262 lines
37 KiB
XML
262 lines
37 KiB
XML
<document id="8AB433BD14F411D81DD19D386E255E5E" ID-CLB-Dataset="6611" ID-DOI="10.5281/zenodo.4502132" ID-GBIF-Dataset="60740210-999d-4526-88e3-464fd93143ac" ID-ISSN="2345-7600" ID-Zenodo-Dep="4502132" ID-ZooBank="4D5F3585-04B2-45AA-BA00-8795AEF68A83" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1612458956132" checkinUser="carolina" docAuthor="Köhler, Frank" docDate="2014" docId="03F3175FEF695E42FE923F2DFD36FECA" docLanguage="en" docName="RafflesBZool.62.115-123.pdf" docOrigin="Raffles Bulletin of Zoology 62" docStyle="DocumentStyle:F2D34FF25DBF9998BB8DF6F24A5B097A.5:RafflesBZool.2014-2018.journal_article" docStyleId="F2D34FF25DBF9998BB8DF6F24A5B097A" docStyleName="RafflesBZool.2014-2018.journal_article" docStyleVersion="5" docTitle="Rhagada abbasi Köhler 2014, n. sp." docType="treatment" docUuid="12721561-AD72-4DF4-9BDC-221CD7E2306C" docUuidSource="ZooBank" docVersion="10" lastPageNumber="121" masterDocId="FFCA6F27EF6F5E45FFDE3C2BFFD2FFFE" masterDocTitle="On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands" masterLastPageNumber="123" masterPageNumber="115" pageNumber="121" updateTime="1698926749818" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:titleInfo id="C189768AE269809AD54FB70C0DEA6DB0">
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<mods:title id="08CA615DF3ACE02FC42F24730604C3B1">On the land snail Rhagada Albers, 1860 (Gastropoda: Camaenidae) from across the Lesser Sunda Islands</mods:title>
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</mods:titleInfo>
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<mods:name id="218A6FDCA8D400A40EACAEFED5471FCA" type="personal">
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<mods:roleTerm id="EA66CC08E84B7FD44EF709B9604CF01D">Author</mods:roleTerm>
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<mods:namePart id="6B820B02D18CBFD014AADD31843BF7D6">Köhler, Frank</mods:namePart>
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<mods:typeOfResource id="36EF8A7B29C90B9F24B87C59194F58B3">text</mods:typeOfResource>
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<mods:title id="04B256974B8E20071C762603BE6D011F">Raffles Bulletin of Zoology</mods:title>
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<mods:part id="2A85B6F020AD29019FC77EF699BF60B5">
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<mods:date id="EF0C64E19EFCFACAF3428F1185E47E1E">2014</mods:date>
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<mods:number id="0E9F0E62A7EA7BEAFA31B617FB32B43E">2014-03-17</mods:number>
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<mods:number id="50B1454D2A54F118E65D88CF4A68266C">62</mods:number>
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<mods:start id="68F698AB6F8FAF64FF31D88BFB846F95">115</mods:start>
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<mods:classification id="189EAAC5BBCFC961ABA4532F45B63041">journal article</mods:classification>
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<mods:identifier id="880F56347786CE87096CF10FFFB9D5AD" type="DOI">10.5281/zenodo.4502132</mods:identifier>
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<treatment id="03F3175FEF695E42FE923F2DFD36FECA" ID-DOI="http://doi.org/10.5281/zenodo.4502074" ID-GBIF-Taxon="182237999" ID-Zenodo-Dep="4502074" LSID="urn:lsid:zoobank.org:act:12721561-AD72-4DF4-9BDC-221CD7E2306C" httpUri="http://treatment.plazi.org/id/03F3175FEF695E42FE923F2DFD36FECA" lastPageId="7" lastPageNumber="121" pageId="6" pageNumber="121">
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<subSubSection id="C340F5C2EF695E43FE923F2DFD91FCE0" box="[332,579,774,798]" pageId="6" pageNumber="121" type="nomenclature">
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<paragraph id="8BE5A649EF695E43FE923F2DFD91FCE0" blockId="6.[332,579,774,830]" box="[332,579,774,798]" pageId="6" pageNumber="121">
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<heading id="D0AD1125EF695E43FE923F2DFD91FCE0" box="[332,579,774,798]" centered="true" fontSize="10" level="2" pageId="6" pageNumber="121" reason="3">
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<taxonomicName id="4C5ADDCAEF695E43FE923F2DFE2DFCE0" ID-CoL="6X63X" authority="Köhler, 2014" authorityName="Köhler" authorityYear="2014" box="[332,511,774,798]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="species" species="abbasi" status="sp. nov.">
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<emphasis id="B92E7A5BEF695E43FE923F2DFE2DFCE0" bold="true" box="[332,511,774,798]" italics="true" pageId="6" pageNumber="121">Rhagada abbasi</emphasis>
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</taxonomicName>
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<taxonomicNameLabel id="A21DC720EF695E43FDD63F2CFD91FCE0" box="[520,579,775,798]" pageId="6" pageNumber="121" rank="species">n. sp.</taxonomicNameLabel>
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||
</heading>
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||
</paragraph>
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||
</subSubSection>
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||
<subSubSection id="C340F5C2EF695E43FEB23F0CFDF1FCC0" box="[364,547,807,830]" pageId="6" pageNumber="121" type="description">
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<paragraph id="8BE5A649EF695E43FEB23F0CFDF1FCC0" blockId="6.[332,579,774,830]" box="[364,547,807,830]" pageId="6" pageNumber="121">
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(
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<figureCitation id="1361BACCEF695E43FEAA3F0CFE67FCC0" box="[372,437,807,830]" captionStart="Fig" captionStartId="1.[113,148,1918,1939]" captionTargetBox="[113,752,690,1894]" captionTargetId="figure-175@1.[113,752,690,1894]" captionTargetPageId="1" captionText="Fig. 1. Shells (1.5× of natural size). A, Rhagada solorensis, Solor Island, AM C.471154; B, Rhagada setzeri setzeri, Alor Island, AM C.471156; C, D, Rhagada setzeri atauroensis n. subsp., Atauro Island: C, Paratype AM C.468711; D, Paratype AM C.468709; E–H, Rhagada abbasi n. sp., Lembata Island: E, Holotype AM C.478280; F–H: Paratypes AM C.471153, showing variation in sculpture and colour pattern. Scale bar = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.4502136" httpUri="https://zenodo.org/record/4502136/files/figure.png" pageId="6" pageNumber="121">Figs 1</figureCitation>
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E–H, 2D)
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</paragraph>
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</subSubSection>
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<subSubSection id="C340F5C2EF695E43FF563F4DFD42FBD9" pageId="6" pageNumber="121" type="materials_examined">
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<paragraph id="8BE5A649EF695E43FF563F4DFDE0FC69" blockId="6.[136,775,870,919]" pageId="6" pageNumber="121">
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<emphasis id="B92E7A5BEF695E43FF563F4DFEDDFC85" bold="true" box="[136,271,870,891]" pageId="6" pageNumber="121">Type locality.</emphasis>
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<materialsCitation id="3B32AC14EF695E43FECA3F4DFDFCFC69" ID-GBIF-Occurrence="3114366314" country="Indonesia" latitude="-8.416667" location="Lembata Island" longLatPrecision="1301" longitude="123.46667" pageId="6" pageNumber="121" specimenCount="1" stateProvince="Nusa Tenggara Timur">
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<collectingCountry id="F34DE6D9EF695E43FECA3F4DFEA6FC85" box="[276,372,870,891]" name="Indonesia" pageId="6" pageNumber="121">Indonesia</collectingCountry>
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,
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<collectingRegion id="499E68ABEF695E43FEA33F4DFD78FC85" box="[381,682,870,891]" country="Indonesia" name="Nusa Tenggara Timur" pageId="6" pageNumber="121">Nusa Tenggara Timur Province</collectingRegion>
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,
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<location id="8E85F092EF695E43FD6D3F4DFF11FC69" LSID="urn:lsid:plazi:treatment:03F3175FEF695E42FE923F2DFD36FECA:8E85F092EF695E43FD6D3F4DFF11FC69" country="Indonesia" latitude="-8.416667" longLatPrecision="1301" longitude="123.46667" name="Lembata Island" pageId="6" pageNumber="121" stateProvince="Nusa Tenggara Timur">Lembata Island</location>
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(= Lomblem), –
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<geoCoordinate id="EE6EC08EEF695E43FEB73FA9FE65FC69" box="[361,439,898,919]" degrees="8" direction="south" minutes="25" orientation="latitude" pageId="6" pageNumber="121" precision="925" value="-8.416667">8° 25' S</geoCoordinate>
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,
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<geoCoordinate id="EE6EC08EEF695E43FE1A3FA9FDFCFC69" box="[452,558,898,919]" degrees="123" direction="east" minutes="28" orientation="longitude" pageId="6" pageNumber="121" precision="925" value="123.46667">123° 28' E</geoCoordinate>
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</materialsCitation>
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.
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</paragraph>
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<paragraph id="8BE5A649EF695E43FF563F97FD5AFC2F" blockId="6.[136,648,956,977]" box="[136,648,956,977]" pageId="6" pageNumber="121">
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<materialsCitation id="3B32AC14EF695E43FF563F97FD5AFC2F" ID-GBIF-Occurrence="3114366302" box="[136,648,956,977]" collectionCode="AM" pageId="6" pageNumber="121" specimenCount="1" typeStatus="holotype">
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<emphasis id="B92E7A5BEF695E43FF563F97FF39FC2F" bold="true" box="[136,235,956,977]" pageId="6" pageNumber="121">
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<typeStatus id="54E118EBEF695E43FF563F97FF35FC2F" box="[136,231,956,977]" pageId="6" pageNumber="121" type="holotype">Holotype</typeStatus>
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.
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</emphasis>
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<collectionCode id="ED4B3E8CEF695E43FF2C3F97FECBFC2F" box="[242,281,956,977]" country="Australia" name="Australian Museum" pageId="6" pageNumber="121" type="Museum">AM</collectionCode>
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C.478280 (dissected specimen, wet).
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</materialsCitation>
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</paragraph>
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<paragraph id="8BE5A649EF695E43FF563FDDFD42FBD9" blockId="6.[136,775,1014,1063]" pageId="6" pageNumber="121">
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<materialsCitation id="3B32AC14EF695E43FF563FDDFD5EFBD9" ID-GBIF-Occurrence="3114366303" collectionCode="AM" pageId="6" pageNumber="121" specimenCode="ZMB 117967" specimenCount="1" typeStatus="paratype">
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<emphasis id="B92E7A5BEF695E43FF563FDDFF26FBF5" bold="true" box="[136,244,1014,1035]" pageId="6" pageNumber="121">
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<typeStatus id="54E118EBEF695E43FF563FDDFF22FBF5" box="[136,240,1014,1035]" pageId="6" pageNumber="121" type="paratype">Paratypes</typeStatus>
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.
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</emphasis>
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<collectionCode id="ED4B3E8CEF695E43FF243FDDFEF3FBF5" box="[250,289,1014,1035]" country="Australia" name="Australian Museum" pageId="6" pageNumber="121" type="Museum">AM</collectionCode>
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C.471153 (4 wet, 7 dry),
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<collectionCode id="ED4B3E8CEF695E43FDC33FDDFD96FBF5" box="[541,580,1014,1035]" country="Australia" name="Australian Museum" pageId="6" pageNumber="121" type="Museum">AM</collectionCode>
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C.468979 (2 wet, 5 dry),
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<collectionCode id="ED4B3E8CEF695E43FF623839FF31FBD9" box="[188,227,1042,1063]" country="Australia" name="Australian Museum" pageId="6" pageNumber="121" type="Museum">AM</collectionCode>
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C.478281 (50 dry),
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<specimenCode id="DBFC0E32EF695E43FE703839FDE3FBD9" box="[430,561,1042,1063]" collectionCode="ZMB" country="Germany" httpUri="http://grbio.org/cool/8syf-d21i" name="Museum für Naturkunde Berlin (Zoological Collections)" pageId="6" pageNumber="121" type="Museum">ZMB 117967</specimenCode>
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(50 dry)
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</materialsCitation>
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.
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</paragraph>
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</subSubSection>
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<subSubSection id="C340F5C2EF695E43FF56387AFDB3FB76" pageId="6" pageNumber="121" type="etymology">
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<paragraph id="8BE5A649EF695E43FF56387AFDB3FB76" blockId="6.[136,775,1105,1160]" pageId="6" pageNumber="121">
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<emphasis id="B92E7A5BEF695E43FF56387AFEDAFB96" bold="true" box="[136,264,1105,1128]" pageId="6" pageNumber="121">Etymology.</emphasis>
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In honour of John Abbas,
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<collectingRegion id="499E68ABEF695E43FDEF387AFDADFB96" box="[561,639,1105,1128]" country="Indonesia" name="Jakarta Raya" pageId="6" pageNumber="121">Jakarta</collectingRegion>
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, who kindly provided material from Lembata and Solor.
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</paragraph>
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</subSubSection>
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<subSubSection id="C340F5C2EF695E43FF56389AFE3FF936" pageId="6" pageNumber="121" type="description">
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<paragraph id="8BE5A649EF695E43FF56389AFEB0FA36" blockId="6.[136,775,1201,1480]" pageId="6" pageNumber="121">
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<emphasis id="B92E7A5BEF695E43FF56389AFEC1FB36" bold="true" box="[136,275,1201,1224]" pageId="6" pageNumber="121">Description.</emphasis>
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Shell moderate in size (D = 15–
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<quantity id="4CA20BACEF695E43FDAD389AFD13FB36" box="[627,705,1201,1224]" metricMagnitude="-2" metricUnit="m" metricValue="1.5" pageId="6" pageNumber="121" unit="mm" value="15.0">15 mm</quantity>
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;
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<tableCitation id="C6D893F2EF695E43FD12389AFF46FB16" captionStart="Table 1" captionStartId="4.[136,189,191,212]" captionTargetPageId="4" captionText="Table 1. Shell measurements for N shells of different Rhagada species and subspecies. H = height; D = diameter; W = no. of whorls." httpUri="http://table.plazi.org/id/DF25F6C1EF6B5E41FF563C94FA79FF2A" pageId="6" pageNumber="121" tableUuid="DF25F6C1EF6B5E41FF563C94FA79FF2A">Table 1</tableCitation>
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), broadly subglobose, with variable reddish-brown banding on whitish to creamish background, with sculpture of faint to well-developed axial ribs, umbilicus forming a chink, lip thickened, slightly reflected (
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<figureCitation id="1361BACCEF695E43FE2F391AFDFDFAB6" box="[497,559,1329,1352]" captionStart="Fig" captionStartId="1.[113,148,1918,1939]" captionTargetBox="[113,752,690,1894]" captionTargetId="figure-175@1.[113,752,690,1894]" captionTargetPageId="1" captionText="Fig. 1. Shells (1.5× of natural size). A, Rhagada solorensis, Solor Island, AM C.471154; B, Rhagada setzeri setzeri, Alor Island, AM C.471156; C, D, Rhagada setzeri atauroensis n. subsp., Atauro Island: C, Paratype AM C.468711; D, Paratype AM C.468709; E–H, Rhagada abbasi n. sp., Lembata Island: E, Holotype AM C.478280; F–H: Paratypes AM C.471153, showing variation in sculpture and colour pattern. Scale bar = 10 mm." figureDoi="http://doi.org/10.5281/zenodo.4502136" httpUri="https://zenodo.org/record/4502136/files/figure.png" pageId="6" pageNumber="121">Fig. 1</figureCitation>
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E–H). Penial verge smooth with pointed tip and well-developed sperm groove, inner penial wall with well-developed longitudinal pilasters, epiphallus with longitudinal pilasters along inner wall, short flagellum (
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<figureCitation id="1361BACCEF695E43FF23399AFE84FA36" box="[253,342,1457,1480]" captionStart="Fig" captionStartId="2.[136,171,2006,2027]" captionTargetBox="[143,1467,293,1913]" captionTargetId="figure-6@2.[129,1480,291,1924]" captionTargetPageId="2" captionText="Fig. 2. Penial anatomy, with penial chamber opened showing internal anatomy. A, Rhagada solorensis, Solor Island, AM C.471154; B, Rhagada marghitae, Nusa Penida AM C.327851; C, Rhagada setzeri atauroensis n. subsp., Atauro Island, holotype AM C.478282; D, Rhagada abbasi n. sp., Lembata Island, holotype AM C.478280. Abbreviations: ep = epiphallus; fl = epiphallic flagellum; gp = genital pore; lwp = longitudinal wall pilaster; pv = penial verge; rm = penial retractor muscle; va = vagina; vd = vas deferens. Scale bars = 2 mm." figureDoi="http://doi.org/10.5281/zenodo.4502138" httpUri="https://zenodo.org/record/4502138/files/figure.png" pageId="6" pageNumber="121">Fig. 2D</figureCitation>
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).
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</paragraph>
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<paragraph id="8BE5A649EF695E43FF5639DAFE3FF936" blockId="6.[136,775,1521,1736]" pageId="6" pageNumber="121">
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<emphasis id="B92E7A5BEF695E43FF5639DAFE59F9F6" bold="true" box="[136,395,1521,1544]" pageId="6" pageNumber="121">Comparative remarks.</emphasis>
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Shell smaller than
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<taxonomicName id="4C5ADDCAEF695E43FD8739DAFD30F9F6" baseAuthorityName="Martens" baseAuthorityYear="1863" box="[601,738,1521,1544]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="species" species="solorensis">
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<emphasis id="B92E7A5BEF695E43FD8739DAFD30F9F6" box="[601,738,1521,1544]" italics="true" pageId="6" pageNumber="121">R. solorensis</emphasis>
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</taxonomicName>
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,
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<taxonomicName id="4C5ADDCAEF695E43FD2E39DAFF20F9D6" authorityName="Martens" authorityYear="1891" class="Gastropoda" family="Camaenidae" genus="Helix" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="species" species="floresiana">
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<emphasis id="B92E7A5BEF695E43FD2E39DAFF20F9D6" italics="true" pageId="6" pageNumber="121">R. floresiana</emphasis>
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</taxonomicName>
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, and
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<taxonomicName id="4C5ADDCAEF695E43FEEB3A3AFE3FF9D6" authority="setzeri" box="[309,493,1553,1576]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="subSpecies" species="setzeri" subSpecies="setzeri">
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<emphasis id="B92E7A5BEF695E43FEEB3A3AFE3FF9D6" box="[309,493,1553,1576]" italics="true" pageId="6" pageNumber="121">R. setzeri setzeri</emphasis>
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</taxonomicName>
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, but similar to.
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<taxonomicName id="4C5ADDCAEF695E43FD783A3AFED1F9B6" authorityName="Köhler" authorityYear="2014" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="subSpecies" species="setzeri" subSpecies="atauroensis">
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<emphasis id="B92E7A5BEF695E43FD783A3AFED1F9B6" italics="true" pageId="6" pageNumber="121">R setzeri atauroensis</emphasis>
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</taxonomicName>
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. Most typical features are smooth penial verge with pointed tip and well-developed longitudinal pilasters of penial wall. Well-differentiated from
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<taxonomicName id="4C5ADDCAEF695E43FD963A5AFD07F976" baseAuthorityName="Martens" baseAuthorityYear="1863" box="[584,725,1649,1672]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="species" species="solorensis">
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<emphasis id="B92E7A5BEF695E43FD963A5AFD07F976" box="[584,725,1649,1672]" italics="true" pageId="6" pageNumber="121">R. solorensis</emphasis>
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</taxonomicName>
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and
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<taxonomicName id="4C5ADDCAEF695E43FF563ABAFF3EF956" authorityName="Maassen" authorityYear="2009" box="[136,236,1681,1704]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="species" species="setzeri">
|
||
<emphasis id="B92E7A5BEF695E43FF563ABAFF3EF956" box="[136,236,1681,1704]" italics="true" pageId="6" pageNumber="121">R. setzeri</emphasis>
|
||
</taxonomicName>
|
||
in a molecular phylogeny based on mitochondrial COI and 16S sequences (
|
||
<figureCitation id="1361BACCEF695E43FE433A9AFE0DF936" box="[413,479,1713,1736]" captionStart="Fig" captionStartId="5.[113,148,2006,2027]" captionTargetBox="[157,1404,941,1982]" captionTargetId="figure-153@5.[151,1414,931,1991]" captionTargetPageId="5" captionText="Fig. 3. Bayesian consensus phylogram based on analyses of concatenated 16S and COI sequences, adapted from Köhler & Criscione (2013). Lesser Sunda species Rhagada solorensis, R. setzeri setzeri, R. setzeri atauroensis n. subsp., and R. abbasi n. sp. form the sister clade of Rhagada from the Australian Pilbara. Clades from the Australian Kimberley and Pilbara regions collapsed; outgroup pruned from tree; see Köhler & Criscione (2013) for further details." figureDoi="http://doi.org/10.5281/zenodo.4502142" httpUri="https://zenodo.org/record/4502142/files/figure.png" pageId="6" pageNumber="121">Fig. 3</figureCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C340F5C2EF695E42FF563ADAFD36FECA" lastPageId="7" lastPageNumber="122" pageId="6" pageNumber="121" type="discussion">
|
||
<paragraph id="8BE5A649EF695E43FF563ADAFBD0FBA0" blockId="6.[136,775,1777,2088]" lastBlockId="6.[835,1475,775,1118]" pageId="6" pageNumber="121">
|
||
<emphasis id="B92E7A5BEF695E43FF563ADAFED5F8F6" bold="true" box="[136,263,1777,1800]" pageId="6" pageNumber="121">Discussion.</emphasis>
|
||
Previous taxonomic treatments of Lesser Sunda species were predominantly based on shell characters, which were usually inferred from small series of specimens. The better known Australian
|
||
<taxonomicName id="4C5ADDCAEF695E43FE583B7AFE37F896" authorityName="Albers" authorityYear="1860" box="[390,485,1873,1896]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="genus">
|
||
<emphasis id="B92E7A5BEF695E43FE583B7AFE37F896" box="[390,485,1873,1896]" italics="true" pageId="6" pageNumber="121">Rhagada</emphasis>
|
||
</taxonomicName>
|
||
species were shown to have rather conserved shells with regard to general shape and size while exhibiting considerable variation in shell sculpture, umbilical morphology and colour (
|
||
<bibRefCitation id="EFCBDBB8EF695E43FDD23B9AFD72F836" author="Kohler F" box="[524,672,1969,1992]" pageId="6" pageNumber="121" pagination="203 - 406" refId="ref6101" refString="Kohler F (2011) The camaenid species of the Kimberley Islands, Western Australia (Stylommatophora: Helicoidea). Malacologia, 54: 203 - 406." type="journal article" year="2011">Köhler, 2011</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFCBDBB8EF695E43FD713B9AFED5F816" author="Johnson MS & Hamilton ZR & Teale ROY & Kendrick PG" pageId="6" pageNumber="121" pagination="316 - 327" refId="ref6053" refString="Johnson MS, Hamilton ZR, Teale ROY & Kendrick PG (2012) Endemic evolutionary radiation of Rhagada land snails (Pulmonata: Camaenidae) in a continental archipelago in northern Western Australia. Biological Journal of the Linnean Society, 106: 316 - 327." type="journal article" year="2012">Johnson et al., 2012</bibRefCitation>
|
||
). Extreme amounts of intraspecific variation in shells were observed in some island populations, but are governed by strong selection in an extreme environment (
|
||
<bibRefCitation id="EFCBDBB8EF695E43FC953F2CFBDFFCE0" author="Stankowski S" box="[843,1037,775,798]" pageId="6" pageNumber="121" pagination="756 - 769" refId="ref6847" refString="Stankowski S (2011) Extreme, continuous variation in an island snail: local diversification and association of shell form with the current environment. Biological Journal of the Linnean Society, 104: 756 - 769." type="journal article" year="2011">Stankowski, 2011</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFCBDBB8EF695E43FBC63F2CFB80FCE0" author="Stankowski S" box="[1048,1106,775,798]" pageId="6" pageNumber="121" pagination="2726 - 2741" refId="ref6886" refString="Stankowski S (2013) Ecological speciation in an island snail: evidence for the parallel evolution of a novel ecotype and maintenance by ecologically dependent postzygotic isolation. Molecular Ecology, 22: 2726 - 2741." type="journal article" year="2013">2013</bibRefCitation>
|
||
). By contrast, the species from the
|
||
<collectingRegion id="499E68ABEF695E43FC9D3F0CFBF6FCC0" box="[835,1060,807,830]" country="Indonesia" name="Nusa Tenggara Timur" pageId="6" pageNumber="121">Lesser Sunda Islands</collectingRegion>
|
||
remain poorly documented in terms of available study material. However, where large numbers of specimens are available for study (i.e.,
|
||
<taxonomicName id="4C5ADDCAEF695E43FB0A3F4CFA60FC80" authorityName="Köhler" authorityYear="2014" box="[1236,1458,871,894]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="subSpecies" species="setzeri" subSpecies="atauroensis">
|
||
<emphasis id="B92E7A5BEF695E43FB0A3F4CFA60FC80" box="[1236,1458,871,894]" italics="true" pageId="6" pageNumber="121">R. setzeri atauroensis</emphasis>
|
||
</taxonomicName>
|
||
), the intraspecific variation of shells was found to correspond well with levels observed in Australian species. There is no reason to assume that other Lesser Sunda species exhibit much lower amounts of intraspecific variability. Consequently, traditionally emphasised shell characters, such as sculpture, umbilical morphology and banding pattern, may be of little taxonomic utility.
|
||
</paragraph>
|
||
<paragraph id="8BE5A649EF695E43FC9D38ACFB6DF840" blockId="6.[835,1475,1159,1982]" pageId="6" pageNumber="121">
|
||
Recent studies in Australian camaenids, including
|
||
<taxonomicName id="4C5ADDCAEF695E43FA8538ACFA69FB60" authorityName="Albers" authorityYear="1860" box="[1371,1467,1159,1182]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="genus">
|
||
<emphasis id="B92E7A5BEF695E43FA8538ACFA69FB60" box="[1371,1467,1159,1182]" italics="true" pageId="6" pageNumber="121">Rhagada</emphasis>
|
||
</taxonomicName>
|
||
, showed that comparative penial anatomy (
|
||
<figureCitation id="1361BACCEF695E43FAE9388CFAADFB40" box="[1335,1407,1191,1214]" captionStart="Fig" captionStartId="2.[136,171,2006,2027]" captionTargetBox="[143,1467,293,1913]" captionTargetId="figure-6@2.[129,1480,291,1924]" captionTargetPageId="2" captionText="Fig. 2. Penial anatomy, with penial chamber opened showing internal anatomy. A, Rhagada solorensis, Solor Island, AM C.471154; B, Rhagada marghitae, Nusa Penida AM C.327851; C, Rhagada setzeri atauroensis n. subsp., Atauro Island, holotype AM C.478282; D, Rhagada abbasi n. sp., Lembata Island, holotype AM C.478280. Abbreviations: ep = epiphallus; fl = epiphallic flagellum; gp = genital pore; lwp = longitudinal wall pilaster; pv = penial verge; rm = penial retractor muscle; va = vagina; vd = vas deferens. Scale bars = 2 mm." figureDoi="http://doi.org/10.5281/zenodo.4502138" httpUri="https://zenodo.org/record/4502138/files/figure.png" pageId="6" pageNumber="121">Fig. 2</figureCitation>
|
||
) may provide valuable taxonomic information even in species that possess conserved shells (e.g.,
|
||
<bibRefCitation id="EFCBDBB8EF695E43FB6938CCFA6CFB00" author="Kohler F & Johnson MS" box="[1207,1470,1255,1278]" pageId="6" pageNumber="121" pagination="337 - 362" refId="ref6157" refString="Kohler F & Johnson MS (2012) Species limits in molecular phylogenies: a cautionary tale from Australian land snails (Camaenidae: Amplirhagada). Zoological Journal of the Linnean Society, 165: 337 - 362." type="journal article" year="2012">Köhler & Johnson, 2012</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFCBDBB8EF695E43FC9D392CFBBBFAE0" author="Criscione F & Kohler F" box="[835,1129,1287,1310]" pageId="6" pageNumber="121" pagination="389 - 405" refId="ref5629" refString="Criscione F & Kohler F (2013 a) Conserved shell disguises diversity in Mesodontrachia land snails from the Australian Monsoon Tropics (Gastropoda: Camaenidae). Zoologica Scripta, 42: 389 - 405." type="journal article" year="2013">Criscione & Köhler, 2013a</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFCBDBB8EF695E43FBAB392CFB6EFAE0" author="Criscione F & Kohler F" box="[1141,1212,1287,1310]" pageId="6" pageNumber="121" pagination="101 - 115" refId="ref5666" refString="Criscione F & Kohler F (2013 b) Six new species of Australocosmica Kohler, 2011 from the Kimberley Islands, Western Australia (Mollusca: Pulmonata: Camaenidae). Zootaxa, 3608: 101 - 115." type="journal article" year="2013">2013b</bibRefCitation>
|
||
). The penial anatomy of two already described species (
|
||
<taxonomicName id="4C5ADDCAEF695E43FB4B390CFACDFAC0" baseAuthorityName="Martens" baseAuthorityYear="1863" box="[1173,1311,1319,1342]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="species" species="solorensis">
|
||
<emphasis id="B92E7A5BEF695E43FB4B390CFACDFAC0" box="[1173,1311,1319,1342]" italics="true" pageId="6" pageNumber="121">R. solorensis</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5ADDCAEF695E43FAF0390CFA6BFAC0" authorityName="Falconieri" authorityYear="1995" box="[1326,1465,1319,1342]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="species" species="marghitae">
|
||
<emphasis id="B92E7A5BEF695E43FAF0390CFA6BFAC0" box="[1326,1465,1319,1342]" italics="true" pageId="6" pageNumber="121">R. marghitae</emphasis>
|
||
</taxonomicName>
|
||
) and two new taxa (
|
||
<taxonomicName id="4C5ADDCAEF695E43FBD1396CFB26FAA0" authority="Köhler, 2014" authorityName="Köhler" authorityYear="2014" box="[1039,1268,1351,1374]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="subSpecies" species="setzeri" status="sp. nov." subSpecies="atauroensis">
|
||
<emphasis id="B92E7A5BEF695E43FBD1396CFB26FAA0" box="[1039,1268,1351,1374]" italics="true" pageId="6" pageNumber="121">R. setzeri atauroensis</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A21DC720EF695E43FB25396CFAE5FAA0" box="[1275,1335,1351,1374]" pageId="6" pageNumber="121" rank="species">n. sp.</taxonomicNameLabel>
|
||
,
|
||
<taxonomicName id="4C5ADDCAEF695E43FA9C396CFA75FAA0" authority="Köhler, 2014" authorityName="Köhler" authorityYear="2014" box="[1346,1447,1351,1374]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="species" species="abbasi" status="sp. nov.">
|
||
<emphasis id="B92E7A5BEF695E43FA9C396CFA75FAA0" box="[1346,1447,1351,1374]" italics="true" pageId="6" pageNumber="121">R. abbasi</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A21DC720EF695E43FA70396CFCB0FA80" pageId="6" pageNumber="121" rank="species">n. sp.</taxonomicNameLabel>
|
||
) provided useful taxonomic information complementing the shell-based taxonomy. Molecular phylogenetic analyses based on mitochondrial sequences of the COI and 16S genes showed that
|
||
<taxonomicName id="4C5ADDCAEF695E43FBCF39ECFBA3FA20" authorityName="Albers" authorityYear="1860" box="[1041,1137,1479,1502]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="genus">
|
||
<emphasis id="B92E7A5BEF695E43FBCF39ECFBA3FA20" box="[1041,1137,1479,1502]" italics="true" pageId="6" pageNumber="121">Rhagada</emphasis>
|
||
</taxonomicName>
|
||
species from the
|
||
<collectingRegion id="499E68ABEF695E43FAEC39ECFC5DFA00" country="Indonesia" name="Nusa Tenggara Timur" pageId="6" pageNumber="121">Lesser Sunda Islands</collectingRegion>
|
||
form a clade, which represents the sister group of all known species from the Australian Pilbara region (
|
||
<figureCitation id="1361BACCEF695E43FAA63A2CFA6EF9E0" box="[1400,1468,1543,1566]" captionStart="Fig" captionStartId="5.[113,148,2006,2027]" captionTargetBox="[157,1404,941,1982]" captionTargetId="figure-153@5.[151,1414,931,1991]" captionTargetPageId="5" captionText="Fig. 3. Bayesian consensus phylogram based on analyses of concatenated 16S and COI sequences, adapted from Köhler & Criscione (2013). Lesser Sunda species Rhagada solorensis, R. setzeri setzeri, R. setzeri atauroensis n. subsp., and R. abbasi n. sp. form the sister clade of Rhagada from the Australian Pilbara. Clades from the Australian Kimberley and Pilbara regions collapsed; outgroup pruned from tree; see Köhler & Criscione (2013) for further details." figureDoi="http://doi.org/10.5281/zenodo.4502142" httpUri="https://zenodo.org/record/4502142/files/figure.png" pageId="6" pageNumber="121">Fig. 3</figureCitation>
|
||
; Köhler & Criscione, 2013). Amongst these species,
|
||
<taxonomicName id="4C5ADDCAEF695E43FABF3A0CFC6CF9A0" authorityName="Köhler" authorityYear="2014" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="subSpecies" species="setzeri" subSpecies="atauroensis">
|
||
<emphasis id="B92E7A5BEF695E43FABF3A0CFC6CF9A0" italics="true" pageId="6" pageNumber="121">R. setzeri atauroensis</emphasis>
|
||
</taxonomicName>
|
||
forms the sister group of
|
||
<taxonomicName id="4C5ADDCAEF695E43FB083A6CFA55F9A0" authority="setzeri" box="[1238,1415,1607,1630]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="subSpecies" species="setzeri" subSpecies="setzeri">
|
||
<emphasis id="B92E7A5BEF695E43FB083A6CFA55F9A0" box="[1238,1415,1607,1630]" italics="true" pageId="6" pageNumber="121">R. setzeri setzeri</emphasis>
|
||
</taxonomicName>
|
||
from Alor. Both species are differentiated by mean p-distances of 4% in COI and 2% in 16S (
|
||
<tableCitation id="C6D893F2EF695E43FB473AACFB3DF960" box="[1177,1263,1671,1694]" captionStart="Table 2" captionStartId="6.[136,189,190,211]" captionTargetBox="[147,1457,275,706]" captionText="Table 2. Estimates of evolutionary divergence by means of uncorrected p-distances (base differences per site) within and between species" httpUri="http://table.plazi.org/id/DF25F6C1EF695E43FF563C95FDA8FF0E" pageId="6" pageNumber="121" tableUuid="DF25F6C1EF695E43FF563C95FDA8FF0E">Table 2</tableCitation>
|
||
). Other species are differentiated by p-distances of 11–16% in COI and 8–9% in 16S. These distances correspond well with amounts of interspecific variation found in Australian
|
||
<taxonomicName id="4C5ADDCAEF695E43FAD33ACCFABFF900" authorityName="Albers" authorityYear="1860" box="[1293,1389,1767,1790]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="genus">
|
||
<emphasis id="B92E7A5BEF695E43FAD33ACCFABFF900" box="[1293,1389,1767,1790]" italics="true" pageId="6" pageNumber="121">Rhagada</emphasis>
|
||
</taxonomicName>
|
||
species (about 4%;
|
||
<bibRefCitation id="EFCBDBB8EF695E43FC1C3B2CFB4EF8E0" author="Johnson MS & Hamilton ZR & Teale ROY & Kendrick PG" box="[962,1180,1799,1822]" pageId="6" pageNumber="121" pagination="316 - 327" refId="ref6053" refString="Johnson MS, Hamilton ZR, Teale ROY & Kendrick PG (2012) Endemic evolutionary radiation of Rhagada land snails (Pulmonata: Camaenidae) in a continental archipelago in northern Western Australia. Biological Journal of the Linnean Society, 106: 316 - 327." type="journal article" year="2012">Johnson et al., 2012</bibRefCitation>
|
||
) as well as other camaenid genera, such as
|
||
<taxonomicName id="4C5ADDCAEF695E43FC323B0CFB86F8C0" authorityName="Iredale" authorityYear="1939" box="[1004,1108,1831,1854]" class="Gastropoda" family="Camaenidae" genus="Exiligada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="genus">
|
||
<emphasis id="B92E7A5BEF695E43FC323B0CFB86F8C0" box="[1004,1108,1831,1854]" italics="true" pageId="6" pageNumber="121">Exiligada</emphasis>
|
||
</taxonomicName>
|
||
(about 3%;
|
||
<bibRefCitation id="EFCBDBB8EF695E43FB083B0CFA69F8C0" author="Criscione F & Law ML & Kohler F" box="[1238,1467,1831,1854]" pageId="6" pageNumber="121" pagination="689 - 722" refId="ref5706" refString="Criscione F, Law ML & Kohler F (2012) Land snail diversity in the monsoon tropics of Northern Australia: revision of the genus Exiligada Iredale, 1939 (Mollusca: Pulmonata: Camaenidae), with description of 13 new species. Zoological Journal of the Linnean Society, 166: 689 - 722." type="journal article" year="2012">Criscione et al., 2012</bibRefCitation>
|
||
) and
|
||
<taxonomicName id="4C5ADDCAEF695E43FCAA3B6CFBDDF8A0" authorityName="Iredale" authorityYear="1933" box="[884,1039,1863,1886]" class="Gastropoda" family="Camaenidae" genus="Amplirhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="genus">
|
||
<emphasis id="B92E7A5BEF695E43FCAA3B6CFBDDF8A0" box="[884,1039,1863,1886]" italics="true" pageId="6" pageNumber="121">Amplirhagada</emphasis>
|
||
</taxonomicName>
|
||
(about 6%;
|
||
<bibRefCitation id="EFCBDBB8EF695E43FB413B6CFA64F8A0" author="Kohler F & Johnson MS" box="[1183,1462,1863,1886]" pageId="6" pageNumber="121" pagination="337 - 362" refId="ref6157" refString="Kohler F & Johnson MS (2012) Species limits in molecular phylogenies: a cautionary tale from Australian land snails (Camaenidae: Amplirhagada). Zoological Journal of the Linnean Society, 165: 337 - 362." type="journal article" year="2012">Köhler & Johnson, 2012</bibRefCitation>
|
||
). However, extreme intraspecific distances of up to 10% have occasionally also been observed in camaenid snails (
|
||
<bibRefCitation id="EFCBDBB8EF695E43FAA43BACFCA9F840" author="Chiba S" pageId="6" pageNumber="121" pagination="460 - 471" refId="ref5598" refString="Chiba S (1999) Accelerated evolution of land snails Mandarina in the oceanic Bonin Islands: evidence from mitochondrial DNA sequences. Evolution, 53: 460 - 471." type="journal article" year="1999">Chiba, 1999</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFCBDBB8EF695E43FC573B8CFB61F840" author="Criscione F & Kohler F" box="[905,1203,1959,1982]" pageId="6" pageNumber="121" pagination="389 - 405" refId="ref5629" refString="Criscione F & Kohler F (2013 a) Conserved shell disguises diversity in Mesodontrachia land snails from the Australian Monsoon Tropics (Gastropoda: Camaenidae). Zoologica Scripta, 42: 389 - 405." type="journal article" year="2013">Criscione & Köhler, 2013a</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8BE5A649EF695E42FC9D3BCCFD36FECA" blockId="6.[835,1473,2023,2078]" lastBlockId="7.[113,752,189,308]" lastPageId="7" lastPageNumber="122" pageId="6" pageNumber="121">
|
||
Progress has been made towards a more complete documentation of the diversity of
|
||
<taxonomicName id="4C5ADDCAEF695E43FB02342CFA90F7E0" authorityName="Albers" authorityYear="1860" box="[1244,1346,2055,2078]" class="Gastropoda" family="Camaenidae" genus="Rhagada" kingdom="Animalia" order="Stylommatophora" pageId="6" pageNumber="121" phylum="Mollusca" rank="genus">
|
||
<emphasis id="B92E7A5BEF695E43FB02342CFA90F7E0" box="[1244,1346,2055,2078]" italics="true" pageId="6" pageNumber="121">Rhagada</emphasis>
|
||
</taxonomicName>
|
||
across the
|
||
<collectingRegion id="499E68ABEF685E42FFAF3C96FE8EFF2A" box="[113,348,189,212]" country="Indonesia" name="Nusa Tenggara Timur" pageId="7" pageNumber="122">Lesser Sunda Islands</collectingRegion>
|
||
. However, complete sampling on all
|
||
<collectingRegion id="499E68ABEF685E42FFAF3CF6FE8BFF0A" box="[113,345,221,244]" country="Indonesia" name="Nusa Tenggara Timur" pageId="7" pageNumber="122">Lesser Sunda Islands</collectingRegion>
|
||
is required to address the remaining taxonomic issues and to improve our understanding of the evolution of this group throughout the entire archipelago.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |