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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.615.8125" ID-GBIF-Dataset="6aefe969-b764-4a4d-8802-75e7bdc17379" ID-PMC="PMC5027776" ID-Pensoft-Pub="1313-2970-615-33" ID-PubMed="27667939" ID-ZBK="7967128BEDED4C19A14E680F3109F676" ModsDocAuthor="" ModsDocDate="2016" ModsDocID="1313-2970-615-33" ModsDocOrigin="ZooKeys 615" ModsDocTitle="Borniopsismortoni sp. n. (Heterodonta, Galeommatoidea, Galeommatidaesensu lato), a new bivalve commensal with a synaptid sea cucumber from Japan" checkinTime="1473279560164" checkinUser="pensoft" docAuthor="Goto, Ryutaro &amp; Ishikawa, Hiroshi" docDate="2016" docId="12E60A4BE8E77D60F27420CBD150A4D1" docLanguage="en" docName="ZooKeys 615: 33-45" docOrigin="ZooKeys 615" docSource="http://dx.doi.org/10.3897/zookeys.615.8125" docTitle="Borniopsis mortoni Goto &amp; Ishikawa, 2016, sp. n." docType="treatment" docUuid="1782E002-A7F1-42D1-A1EC-0824A89B5A3E" docUuidSource="ZooBank" docVersion="6" lastPageNumber="39" masterDocId="FFEC552AFFAEFF9A38290E2CCD08FFDB" masterDocTitle="Borniopsismortoni sp. n. (Heterodonta, Galeommatoidea, Galeommatidaesensu lato), a new bivalve commensal with a synaptid sea cucumber from Japan" masterLastPageNumber="45" masterPageNumber="33" pageNumber="34" updateTime="1668163477717" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Borniopsismortoni sp. n. (Heterodonta, Galeommatoidea, Galeommatidaesensu lato), a new bivalve commensal with a synaptid sea cucumber from Japan</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Goto, Ryutaro</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ishikawa, Hiroshi</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2016</mods:date>
<mods:detail type="volume">
<mods:number>615</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>33</mods:start>
<mods:end>45</mods:end>
</mods:extent>
</mods:part>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.615.8125</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.615.8125</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-615-33</mods:identifier>
<mods:identifier type="ZBK">7967128BEDED4C19A14E680F3109F676</mods:identifier>
<mods:identifier type="ZooBank">7967128BEDED4C19A14E680F3109F676</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="127880676" LSID="urn:lsid:zoobank.org:act:1782E002-A7F1-42D1-A1EC-0824A89B5A3E" httpUri="http://treatment.plazi.org/id/12E60A4BE8E77D60F27420CBD150A4D1" lastPageId="6" lastPageNumber="39" pageId="1" pageNumber="34">
<subSubSection pageId="1" pageNumber="34" type="multiple">
<paragraph pageId="1" pageNumber="34">Taxon classification Animalia Veneroida Galeommatidae</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="34" type="nomenclature">
<paragraph pageId="1" pageNumber="34">
<taxonomicName LSID="http://zoobank.org/1782E002-A7F1-42D1-A1EC-0824A89B5A3E" class="Bivalvia" family="Galeommatidae" genus="Borniopsis" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Veneroida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
<taxonomicNameLabel pageId="1" pageNumber="34">sp. n.</taxonomicNameLabel>
Figs 1, 2, 3, 4
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="34" type="materials_examined">
<paragraph pageId="1" pageNumber="34">Material examined.</paragraph>
<paragraph pageId="1" pageNumber="34">
Holotype (Figs 1, 2): UMMZ 305035 (SL 4.1 mm, SH 2.8 mm). Paratype 1 (Fig. 3): UMMZ 305036-1 (SL 3.9 mm, SH 2.7 mm), paratype 2: UMMZ 30536-2 (SL 2.4 mm, SH 1.8 mm), paratype 3: NSMT-Mo 78968 (SL 3.7 mm, SH 2.7 mm) and paratype 4 (Fig. 1B): NSMT-Mo 78969 (SL 3.3 mm, SH 2.3 mm). Non-type specimens: four individuals of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
(SL 2.4, 3.3, 3.8, 3.3 mm). All specimens were collected in the mud flats at the mouth of the Souzu River, Ainan-cho, Ehime Prefecture, southwestern Shikoku Island, Japan (
<geoCoordinate direction="north" orientation="latitude" precision="925" value="32.95">32°57'N</geoCoordinate>
,
<geoCoordinate direction="east" orientation="longitude" precision="925" value="132.55">132°33'E</geoCoordinate>
). Comparative species (Fig. 4):
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis tsurumaru" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="tsurumaru">Borniopsis tsurumaru</taxonomicName>
, SBMNH 149526;
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ariakensis" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="ariakensis">Borniopsis ariakensis</taxonomicName>
, SBMNH 35056;
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ochetostomae" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="ochetostomae">Borniopsis ochetostomae</taxonomicName>
, SBMNH149525; and
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis maipoensis" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="maipoensis">Borniopsis maipoensis</taxonomicName>
, SBMNH 35126.
</paragraph>
<caption pageId="1" pageNumber="34">
<paragraph pageId="1" pageNumber="34">
Figure 1.
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
and its host. A A crawling individual of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
. B Right shell valve of Paratype 4 (NSMT Mo 78969) bleached to remove the periostracum for observation of shell surface sculpture C, D
<taxonomicName class="Holothuroidea" family="Synaptidae" genus="Patinapta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Patinapta ooplax" order="Apodida" pageId="1" pageNumber="34" phylum="Echinodermata" rank="species" species="ooplax">Patinapta ooplax</taxonomicName>
with multiple individuals of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
attached E, F
<taxonomicName class="Holothuroidea" family="Synaptidae" genus="Patinapta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Patinapta ooplax" order="Apodida" pageId="1" pageNumber="34" phylum="Echinodermata" rank="species" species="ooplax">Patinapta ooplax</taxonomicName>
in situ in mud flats with
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
attached (arrowed). Photo credits: H. Ishikawa:
<normalizedToken originalValue="A–C">A-C</normalizedToken>
, E, F; Y. Hamamura: D. Scale bar: 1 mm.
</paragraph>
</caption>
<caption pageId="1" pageNumber="34">
<paragraph pageId="1" pageNumber="34">
Figure 2. Holotype of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
(UMMZ 305035) (SL 4.1 mm). A, C, E Right valve B, D Left valve F, G Hinge structure of left and right valves. Abbreviations: aa, anterior adductor muscle; ac, anterior cardinal tooth; f, foot; gn, gonad; il, internal ligament; lid, inner demibranch of left side; pa, posterior adductor muscle; rid, inner demibranch of right side; st, stomach. Scale bars 1 mm. Photo credits: R. Goto:
<normalizedToken originalValue="A–G">A-G</normalizedToken>
.
</paragraph>
</caption>
<caption pageId="1" pageNumber="34">
<paragraph pageId="1" pageNumber="34">
Figure 3. Paratype 1 of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
(UMMZ 305035) (SL 2.4 mm). A, C Right valve B, D Left valve E, F Hinge structure of right and left valves. Abbreviations: ac, anterior cardinal tooth; il, internal ligament. Scale bars 0.5 mm. Photo credits: R. Goto:
<normalizedToken originalValue="A–F">A-F</normalizedToken>
.
</paragraph>
</caption>
<caption pageId="1" pageNumber="34">
<paragraph pageId="1" pageNumber="34">
Figure 4. A
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis tsurumaru" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="tsurumaru">Borniopsis tsurumaru</taxonomicName>
(SBMNH 149526) B
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ariakensis" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="ariakensis">Borniopsis ariakensis</taxonomicName>
(SBMNH 35056) C
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ochetostomae" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="ochetostomae">Borniopsis ochetostomae</taxonomicName>
(SBMNH 149525) and D
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis maipoensis" order="Galeommatida" pageId="1" pageNumber="34" phylum="Mollusca" rank="species" species="maipoensis">Borniopsis maipoensis</taxonomicName>
(SBMNH 35126). Photo credits: R. Goto:
<normalizedToken originalValue="A–C">A-C</normalizedToken>
; P. Valentich-Scott: D. Scale bars 1 mm.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="1" pageNumber="34" type="type locality">
<paragraph pageId="1" pageNumber="34">Type locality and habitat.</paragraph>
<paragraph pageId="1" pageNumber="34">
Mud flats at the mouth of the Souzu River, Ainan-cho, Ehime, southwestern Shikoku Island, Japan (
<geoCoordinate direction="north" orientation="latitude" precision="925" value="32.95">32°57'N</geoCoordinate>
,
<geoCoordinate direction="east" orientation="longitude" precision="925" value="132.55">132°33'E</geoCoordinate>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="4" lastPageNumber="37" pageId="1" pageNumber="34" type="description">
<paragraph pageId="1" pageNumber="34">Description.</paragraph>
<paragraph lastPageId="4" lastPageNumber="37" pageId="1" pageNumber="34">
Shell (Figs 1-3): Shell small (up to 4.1 mm), thin, slightly inflated, shape elongate-ovate and equivalve; inequilateral, longer anteriorly. Anterior and posterior margins rounded, ventral margins slightly rounded. Umbo small. Beak small, prosogyrate, situated 2/3 of way toward posterior. Each valve covered by tan to dark brown periostracum with black deposits, often eroded around beaks (Figs 2, 3). Shell surface underneath periostracum smooth and whitish with pearly luster (Fig. 1B).
<pageBreakToken pageId="2" pageNumber="35" start="start">Sculpture</pageBreakToken>
consisting of fine, dense and a few strong, widely spaced commarginal growth striae visible even on periostracum and very faint dense radial striae only visible underneath periostracum (Fig. 1B). Hinge of each valve consisting of a single stout cardinal tooth in front of umbo and well-developed oblique internal ligament posterior to umbo (Figs 2F, G; 3E, F). Soft parts (Figs 1, 2): Mantle not reflected,
<pageBreakToken pageId="3" pageNumber="36" start="start">without</pageBreakToken>
prominent tentacles, edges narrowly extend beyond margin of shell, with numerous short papillae regularly arranged. Both anterior and posterior adductor muscles elongate-ovate, subequal and situated in the middle of dorsal and ventral margin. Ctenidia with gill axis nearly vertical, flat, consisting of single demibranch with both ascending and descending lamellae, joined anteriorly to inner and outer
<pageBreakToken pageId="4" pageNumber="37" start="start">labial</pageBreakToken>
palps. Labial palps leaf-shaped. Foot slender, of moderate size, with small heel; byssal glands located just in front of heel. Gonads situated from middle to posterior in visceral mass just below umbo. Stomach and digestive gland large, occupying anterior part of visceral mass.
</paragraph>
</subSubSection>
<subSubSection pageId="4" pageNumber="37" type="distribution">
<paragraph pageId="4" pageNumber="37">Distribution.</paragraph>
<paragraph pageId="4" pageNumber="37">Only known from the type locality.</paragraph>
</subSubSection>
<subSubSection pageId="4" pageNumber="37" type="host">
<paragraph pageId="4" pageNumber="37">Host.</paragraph>
<paragraph pageId="4" pageNumber="37">
<taxonomicName class="Holothuroidea" family="Synaptidae" genus="Patinapta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Patinapta ooplax" order="Apodida" pageId="4" pageNumber="37" phylum="Echinodermata" rank="species" species="ooplax">Patinapta ooplax</taxonomicName>
(
<taxonomicName lsidName="Galeommatoidea" pageId="4" pageNumber="37" phylum="Echinodermata" rank="phylum">Echinodermata</taxonomicName>
:
<taxonomicName class="Holothuroidea" lsidName="Galeommatoidea" pageId="4" pageNumber="37" rank="class">Holothuroidea</taxonomicName>
:
<taxonomicName lsidName="Galeommatoidea" order="Apodida" pageId="4" pageNumber="37" rank="order">Apodida</taxonomicName>
:
<taxonomicName family="Synaptidae" lsidName="" pageId="4" pageNumber="37" rank="family">Synaptidae</taxonomicName>
) (Fig. 1
<normalizedToken originalValue="C–F">C-F</normalizedToken>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="5" lastPageNumber="38" pageId="4" pageNumber="37" type="host">
<paragraph pageId="4" pageNumber="37">Host association.</paragraph>
<paragraph lastPageId="5" lastPageNumber="38" pageId="4" pageNumber="37">
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="4" pageNumber="37" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
attaches to the body surface of
<taxonomicName class="Holothuroidea" family="Synaptidae" genus="Patinapta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Patinapta ooplax" order="Apodida" pageId="4" pageNumber="37" phylum="Echinodermata" rank="species" species="ooplax">Patinapta ooplax</taxonomicName>
by both its foot and byssal threads (Fig. 1
<normalizedToken originalValue="C–F">C-F</normalizedToken>
). Individuals were patchily distributed in the mud flats. Within the particular patches we sampled, approximately 70% of synaptids served as hosts for
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="4" pageNumber="37" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
. Several
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="4" pageNumber="37" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
often occured on one the same host. At the maximum, more than 10 individuals were attached to a single host. Two
<taxonomicName class="Holothuroidea" family="Synaptidae" genus="Patinapta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Patinapta ooplax" order="Apodida" pageId="5" pageNumber="38" phylum="Echinodermata" rank="species" species="ooplax">
<pageBreakToken pageId="5" pageNumber="38" start="start">Patinapta</pageBreakToken>
ooplax
</taxonomicName>
infested by
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
also harbored the endoparasitic eulimid gastropod
<taxonomicName class="Gastropoda" family="Eulimidae" genus="Hypermastus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hypermastus lacteus" order="Littorinimorpha" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="lacteus">Hypermastus lacteus</taxonomicName>
(A. Adams, 1864).
</paragraph>
</subSubSection>
<subSubSection pageId="5" pageNumber="38" type="etymology">
<paragraph pageId="5" pageNumber="38">Etymology.</paragraph>
<paragraph pageId="5" pageNumber="38">
The specific name is dedicated to Dr. Brian Morton who has made great contributions to marine biology, marine ecology and malacology. He discovered many interesting commensal galeommatoidean species from Hong Kong, some of which now belong to the genus
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="genus">Borniopsis</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection lastPageId="6" lastPageNumber="39" pageId="5" pageNumber="38" type="remarks">
<paragraph pageId="5" pageNumber="38">Remarks.</paragraph>
<paragraph pageId="5" pageNumber="38">
The genus
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="genus">Borniopsis</taxonomicName>
has been variously assigned to the
<taxonomicName family="Kelliidae" lsidName="" pageId="5" pageNumber="38" rank="family">Kelliidae</taxonomicName>
(
<bibRefCitation author="Morton, B" journalOrPublisher="Asian Marine Biology" pageId="11" pageNumber="44" pagination="129 - 160" title="The Hong Kong Galeommatacea (Mollusca: Bivalvia) and their hosts, with descriptions of new species." volume="6" year="1989">Morton and Scott 1989</bibRefCitation>
),
<taxonomicName family="Lasaeidae" lsidName="" pageId="5" pageNumber="38" rank="family">Lasaeidae</taxonomicName>
(
<bibRefCitation author="Bieler, R" journalOrPublisher="Malacologia" pageId="9" pageNumber="42" pagination="113 - 184" title="Classification of bivalve families." volume="52" year="2010">Bieler et al. 2010</bibRefCitation>
), and the subfamily
<taxonomicName lsidName="" pageId="5" pageNumber="38" rank="subfamily" subfamily="Montacutinae">Montacutinae</taxonomicName>
within
<taxonomicName family="Galeommatidae" lsidName="" pageId="5" pageNumber="38" rank="family">Galeommatidae</taxonomicName>
sensu lato (
<bibRefCitation author="Huber, M" journalOrPublisher="ConchBooks, Harxheim" pageId="10" pageNumber="43" title="Compendium of Bivalves 2. A full-color guide to the remaining seven families. A systematic listing of 8500 bivalve species and 10500 synonyms." year="2015">Huber 2015</bibRefCitation>
). However, these family- or subfamily-level groupings are ill-defined when a range of characters and taxa are considered (
<bibRefCitation author="Ponder, WF" editor="Beesley, PL" journalOrPublisher="CSIRO Publishing, Melbourne" pageId="11" pageNumber="44" pagination="316 - 318" title="Superfamily Galeommatoidea." volumeTitle="Mollusca: the Southern Synthesis" year="1998">Ponder 1998</bibRefCitation>
). Indeed, the molecular analyses conducted by
<bibRefCitation author="Goto, R" journalOrPublisher="BMC Evolutionary Biology" pageId="10" pageNumber="43" title="Molecular phylogeny of the bivalve superfamily Galeommatoidea (Heterodonta, Veneroida) reveals dynamic evolution of symbiotic life style and interphylum host switching." url="10.1186/1471-2148-12-172" volume="12" year="2012">Goto et al. (2012)</bibRefCitation>
showed that each of these groups is actually polyphyletic. In this study, we assigned the genus
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="genus">Borniopsis</taxonomicName>
tentatively to
<taxonomicName family="Galeommatidae" lsidName="" pageId="5" pageNumber="38" rank="family">Galeommatidae</taxonomicName>
sensu lato, which
<bibRefCitation author="Ponder, WF" editor="Beesley, PL" journalOrPublisher="CSIRO Publishing, Melbourne" pageId="11" pageNumber="44" pagination="316 - 318" title="Superfamily Galeommatoidea." volumeTitle="Mollusca: the Southern Synthesis" year="1998">Ponder (1998)</bibRefCitation>
defined by the same diagnosis that was applied to the superfamily
<taxonomicName lsidName="Galeommatoidea" pageId="5" pageNumber="38" rank="superfamily" superfamily="Galeommatoidea">Galeommatoidea</taxonomicName>
, as did
<bibRefCitation author="Huber, M" journalOrPublisher="ConchBooks, Harxheim" pageId="10" pageNumber="43" title="Compendium of Bivalves 2. A full-color guide to the remaining seven families. A systematic listing of 8500 bivalve species and 10500 synonyms." year="2015">Huber (2015)</bibRefCitation>
. Further taxonomic assignment of this genus within
<taxonomicName lsidName="Galeommatoidea" pageId="5" pageNumber="38" rank="superfamily" superfamily="Galeommatoidea">Galeommatoidea</taxonomicName>
(or
<taxonomicName family="Galeommatidae" lsidName="" pageId="5" pageNumber="38" rank="family">Galeommatidae</taxonomicName>
sensu lato) should be delayed until its family-level (or subfamily-level) classification is revised.
</paragraph>
<paragraph pageId="5" pageNumber="38">
As with
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
, both
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis tsurumaru" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="tsurumaru">Borniopsis tsurumaru</taxonomicName>
and
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ariakensis" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="ariakensis">Borniopsis ariakensis</taxonomicName>
have a symbiotic relationship with synaptid sea cucumbers (
<bibRefCitation author="Morton, B" journalOrPublisher="Hong Kong University Press, Hong Kong" pageId="11" pageNumber="44" title="Partnerships in the Sea: Hong Kong's Marine Symbioses." url="http://www.jstor.org/stable/j.ctt2jc6c7" year="1988">Morton 1988</bibRefCitation>
,
<bibRefCitation author="Morton, B" journalOrPublisher="Asian Marine Biology" pageId="11" pageNumber="44" pagination="129 - 160" title="The Hong Kong Galeommatacea (Mollusca: Bivalvia) and their hosts, with descriptions of new species." volume="6" year="1989">Morton and Scott 1989</bibRefCitation>
,
<bibRefCitation author="Luetzen, H" journalOrPublisher="Journal of Morphology" pageId="10" pageNumber="43" pagination="407 - 420" title="Morphology, structure of dimorphic sperm, and reproduction in the hermaphroditic commensal bivalve Pseudopythinatsrumaru (Galeommatoidea: Kellidae)." url="10.1002/jmor.10158" volume="262" year="2004">
<normalizedToken originalValue="Lützen">Luetzen</normalizedToken>
et al. 2004
</bibRefCitation>
,
<bibRefCitation author="Kai, T" journalOrPublisher="Journal of Crustacean Biology" pageId="10" pageNumber="43" pagination="342 - 351" title="Description of zoeae and habitat of Elamenopsis ariakensis (Brachyura: Hymenosomatidae) living within the burrows of the sea cucumber Protankyrabidentata." url="10.1163/20021975-99990379" volume="28" year="2008">Kai and Henmi 2008</bibRefCitation>
). However, the particular host species differ between them -
<taxonomicName class="Holothuroidea" family="Synaptidae" genus="Patinapta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Patinapta ooplax" order="Apodida" pageId="5" pageNumber="38" phylum="Echinodermata" rank="species" species="ooplax">Patinapta ooplax</taxonomicName>
(host solely for
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
) and
<taxonomicName class="Holothuroidea" family="Synaptidae" genus="Protankyra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Protankyra bidentata" order="Apodida" pageId="5" pageNumber="38" phylum="Echinodermata" rank="species" species="bidentata">Protankyra bidentata</taxonomicName>
(Woodward &amp; Barrett, 1858) (host for both
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis tsurumaru" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="tsurumaru">Borniopsis tsurumaru</taxonomicName>
and
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ariakensis" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="ariakensis">Borniopsis ariakensis</taxonomicName>
) (
<bibRefCitation author="Morton, B" journalOrPublisher="Asian Marine Biology" pageId="11" pageNumber="44" pagination="129 - 160" title="The Hong Kong Galeommatacea (Mollusca: Bivalvia) and their hosts, with descriptions of new species." volume="6" year="1989">Morton and Scott 1989</bibRefCitation>
,
<bibRefCitation author="Luetzen, H" journalOrPublisher="Journal of Morphology" pageId="10" pageNumber="43" pagination="407 - 420" title="Morphology, structure of dimorphic sperm, and reproduction in the hermaphroditic commensal bivalve Pseudopythinatsrumaru (Galeommatoidea: Kellidae)." url="10.1002/jmor.10158" volume="262" year="2004">
<normalizedToken originalValue="Lützen">Luetzen</normalizedToken>
et al. 2004
</bibRefCitation>
).
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
always attaches directly onto the body surface of the host (this study), whereas
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis tsurumaru" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="tsurumaru">Borniopsis tsurumaru</taxonomicName>
can attach to the body surface of the host, or the wall of the
<normalizedToken originalValue="host’s">host's</normalizedToken>
burrow, or to the carapace of commensal crabs living in the same burrows (
<bibRefCitation author="Morton, B" journalOrPublisher="Hong Kong University Press, Hong Kong" pageId="11" pageNumber="44" title="Partnerships in the Sea: Hong Kong's Marine Symbioses." url="http://www.jstor.org/stable/j.ctt2jc6c7" year="1988">Morton 1988</bibRefCitation>
,
<bibRefCitation author="Morton, B" journalOrPublisher="Asian Marine Biology" pageId="11" pageNumber="44" pagination="129 - 160" title="The Hong Kong Galeommatacea (Mollusca: Bivalvia) and their hosts, with descriptions of new species." volume="6" year="1989">Morton and Scott 1989</bibRefCitation>
,
<bibRefCitation author="Luetzen, H" journalOrPublisher="Journal of Morphology" pageId="10" pageNumber="43" pagination="407 - 420" title="Morphology, structure of dimorphic sperm, and reproduction in the hermaphroditic commensal bivalve Pseudopythinatsrumaru (Galeommatoidea: Kellidae)." url="10.1002/jmor.10158" volume="262" year="2004">
<normalizedToken originalValue="Lützen">Luetzen</normalizedToken>
et al. 2004
</bibRefCitation>
,
<bibRefCitation author="Kai, T" journalOrPublisher="Journal of Crustacean Biology" pageId="10" pageNumber="43" pagination="342 - 351" title="Description of zoeae and habitat of Elamenopsis ariakensis (Brachyura: Hymenosomatidae) living within the burrows of the sea cucumber Protankyrabidentata." url="10.1163/20021975-99990379" volume="28" year="2008">Kai and Henmi 2008</bibRefCitation>
,
<bibRefCitation author="Goto, R" journalOrPublisher="BMC Evolutionary Biology" pageId="10" pageNumber="43" title="Molecular phylogeny of the bivalve superfamily Galeommatoidea (Heterodonta, Veneroida) reveals dynamic evolution of symbiotic life style and interphylum host switching." url="10.1186/1471-2148-12-172" volume="12" year="2012">Goto et al. 2012</bibRefCitation>
). Furthermore, the number of bivalves per host is much higher in
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
(several to more than 10) than
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis tsurumaru" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="tsurumaru">Borniopsis tsurumaru</taxonomicName>
and
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ariakensis" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="ariakensis">Borniopsis ariakensis</taxonomicName>
(usually one) (
<bibRefCitation author="Luetzen, H" journalOrPublisher="Journal of Morphology" pageId="10" pageNumber="43" pagination="407 - 420" title="Morphology, structure of dimorphic sperm, and reproduction in the hermaphroditic commensal bivalve Pseudopythinatsrumaru (Galeommatoidea: Kellidae)." url="10.1002/jmor.10158" volume="262" year="2004">
<normalizedToken originalValue="Lützen">Luetzen</normalizedToken>
et al. 2004
</bibRefCitation>
, Goto, Ishikawa, and Hamamura, personal observations).
</paragraph>
<paragraph pageId="5" pageNumber="38">
The shells of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis tsurumaru" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="tsurumaru">Borniopsis tsurumaru</taxonomicName>
and
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ariakensis" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="ariakensis">Borniopsis ariakensis</taxonomicName>
are much larger (up to 11-12 mm in SL) than those of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
(up to 4.1 mm) (
<bibRefCitation author="Morton, B" journalOrPublisher="Asian Marine Biology" pageId="11" pageNumber="44" pagination="129 - 160" title="The Hong Kong Galeommatacea (Mollusca: Bivalvia) and their hosts, with descriptions of new species." volume="6" year="1989">Morton and Scott 1989</bibRefCitation>
) (Fig. 4A, B). Probably, this corresponds with the size of the host because
<taxonomicName class="Holothuroidea" family="Synaptidae" genus="Protankyra" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Protankyra bidentata" order="Apodida" pageId="5" pageNumber="38" phylum="Echinodermata" rank="species" species="bidentata">Protankyra bidentata</taxonomicName>
is much larger than
<taxonomicName class="Holothuroidea" family="Synaptidae" genus="Patinapta" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Patinapta ooplax" order="Apodida" pageId="5" pageNumber="38" phylum="Echinodermata" rank="species" species="ooplax">Patinapta ooplax</taxonomicName>
. The shells of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis tsurumaru" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="tsurumaru">Borniopsis tsurumaru</taxonomicName>
and
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ariakensis" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="ariakensis">Borniopsis ariakensis</taxonomicName>
are thicker and more inflated than those of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
(
<bibRefCitation author="Luetzen, H" journalOrPublisher="Journal of Morphology" pageId="10" pageNumber="43" pagination="407 - 420" title="Morphology, structure of dimorphic sperm, and reproduction in the hermaphroditic commensal bivalve Pseudopythinatsrumaru (Galeommatoidea: Kellidae)." url="10.1002/jmor.10158" volume="262" year="2004">
<normalizedToken originalValue="Lützen">Luetzen</normalizedToken>
et al. 2004
</bibRefCitation>
; this study). In addition, the shells of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
are always covered by a dark brown periostracum, whereas those of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis tsurumaru" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="tsurumaru">Borniopsis tsurumaru</taxonomicName>
and
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ariakensis" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="ariakensis">Borniopsis ariakensis</taxonomicName>
are often whitish, although some are dark brown. The umbones of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis tsurumaru" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="tsurumaru">Borniopsis tsurumaru</taxonomicName>
and
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ariakensis" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="ariakensis">Borniopsis ariakensis</taxonomicName>
are more protruding than those of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
(Fig. 4A, B). A molecular analysis is needed to understand whether these three synaptid-associated species are monophyletic or not. In addition, morphological variation of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis tsurumaru" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="tsurumaru">Borniopsis tsurumaru</taxonomicName>
and
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ariakensis" order="Galeommatida" pageId="5" pageNumber="38" phylum="Mollusca" rank="species" species="ariakensis">Borniopsis ariakensis</taxonomicName>
is apparently continuous (Goto, Ishikawa and Hamamura, pers. obs.) so molecular testing should be employed to investigate whether they can be distinguished genetically or not.
</paragraph>
<paragraph pageId="6" pageNumber="39">
<pageBreakToken pageId="6" pageNumber="39" start="start">The</pageBreakToken>
present new species also closely resembles
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ochetostomae" order="Galeommatida" pageId="6" pageNumber="39" phylum="Mollusca" rank="species" species="ochetostomae">Borniopsis ochetostomae</taxonomicName>
and
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis maipoensis" order="Galeommatida" pageId="6" pageNumber="39" phylum="Mollusca" rank="species" species="maipoensis">Borniopsis maipoensis</taxonomicName>
in having an elongate ovate shell covered by a brownish periostracum (Fig. 4C, D). However,
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ochetostomae" order="Galeommatida" pageId="6" pageNumber="39" phylum="Mollusca" rank="species" species="ochetostomae">Borniopsis ochetostomae</taxonomicName>
is much larger (up to 11 mm) than
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="6" pageNumber="39" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
and its beak is located more centrally than that of
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="6" pageNumber="39" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
(
<bibRefCitation author="Morton, B" journalOrPublisher="Asian Marine Biology" pageId="11" pageNumber="44" pagination="129 - 160" title="The Hong Kong Galeommatacea (Mollusca: Bivalvia) and their hosts, with descriptions of new species." volume="6" year="1989">Morton and Scott 1989</bibRefCitation>
,
<bibRefCitation author="Jespersen, A" journalOrPublisher="Zoomorphology" pageId="10" pageNumber="43" pagination="159 - 172" title="Ultrastructure of dimorphic sperm and seminal receptacle in the hermaphrodites Barrimysiasiphonosomae and Pseudopythinaochetostomae (Bivalvia, Galeommatoidea)." url="10.1007/s00435-001-0053-5" volume="121" year="2002">Jespersen et al. 2002</bibRefCitation>
, this study) (Fig. 4C). On the other hand,
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis maipoensis" order="Galeommatida" pageId="6" pageNumber="39" phylum="Mollusca" rank="species" species="maipoensis">Borniopsis maipoensis</taxonomicName>
is rather smaller (up to 3 mm) and more rounded than
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="6" pageNumber="39" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
(
<bibRefCitation author="Morton, B" journalOrPublisher="Asian Marine Biology" pageId="11" pageNumber="44" pagination="129 - 160" title="The Hong Kong Galeommatacea (Mollusca: Bivalvia) and their hosts, with descriptions of new species." volume="6" year="1989">Morton and Scott 1989</bibRefCitation>
, this study) (Fig. 4D). In addition,
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis maipoensis" order="Galeommatida" pageId="6" pageNumber="39" phylum="Mollusca" rank="species" species="maipoensis">Borniopsis maipoensis</taxonomicName>
has two distinct papillae on the dorsal surface of the foot (
<bibRefCitation author="Morton, B" journalOrPublisher="Asian Marine Biology" pageId="11" pageNumber="44" pagination="129 - 160" title="The Hong Kong Galeommatacea (Mollusca: Bivalvia) and their hosts, with descriptions of new species." volume="6" year="1989">Morton and Scott 1989</bibRefCitation>
), whereas we did not observe such papillae on
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="6" pageNumber="39" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
(Fig. 1A). And lastly, the hosts for these three species are different -
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis mortoni" order="Galeommatida" pageId="6" pageNumber="39" phylum="Mollusca" rank="species" species="mortoni">Borniopsis mortoni</taxonomicName>
(holothurian hosts),
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis ochetostomae" order="Galeommatida" pageId="6" pageNumber="39" phylum="Mollusca" rank="species" species="ochetostomae">Borniopsis ochetostomae</taxonomicName>
(echiuran hosts) and
<taxonomicName class="Bivalvia" family="Lasaeidae" genus="Borniopsis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Borniopsis maipoensis" order="Galeommatida" pageId="6" pageNumber="39" phylum="Mollusca" rank="species" species="maipoensis">Borniopsis maipoensis</taxonomicName>
(probably tanaid hosts) (
<bibRefCitation author="Morton, B" journalOrPublisher="Asian Marine Biology" pageId="11" pageNumber="44" pagination="129 - 160" title="The Hong Kong Galeommatacea (Mollusca: Bivalvia) and their hosts, with descriptions of new species." volume="6" year="1989">Morton and Scott 1989</bibRefCitation>
, this study).
</paragraph>
</subSubSection>
</treatment>
</document>
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