-id.net/wiki/Plectostoma_laidlawi</p><p><xref ref-type="fig" rid="F17">Figures 17O</xref><xref ref-type="fig" rid="F48">, 48</xref><xref ref-type="fig" rid="F49">, 49</xref><xref ref-type="app" rid="APP1">, Appendix 15</xref></p><list list-type="simple" list-content="nomenclature-citation-list"><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma laidlawi</named-content><named-content content-type="comment"> Sykes, 1902a: 22 (original description).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma laidlawi</named-content><named-content content-type="comment"> Sykes, <xref rid="B116" ref-type="bibr">Sykes (1902b</xref>: Plate 3 - figures 13 & 14).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma laidlawi</named-content><named-content content-type="comment"> Sykes, <xref rid="B73" ref-type="bibr">von Moellendorff (1902</xref>: 143).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma laidlawi</named-content><named-content content-type="comment"> Sykes, <xref rid="B61" ref-type="bibr">Laidlaw (1928</xref>: 36).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma laidlawi</named-content><named-content content-type="comment"> Sykes, <xref rid="B6" ref-type="bibr">van Benthem Jutting (1952</xref>: 41).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma laidlawi</named-content><named-content content-type="comment"> Sykes, <xref rid="B55" ref-type="bibr">Ito et al. (2009</xref>: 58).</named-content></p></list-item><list-item><p content-type="nomenclature-citation"><named-content content-type="taxon-name">Opisthostoma laidlawi</named-content><named-content content-type="comment"> Sykes, <xref rid="B94" ref-type="bibr">Sasaki (2010</xref>: 126, figure 2.7B).</named-content></p></list-item></list><sec sec-type="treatment-Type material"><title>Type material.</title><p>Holotype: Not Seen. Paratype: ZMA 136014(1) (Seen).</p></sec><sec sec-type="treatment-Other examined materials"><title>Other examined materials.</title><p>BMNH C_ACC1825(1), BOR 5510(>25), BOR 5571(2), RMNH 156268(1), ZMA 162095(2), ZMA 162097(>25), ZMA 162099(>10), ZMA 162100(>25), ZMA 162096(1), V5558(1), V 7878(3), V 8345(>100), V 8476(>25), V 8670(>25), V 8693(>50), V 8950(>50), V 9044(>25), V 9109(>25), V 9125(>25), V 9359(6).</p></sec><sec sec-type="treatment-Diagnosis"><title>Diagnosis.</title><p>Shares with <italic><named-content content-type="taxon-name">Plectostoma annandalei</named-content></italic>, <italic><named-content content-type="taxon-name">Plectostoma tenggekensis</named-content></italic>, and <italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic> the general shell form in terms of spire and tuba shape, but differs by having a distinctly convex whorl periphery and straight ribs.</p></sec><sec sec-type="treatment-Description"><title>Description.</title><p><bold>Apex.</bold> Shape: moderately to distinctly convex.</p><p><bold>Spire.</bold> Height: 1.6–2.3 mm. Width: 1.3–1.8 mm. Number of whorls: 3 5/8–4. Apical spire shape: depressed conical. Basal spire shape: conical to ovoid. Whorl periphery: distinctly convex. Umbilicus: open.</p><p><bold>Constriction.</bold> Parietal teeth: two. Basal teeth: none.</p><p><bold>Tuba.</bold> Coiling direction: type 2 and aperture visible in right lateral view. Tuba whorl length in proportion to spire last whorl: <italic>ca.</italic> 5/8–7/8. Proportion of tuba that attaches to spire: varies from completely attached to as much as half of the tuba detached from spire.</p><p><bold>Aperture and peristome.</bold> Peristome: double peristomes. Outer peristome shape: similar to inner peristome, projected all around, except the posterior part, where the two lateral sides are slightly more projected than the anterior side or left lateral sides slightly more projected than the anterior and right lateral side.</p><p><bold>Spiral lines.</bold> Thick lines: present. Thin lines: present.</p><p><bold>Radial ribs.</bold> Rib density: 5–6 ribs per mm. Rib intensity: thin. Shape: straight to slightly curved. Inclination: moderately prosoclin.</p><fig id="F48" orientation="portrait" position="float"><label>Figures 48.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic> (Sykes, 1902) (Form BOR 5510). <bold>A</bold> frontal view <bold>B</bold> left lateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g048"></graphic></fig><fig id="F49" orientation="portrait" position="float"><label>Figures 49.</label><caption><p><italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic> (Sykes, 1902) (Form BOR 5571). <bold>A</bold> frontal view <bold>B</bold> leftlateral view <bold>C</bold> back view <bold>D</bold> right lateral view <bold>E</bold> top view <bold>F</bold> bottom view <bold>G</bold> parietal part of constriction inner whorl <bold>H</bold> basal part of constriction inner whorl. Scale bar = 1 mm (for <bold>A–F</bold>).</p></caption><graphic xlink:href="zookeys-393-001-g049"></graphic></fig></sec><sec sec-type="treatment-Distribution"><title>Distribution.</title><p>Type locality. “Kelantan, Malay Peninsula”, collected by J. Waterstradt (Sykes 1902). According to <xref rid="B125" ref-type="bibr">Waterstradt’s (1902)</xref> itinerary, he visited one of the limestone hills at Kampung Pulai. The Pulai Princess cave (<named-content content-type="dwc:verbatimCoordinates">4°47'38"N, 101°56'31"E</named-content>) fits perfectly with Waterstradt’s descriptions on the local population, temples and hill’s topography; he also described that he and his collectors sampled <pmc-comment>PageBreak</pmc-comment><pmc-comment>PageBreak</pmc-comment>shells here (<xref rid="B125" ref-type="bibr">Waterstradt 1902</xref>: 9–10). We conclude that Pulai Princess Cave must be the type locality of <italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic>. However, a recent survey at the highly degraded Bukit Pulai Princess cave failed to retrieve any shells of <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic>.</p><p>Distribution range. This species occurs on the limestone hills along Berok River and Nenggiri River (<xref ref-type="fig" rid="F18">Figure 18B</xref>).</p></sec><sec sec-type="treatment-Conservation status"><title>Conservation status.</title><p>Vulnerable (D2 ver. 10.1). Living populations were present on only two limestone hills during a survey in May 2011. All the limestone hills for which this species is known are surrounded by degraded forest and oil palm plantation.</p></sec><sec sec-type="treatment-Discussion"><title>Discussion.</title><p>The morphology of this species is quite variable in shell size and degree of attachment of the tuba to the spire. The distribution range partially overlaps with that of <italic><named-content content-type="taxon-name">Plectostoma davisoni</named-content></italic>. It is possible that the species actually consists of two or more cryptic species, and thus more genetic data are needed.</p></sec></sec></sec><sec><title>Author contributions</title><p>Conceived and designed the experiments: Thor-Seng Liew. Performed the experiments: Thor-Seng Liew. Analyzed the data: Thor-Seng Liew, Jaap Jan Vermeulen. Contributed reagents/materials/analysis tools: Thor-Seng Liew, Jaap Jan Vermeulen, Mohammad Effendi bin Marzuki, Menno Schilthuizen. Wrote the paper: Thor-Seng Liew, Jaap Jan Vermeulen, Menno Schilthuizen, Mohammad Effendi bin Marzuki.</p></sec><sec sec-type="supplementary-material"><title>Supplementary Material</title><supplementary-material id="zookeys.393.6717-treatment1" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment1.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment2" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">dindingensis</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment2.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment3" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">mengaburensis</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment3.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment4" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">sinyumensis</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment4.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment5" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">umbilicatum</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment5.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment6" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">siphonostomum</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment6.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment7" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">panhai</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment7.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment8" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">christae</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment8.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment9" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">crassipupa</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment9.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment10" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">tonkinianum</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment10.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment11" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">whitteni</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment11.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment12" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">sciaphilum</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment12.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment13" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">senex</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment13.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment14" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">turriforme</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment14.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment15" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">palinhelix</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment15.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment16" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">retrovertens</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment16.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment17" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">ikanensis</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment17.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment18" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">kayiani</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment18.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment19" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">davisoni</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment19.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment20" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">relauensis</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment20.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment21" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">kakiense</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment21.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment22" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">kubuensis</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment22.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment23" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">laemodes</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment23.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment24" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">salpidomon</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment24.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment25" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">kitteli</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment25.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment26" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">klongsangensis</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment26.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment27" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">charasense</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment27.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment28" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">tohchinyawi</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment28.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment29" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">annandalei</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment29.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment30" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">praeco</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment30.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment31" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">tenggekensis</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment31.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material><supplementary-material id="zookeys.393.6717-treatment32" content-type="local-data"><caption><title>XML Treatment for
<named-content content-type="genus">Plectostoma</named-content><named-content content-type="species">laidlawi</named-content></title></caption><media xlink:href="zookeys.393.6717-treatment32.xml" mimetype="text" mime-subtype="xml"></media></supplementary-material></sec></body><back><ack><title>Acknowledgements</title><p>We thank Geoffrey Davison, Reuben Clements, Yansen Chen, Wim Maassen, and A.J. de Winter for help and fruitful discussion. Petherine Jimbau (BOR), Jonathan Ablett and Fred Naggs (BMNH), and Jeroen Goud, Rob Moolenbeek and Bram van der Bijl (Naturalis) made the museum collections accessible for TSL. TSL thanks Universiti Malaysia Sabah and the Ministry of Higher Education (Malaysia) for support. TSL thanks Economic Planning Unit, Prime Minister’s Department (UPE: 40/200/19/2524), State Planning Unit, Chief Minister’s Department, Sarawak ((47) UPN/S/G1/I/10.1 Vol.27), Forest Department Sarawak (Research Permit NPW.907.4.4(V)-19; Park Permit No. 07/2010; Export Permit No. 09003), and Department of Wildlife and National Parks (Mr. Burhanuddin Mohd Nor). This project is supported by ALW-NWO (819.01.012) and EOL Rubenstein Fellowship (2010) to TSL. We thank Frank Koehler, Zoltán Fehér and an anonymous reviewer for useful comments which improved this paper.</p><pmc-comment>PageBreak</pmc-comment>
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</ref-list><app-group><app id="APP1"><title>Appendix</title><p>(<ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.6084/m9.figshare.830412">http://dx.doi.org/10.6084/m9.figshare.830412</ext-link>)</p><p><bold>Appendix 1.</bold> Video tutorials of cybertaxonomy procedures.</p><p><bold>Appendix 2.</bold> Details for collection data.</p><p><bold>Appendix 3.</bold> Spiral striation profile on the shell spire of 29 <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species.</p><p><bold>Appendix 4.</bold> Radial rib spacing profile on the shell spire of 29 <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species.</p><p><bold>Appendix 5.</bold> A KML file consisting of collection data, image links and distribution data.</p><p><bold>Appendix 6.</bold> A Blender file consisting of 17 shell 3D models of 2 species, Part 1 of 10.</p><p><italic><named-content content-type="taxon-name">Plectostoma christae</named-content></italic> (Maassen, 2001)</p><p>- V 12702 (1), BOR 5505 (1), BOR 5506 (2), BOR 5572(1), V 9285 (2).</p><p><italic><named-content content-type="taxon-name">Plectostoma crassipupa</named-content></italic> (van Benthem Jutting, 1952)</p><p>- BOR 5512 (1), V 8392 (2), V 9097 (3), V 9326 (2), V 9353 (2).</p><p><bold>Appendix 7.</bold> A Blender file consisting of 18 shell 3D models of 6 species, Part 2 of 10.</p><p><italic><named-content content-type="taxon-name">Plectostoma dindingensis</named-content></italic> sp. n.</p><p>- BOR 5612 (3), BOR 5642 (1).</p><p><italic><named-content content-type="taxon-name">Plectostoma mengaburensis</named-content></italic> sp. n.</p><p>- BOR 5643 (1), V 8822 (1).</p><p><italic><named-content content-type="taxon-name">Plectostoma panhai</named-content></italic> (Maassen, 2001)</p><p>- ZMA 138438 (1), RMNH 81810 (2).</p><p><italic><named-content content-type="taxon-name">Plectostoma sciaphilum</named-content></italic> (van Benthem Jutting, 1952)</p><p>- ZMA 136050 (3).</p><p><italic><named-content content-type="taxon-name">Plectostoma senex</named-content></italic> (van Benthem Jutting, 1952)</p><p>- BOR 5628 (2), BOR 5603 (1).</p><p><italic><named-content content-type="taxon-name">Plectostoma sinyumensis</named-content></italic> (Maassen, 2001)</p><p>- BOR 5623 (2), BOR 5537 (1).</p><p><bold>Appendix 8.</bold> A Blender file consisting of 14 shell 3D models of 5 species, Part 3 of 10.</p><p><italic><named-content content-type="taxon-name">Plectostoma siphonostomum</named-content></italic> (van Benthem Jutting, 1952)</p><p>- BOR 5521 (1), BOR 5557 (1).</p><p><italic><named-content content-type="taxon-name">Plectostoma whitteni</named-content></italic> sp. n.</p><p>- BOR 5644 (1), V 8885 (2).</p><p><italic><named-content content-type="taxon-name">Plectostoma tonkinianum</named-content></italic> (Dautzenberg & Fischer, 1905)</p><p>- V 11502 (1), V 9940 (1).</p><p><italic><named-content content-type="taxon-name">Plectostoma turriforme</named-content></italic> (van Benthem Jutting, 1952)</p><p>- BOR 5509 (1), ZMA 136069 (2).</p><pmc-comment>PageBreak</pmc-comment>
<p><italic><named-content content-type="taxon-name">Plectostoma umbilicatum</named-content></italic> (van Benthem Jutting, 1952)</p><p>- BOR 5503 (2), BOR 5625 (2).</p><p><bold>Appendix 9.</bold> A Blender file consisting of 19 shell 3D models of 1 species, Part 4 of 10.</p><p><italic><named-content content-type="taxon-name">Plectostoma davisoni</named-content></italic> sp. n.</p><p>- BOR 5626 (2), BOR 5641 (2), BOR 5646 (1), V 14242 (1), V 8265 (3), V 8652 (3), V 9206 (1), V 9243 (2), ZMA 162071 (4).</p><p><bold>Appendix 10.</bold> A Blender file consisting of 19 shell 3D models of 1 species, Part 5 of 10.</p><p><italic><named-content content-type="taxon-name">Plectostoma davisoni</named-content></italic> sp. n.</p><p>- V 9340 (2), ZMA 162069 (5), ZMA 162070 (3), ZMA 162146 (3), ZMA 162147 (6).</p><p><bold>Appendix 11.</bold> A Blender file consisting of 18 shell 3D models of 4 species, Part 6 of 10.</p><p><italic><named-content content-type="taxon-name">Plectostoma kayiani</named-content></italic> sp. n.</p><p>- RMNH 330803 (1), V 8883 (2)</p><p><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n. Form BOR 5507</p><p>- BOR 5645 (1), V 9320 (4), V9446 (4).</p><p><italic><named-content content-type="taxon-name">Plectostoma kakiense</named-content></italic> (Tomlin, 1948)</p><p>- BOR 5516 (1), BOR 5517 (2).</p><p><italic><named-content content-type="taxon-name">Plectostoma kitteli</named-content></italic> (Maassen, 2002)</p><p>- RMNH 92956 (2), V 12697 (1).</p><p><bold>Appendix 12.</bold> A Blender file consisting of 14 shell 3D models of 3 species, Part 7 of 10.</p><p><italic><named-content content-type="taxon-name">Plectostoma kubuensis</named-content></italic> sp. n.</p><p>- BOR 5518 (1), BOR 5519 (3), BOR 5648 (1).</p><p><italic><named-content content-type="taxon-name">Plectostoma laemodes</named-content></italic> (van Benthem Jutting, 1961)</p><p>- RMNH 156267 (3), V 8490 (2), ZMA 136011 (1).</p><p><italic><named-content content-type="taxon-name">Plectostoma palinhelix</named-content></italic> (van Benthem Jutting, 1952)</p><p>- BOR 5520 (1), V 5104 (2).</p><p><bold>Appendix 13.</bold> A Blender file consisting of 11 shell 3D models of 3 species, Part 8 of 10.</p><p><italic><named-content content-type="taxon-name">Plectostoma relauensis</named-content></italic> sp. n.</p><p>- BOR 5511 (2), BOR 5647 (1), V 8196 (2).</p><p><italic><named-content content-type="taxon-name">Plectostoma retrovertens</named-content></italic> (Tomlin, 1938)</p><p>- V 5142 (2), BOR 5559 (1).</p><p><italic><named-content content-type="taxon-name">Plectostoma salpidomon</named-content></italic> (van Benthem Jutting, 1952)</p><p>- BOR 5539 (1), V 8658 (2).</p><p><bold>Appendix 14.</bold> A Blender file consisting of 13 shell 3D models of 5 species, Part 9 of 10.</p><p><italic><named-content content-type="taxon-name">Plectostoma tohchinyawi</named-content></italic> sp. n.</p><p>- BOR 5533 (2), BOR 5649 (1), V 8811 (2).</p><pmc-comment>PageBreak</pmc-comment>
<p><italic><named-content content-type="taxon-name">Plectostoma charasense</named-content></italic> (Tomlin, 1948)</p><p>- ZMA 162063 (1), ZMA 162064 (2).</p><p><italic><named-content content-type="taxon-name">Plectostoma praeco</named-content></italic> van Benthem Jutting, 1961</p><p>- ZMA 162150 (1), ZMA 162151 (1), ZMA 162152 (1).</p><p><italic><named-content content-type="taxon-name">Plectostoma ikanensis</named-content></italic> sp. n. Form BOR 5504</p><p>- BOR 5504 (1).</p><p><italic><named-content content-type="taxon-name">Plectostoma tenggekensis</named-content></italic> sp. n.</p><p>- BOR 5596 (1), BOR 5649 (1).</p><p><bold>Appendix 15.</bold> A Blender file consisting of 10 shell 3D models of 1 species, Part 10 of 10.</p><p><italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic> (Sykes, 1902) Form BOR 5510</p><p>- BOR 5510 (1), V 5558 (1), ZMA 136014 (1).</p><p><italic><named-content content-type="taxon-name">Plectostoma laidlawi</named-content></italic> (Sykes, 1902) Form BOR 5571</p><p>- BOR 5571 (1), V 7878 (2), ZMA 162099 (4).</p><p><bold>Appendix 16.</bold> 51 COI sequences for 19 <italic><named-content content-type="taxon-name">Plectostoma</named-content></italic> species in FASTA format. Each sequence is labeled with BOLD process ID, GenBank accession no, sample ID and species name.</p><p><bold>Appendix 17.</bold> DNA sequence alignment for phylogenetic analysis.</p><p><bold>Appendix 18.</bold> Shell colouration of <italic><named-content content-type="taxon-name">Opisthostoma</named-content></italic> species.</p></app></app-group></back></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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