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<mods:title id="A3BBCDC31E4179A74958D039EBFC6F69">A review of the land snail genus Alycaeus (Gastropoda, Alycaeidae) in Peninsular Malaysia</mods:title>
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<subSubSection id="5C9F0BD4F00C8A7F681DE78183C48480" pageId="30" pageNumber="31" type="nomenclature">
<paragraph id="98530CA7514DD8CAC7C19012E67A4B37" pageId="30" pageNumber="31">
<taxonomicName id="EF11898DFA1E14A6C1658902D2F7ED69" ID-CoL="7PYGR" authority="Sykes, 1902" authorityName="Sykes" authorityYear="1902" class="Gastropoda" family="Alycaeidae" genus="Alycaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Alycaeus kelantanensis" order="Architaenioglossa" pageId="30" pageNumber="31" phylum="Mollusca" rank="species" species="kelantanensis">Alycaeus kelantanensis Sykes, 1902</taxonomicName>
Figures 7
<normalizedToken id="1743C0E9205276163FB7C637DBF05F99" originalValue="PQ">P-Q</normalizedToken>
, 21, 31K
</paragraph>
</subSubSection>
<subSubSection id="5F07E25B313DA30FF5C169FA8C16BB6A" pageId="31" pageNumber="32" type="reference_group">
<paragraph id="39CDAA677A5A4371C0D5DBAD244AE559" pageId="31" pageNumber="32">
<taxonomicName id="0813E2997DFAA5036744B38B774F3F05" class="Gastropoda" family="Alycaeidae" genus="Alycaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Alycaeus kelantanense" order="Architaenioglossa" pageId="31" pageNumber="32" phylum="Mollusca" rank="species" species="kelantanense">
<pageBreakToken id="61FD48E48DD35CAB5F40B72F90BF9C3D" pageId="31" pageNumber="32" start="start">Alycaeus</pageBreakToken>
kelantanense
</taxonomicName>
Sykes, 1902: 60-62, Figure 11-12.
</paragraph>
<paragraph id="00241A1C9CE76CA5AA533F7098941D87" pageId="31" pageNumber="32">
<taxonomicName id="C298C56098CE441901742EF52FA24384" class="Gastropoda" family="Alycaeidae" genus="Alycaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Alycaeus kelantanensis" order="Architaenioglossa" pageId="31" pageNumber="32" phylum="Mollusca" rank="species" species="kelantanensis">Alycaeus kelantanensis</taxonomicName>
:
<bibRefCitation id="30099B359F07B6F1ED38C1C397672B90" author="Moellendorff, von OF" journalOrPublisher="Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft" pageId="52" pageNumber="53" pagination="135 - 149" title="Binnenmollusken aus Hinterindien 1. Landschnecken von Kelantan, Ostkueste der Halbinsel Malacca." url="http://www.biodiversitylibrary.org/part/145333/summary" volume="34" year="1902">
<normalizedToken id="E56415C73527117C34556A5D52C072EA" originalValue="Möllendorff">Moellendorff</normalizedToken>
1902
</bibRefCitation>
: 135-149;
<bibRefCitation id="384029446546973C0C806A539560CB91" author="Laidlaw, FF" journalOrPublisher="Journal of the Malayan Branch of the Royal Asiatic Society" pageId="51" pageNumber="52" pagination="25 - 37" title="A list of the Land and Fresh-water Mollusca of the Malay Peninsula with notes." url="http://www.jstor.org/stable/41559818" volume="6" year="1928">Laidlaw 1928</bibRefCitation>
: 35;
<bibRefCitation id="AB4542C461611BD97668120C3CFD30D1" author="Berry, AJ" journalOrPublisher="Bulletin of the National Museum Singapore" pageId="49" pageNumber="50" pagination="27 - 30" title="A collection of land mollusca from limestone in Ulu Kelantan." url="http://blsea.nus.edu.sg/en/persistent/584163f8-61e7-4329-8b04-d62e96e76796" volume="33" year="1965">Berry 1965</bibRefCitation>
: 28;
<bibRefCitation id="E3F345B20A5E85762E23931C43BCA5D9" author="Schilthuizen, M" journalOrPublisher="Malayan Nature Journal" pageId="53" pageNumber="54" pagination="351 - 354" title="A note on the ecology of West-Malaysian calcicolous snails (Mollusca: Gastropoda: Diplommatinidae, Cyclophoridae, Vertiginidae, Streptaxidae)." volume="53" year="1999">Schilthuizen et al. 1999</bibRefCitation>
: 353;
<bibRefCitation id="C2506D4F7B63AE2F78829E93CE80559C" pageId="31" pageNumber="32">Maassen 2001</bibRefCitation>
: 22;
<bibRefCitation id="4CC24EF24A61056BAA6B0A0CCF1D1ABA" author="Clements, R" journalOrPublisher="Biological Conservation" pageId="49" pageNumber="50" pagination="2751 - 2764" title="Using biogeographical patterns of endemic land snails to improve conservation planning for limestone karsts." url="https://doi.org/10.1016/j.biocon.2008.08.011" volume="141" year="2008">Clements et al. 2008</bibRefCitation>
: 2760;
<bibRefCitation id="590687C1ADBA75C6C18314A31B56E75A" author="Liew, TS" editor="Che, Omar I" journalOrPublisher="The Bulletin of the Raffles Museum" pageId="51" pageNumber="52" title="Penerbit Universiti Malaysia Kelantan, Kota Bharu" url="https://doi.org/10.13140/RG.2.1.4311.5364" year="2010">Liew 2010</bibRefCitation>
: 101;
<bibRefCitation id="91CE783095DEE217BC0F204882551293" author="Tarruella, A" journalOrPublisher="Spira" pageId="53" pageNumber="54" pagination="71 - 76" title="Listado taxonomico ilustrado de la familia Cyclophoridae J. E. Gray, 1847 (Mollusca: Gastropoda): Parte 2. La subfamilia Alycaeinae J. E. Gray, 1850." url="http://www.molluscat.com/assets/spira_4_12_7.pdf" volume="4" year="2011">
Tarruella and
<normalizedToken id="AC5F70FE5BAECA13D8D62DA723D8B5DF" originalValue="Domènech">Domenech</normalizedToken>
2011
</bibRefCitation>
: 72, figure 2C.
</paragraph>
</subSubSection>
<subSubSection id="6E202A5F9FD22682E5029C60658C186C" pageId="31" pageNumber="32" type="type locality">
<paragraph id="EF5CC04205B8CAA489471EBC9A129975" pageId="31" pageNumber="32">Type locality.</paragraph>
<paragraph id="3B0006A6F240D1D1CB38E7E59778A6E6" pageId="31" pageNumber="32">Kelantan, Malay Peninsula.</paragraph>
</subSubSection>
<subSubSection id="651B0A02DC145664894F4FB83903A863" pageId="31" pageNumber="32" type="type material">
<paragraph id="0077CA23CE32DD298D08D75C966A6847" pageId="31" pageNumber="32">Type material.</paragraph>
<paragraph id="497369A7C2408DCF76F6DD86B901138A" pageId="31" pageNumber="32">Syntype. Kelantan: RMNH 153594/1 (Seen).</paragraph>
</subSubSection>
<subSubSection id="0095764E71D0BD7F1D0D321632533A36" lastPageId="32" lastPageNumber="33" pageId="31" pageNumber="32" type="materials_examined">
<paragraph id="49CB271F439C787650240606D93D1D77" pageId="31" pageNumber="32">Other examined materials.</paragraph>
<paragraph id="256BAD6239AFF7C23DA2F4AA326FD839" lastPageId="32" lastPageNumber="33" pageId="31" pageNumber="32">
KTN 01 Gunung Reng, Kelantan: BOR/MOL 6837/1. KTN 45 unnamed hill at Kampung Bayu, Kelantan: BOR/MOL 6236/6, BOR/MOL 12974/1. KTN 66 Batu Tongkat, Kelantan: ZRC 1975.2.21.838-875/44. KTN 77 Bukit Sejuk, Kelantan: BOR/MOL 6199/3, BOR/MOL 6231/1. KTN 95 Gua Panjang, Kelantan: BOR/MOL 6200/6, BOR/MOL 6829/1, BOR/MOL 8329/13, BOR/MOL 8330/5. KTN 109 part of Gua Panjang, Kelantan: BOR/MOL 12973/1. KTN 134 Gua Batu Boh, Kelantan: BOR/MOL 6828/1, BOR/MOL 8328/3, BOR/MOL 8409/3. KTN 135 Gua Serai, Kelantan: BOR/MOL 242/1. KTN 136 Gua Musang, Kelantan: ZRC 1975.2.21.250-349/102, ZRC 1975.2.24.452-476/25, ZRC 1975.2.24.309-351/43. KTN 139 unnamed hill, Kelantan: BOR/MOL 6197/5. KTN 148 'Hill 001', Kelantan: BOR/MOL 12976/4. KTN 149 unnamed hill, Kelantan: BOR/MOL 6833/1, BOR/MOL 8333/11. KTN 176 Batu Lesong, Kelantan: BOR/MOL 6830/1, BOR/MOL 8331/7, BOR/MOL 8332/8. Mykarst-168, Kelantan: BOR/MOL 6189/2, BOR/MOL 6838/2. Mykarst-186 Gua Madu, Kelantan: BOR/MOL 6234/1, BOR/MOL 6825/1, BOR/MOL 8327/7, ZRC 1975.2.21.350-430/91. Limestone outcrop 3 km north of Felda Ciku 5 settlement,
<pageBreakToken id="57D5F56C05893004E2682349496A2F58" pageId="32" pageNumber="33" start="start">Kelantan</pageBreakToken>
: BOR/MOL 6238/3. Bukit Gua Peraling, Kelantan: ZRC 2.24.440-451/12. Kelantan: RMNH 153593/3, ZMA 33451/2. PHG 40 Gua Layang, Pahang: BOR/MOL 6822/1, BOR/MOL 8325/2, BOR/MOL 8326/7. PHG 114
<normalizedToken id="107C7AD0979C9DBE35C925711AF9BCC2" originalValue="Boomerang">'Boomerang'</normalizedToken>
Hill, Pahang: BOR/MOL 6188/2. Unnamed limestone hill north of Jalan Lipis-Gua Musang Lama, Felda Chegar Perah I and II, Pahang: BOR/MOL 6233/1, BOR/MOL 6814/1, BOR/MOL 8321/5, BOR/MOL 8322/3.
</paragraph>
</subSubSection>
<subSubSection id="D4DD743CA733E65543EAB409FEEE1670" pageId="32" pageNumber="33" type="description">
<paragraph id="03D8DEF83C4CF832647E092B11E13631" pageId="32" pageNumber="33">Description.</paragraph>
<paragraph id="7167E92EE3FFC38827DE959E67A00151" pageId="32" pageNumber="33">Protoconch. Smooth.</paragraph>
<paragraph id="C3F766F6DCFEDBC79CDD37E958D22515" pageId="32" pageNumber="33">Shell shape. Conical. Shell height: 3.33-5.04 mm. Shell width: 3.36-5.04 mm.</paragraph>
<paragraph id="07E96FDE425D791461535C63079F6260" pageId="32" pageNumber="33">Spire. Spire height: 1.08-1.76 mm. Spire width: 1.40-2.25 mm. Number of whorls: up to 4. Spire shape: oblong conical. Whorl periphery rounded. Umbilicus open.</paragraph>
<paragraph id="A8805FB60C734E8B153D63C1046B1C66" pageId="32" pageNumber="33">Whorl constriction. At about 3 ⅝ whorls posterior of protoconch.</paragraph>
<paragraph id="AA70D21F3AA5B49312090AFEC5F829E8" pageId="32" pageNumber="33">Breathing tube. Length: 0.38-0.67 mm.</paragraph>
<paragraph id="FD32F6034B265448DB23D189BA02B609" pageId="32" pageNumber="33">Aperture and peristome. Aperture circular, moderately expanded to very expanded. Aperture height: 1.82-2.73 mm. Aperture width: 1.69-2.46 mm. Peristome double, thickened, folded posteriorly into a wing-like upper palatal structure, notched at suture. Peristome orientation 26-34° oblique with respect to the coiling axis.</paragraph>
<paragraph id="F801D5CB65ADC0FBFCAE9AD9F3533DB6" pageId="32" pageNumber="33">Spiral lines. Distinct. Regularly spaced. Approximately 15-28 lines per 1 mm.</paragraph>
<paragraph id="578B717BDE1FACC14D6E0847BA3FF6A3" pageId="32" pageNumber="33">Radial ribs running anterior of breathing tube. Radial ribs pronounced, evenly spaced. Approximately 12-18 ribs per 1 mm.</paragraph>
<paragraph id="5160C79AF42DA519C8EF484084A22CD7" pageId="32" pageNumber="33">Radial ribs running perpendicular to breathing tube. Radial ribs pronounced, white and thicker than those anterior of breathing tube, evenly spaced. Approximately 12-29 ribs per 1 mm.</paragraph>
<paragraph id="920C29EBF4E384A876DA57CE1A4763D3" pageId="32" pageNumber="33">Radial ribs running posterior of breathing tube. Radial ribs absent immediately posterior of constriction, becoming pronounced and evenly spaced at the middle, absent again towards the aperture. Approximately 11-20 ribs per 1 mm.</paragraph>
<paragraph id="3AE17B01232E0707147EF374EAC84849" pageId="32" pageNumber="33">Operculum. Concave, rounded. Exterior covered with thin calcareous layer, with very indistinct appressed multilamellae. Interior covered with proteinaceous coating, smooth, with indistinct mamilla.</paragraph>
<paragraph id="348731A50C572BA07590B459D9E3BF6A" pageId="32" pageNumber="33">Shell colour. Whorls white throughout. First 3 whorls occasionally yellow.</paragraph>
<paragraph id="6B2A3F092D7054ABD11C16CD0740A0C0" pageId="32" pageNumber="33">Living animal. Body cream-white. Head pinkish. Tentacles yellow, with red tips.</paragraph>
</subSubSection>
<subSubSection id="C96C34DE0EB24F316FAD671F56AEE7A7" pageId="32" pageNumber="33" type="habitat">
<paragraph id="0702482A39D6B12F6FA6D1CCBDCF6954" pageId="32" pageNumber="33">Habitat and ecology.</paragraph>
<paragraph id="A00B0E07B05E3AC494CF13F69C3E5C71" pageId="32" pageNumber="33">Lives in rock crevices and limestone solution holes. In moist, shady forests. Occur on both limestone and non-limestone areas.</paragraph>
</subSubSection>
<subSubSection id="8A170F24C6D1277A581B893DA3253F48" pageId="32" pageNumber="33" type="distribution">
<paragraph id="7BE58F2689EB9C9DAC9B93E4114BE98F" pageId="32" pageNumber="33">Distribution range.</paragraph>
<paragraph id="DCCF24D912C6A1AB2EA99C4B488ED6B8" pageId="32" pageNumber="33">Widespread across southern Kelantan.</paragraph>
</subSubSection>
<subSubSection id="B0F300316EF24FD4B24924BE01DCF5E4" pageId="32" pageNumber="33" type="differential diagnosis">
<paragraph id="1708FB28017E259265B6AE9BA4978C5B" pageId="32" pageNumber="33">Differential diagnosis.</paragraph>
<paragraph id="2CEFD2678202AB52FD2E6DC48672F7A8" pageId="32" pageNumber="33">
<taxonomicName id="6FED22C8FE8BD62E9CE9076B18BD5C3A" class="Gastropoda" family="Alycaeidae" genus="Alycaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Alycaeus kelantanensis" order="Architaenioglossa" pageId="32" pageNumber="33" phylum="Mollusca" rank="species" species="kelantanensis">Alycaeus kelantanensis</taxonomicName>
varies in shell size and whorl convexity but can nevertheless be distinguished from other similar sized
<taxonomicName id="642AD4DDE9C59EA35AC3C2EAA1FCBD67" class="Gastropoda" family="Alycaeidae" genus="Alycaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Alycaeus" order="Architaenioglossa" pageId="32" pageNumber="33" phylum="Mollusca" rank="genus">Alycaeus</taxonomicName>
by its rather angular and winged double peristome, pronounced and widely-spaced radial ribs and spiral lines, as well as the red-tipped yellow tentacles of its soft body.
<taxonomicName id="4545EA35179E5C1A0B87FBF99EC3D687" class="Gastropoda" family="Alycaeidae" genus="Alycaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Alycaeus kelantanensis" order="Architaenioglossa" pageId="32" pageNumber="33" phylum="Mollusca" rank="species" species="kelantanensis">Alycaeus kelantanensis</taxonomicName>
is similar to
<taxonomicName id="D4405FA5055C070A3283CB55D7AC37AB" lsidName="A. clementsi" pageId="32" pageNumber="33" rank="species" species="clementsi">A. clementsi</taxonomicName>
sp. n. in shell colour but differs in being consistently smaller shelled (smaller by about 2.13 mm in shell height, 2.32 mm in shell width), having smooth operculum exterior and a more angular double peristome.
</paragraph>
</subSubSection>
<subSubSection id="6FF602D7D7D7548C3AE8A3B828AFBBD0" lastPageId="33" lastPageNumber="34" pageId="32" pageNumber="33" type="discussion">
<paragraph id="C0139A17EDD9A17DA23C80AAA8FE13DF" pageId="32" pageNumber="33">Discussion.</paragraph>
<paragraph id="A578321D496F3A39E85F0323F538875D" lastPageId="33" lastPageNumber="34" pageId="32" pageNumber="33">
The type specimen for
<taxonomicName id="252ED008F2A7AD1B16AA27B180AF1DEC" lsidName="A. kelantanensis" pageId="32" pageNumber="33" rank="species" species="kelantanensis">A. kelantanensis</taxonomicName>
was collected by John Waterstradt, likely from the Pulai Princess Cave hill (
<geoCoordinate id="5592F35035CD6C38BF8CD6778830ED38" direction="north" orientation="latitude" precision="15" value="4.793889">4°47'38&quot;N</geoCoordinate>
,
<geoCoordinate id="D5DE698F319EFB26EA86138C92D312C1" direction="east" orientation="longitude" precision="15" value="101.94195">101°56'31&quot;E</geoCoordinate>
), south of Gua Musang, Kelantan (after
<bibRefCitation id="DB8AA28FE0DDAF27084EB450A8CA7175" author="Liew, T" journalOrPublisher="ZooKeys" pageId="51" pageNumber="52" pagination="1 - 107" title="A cybertaxonomic revision of the micro-landsnail genus Plectostoma Adam (Mollusca, Caenogastropoda, Diplommatinidae), from Peninsular Malaysia, Sumatra and Indochina." url="https://doi.org/10.3897/zookeys.393.6717" volume="393" year="2014">Liew et al. 2014</bibRefCitation>
,
<bibRefCitation id="2ED852703536BEFFCB954535A1E97E90" author="Waterstradt, J" journalOrPublisher="Journal of the Straits Branch of the Royal Asiatic Society" pageId="54" pageNumber="55" pagination="1 - 27" title="Kelantan and my trip to Gunong Tahan." url="http://www.biodiversitylibrary.org/part/145327/summary" volume="37" year="1902">Waterstradt 1902</bibRefCitation>
).
<taxonomicName id="4552C8A6A3DCEE788DA10797FF2FCBC7" class="Gastropoda" family="Alycaeidae" genus="Alycaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Alycaeus kelantanensis" order="Architaenioglossa" pageId="32" pageNumber="33" phylum="Mollusca" rank="species" species="kelantanensis">Alycaeus kelantanensis</taxonomicName>
shells from
<pageBreakToken id="7042DAEB56723335155C865C1D8A847C" pageId="33" pageNumber="34" start="start">across</pageBreakToken>
its range are consistent in their diagnostic characters but varies in shell size within and between populations. Aside from the locations listed above, this species has also been recorded from limestone hills near Kuala Betis, 25 km west of Gua Musang, Kelantan (
<bibRefCitation id="066685006116EFCAA4A811E23DD374E0" author="Berry, AJ" journalOrPublisher="Bulletin of the National Museum Singapore" pageId="49" pageNumber="50" pagination="27 - 30" title="A collection of land mollusca from limestone in Ulu Kelantan." url="http://blsea.nus.edu.sg/en/persistent/584163f8-61e7-4329-8b04-d62e96e76796" volume="33" year="1965">Berry 1965</bibRefCitation>
) and in the non-limestone Lojing Highlands, Kelantan (
<bibRefCitation id="16A46F6B6F4C608218004751AFE3EBEE" author="Liew, TS" editor="Che, Omar I" journalOrPublisher="The Bulletin of the Raffles Museum" pageId="51" pageNumber="52" title="Penerbit Universiti Malaysia Kelantan, Kota Bharu" url="https://doi.org/10.13140/RG.2.1.4311.5364" year="2010">Liew 2010</bibRefCitation>
).
</paragraph>
<caption id="9D57E38DD43C8994F5B6C817DE6DCA8C" pageId="33" pageNumber="34">
<paragraph id="C2D49698A8AFFD7478486E60526744C3" pageId="33" pageNumber="34">
Figure 21.
<taxonomicName id="11542C3E4DB7E7BF54030711C70C0E93" class="Gastropoda" family="Alycaeidae" genus="Alycaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Alycaeus kelantanensis" order="Architaenioglossa" pageId="33" pageNumber="34" phylum="Mollusca" rank="species" species="kelantanensis">Alycaeus kelantanensis</taxonomicName>
Sykes, 1902.
<normalizedToken id="712CF640407358445733955D09123967" originalValue="AE">A-E</normalizedToken>
Shell of syntype, RMNH 153594
<normalizedToken id="9F40F9A0584626B835E015179356C228" originalValue="FJ">F-J</normalizedToken>
Shell of BOR/MOL 8327
<normalizedToken id="C08C13EA25738564E6238001FC6FF498" originalValue="KM">K-M</normalizedToken>
Close up of shell of BOR/MOL 8327
<normalizedToken id="0FEB802E8444DDB6B66E0034DF4CAD49" originalValue="NQ">N-Q</normalizedToken>
Operculum of BOR/MOL 6825
<normalizedToken id="ACC0204C2C5F46FCE3EEC3F8E73F9D65" originalValue="RV">R-V</normalizedToken>
Shell of BOR/MOL 12973
<normalizedToken id="CFAE794AD2F579BAEC853AEC690D8172" originalValue="WAA">W-AA</normalizedToken>
Shell of BOR/MOL 12974. Scale bars:
<normalizedToken id="E709C08A7BE58D178AF02A809564D0A9" originalValue="AJ">A-J</normalizedToken>
, K,
<normalizedToken id="EFFFD6CC9ED92B65522A16C589D831E1" originalValue="RAA">R-AA</normalizedToken>
1 mm; L, M,
<normalizedToken id="A1684E0805522AAF1CFCBC91C5424312" originalValue="NQ">N-Q</normalizedToken>
0.5 mm. Photographs
<normalizedToken id="3D59D71786A7B66AC6C399F2C4A38067" originalValue="AE">A-E</normalizedToken>
by Thor Seng Liew. Photographs
<normalizedToken id="1E61B43B06FE772DE26AC1FACD6B57E5" originalValue="FAA">F-AA</normalizedToken>
by Junn Kitt Foon.
</paragraph>
</caption>
</subSubSection>
</treatment>
</document>