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<mods:title id="15231C7345A2C401031CE0768B1279EE">A molecular and conchological dissection of the &quot; scaly &quot; Georissa of Malaysian Borneo (Gastropoda, Neritimorpha, Hydrocenidae)</mods:title>
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<mods:namePart id="DAE20AF179876B91133DD135D787AFFE">Khalik, Mohd Zacaery</mods:namePart>
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<mods:namePart id="1CE806D1356564F388DE09E52D3CB0C9">Hendriks, Kasper</mods:namePart>
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<mods:namePart id="065AE89518A6AAEE130ADB652C7EB58F">Vermeulen, Jaap J.</mods:namePart>
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<mods:namePart id="44001259CC152B4C652A056B6E4635F8">Schilthuizen, Menno</mods:namePart>
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<treatment id="0489A22D9C569A311C9436A606D45126" ID-GBIF-Taxon="146053692" LSID="urn:lsid:plazi:treatment:0489A22D9C569A311C9436A606D45126" httpUri="http://treatment.plazi.org/id/0489A22D9C569A311C9436A606D45126" lastPageNumber="1" pageId="0" pageNumber="1">
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<paragraph id="AAF4842138898E00A52C90A7275E1A22" pageId="0" pageNumber="1">
<taxonomicName id="CFE3EA002A8B0DD1A1B49EC6965629BF" ID-CoL="3FRYB" authority="Thompson &amp; Dance, 1983" authorityName="Thompson &amp; Dance" authorityYear="1983" class="Gastropoda" family="Hydrocenidae" genus="Georissa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Georissa pyrrhoderma" order="Cycloneritida" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="pyrrhoderma">Georissa pyrrhoderma Thompson &amp; Dance, 1983</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="DC24A3602E83451363839F953159275D" pageId="0" pageNumber="1" type="reference_group">
<paragraph id="0D782F6AA5DB3344A21502830983BF78" pageId="0" pageNumber="1">
<taxonomicName id="CD5DA78217794B500789291482B035B0" class="Gastropoda" family="Hydrocenidae" genus="Georissa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Georissa pyrrhoderma" order="Cycloneritida" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="pyrrhoderma">Georissa pyrrhoderma</taxonomicName>
Thompson &amp; Dance, 1983: 123, fig. 64.
</paragraph>
<paragraph id="E58BE10DDC88314098E746405FD4359E" pageId="0" pageNumber="1">
<taxonomicName id="11BF0F93A4C94608C4C02DA0B5475662" class="Gastropoda" family="Hydrocenidae" genus="Georissa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Georissa pyrrhoderma" order="Cycloneritida" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="pyrrhoderma">Georissa pyrrhoderma</taxonomicName>
van Benthem-Jutting, in
<bibRefCitation id="3755617BB041E0CD3A1BF2B5D07FE819" author="Beron, P" journalOrPublisher="Historia Naturalis Bulgarica" pageId="0" pageNumber="1" pagination="169 - 210" title="Comparative study of the invertebrate cave faunas of Southeast Asia and New Guinea." url="http://www.nmnhs.com/historia-naturalis-bulgarica/pdfs/000365000212015.pdf" volume="21" year="2015">Beron 2015</bibRefCitation>
: 181.
</paragraph>
</subSubSection>
<subSubSection id="59E9C547D6ADE53396DA433270575D0F" pageId="0" pageNumber="1" type="type locality">
<paragraph id="18AAC751E97B8158EA60D7BB9825F492" pageId="0" pageNumber="1">Type locality.</paragraph>
<paragraph id="87E5E79554EAFFD7C9E755AD92D3B291" pageId="0" pageNumber="1">
Borneo, Sarawak, First Division, western side of Gunong Selabor, Semabang entrance to Lobang Batu Cave (
<geoCoordinate id="F57F274FA478F6CCF64AA5773FF121C5" direction="north" orientation="latitude" precision="925" value="0.9166667">00°55'N</geoCoordinate>
,
<geoCoordinate id="C228415D9CF567739F716606FE075D2E" direction="east" orientation="longitude" precision="925" value="110.416664">110°25'E</geoCoordinate>
).
</paragraph>
</subSubSection>
<subSubSection id="D0A7F325C9229B01EFFA164BD41EF7DE" pageId="0" pageNumber="1" type="type material">
<paragraph id="84AE90372ED0F15A50E1A0AEF5192FD7" pageId="0" pageNumber="1">Type material.</paragraph>
<paragraph id="63D95242AAF8AAD1F90E4C9E0CDF56D0" pageId="0" pageNumber="1">Holotype. Borneo, Sarawak, First Division, western side of Gunong Selabor, Semabang entrance to Lobang Batu Cave: UF36183 (not seen). Paratypes. Borneo, Sarawak, First Division, western side of Gunong Selabor, Semabang entrance to Lobang Batu Cave: UF 36184, UF 36185 (not seen).</paragraph>
</subSubSection>
<subSubSection id="2754C6B2FC6170ACE34368A2FA6F2103" pageId="0" pageNumber="1" type="materials_examined">
<paragraph id="75FE5577E27FDF22AFFCC4F4E439EEC0" pageId="0" pageNumber="1">Other materials.</paragraph>
<paragraph id="CD7D8E130B99905C38B3C30B01980009" pageId="0" pageNumber="1">
Gunong Silabur, Serian, Sarawak, Malaysia (
<geoCoordinate id="851274A5CE0213F57881E57CB95B6BE2" direction="north" orientation="latitude" precision="9" value="0.95750004">00°57.45'N</geoCoordinate>
,
<geoCoordinate id="64AB3943422E270D2FF905DBB319A1BD" direction="east" orientation="longitude" precision="9" value="110.503334">110°30.20'E</geoCoordinate>
): MZU/MOL 17.09, MZU/MOL 17.10, MZU/MOL 17.11, MZU/MOL 17.12, MZU/MOL 17.13, MZU/MOL 17.14, MZU/MOL 17.15, MZU/MOL 17.16, MZU/MOL 17.17, MZU/MOL 17.18, MZU/MOL 17.19, MZU/MOL 17.20, MZU/MOL 17.21, MZU/MOL 17.22, MZU/MOL 17.23, MZU/MOL 17.24.
</paragraph>
</subSubSection>
<subSubSection id="43502EAF08075D2635785D3416869AC1" pageId="0" pageNumber="1" type="description">
<paragraph id="DF67948557B5ED0CACC040FE53EA69B6" pageId="0" pageNumber="1">Description.</paragraph>
<paragraph id="2775A1C9F09747FF95FC7C7080370FA3" pageId="0" pageNumber="1">
Protoconch. Color: red to brown. Sculpture pattern: smooth to meshed, with ellipsoid mesh shape. Mesh width: 11-26
<normalizedToken id="C0755D1BB446FE8DB7A932F27E5593C0" originalValue="µm">µm</normalizedToken>
. Teleoconch. Color: brown to red. First whorl: shouldered, slightly curved above the shoulder, flat, cylindrical below the shoulder. Subsequent whorls: initially shouldered, but soon grading into uniformly rounded and quickly expanding whorls, with a deeply impressed suture; number of whorls: 2
<normalizedToken id="040FA7985080D25FA3B3F41B247D5C33" originalValue="¼">1/4</normalizedToken>
-2
<normalizedToken id="54E453B6F211EFACC3AA6C79A3E218B9" originalValue="½">1/2</normalizedToken>
. SH: 1.16-1.31 mm, SW: 1.12-1.20 mm, SI: 1.03-1.09. Shell sculpture. Radial sculpture: absent, only weak to strong growth lines are visible. Spiral sculpture: present, strong spiral sculpture. Scales: a single series of low, small and acute, unevenly spaced scales above the periphery, occasionally, in the vicinity of the aperture, subordinate series of minute scales accompany the major series. Aperture. Shape: rounded, tilted below the palatal side. Basal side: rounded, strongly angular at the columellar region. Parietal side: straight, palatal edge attached to the body whorl. AH: 0.58-0.63 mm, AW: 0.75-0.85 mm, AI: 0.73-0.81.
</paragraph>
<caption id="3477BC970B47390A8B7A9B1FBADAFD34" pageId="0" pageNumber="1">
<paragraph id="750E4061FFB6B006F710F531CB1CE3F4" pageId="0" pageNumber="1">
Figure 15.
<taxonomicName id="3813DED7F090C2397A5EF24392557709" class="Gastropoda" family="Hydrocenidae" genus="Georissa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Georissa pyrrhoderma" order="Cycloneritida" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="pyrrhoderma">Georissa pyrrhoderma</taxonomicName>
Thompson &amp; Dance, 1983.
<normalizedToken id="FA93D3B60A5430C626DA47C169FE3AE5" originalValue="AC">A-C</normalizedToken>
MZU/MOL 17.10
<normalizedToken id="EEBCCCB437B0CA70B6146AE4DC2FB8E0" originalValue="DK">D-K</normalizedToken>
MZU/MOL 17.09. A, D Shell apertural view B Shell side view C Shell rear view
<normalizedToken id="315F78D99913C39E3EC82CFB5A4DE0DC" originalValue="EF">E-F</normalizedToken>
Shell cross-section from 3D model
<normalizedToken id="9EDFD43AA7D76510FC8A7E205D2C3C93" originalValue="GH">G-H</normalizedToken>
Operculum frontal and ventral view I Shell top view J Protoconch side view K Close up of protoconch from top at 1000
<normalizedToken id="61E1E6C41E74B90D6E0A278EDD77F6A7" originalValue="×">x</normalizedToken>
magnification. Scale bars: 500
<normalizedToken id="5E453FC222CAA92942B8D63439C7358B" originalValue="µm">µm</normalizedToken>
(
<normalizedToken id="85BAADB3D04758904291260115223F72" originalValue="AI">A-I</normalizedToken>
); 200
<normalizedToken id="C702CB5CE2FBE3CF011167010ED9B039" originalValue="µm">µm</normalizedToken>
(J); 10
<normalizedToken id="88D33B218488EF2D6C43153374D14A5C" originalValue="µm">µm</normalizedToken>
(K).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="E2B092BCA4A9FDA240C0F1F53EC9CE3A" pageId="0" pageNumber="1" type="cross diagnosis">
<paragraph id="8126C3780BF55C68737298E416BED192" pageId="0" pageNumber="1">Cross diagnosis.</paragraph>
<paragraph id="DF8B6A9E200C6FD10D53C803171AC68C" pageId="0" pageNumber="1">
<taxonomicName id="3D7486E16290FD8039D6D184C7657896" class="Gastropoda" family="Hydrocenidae" genus="Georissa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Georissa pyrrhoderma" order="Cycloneritida" pageId="0" pageNumber="1" phylum="Mollusca" rank="species" species="pyrrhoderma">Georissa pyrrhoderma</taxonomicName>
has a shell habitus that is similar to
<taxonomicName id="5649BC94D29BA626A4360964C242B520" lsidName="G. kobelti" pageId="0" pageNumber="1" rank="species" species="kobelti">G. kobelti</taxonomicName>
,
<taxonomicName id="586F57CA12FA0D9924568DAD80CE1DDF" lsidName="G. hosei" pageId="0" pageNumber="1" rank="species" species="hosei">G. hosei</taxonomicName>
, and
<taxonomicName id="74094591392A1E17BB19336ED99051B3" lsidName="G. sepulutensis" pageId="0" pageNumber="1" rank="species" species="sepulutensis">G. sepulutensis</taxonomicName>
. The latter two species are high variable and are morphologically, especially in sculpture, closely related to
<taxonomicName id="E2B4F11E3CB20D81F5D5D526BFBCAB81" lsidName="G. pyrrhoderma" pageId="0" pageNumber="1" rank="species" species="pyrrhoderma">G. pyrrhoderma</taxonomicName>
. Therefore,
<taxonomicName id="A4239A8296176CAE52E32959537E29CB" lsidName="G. pyrrhoderma" pageId="0" pageNumber="1" rank="species" species="pyrrhoderma">G. pyrrhoderma</taxonomicName>
is nearly indistinguishable from certain forms of these other species.
</paragraph>
</subSubSection>
<subSubSection id="CDB622031D24D19C15B57E555618E9AD" pageId="0" pageNumber="1" type="distribution">
<paragraph id="DD7727CED2192E756BD04EA2D0186A91" pageId="0" pageNumber="1">Distribution.</paragraph>
<paragraph id="064A9A7BD59F209562A51877E9290B7F" pageId="0" pageNumber="1">Only known from the type locality, Gunung Silabur, Serian, Sarawak, Malaysia.</paragraph>
</subSubSection>
<subSubSection id="881F5B19F16A5F88056CCE102CB6D833" pageId="0" pageNumber="1" type="molecular analysis">
<paragraph id="3EBCE7C835DF8CE0FD3D5333962CB94F" pageId="0" pageNumber="1">Molecular analysis.</paragraph>
<paragraph id="57E16453764F23BE836A2A479A7E8E98" pageId="0" pageNumber="1">
In the ML and Bayesian analyses, all
<taxonomicName id="16083A0F7807AFEDD0F1DD33DD0F7D21" lsidName="G. pyrrhoderma" pageId="0" pageNumber="1" rank="species" species="pyrrhoderma">G. pyrrhoderma</taxonomicName>
(16S: n = 28; CO1: n = 26) individuals group together in one clade with 99% BS and 100% PP. Its sister clade is
<taxonomicName id="7EBA6F45B9B8DF5F1398583FAEC90219" lsidName="G. scalinella" pageId="0" pageNumber="1" rank="species" species="scalinella">G. scalinella</taxonomicName>
+
<taxonomicName id="B7CB431DAC22C150353E21E35C98D91F" lsidName="G. kinabatanganensis" pageId="0" pageNumber="1" rank="species" species="kinabatanganensis">G. kinabatanganensis</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="68D0C583316C7DBBA0F745680033A58C" pageId="0" pageNumber="1" type="discussion">
<paragraph id="8BBAAD8468742CC2458C4F04BC020E8A" pageId="0" pageNumber="1">Discussion.</paragraph>
<paragraph id="5981B6DAD32EBE6C144F0D82D5A7A142" pageId="0" pageNumber="1">
In the original description,
<bibRefCitation id="A410A422D9131BAA0D32404A23F84A5D" author="Thompson, FG" journalOrPublisher="Bulletin of the Florida State Museum Biological Sciences" pageId="0" pageNumber="1" pagination="101 - 152" title="Non-marine mollusks of Borneo. II Pulmonata: Pupillidae, Clausiliidae. III Prosobranchia: Hydrocenidae, Helicinidae." url="https://www.floridamuseum.ufl.edu/files/2114/7180/1931/Vol-29-No-3.PDF" volume="29" year="1983">Thompson and Dance (1983)</bibRefCitation>
did not compare
<taxonomicName id="CB6ECFE555D6D5194BBA0F402FA1F119" lsidName="G. pyrrhoderma" pageId="0" pageNumber="1" rank="species" species="pyrrhoderma">G. pyrrhoderma</taxonomicName>
with members of their
<taxonomicName id="B01DE0E7F36BDDB6D92AC1F8874CA05A" lsidName="G hosei" pageId="0" pageNumber="1" rank="species" species="hosei">hosei</taxonomicName>
-group (which our molecular analyses show it belongs in). Instead, they considered it allied to
<taxonomicName id="373EAD07961585227D6AE09E534A9B03" lsidName="G. borneensis" pageId="0" pageNumber="1" rank="species" species="borneensis">G. borneensis</taxonomicName>
. Possibly this misalignment was caused by the fact that the type specimens appear to lack the series of scales that is present on most of the specimens we collected. Nonetheless, given the restricted range of collection localities at Gunung Silabur and the degree of variability in our material, we consider our and Thompson and
<normalizedToken id="E15BA9A07BFBAC7788FD3C2B512A2BF1" originalValue="Dances">Dance's</normalizedToken>
material as conspecific. The paratypes specimen NHMUK 1984005 (Semabang entrance to Lobang Batu Cave, W. side of Gunong Selabor, 1st. Div., Sarawak, Malaysia: seen) is
<taxonomicName id="89D328002275897D41DCD25072C65361" lsidName="G. silaburensis" pageId="0" pageNumber="1" rank="species" species="silaburensis">G. silaburensis</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
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