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<document ID-DOI="http://doi.org/10.5281/zenodo.6490793" ID-GBIF-Dataset="3cbc5830-b5eb-48c5-ab06-b6af21676fc4" ID-GBIF-Taxon="157487358" ID-Zenodo-Dep="6490793" checkinTime="1559991059721" checkinUser="plazi" docAuthor="Schilthuizen, Menno, Clavera, Anna Pazos, Khoo, Min Sheng, Bondar, Carin A., Elder, Charles H. S., Bouma, Aglaia M., Eddington, Taly D., Reisinger, Christian, Cosentino, Emanuela, Rossato, Marzia &amp; Delledonne, Massimo" docDate="2019" docId="033087ECD048EB65FCE44D11C0B7F94B" docLanguage="en" docName="jMolluscanStudies.85.1.35-40.pdf" docOrigin="Journal of Molluscan Studies 85" docStyle="DocumentStyle{}" docTitle="Microparmarion exquadratus Schilthuizen &amp; Clavera &amp; Khoo &amp; Bondar &amp; Elder &amp; Bouma &amp; Eddington &amp; Reisinger &amp; Cosentino &amp; Rossato &amp; Delledonne 2019, new species" docType="treatment" docVersion="5" lastPageId="3" lastPageNumber="38" masterDocId="FF09FF94D049EB66FFB34F1FC442FFBD" masterDocTitle="Bringing the lab to the field: a new lowland Microparmarion semi-slug (Gastropoda: Ariophantidae) described and DNA-barcoded in the forest" masterLastPageNumber="40" masterPageNumber="35" pageId="1" pageNumber="36" updateTime="1650976690827" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Bringing the lab to the field: a new lowland Microparmarion semi-slug (Gastropoda: Ariophantidae) described and DNA-barcoded in the forest</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Schilthuizen, Menno</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Clavera, Anna Pazos</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Khoo, Min Sheng</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bondar, Carin A.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Elder, Charles H. S.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bouma, Aglaia M.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Eddington, Taly D.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Reisinger, Christian</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Cosentino, Emanuela</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Rossato, Marzia</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Delledonne, Massimo</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>Journal of Molluscan Studies</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2019</mods:date>
<mods:detail type="pubDate">
<mods:number>2018-11-17</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>85</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>35</mods:start>
<mods:end>40</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1093/mollus/eyy052</mods:identifier>
<mods:identifier type="GBIF-Dataset">3cbc5830-b5eb-48c5-ab06-b6af21676fc4</mods:identifier>
<mods:identifier type="Zenodo-Dep">3241756</mods:identifier>
</mods:mods>
<treatment ID-DOI="http://doi.org/10.5281/zenodo.6490793" ID-GBIF-Taxon="157487358" ID-Zenodo-Dep="6490793" LSID="urn:lsid:plazi:treatment:033087ECD048EB65FCE44D11C0B7F94B" httpUri="http://treatment.plazi.org/id/033087ECD048EB65FCE44D11C0B7F94B" lastPageId="3" lastPageNumber="38" pageId="1" pageNumber="36">
<subSubSection pageId="1" pageNumber="36" type="nomenclature">
<paragraph blockId="1.[855,1466,526,622]" pageId="1" pageNumber="36">
<heading bold="true" fontSize="10" level="1" pageId="1" pageNumber="36" reason="0">
<emphasis bold="true" pageId="1" pageNumber="36">
<taxonomicName authority="Schilthuizen et al." authorityName="Schilthuizen &amp; Clavera &amp; Khoo &amp; Bondar &amp; Elder &amp; Bouma &amp; Eddington &amp; Reisinger &amp; Cosentino &amp; Rossato &amp; Delledonne" authorityYear="2019" box="[855,1459,526,550]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="1" pageNumber="36" phylum="Mollusca" rank="species" species="exquadratus" status="new species">
<emphasis bold="true" box="[855,1226,526,550]" italics="true" pageId="1" pageNumber="36">Microparmarion exquadratus</emphasis>
Schilthuizen
<emphasis bold="true" box="[1394,1466,526,550]" italics="true" pageId="1" pageNumber="36">et al.,</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1087,1235,558,582]" pageId="1" pageNumber="36" rank="species">new species</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection box="[1115,1207,600,622]" pageId="1" pageNumber="36" type="description">
<paragraph blockId="1.[855,1466,526,622]" box="[1115,1207,600,622]" pageId="1" pageNumber="36">
(
<figureCitation box="[1121,1201,600,622]" captionStart-0="Figure 2" captionStart-1="Figure 3" captionStart-2="Figure 4" captionStartId-0="2.[151,215,1686,1705]" captionStartId-1="3.[151,215,973,992]" captionStartId-2="3.[840,904,915,934]" captionTargetBox-0="[203,1430,152,1662]" captionTargetBox-1="[154,789,152,949]" captionTargetBox-2="[843,1478,152,891]" captionTargetId-0="figure@2.[203,1430,152,1662]" captionTargetId-1="figure@3.[154,789,152,949]" captionTargetId-2="figure@3.[843,1478,152,891]" captionTargetPageId-0="2" captionTargetPageId-1="3" captionTargetPageId-2="3" captionText-0="Figure 2. External morphology of (presumably) adult Microparmarion exquadratus n. sp. A preserved individual (syntype SP.13561) showing left lateral (A), right lateral (B), ventral (C) and dorsal (D) views. Six of the syntypes (EJ) in living state as photographed in the field in Tawau Hills Park, Sabah." captionText-1="Figure 3. Genital system of Microparmarion exquadratus n. sp. (syntype SP13560)." captionText-2="Figure 4. Dart sac of Microparmarion exquadratus n. sp. (syntype SP13560), cleared with clove oil and showing dart (shaded) in position." httpUri-0="https://zenodo.org/record/3241760/files/figure.png" httpUri-1="https://zenodo.org/record/3241764/files/figure.png" httpUri-2="https://zenodo.org/record/3241762/files/figure.png" pageId="1" pageNumber="36">Figs 24</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="36" type="materials_examined">
<paragraph blockId="1.[840,1482,659,894]" pageId="1" pageNumber="36">
<emphasis box="[840,891,659,680]" italics="true" pageId="1" pageNumber="36">Types</emphasis>
:
<materialsCitation ID-GBIF-Occurrence="2251688752" collectingDate="2018-03-06" collectingDateMax="2018-03-07" collectingDateMin="2018-03-06" collectingMethod="collected at night" collectionCode="SP" collectorName="taxon expedition participants" country="Malaysia" elevation="240" latitude="4.399317" location="vicinity of park headquarters, Tawau Hills Park" longLatPrecision="1" longitude="117.8893" pageId="1" pageNumber="36" specimenCode="SP13559, SP13551, SP13552-SP13554, SP13555-SP13557, SP13561, SP13560" specimenCount="10" specimenCount-adult="6" specimenCount-subadult="4" stateProvince="Sabah" typeStatus="holotype, paratype">
<typeStatus box="[906,1003,659,681]" pageId="1" pageNumber="36">Holotype</typeStatus>
:
<specimenCount box="[1015,1066,659,681]" pageId="1" pageNumber="36" type="adult">adult</specimenCount>
in 70% ethanol (
<specimenCode box="[1246,1339,659,681]" pageId="1" pageNumber="36">SP13559</specimenCode>
).
<typeStatus box="[1357,1458,659,681]" pageId="1" pageNumber="36">Paratypes</typeStatus>
:
<specimenCount pageId="1" pageNumber="36" type="subadult">1 subadult</specimenCount>
(
<specimenCode box="[952,1044,686,708]" pageId="1" pageNumber="36">SP13551</specimenCode>
), dissected and subsampled for DNA, in 70% ethanol;
<specimenCount box="[1032,1153,712,734]" pageId="1" pageNumber="36" type="subadult">3 subadults</specimenCount>
(
<specimenCode box="[1175,1367,712,734]" pageId="1" pageNumber="36">SP13552SP13554</specimenCode>
) in 100% ethanol, subsampled for DNA;
<specimenCount box="[1179,1263,739,761]" pageId="1" pageNumber="36" type="adult">4 adults</specimenCount>
(
<specimenCode box="[1283,1478,738,761]" pageId="1" pageNumber="36">SP13555SP13557</specimenCode>
,
<specimenCode box="[840,929,766,788]" pageId="1" pageNumber="36">SP13561</specimenCode>
) in 70% ethanol;
<specimenCount box="[1121,1193,766,788]" pageId="1" pageNumber="36" type="adult">1 adult</specimenCount>
<specimenCode box="[1203,1292,766,788]" pageId="1" pageNumber="36">SP13560</specimenCode>
dissected, in 70% ethanol. All material from
<location LSID="urn:lsid:plazi:treatment:033087ECD048EB65FCE44D11C0B7F94B:8E466021D048EB67FB324C07C040FCF4" country="Malaysia" latitude="4.399317" longLatPrecision="1" longitude="117.8893" name="vicinity of park headquarters, Tawau Hills Park" pageId="1" pageNumber="36" stateProvince="Sabah">vicinity of park headquarters, Tawau Hills Park</location>
,
<collectingRegion box="[1039,1104,819,841]" country="Malaysia" name="Sabah" pageId="1" pageNumber="36">Sabah</collectingRegion>
,
<collectingCountry box="[1118,1211,819,841]" name="Malaysia" pageId="1" pageNumber="36">Malaysia</collectingCountry>
,
<geoCoordinate box="[1224,1343,818,841]" degrees="4" direction="north" minutes="23.959" orientation="latitude" pageId="1" pageNumber="36" precision="1" value="4.399317">
4°23.959
<emphasis box="[1316,1323,818,840]" italics="true" pageId="1" pageNumber="36"></emphasis>
N
</geoCoordinate>
,
<geoCoordinate degrees="117" direction="east" minutes="53.358" orientation="longitude" pageId="1" pageNumber="36" precision="1" value="117.8893">
117°53.358
<emphasis box="[1475,1482,818,840]" italics="true" pageId="1" pageNumber="36"></emphasis>
E
</geoCoordinate>
;
<elevation box="[873,934,846,868]" metricMagnitude="2" metricUnit="m" metricValue="2.4" pageId="1" pageNumber="36" unit="m" value="240.0">
<quantity box="[873,934,846,868]" metricMagnitude="2" metricUnit="m" metricValue="2.4" pageId="1" pageNumber="36" unit="m" value="240.0">240 m</quantity>
</elevation>
asl,
<collectingMethod box="[988,1172,846,868]" pageId="1" pageNumber="36">collected at night</collectingMethod>
,
<collectingDate box="[1186,1364,845,868]" pageId="1" pageNumber="36" value="2018-03-06" valueMax="2018-03-07" valueMin="2018-03-06">
<date box="[1186,1364,845,868]" pageId="1" pageNumber="36" value="2018-03-06" valueMax="2018-03-07" valueMin="2018-03-06">67 March 2018</date>
</collectingDate>
; leg.
<collectorName pageId="1" pageNumber="36">taxon expedition participants</collectorName>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="36" type="nomenclature">
<paragraph blockId="1.[840,1482,925,974]" pageId="1" pageNumber="36">
<emphasis box="[840,1021,925,947]" italics="true" pageId="1" pageNumber="36">ƶoobank registration</emphasis>
: http://zoobank.org/B64843A4-7A74-4D07- AD96-96C160E6D80B
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="36" type="etymology">
<paragraph blockId="1.[840,1482,1005,1241]" pageId="1" pageNumber="36">
<emphasis box="[840,924,1005,1026]" italics="true" pageId="1" pageNumber="36">Etymology</emphasis>
: The name was chosen at a naming and voting session during the taxon expedition and refers to the squad of participants that jointly sampled this species during night-time walks in the forest. The taxonomic authority for this species is attributed to all authors of this paper. Following Article 51 C of the Code (
<bibRefCitation author="ICZN" box="[846,971,1139,1161]" journalOrPublisher="International Trust for Zoological Nomenclature, London" pageId="1" pageNumber="36" refId="ref3780" refString="ICZN. 1999. International code of zoological nomenclature. Edn 4. International Trust for Zoological Nomenclature, London." title="International code of zoological nomenclature. Edn 4" type="book" year="1999">ICZN, 1999</bibRefCitation>
), the species can be referred to as
<taxonomicName authority="Schilthuizen et al., 2018" authorityName="Schilthuizen" authorityYear="2018" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="1" pageNumber="36" phylum="Mollusca" rank="species" species="exquadratus">
<emphasis italics="true" pageId="1" pageNumber="36">Microparmarion exquadratus</emphasis>
Schilthuizen
<emphasis box="[1077,1123,1165,1187]" italics="true" pageId="1" pageNumber="36">et al.</emphasis>
, 2018
</taxonomicName>
, provided the full citation of this publication appears in the bibliography or elsewhere in the referring work.
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="36" type="diagnosis">
<paragraph blockId="1.[840,1482,1272,1428]" pageId="1" pageNumber="36">
<emphasis box="[840,919,1272,1293]" italics="true" pageId="1" pageNumber="36">Diagnosis</emphasis>
: Among
<taxonomicName authorityName="Simroth" authorityYear="1893" box="[1014,1147,1272,1293]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="1" pageNumber="36" phylum="Mollusca" rank="genus">
<emphasis box="[1014,1147,1272,1293]" italics="true" pageId="1" pageNumber="36">Microparmarion</emphasis>
</taxonomicName>
species of Borneo,
<emphasis box="[1344,1365,1272,1293]" italics="true" pageId="1" pageNumber="36">M</emphasis>
. ex
<emphasis box="[1401,1481,1272,1293]" italics="true" pageId="1" pageNumber="36">quadratus</emphasis>
is characterized by small size (less than half the size of
<taxonomicName authorityName="Collinge &amp; Godwin Austen" authorityYear="1895" box="[1369,1482,1298,1320]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="1" pageNumber="36" phylum="Mollusca" rank="species" species="pollonerai">
<emphasis box="[1369,1482,1298,1320]" italics="true" pageId="1" pageNumber="36">M. pollonerai</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Collinge &amp; Godwin Austen" authorityYear="1895" box="[887,989,1325,1347]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="1" pageNumber="36" phylum="Mollusca" rank="species" species="simrothi">
<emphasis box="[887,989,1325,1347]" italics="true" pageId="1" pageNumber="36">M. simrothi</emphasis>
</taxonomicName>
), three dark longitudinal stripes on head (shared with
<taxonomicName authorityName="Collinge &amp; Godwin Austen" authorityYear="1895" box="[890,990,1352,1374]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="1" pageNumber="36" phylum="Mollusca" rank="species" species="simrothi">
<emphasis box="[890,990,1352,1374]" italics="true" pageId="1" pageNumber="36">M. simrothi</emphasis>
</taxonomicName>
, but lacking in
<taxonomicName authorityName="Collinge &amp; Godwin Austen" authorityYear="1895" box="[1147,1261,1352,1374]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="1" pageNumber="36" phylum="Mollusca" rank="species" species="pollonerai">
<emphasis box="[1147,1261,1352,1374]" italics="true" pageId="1" pageNumber="36">M. pollonerai</emphasis>
</taxonomicName>
and
<emphasis box="[1313,1418,1352,1374]" italics="true" pageId="1" pageNumber="36">M. litteratus</emphasis>
), dark dorsal stripe on tail, kink in dart sac (shared with
<taxonomicName authorityName="Collinge &amp; Godwin Austen" authorityYear="1895" box="[1320,1433,1378,1400]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="1" pageNumber="36" phylum="Mollusca" rank="species" species="pollonerai">
<emphasis box="[1320,1433,1378,1400]" italics="true" pageId="1" pageNumber="36">M. pollonerai</emphasis>
</taxonomicName>
) and reduced receptaculum.
</paragraph>
</subSubSection>
<subSubSection lastPageId="3" lastPageNumber="38" pageId="1" pageNumber="36" type="description">
<paragraph blockId="1.[840,1483,1458,1961]" pageId="1" pageNumber="36">
<emphasis box="[840,937,1459,1480]" italics="true" pageId="1" pageNumber="36">Description:</emphasis>
Exterior in adult, preserved specimens (
<figureCitation box="[1363,1471,1458,1481]" captionStart="Figure 2" captionStartId="2.[151,215,1686,1705]" captionTargetBox="[203,1430,152,1662]" captionTargetId="figure@2.[203,1430,152,1662]" captionTargetPageId="2" captionText="Figure 2. External morphology of (presumably) adult Microparmarion exquadratus n. sp. A preserved individual (syntype SP.13561) showing left lateral (A), right lateral (B), ventral (C) and dorsal (D) views. Six of the syntypes (EJ) in living state as photographed in the field in Tawau Hills Park, Sabah." httpUri="https://zenodo.org/record/3241760/files/figure.png" pageId="1" pageNumber="36">Fig. 2AD</figureCitation>
): Body length 1821 mm. Shell 8.59 mm long, 5 mm wide. Body dull, greyish pink, 3 black dorsal longitudinal stripes, separated by 2 white stripes starting immediately caudal of black eye tentacles and ending ventrally of frontal edge of mantle. Lateral black stripes are half breadth of dorsal stripe. Mantle may also be dull greyish pink, with dark grey mottling. Keeled tail approximately one-third total body length (56 mm in adults), with 1 black dorsal stripe. Margin of foot has vertical, dark grey striated pattern. Sole of foot off-white. Tail with caudal horn, not very prominent. Shell semi-transparent, smooth, thin, pale yellow, with faint trace of spiral on right side, approximately 2 whorls. Genital opening visible on right side as short vertical slit. Dark pattern on mantle of 2, nearly continuous lines encircling shell, which are separated by pale greyish-pink ridge with about 10 dark spots.
</paragraph>
<paragraph blockId="1.[840,1483,1458,1961]" lastBlockId="2.[151,794,1779,1961]" lastPageId="2" lastPageNumber="37" pageId="1" pageNumber="36">
Live specimens (
<figureCitation box="[1022,1113,1858,1881]" captionStart="Figure 2" captionStartId="2.[151,215,1686,1705]" captionTargetBox="[203,1430,152,1662]" captionTargetId="figure@2.[203,1430,152,1662]" captionTargetPageId="2" captionText="Figure 2. External morphology of (presumably) adult Microparmarion exquadratus n. sp. A preserved individual (syntype SP.13561) showing left lateral (A), right lateral (B), ventral (C) and dorsal (D) views. Six of the syntypes (EJ) in living state as photographed in the field in Tawau Hills Park, Sabah." httpUri="https://zenodo.org/record/3241760/files/figure.png" pageId="1" pageNumber="36">Fig. 2EJ</figureCitation>
): Colour dull orange with deep brown pattern. Mantle lobes meet along curved suture, bulging to left, lying left of midline of body. Shell lobes with slight granulation. Frontal part of mantle with slight longitudinal ridges. Caudal horn triangular, slightly longer than wide, pointing caudally. Eye tentacles, when fully extended, as long as shell.
</paragraph>
<caption httpUri="https://zenodo.org/record/3241760/files/figure.png" pageId="2" pageNumber="37" startId="2.[151,215,1686,1705]" targetBox="[203,1430,152,1662]" targetPageId="2">
<paragraph blockId="2.[151,1482,1686,1730]" pageId="2" pageNumber="37">
<emphasis bold="true" box="[151,241,1686,1705]" pageId="2" pageNumber="37">Figure 2.</emphasis>
External morphology of (presumably) adult
<taxonomicName authority="Schilthuizen &amp; Clavera &amp; Khoo &amp; Bondar &amp; Elder &amp; Bouma &amp; Eddington &amp; Reisinger &amp; Cosentino &amp; Rossato &amp; Delledonne, 2019" authorityName="Schilthuizen &amp; Clavera &amp; Khoo &amp; Bondar &amp; Elder &amp; Bouma &amp; Eddington &amp; Reisinger &amp; Cosentino &amp; Rossato &amp; Delledonne" authorityYear="2019" box="[632,844,1687,1707]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="2" pageNumber="37" phylum="Mollusca" rank="species" species="exquadratus" status="n. sp.">
<emphasis box="[632,844,1687,1707]" italics="true" pageId="2" pageNumber="37">Microparmarion exquadratus</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[851,900,1687,1706]" pageId="2" pageNumber="37" rank="species">n. sp.</taxonomicNameLabel>
A preserved individual (syntype SP.13561) showing left lateral (
<emphasis bold="true" box="[1458,1471,1686,1705]" pageId="2" pageNumber="37">A</emphasis>
), right lateral (
<emphasis bold="true" box="[262,277,1711,1730]" pageId="2" pageNumber="37">B</emphasis>
), ventral (
<emphasis bold="true" box="[363,379,1710,1729]" pageId="2" pageNumber="37">C</emphasis>
) and dorsal (
<emphasis bold="true" box="[490,507,1711,1730]" pageId="2" pageNumber="37">D</emphasis>
) views. Six of the syntypes (
<emphasis bold="true" box="[744,777,1711,1730]" pageId="2" pageNumber="37">EJ</emphasis>
) in living state as photographed in the field in Tawau Hills Park, Sabah.
</paragraph>
</caption>
<paragraph blockId="2.[151,794,1779,1961]" lastBlockId="3.[151,794,1086,1214]" lastPageId="3" lastPageNumber="38" pageId="2" pageNumber="37">
Genitalia (
<figureCitation box="[282,344,1832,1854]" captionStart="Figure 3" captionStartId="3.[151,215,973,992]" captionTargetBox="[154,789,152,949]" captionTargetId="figure@3.[154,789,152,949]" captionTargetPageId="3" captionText="Figure 3. Genital system of Microparmarion exquadratus n. sp. (syntype SP13560)." httpUri="https://zenodo.org/record/3241764/files/figure.png" pageId="2" pageNumber="37">Figs 3</figureCitation>
,
<figureCitation box="[362,374,1832,1854]" captionStart="Figure 4" captionStartId="3.[840,904,915,934]" captionTargetBox="[843,1478,152,891]" captionTargetId="figure@3.[843,1478,152,891]" captionTargetPageId="3" captionText="Figure 4. Dart sac of Microparmarion exquadratus n. sp. (syntype SP13560), cleared with clove oil and showing dart (shaded) in position." httpUri="https://zenodo.org/record/3241762/files/figure.png" pageId="2" pageNumber="37">4</figureCitation>
): Atrium wide. Penis doubly folded into Z-shape, proximally narrow (one-quarter width of atrium), gradually doubling in width, then narrowing to original width distally. Penis sharply folding back on itself as a very narrow tube until half length of proximal part of penis, then folding, broadening, eventually tapering into vas deferens. When following the vas deferens from where it departs from penis, it remains detached from free oviduct until level with middle of dart sac. Vagina short, receptaculum reduced, externally invisible, only discernible after clearing in clove oil. Free oviduct irregularly looped and coiled. Dart sac long, equivalent to combined length of vagina and free oviduct, doubly folded into a Z-shape. Basal portion of dart sac is narrow, containing the dart; middle portion half as long as basal portion, externally somewhat granulate; distal portion inflated, sausage-shaped, 3 times as long as wide, flattened on one side. Dart straight, with gently tapering point, possibly with several low, longitudinal flanges; crown twice as wide as dart.
</paragraph>
<caption httpUri="https://zenodo.org/record/3241762/files/figure.png" pageId="3" pageNumber="38" startId="3.[840,904,915,934]" targetBox="[843,1478,152,891]" targetPageId="3">
<paragraph blockId="3.[840,1483,915,958]" pageId="3" pageNumber="38">
<emphasis bold="true" box="[840,931,915,934]" pageId="3" pageNumber="38">Figure 4.</emphasis>
Dart sac of
<taxonomicName authority="Schilthuizen &amp; Clavera &amp; Khoo &amp; Bondar &amp; Elder &amp; Bouma &amp; Eddington &amp; Reisinger &amp; Cosentino &amp; Rossato &amp; Delledonne, 2019" authorityName="Schilthuizen &amp; Clavera &amp; Khoo &amp; Bondar &amp; Elder &amp; Bouma &amp; Eddington &amp; Reisinger &amp; Cosentino &amp; Rossato &amp; Delledonne" authorityYear="2019" box="[1044,1256,915,935]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="3" pageNumber="38" phylum="Mollusca" rank="species" species="exquadratus" status="n. sp.">
<emphasis box="[1044,1256,915,935]" italics="true" pageId="3" pageNumber="38">Microparmarion exquadratus</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1263,1312,915,934]" pageId="3" pageNumber="38" rank="species">n. sp.</taxonomicNameLabel>
(syntype SP13560), cleared with clove oil and showing dart (shaded) in position.
</paragraph>
</caption>
<caption httpUri="https://zenodo.org/record/3241764/files/figure.png" pageId="3" pageNumber="38" startId="3.[151,215,973,992]" targetBox="[154,789,152,949]" targetPageId="3">
<paragraph blockId="3.[151,792,973,1017]" pageId="3" pageNumber="38">
<emphasis bold="true" box="[151,246,973,992]" pageId="3" pageNumber="38">Figure 3.</emphasis>
Genital system of
<taxonomicName authority="Schilthuizen &amp; Clavera &amp; Khoo &amp; Bondar &amp; Elder &amp; Bouma &amp; Eddington &amp; Reisinger &amp; Cosentino &amp; Rossato &amp; Delledonne, 2019" authorityName="Schilthuizen &amp; Clavera &amp; Khoo &amp; Bondar &amp; Elder &amp; Bouma &amp; Eddington &amp; Reisinger &amp; Cosentino &amp; Rossato &amp; Delledonne" authorityYear="2019" box="[431,647,974,994]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="3" pageNumber="38" phylum="Mollusca" rank="species" species="exquadratus" status="n. sp.">
<emphasis box="[431,647,974,994]" italics="true" pageId="3" pageNumber="38">
<taxonomicName box="[431,549,974,994]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="CoL" kingdom="Animalia" order="Stylommatophora" pageId="3" pageNumber="38" phylum="Mollusca" rank="genus">Microparmarion</taxonomicName>
<taxonomicName box="[561,647,974,994]" pageId="3" pageNumber="38" rank="species" species="exquadratus" status="n. sp.">exquadratus</taxonomicName>
</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[659,713,974,993]" pageId="3" pageNumber="38" rank="species">n. sp.</taxonomicNameLabel>
(syntype SP13560).
</paragraph>
</caption>
<paragraph blockId="3.[151,794,1245,1614]" pageId="3" pageNumber="38">
<emphasis box="[151,273,1245,1266]" italics="true" pageId="3" pageNumber="38">DNA barcode</emphasis>
: We obtained 3 sequences for the
<emphasis box="[678,716,1245,1266]" italics="true" pageId="3" pageNumber="38">COI</emphasis>
DNAbarcoding region. These sequences were identical, with the following exceptions when compared with the generated consensus (i.e. the most frequent nucleotide at each position among the three sequences). The sequence for SP13554 lacked a TA dinucleotide at the 5 end, which might be an artefact; we have replaced these nucleotides by unknown bases in the BOLD sequence. The sequence for SP13553 had an A instead of G at position 268, a T instead of C at position 628, and a C instead of T at position 637. The three sequences form a well-supported clade in the tree (
<figureCitation box="[158,212,1512,1534]" captionStart="Figure 5" captionStartId="4.[151,215,1215,1234]" captionTargetBox="[202,1430,156,1189]" captionTargetId="graphics@4.[202,877,160,1131]" captionTargetPageId="4" captionText="Figure 5. Phylogenetic reconstruction for Bornean Microparmarion based on COI barcode locus, with Parmarion as outgroup. Numbers on nodes are Bayesian posterior probabilities. Registration numbers in parentheses: RMNH, Naturalis Biodiversity Center, Leiden, The Netherlands; SP, Sabah Parks collection; FJ481180 is a GenBank accession number. Scale bar indicates substitutions per site." httpUri="https://zenodo.org/record/3241766/files/figure.png" pageId="3" pageNumber="38">Fig. 5</figureCitation>
), which is clearly separated (by&gt;10% sequence divergence) from all other
<emphasis box="[307,490,1539,1560]" italics="true" pageId="3" pageNumber="38">
<taxonomicName authorityName="Simroth" authorityYear="1893" box="[307,440,1539,1560]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="3" pageNumber="38" phylum="Mollusca" rank="genus">Microparmarion</taxonomicName>
COI
</emphasis>
sequences available (all from Sabah as well). Sequences for
<emphasis box="[474,584,1565,1587]" italics="true" pageId="3" pageNumber="38">M. litteratus</emphasis>
were not available.
<taxonomicName authorityName="Schilthuizen &amp; Clavera &amp; Khoo &amp; Bondar &amp; Elder &amp; Bouma &amp; Eddington &amp; Reisinger &amp; Cosentino &amp; Rossato &amp; Delledonne" authorityYear="2019" box="[151,388,1592,1613]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="3" pageNumber="38" phylum="Mollusca" rank="species" species="exquadratus">
<emphasis box="[151,388,1592,1613]" italics="true" pageId="3" pageNumber="38">Microparmarion exquadratus</emphasis>
</taxonomicName>
has been assigned BIN ADM3588.
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="38" type="distribution">
<paragraph blockId="3.[151,793,1645,1961]" lastBlockId="3.[840,1482,1013,1089]" pageId="3" pageNumber="38">
<emphasis box="[151,249,1645,1666]" italics="true" pageId="3" pageNumber="38">Distribution</emphasis>
: So far only known from lowland dipterocarp forest around the headquarters of Tawau Hills Park (
<emphasis box="[618,633,1672,1693]" italics="true" pageId="3" pageNumber="38">c.</emphasis>
240 m asl). It is surprising to find a
<taxonomicName authorityName="Simroth" authorityYear="1893" box="[353,486,1699,1720]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="3" pageNumber="38" phylum="Mollusca" rank="genus">
<emphasis box="[353,486,1699,1720]" italics="true" pageId="3" pageNumber="38">Microparmarion</emphasis>
</taxonomicName>
at such a low elevation, since the genus is normally found in montane forest—it is therefore also not sympatric with any other known
<taxonomicName authorityName="Simroth" authorityYear="1893" box="[513,646,1752,1773]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="3" pageNumber="38" phylum="Mollusca" rank="genus">
<emphasis box="[513,646,1752,1773]" italics="true" pageId="3" pageNumber="38">Microparmarion</emphasis>
</taxonomicName>
species. There is a slight possibility that the population is nonindigenous to the lowlands, since the park s botanical garden (stocked with plants from elsewhere in the park, including the highlands) is nearby. The semi-slug may thus have been introduced from a higherelevation habitat in the surrounding area. However, we failed to find the species on nearby Mount Lucia (1,300 m asl) and we also did not find it in the botanic garden itself, so we tentatively conclude that it is indigenous and possibly widespread throughout Tawau Hills Park. To confirm this, nocturnal sampling will have to be carried out in other parts of the park.
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="38" type="discussion">
<paragraph blockId="3.[840,1483,1119,1782]" pageId="3" pageNumber="38">
<emphasis box="[840,910,1119,1140]" italics="true" pageId="3" pageNumber="38">Remarks</emphasis>
: In colour pattern,
<taxonomicName authorityName="Schilthuizen &amp; Clavera &amp; Khoo &amp; Bondar &amp; Elder &amp; Bouma &amp; Eddington &amp; Reisinger &amp; Cosentino &amp; Rossato &amp; Delledonne" authorityYear="2019" box="[1106,1238,1119,1141]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="3" pageNumber="38" phylum="Mollusca" rank="species" species="exquadratus">
<emphasis box="[1106,1238,1119,1141]" italics="true" pageId="3" pageNumber="38">M. exquadratus</emphasis>
</taxonomicName>
is somewhat similar to a
<taxonomicName authorityName="Simroth" authorityYear="1893" box="[840,973,1147,1168]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="3" pageNumber="38" phylum="Mollusca" rank="genus">
<emphasis box="[840,973,1147,1168]" italics="true" pageId="3" pageNumber="38">Microparmarion</emphasis>
</taxonomicName>
species found at an elevation of 1,200 m asl in Long Pa Sia, southwestern Sabah, and also to a species photographed (but not collected) at 1,000 m asl in Penrissen, Sarawak (
<bibRefCitation author="SCHILTHUIZEN, M." box="[846,1026,1227,1249]" editor="J. Mohd-Azlan" journalOrPublisher="Natural History Publications (Borneo), Kota Kinabalu, Sabah" pageId="3" pageNumber="38" pagination="85 - 94" refId="ref4282" refString="SCHILTHUIZEN, M. 2017. Land snails and slugs. In: Gunung Penrissen- the roof of aeestern Borneo: life from the headaeaters to the coast (J. Mohd-Azlan, A. A. Tuen &amp; I. Das, eds), pp. 85 - 94. Natural History Publications (Borneo), Kota Kinabalu, Sabah." title="Land snails and slugs" type="book chapter" volumeTitle="Gunung Penrissen- the roof of aeestern Borneo: life from the headaeaters to the coast" year="2017">Schilthuizen, 2017</bibRefCitation>
). However, it is unlikely to be conspecific with either, because it differs in colour pattern (the Penrissen specimen has two, rather than three, head stripes, and the Long Pa Sia specimen lacks the black markings on the mantle lobes) and is larger in size. For the time being, therefore, we consider the
<taxonomicName authorityName="Schilthuizen &amp; Clavera &amp; Khoo &amp; Bondar &amp; Elder &amp; Bouma &amp; Eddington &amp; Reisinger &amp; Cosentino &amp; Rossato &amp; Delledonne" authorityYear="2019" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="3" pageNumber="38" phylum="Mollusca" rank="species" species="exquadratus">
<emphasis italics="true" pageId="3" pageNumber="38">M. exquadratus</emphasis>
</taxonomicName>
to be restricted to Tawau Hills Park. Our phylogenetic reconstruction (
<figureCitation box="[938,992,1387,1409]" captionStart="Figure 5" captionStartId="4.[151,215,1215,1234]" captionTargetBox="[202,1430,156,1189]" captionTargetId="graphics@4.[202,877,160,1131]" captionTargetPageId="4" captionText="Figure 5. Phylogenetic reconstruction for Bornean Microparmarion based on COI barcode locus, with Parmarion as outgroup. Numbers on nodes are Bayesian posterior probabilities. Registration numbers in parentheses: RMNH, Naturalis Biodiversity Center, Leiden, The Netherlands; SP, Sabah Parks collection; FJ481180 is a GenBank accession number. Scale bar indicates substitutions per site." httpUri="https://zenodo.org/record/3241766/files/figure.png" pageId="3" pageNumber="38">Fig. 5</figureCitation>
), although primarily intended to assess the genetic distinctness of the new species, also suggests that the genus
<taxonomicName authorityName="Simroth" authorityYear="1893" box="[840,973,1440,1461]" class="Gastropoda" family="Ariophantidae" genus="Microparmarion" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Stylommatophora" pageId="3" pageNumber="38" phylum="Mollusca" rank="genus">
<emphasis box="[840,973,1440,1461]" italics="true" pageId="3" pageNumber="38">Microparmarion</emphasis>
</taxonomicName>
consists of multiple clades that are not fully congruent with named species. We therefore recommend that the genus be subjected to a formal revision.
</paragraph>
<paragraph blockId="3.[840,1483,1119,1782]" pageId="3" pageNumber="38">We must also stress that our work was conducted under several limitations and constraints. It was part of a full 10-day field course schedule, carried out by untrained citizen scientists in a field station with limited equipment and erratic electricity supply. Specimens could not be exported. Consequently, the extent of our work is less than is customary for descriptions of new semi-slug species: we sequenced only three individuals and dissected only one. Also, all individuals were obtained from a very small geographic area. Nonetheless, the genetic and morphological features are suffi- cient to recognize the species unambiguously.</paragraph>
</subSubSection>
</treatment>
</document>