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<document ID-DOI="http://dx.doi.org/10.3897/subtbiol.11.5966" ID-GBIF-Dataset="c61994de-8f0e-4151-9798-aea3d7fac0bd" ID-GBIF-Taxon="127877125" ID-Pensoft-Pub="1314-2615-11-45" LSID="urn:lsid:zoobank.org:act:A3B380AD-B918-451B-866D-7FA59642A733" ModsDocAuthor="" ModsDocDate="2013" ModsDocID="1314-2615-11-45" ModsDocOrigin="Subterranean Biology 11" ModsDocTitle="New Zospeum species (Gastropoda, Ellobioidea, Carychiidae) from 980 m depth in the Lukina Jama Trojama cave system (Velebit Mts., Croatia)" checkinTime="1425841342458" checkinUser="pensoft" docAuthor="Weigand, Alexander M." docDate="2013" docId="278693CF2B7098EC533D9E7DE9D285E4" docLanguage="en" docName="SubterBiol 11: 45-53" docOrigin="Subterranean Biology 11" docSource="http://dx.doi.org/10.3897/subtbiol.11.5966" docTitle="Zospeum tholussum Weigand, sp. n." docType="treatment" docUuid="A3B380AD-B918-451B-866D-7FA59642A733" docUuidSource="ZooBank" docVersion="5" lastPageNumber="45" masterDocId="FFE2FFA3FFCCFFA3FF8DFFC98D640E1B" masterDocTitle="New Zospeum species (Gastropoda, Ellobioidea, Carychiidae) from 980 m depth in the Lukina Jama - Trojama cave system (Velebit Mts., Croatia)" masterLastPageNumber="53" masterPageNumber="45" pageId="0" pageNumber="45" updateTime="1643696289190" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>New Zospeum species (Gastropoda, Ellobioidea, Carychiidae) from 980 m depth in the Lukina Jama - Trojama cave system (Velebit Mts., Croatia)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Weigand, Alexander M.</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:titleInfo>
<mods:title>Subterranean Biology</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2013</mods:date>
<mods:detail type="volume">
<mods:number>11</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>45</mods:start>
<mods:end>53</mods:end>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/subtbiol.11.5966</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/subtbiol.11.5966</mods:identifier>
<mods:identifier type="Pensoft-Pub">1314-2615-11-45</mods:identifier>
<mods:identifier type="ZooBank">5F4196B5-E479-4F16-A87A-9808FA8521E3</mods:identifier>
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<treatment ID-GBIF-Taxon="127877125" LSID="urn:lsid:zoobank.org:act:A3B380AD-B918-451B-866D-7FA59642A733" httpUri="http://treatment.plazi.org/id/278693CF2B7098EC533D9E7DE9D285E4" lastPageNumber="45" pageId="0" pageNumber="45">
<subSubSection pageId="0" pageNumber="45" type="nomenclature">
<paragraph pageId="0" pageNumber="45">
<taxonomicName LSID="http://zoobank.org/A3B380AD-B918-451B-866D-7FA59642A733" authority="Weigand" class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum tholussum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="tholussum">Zospeum tholussum Weigand</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="45">sp. n.</taxonomicNameLabel>
Figs 1-4
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="material examined">
<paragraph pageId="0" pageNumber="45">Material examined.</paragraph>
<paragraph pageId="0" pageNumber="45">A single living specimen was collected on 31.07.2010 (leg. Jana Bedek), which is designated as the holotype specimen (BOLD-ID BARCA210-13) (Fig. 1, life image; Fig. 2, upper row). Eight shells comprise paratype specimens, of which one was partially broken and used to investigate the form of the columellar fold. Shells were collected on several days during the caving expedition from 29.07. - 03.08.2010 (Fig. 2). All investigated specimens are deposited in the Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main, Germany (museum voucher SMF 341633).</paragraph>
<caption pageId="0" pageNumber="45">
<paragraph pageId="0" pageNumber="45">
Figure 2. Holotype and paratypes of
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum tholussum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="tholussum">Zospeum tholussum</taxonomicName>
. The holotype (former living specimen) is marked with a solid line; five paratype specimens (shells) are surrounded by dotted lines.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="diagnosis">
<paragraph pageId="0" pageNumber="45">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="45">
The general shape of
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum tholussum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="tholussum">Zospeum tholussum</taxonomicName>
sp. n. resembles larger specimens of
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum amoenum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="amoenum">Zospeum amoenum</taxonomicName>
(Frauenfeld, 1856) from which it can be best delineated by the dome-like structured 2nd whorl (Fig. 2), the weak but present columellar fold (Fig. 3) and by means of their DNA barcodes (11.7 %-12.1 % genetic p-distance between both species).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="description">
<paragraph pageId="0" pageNumber="45">Description.</paragraph>
<paragraph pageId="0" pageNumber="45">
The eight conchologically investigated specimens (the holotype and seven paratypes) demonstrate moderate shell variability (Table 1). Shells are generally smooth, very thin and fragile. Fresh shells or those of living specimens are translucent, older ones adopt a milky white color. The number of whorls are 5-6, SH ranges from 1.41-1.81 mm, SW from 0.93-1.12 mm, AH from 0.44-0.54 mm and AW from 0.38-0.46 mm. The ratio of SH/SW is between 1.34-1.62 and AH/AW between 1.05-1.30. The 2nd whorl is remarkably enlarged possessing a height of 2/3-5/6 the height of the 3rd + 4th whorl. The columellar is only weakly developed (Fig. 3) and no teeth (i.e. parietalis, palatalis or columellaris) are present. The DNA barcode of the holotype, when compared with other DNA barcodes of
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="genus">Zospeum</taxonomicName>
species deposited in BOLD, shows its lowest interspecific genetic distance to
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum pretneri" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="pretneri">Zospeum pretneri</taxonomicName>
Bole, 1960 (5.6 %; BOLD-ID: BARCA120-10). This result is well above the barcoding gap of 3.2 % for
<taxonomicName family="Carychiidae" lsidName="" pageId="0" pageNumber="45" rank="family">Carychiidae</taxonomicName>
, which is suitable to separate between-species (&gt; 3.2 %, interspecific) and within-species genetic diversity (&lt;3.2 %, intraspecific) in
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="genus">Zospeum</taxonomicName>
(
<bibRefCitation author="Weigand, AM" journalOrPublisher="Molecular Ecology Resources" pageId="0" pageNumber="45" pagination="255 - 265" title="A new approach to an old conundrum - DNA barcoding sheds new light on phenotypic plasticity and morphological stasis in microsnails (Gastropoda, Pulmonata, Carychiidae)." url="10.1111/j.1755-0998.2010.02937.x" volume="11" year="2011">Weigand et al. 2011</bibRefCitation>
,
<bibRefCitation author="Weigand, AM" journalOrPublisher="BMC Evolutionary Biology" pageId="0" pageNumber="45" title="Evolution of microgastropods (Ellobioidea, Carychiidae): integrating taxonomic, phylogenetic and evolutionary hypotheses." url="10.1186/1471-2148-13-18" volume="13" year="2013">2013</bibRefCitation>
).
</paragraph>
<caption ID-Table-UUID="F60F30150E5DB71A08D4F45CE5EA393C" httpUri="http://table.plazi.org/id/F60F30150E5DB71A08D4F45CE5EA393C" pageId="0" pageNumber="45">
<paragraph pageId="0" pageNumber="45">
Table 1. Conchological measurements for
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum tholussum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="tholussum">Zospeum tholussum</taxonomicName>
. Seven paratypes (Ind. 1-7) and the holotype specimen (Ind. 8) were measured. All measurements are given in millimeter. Additionally, ratios of shell height / shell width (sh/sw) and aperture height / aperture width (ah/aw) were calculated. The range of variability is given for all parameters.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="45">
<table pageId="0" pageNumber="45">
<tr pageId="0" pageNumber="45">
<th colspan="1" pageId="0" pageNumber="45" rowspan="1">Ind.</th>
<th colspan="1" pageId="0" pageNumber="45" rowspan="1">sh/sw</th>
<th colspan="1" pageId="0" pageNumber="45" rowspan="1">ah/aw</th>
<th colspan="1" pageId="0" pageNumber="45" rowspan="1">shell height</th>
<th colspan="1" pageId="0" pageNumber="45" rowspan="1">shell width</th>
<th colspan="1" pageId="0" pageNumber="45" rowspan="1">aperture height</th>
<th colspan="1" pageId="0" pageNumber="45" rowspan="1">aperture width</th>
</tr>
</table>
</paragraph>
<caption pageId="0" pageNumber="45">
<paragraph pageId="0" pageNumber="45">
Figure 3. Columellar fold views of
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum tholussum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="tholussum">Zospeum tholussum</taxonomicName>
. The columellar fold of the broken paratype specimen shell is shown in clockwise rotation.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="habitat">
<paragraph pageId="0" pageNumber="45">Habitat.</paragraph>
<paragraph pageId="0" pageNumber="45">The single living specimen was found in an unnamed large chamber at 980 m depth (85 m long, 70 m wide) with lots of stones, rocks and sand (Fig. 1, right site). A temporal small stream of running water was present close to the collecting site. Air temperature was between 3.3-3.5 °C (depending on the measurement device), water temperature 5.1 °C and air humidity 100 %. Shells were observed beginning from 800 m depth till the bottom of the cave. Shells were generally found on layers of mud (Fig. 1, left site). The first 200 m of the entrance passage of Lukina Jama are permanently covered by varying levels of snow and ice.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="etymology">
<paragraph pageId="0" pageNumber="45">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="45">The Latin word tholus means dome or cupola and refers to the remarkable dome-like shape of the 2nd whorl.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="45" type="remarks">
<paragraph pageId="0" pageNumber="45">Remarks.</paragraph>
<paragraph pageId="0" pageNumber="45">
In addition to the newly described
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum tholussum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="tholussum">Zospeum tholussum</taxonomicName>
, a second
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="genus">Zospeum</taxonomicName>
species is present in the Lukina
<normalizedToken originalValue="JamaTrojama">Jama-Trojama</normalizedToken>
cave system (Fig. 4C). This species can be differentiated from
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum tholussum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="tholussum">Zospeum tholussum</taxonomicName>
by the presence of a tooth, its general shell shape, more prominent columellar fold and absence of the characteristic dome-like structured 2nd whorl. Because only shells were found, no DNA barcodes are available for this species.
</paragraph>
<caption pageId="0" pageNumber="45">
<paragraph pageId="0" pageNumber="45">
Figure 4.
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="genus">Zospeum</taxonomicName>
species. A
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum tholussum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="tholussum">Zospeum tholussum</taxonomicName>
holotype B
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum pretneri" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="pretneri">Zospeum pretneri</taxonomicName>
from locus typicus, which demonstrates the lowest genetic distance to
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum tholussum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="tholussum">Zospeum tholussum</taxonomicName>
C Second
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="genus">Zospeum</taxonomicName>
sp. from the Lukina
<normalizedToken originalValue="JamaTrojama">Jama-Trojama</normalizedToken>
cave system.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="45">
So far,
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum tholussum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="tholussum">Zospeum tholussum</taxonomicName>
is only known from the Lukina
<normalizedToken originalValue="JamaTrojama">Jama-Trojama</normalizedToken>
cave system. However, this cave system is situated in the distribution range of the morphologically related species
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum amoenum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="amoenum">Zospeum amoenum</taxonomicName>
(Frauenfeld, 1856), which inhabits caves of the West Balkan of North Slovenia, West Croatia, Bosnia and Herzegovina and South to Montenegro (
<bibRefCitation author="Bole, J" journalOrPublisher="Razprave" pageId="0" pageNumber="45" pagination="1 - 43" title="Rod Zospeum Bourguignat 1856 (Gastropoda, Ellobiidae) v Jugoslaviji. Slovenska Akademija Znanosti in Umetnosti." volume="17" year="1974">Bole 1974</bibRefCitation>
).
<bibRefCitation author="Absolon, K" journalOrPublisher="Zlata Praha" pageId="0" pageNumber="45" title="Z vyzkumnyh cest po kraseh balkanu." volume="33" year="1916">Absolon (1916)</bibRefCitation>
named but did not describe another morphologically related species -
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum troglobalcanicum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="troglobalcanicum">Zospeum troglobalcanicum</taxonomicName>
. Since
<bibRefCitation author="Weigand, AM" journalOrPublisher="BMC Evolutionary Biology" pageId="0" pageNumber="45" title="Evolution of microgastropods (Ellobioidea, Carychiidae): integrating taxonomic, phylogenetic and evolutionary hypotheses." url="10.1186/1471-2148-13-18" volume="13" year="2013">Weigand et al. (2013)</bibRefCitation>
revealed that
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="genus">Zospeum</taxonomicName>
spp. with an inferred large distribution range are particularly prone to host additional, morphologically unrecognized species, some of the distribution records for
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum amoenum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="amoenum">Zospeum amoenum</taxonomicName>
may very well refer to
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum tholussum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="tholussum">Zospeum tholussum</taxonomicName>
. Moreover, the intraspecific genetic and conchological variability of the new species should be investigated in greater detail, thus to judge about the taxonomic information content of the diagnostic characters applied within this description.
</paragraph>
<paragraph pageId="0" pageNumber="45">
The preliminary information on the habitat of
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum tholussum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="species" species="tholussum">Zospeum tholussum</taxonomicName>
is in congruence with previous findings for
<taxonomicName class="Gastropoda" family="Ellobiidae" genus="Zospeum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Zospeum" order="Ellobiida" pageId="0" pageNumber="45" phylum="Mollusca" rank="genus">Zospeum</taxonomicName>
. Interestingly, the grazing-labyrinth-like structure, in which most of the shells were embedded (Fig. 1, left site), has been already observed during caving expeditions in Northern Spain (
<bibRefCitation author="Jochum, A" journalOrPublisher="MalaCo" pageId="0" pageNumber="45" pagination="400 - 403" title="The microscopic ellobioid, Zospeum Bourguignat, 1856 (Pulmonata, Ellobioidea, Carychiidae) makes a big debut in Basque Country and the province of Burgos (Spain)." volume="8" year="2012">Jochum et al. 2012</bibRefCitation>
). Finally, the presence of a temporal stream at the collecting site of the living specimen further supports a passive dispersal scenario as suggested by
<bibRefCitation author="Weigand, AM" journalOrPublisher="BMC Evolutionary Biology" pageId="0" pageNumber="45" title="Evolution of microgastropods (Ellobioidea, Carychiidae): integrating taxonomic, phylogenetic and evolutionary hypotheses." url="10.1186/1471-2148-13-18" volume="13" year="2013">Weigand et al. (2013)</bibRefCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>