352 lines
42 KiB
XML
352 lines
42 KiB
XML
<document ID-DOI="http://dx.doi.org/10.3897/zookeys.160.2290" ID-GBIF-Dataset="f93df35f-2101-43af-97e3-9fa3fe24a2ee" ID-PMC="PMC3253628" ID-Pensoft-Pub="1313-2970-160-1" ID-PubMed="22303117" ModsDocAuthor="" ModsDocDate="2011" ModsDocID="1313-2970-160-1" ModsDocOrigin="ZooKeys 160" ModsDocTitle="A new horsehair worm, Chordodes formosanus sp. n. (Nematomorpha, Gordiida) from Hierodula mantids of Taiwan and Japan with redescription of a closely related species, Chordodes japonensis" checkinTime="1451249424265" checkinUser="pensoft" docAuthor="Chiu, Ming-Chung, Huang, Chin-Gi, Wu, Wen-Jer & Shiao, Shiuh-Feng" docDate="2011" docId="E81C925F3756B77471D97C7B031DB5EE" docLanguage="en" docName="ZooKeys 160: 1-22" docOrigin="ZooKeys 160" docSource="http://dx.doi.org/10.3897/zookeys.160.2290" docTitle="Chordodes japonensis Inoue 1952" docType="treatment" docVersion="3" lastPageNumber="14" masterDocId="ED57805BFFEBFFEAB119FF8DFFA0CD3A" masterDocTitle="A new horsehair worm, Chordodes formosanus sp. n. (Nematomorpha, Gordiida) from Hierodula mantids of Taiwan and Japan with redescription of a closely related species, Chordodes japonensis" masterLastPageNumber="22" masterPageNumber="1" pageNumber="9" updateTime="1668153101227" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>A new horsehair worm, Chordodes formosanus sp. n. (Nematomorpha, Gordiida) from Hierodula mantids of Taiwan and Japan with redescription of a closely related species, Chordodes japonensis</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Chiu, Ming-Chung</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Huang, Chin-Gi</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Wu, Wen-Jer</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Shiao, Shiuh-Feng</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>ZooKeys</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2011</mods:date>
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<mods:detail type="volume">
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<mods:number>160</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>1</mods:start>
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<mods:end>22</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:location>
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<mods:url>http://dx.doi.org/10.3897/zookeys.160.2290</mods:url>
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</mods:location>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.160.2290</mods:identifier>
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<mods:identifier type="Pensoft-Pub">1313-2970-160-1</mods:identifier>
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</mods:mods>
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<treatment ID-GBIF-Taxon="152032838" LSID="urn:lsid:plazi:treatment:E81C925F3756B77471D97C7B031DB5EE" httpUri="http://treatment.plazi.org/id/E81C925F3756B77471D97C7B031DB5EE" lastPageId="13" lastPageNumber="14" pageId="8" pageNumber="9">
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<subSubSection pageId="8" pageNumber="9" type="nomenclature">
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<paragraph pageId="8" pageNumber="9">
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<taxonomicName LSID="http://species-id.net/wiki/Chordodes_japonensis" authority="Inoue, 1952" authorityName="Inoue" authorityYear="1952" class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="8" pageNumber="9" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis Inoue, 1952</taxonomicName>
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9" type="material examined">
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<paragraph pageId="8" pageNumber="9">Material examined.</paragraph>
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<paragraph lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
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Examined male collected with its host from Miyazaki Prefecture, Japan (
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<geoCoordinate direction="north" orientation="latitude" precision="1" value="32.17264">32°10'21.50"N</geoCoordinate>
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,
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<geoCoordinate direction="east" orientation="longitude" precision="1" value="131.46014">131°27'36.53"E</geoCoordinate>
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) by Yasukuni Ono on 5-XI-2010. Partial body of horsehair worm deposited with its host at Department of Entomology,
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<pageBreakToken pageId="9" pageNumber="10" start="start">National</pageBreakToken>
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Taiwan University, Taipei, Taiwan. Accession number of partial COI sequence in GenBank: JF808206.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="9" pageNumber="10" type="host">
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<paragraph pageId="9" pageNumber="10">Host.</paragraph>
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<paragraph pageId="9" pageNumber="10">
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Chinese mantids,
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<taxonomicName class="Insecta" family="Mantidae" genus="Tenodera" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tenodera sinensis" order="Mantodea" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="sinensis">Tenodera sinensis</taxonomicName>
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(
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<taxonomicName class="Insecta" family="Lycidae" genus="Mantodea" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Mantodea" order="Coleoptera" pageId="9" pageNumber="10" phylum="Arthropoda" rank="genus">Mantodea</taxonomicName>
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:
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<taxonomicName family="Mantidae" lsidName="" pageId="9" pageNumber="10" rank="family">Mantidae</taxonomicName>
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), which are sometimes classified as
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<taxonomicName class="Insecta" family="Mantidae" genus="Tenodera" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tenodera aridifolia" order="Mantodea" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="aridifolia">Tenodera aridifolia</taxonomicName>
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.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="9" pageNumber="10" type="redescription">
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<paragraph pageId="9" pageNumber="10">Redescription.</paragraph>
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<paragraph pageId="9" pageNumber="10">(Fig. 6)</paragraph>
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<paragraph pageId="9" pageNumber="10">Male adult (n = 1). Body length 220 mm, width (widest) 0.94 mm (after dehydration). In alcohol-preserved specimens, body rough and flat with dorsal and ventral grooves, dark-brown with a darkly pigmented line on ventral side.</paragraph>
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<paragraph pageId="9" pageNumber="10">
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Posterior end (Fig. 6A) not lobed, with short spines (ca. 5-13
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<normalizedToken originalValue="μm">μm</normalizedToken>
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) among areoles on margin. Cloacal opening subterminal, oval 44
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<normalizedToken originalValue="μm">μm</normalizedToken>
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long and 25
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<normalizedToken originalValue="μm">μm</normalizedToken>
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wide, with circumcloacal spines. A pair of oval regions without areoles posterior to cloacal opening, each with scattered bristles extending as two rows of ventral strips (115-120
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<normalizedToken originalValue="μm">μm</normalizedToken>
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wide) structured by cord-like folds or flat areoles. Paired oval bristlefields (82
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<normalizedToken originalValue="μm">μm</normalizedToken>
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wide and 231
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<normalizedToken originalValue="μm">μm</normalizedToken>
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long) bearing numerous branched and unbranched bristles on borders between flat areoles and normal areoles on lateral side of cloacal opening. Anterior end tapered, same color as body, with white tip (white cap) but no dark collar under stereomicroscopy. Under SEM, anterior end (Fig. 6F) smooth with short, thick bristles and small spines; mouth open on cone at anterior extremity.
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</paragraph>
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<paragraph pageId="9" pageNumber="10">
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Entire body covered by areoles with slightly cord-like folds in between. Areoles characterized into six types (simple, tubercle, thorn, circumcluster, and two types of crowned areoles). Simple areoles (Fig. 6C), most abundant, covering entire body surface except anterior end and ventral side of posterior end; each 5-11
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<normalizedToken originalValue="μm">μm</normalizedToken>
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in diameter, more or less circular or oval, surface uneven, some areas with short bristles. Simple areoles varying in height and some significantly elevated areoles in clusters of two to five, appearing as bulging areoles but darker under light microscopy. Tubercle areoles (Fig. 6C) scattered among simple areoles, each similarly shaped to simple areole but with a tubercle (about 7
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<normalizedToken originalValue="μm">μm</normalizedToken>
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long) on apically concave center. Thorn areoles (Fig. 6E) similar to tubercle areoles but with a long solid thorn (15
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<normalizedToken originalValue="μm">μm</normalizedToken>
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long) instead of tubercle; number of thorn areoles much fewer than tubercle areoles. Crowned areoles (Figs 6B, D) clustered in pairs with a central tubercle in between and surrounded by 7-14 circumcluster areoles with short filaments on apical surface; scattered over trunk; each with medium filaments (12-20
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<normalizedToken originalValue="μm">μm</normalizedToken>
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) originating from apical center and sidelong to edges. Crowned areoles roughly arranged in two lines on ventral and dorsal midlines, bearing significantly longer filaments (most around 100
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<normalizedToken originalValue="μm">μm</normalizedToken>
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, none> 150
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<normalizedToken originalValue="μm">μm</normalizedToken>
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) (Figs 6B, C).
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</paragraph>
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<caption pageId="9" pageNumber="10">
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<paragraph pageId="9" pageNumber="10">
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Figure 6. Male adult of
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="9" pageNumber="10" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
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. A Posterior end of male B ventral side of male body C cuticular surface with four types of areole D short-crowned areoles E thorn areole F anterior end. Bri, bristle; Cir, circumcluster areole; Clo, cloacal opening; Lc, long crowned areole; M, mouth; Sc, short-crowned areole; Sim, simple areoles; Tu, tubercle areole.
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10" type="phylogeny">
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<paragraph pageId="9" pageNumber="10">Phylogeny.</paragraph>
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<paragraph pageId="9" pageNumber="10">
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A phylogenetic tree of the 40 samples of horsehair worms collected from three species of mantids
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<taxonomicName class="Insecta" family="Mantidae" genus="Hierodula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hierodula formosana" order="Mantodea" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="formosana">Hierodula formosana</taxonomicName>
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,
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<taxonomicName class="Insecta" family="Mantidae" genus="Hierodula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hierodula patellifera" order="Mantodea" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="patellifera">Hierodula patellifera</taxonomicName>
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, and
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<taxonomicName class="Insecta" family="Mantidae" genus="Tenodera" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tenodera sinensis" order="Mantodea" pageId="9" pageNumber="10" phylum="Arthropoda" rank="species" species="sinensis">Tenodera sinensis</taxonomicName>
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from Taiwan and Japan with one outgroup (
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Paragordius" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paragordius" order="Gordioidea" pageId="9" pageNumber="10" phylum="Nematomorpha" rank="genus">Paragordius</taxonomicName>
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sp.) is shown in Fig. 7.
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</paragraph>
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<paragraph lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10">
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Comparison of the 40 horsehair worm samples (39
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="9" pageNumber="10" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
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and one
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="9" pageNumber="10" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
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) revealed there were 31 haplotypes with 432 invariable sites, 64 singletons, and 32 parsimoniously informative sites. Newly sequenced COI data were deposited in the GenBank database (see Table 1 for accession nos.). Samples from the host mantids of
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<taxonomicName class="Insecta" family="Mantidae" genus="Hierodula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hierodula" order="Mantodea" pageId="9" pageNumber="10" phylum="Arthropoda" rank="genus">Hierodula</taxonomicName>
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and
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<taxonomicName class="Insecta" family="Mantidae" genus="Tenodera" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tenodera" order="Mantodea" pageId="9" pageNumber="10" phylum="Arthropoda" rank="genus">Tenodera</taxonomicName>
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(which are considered two species,
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="9" pageNumber="10" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
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sp. n. and
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="9" pageNumber="10" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
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, respectively)were significantly separated into two groups based
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<pageBreakToken pageId="10" pageNumber="11" start="start">on</pageBreakToken>
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the phylogenetic tree restructured by the NJ method (Fig. 7). The genetic distance between these two groups was 0.16840. This result supports that theybelong to different species as we suggested based on morphology. The phylogenetic tree also revealed a polytomic topology among the 39 horsehair worms parasitizing
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<taxonomicName class="Insecta" family="Mantidae" genus="Hierodula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hierodula" order="Mantodea" pageId="10" pageNumber="11" phylum="Arthropoda" rank="genus">Hierodula</taxonomicName>
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. Although some clades were observed, they were not highly supported due to low bootstrap values and short genetic distances; the mean genetic distances among them was 0.00979 with a range of 0.000-0.01922. Since the genetic distance between the larvae collected in the field and
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="10" pageNumber="11" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
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was only 0.00759, we suggest that those larvae belong to the same species.
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</paragraph>
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<caption pageId="10" pageNumber="11">
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<paragraph pageId="10" pageNumber="11">
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Figure 7. Neighbor-joining tree of
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="10" pageNumber="11" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
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sp. n. and
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="10" pageNumber="11" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
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with genetic distances between each group. Abbreviations in the table indicate the horsehair
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<normalizedToken originalValue="worms’">worms'</normalizedToken>
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mantid hosts (Hf,
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<taxonomicName class="Insecta" family="Mantidae" genus="Hierodula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hierodula formosana" order="Mantodea" pageId="10" pageNumber="11" phylum="Arthropoda" rank="species" species="formosana">Hierodula formosana</taxonomicName>
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; Hp,
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<taxonomicName class="Insecta" family="Mantidae" genus="Hierodula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hierodula patellifera" order="Mantodea" pageId="10" pageNumber="11" phylum="Arthropoda" rank="species" species="patellifera">Hierodula patellifera</taxonomicName>
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; Ts,
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<taxonomicName class="Insecta" family="Mantidae" genus="Tenodera" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tenodera sinensis" order="Mantodea" pageId="10" pageNumber="11" phylum="Arthropoda" rank="species" species="sinensis">Tenodera sinensis</taxonomicName>
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) and collecting localities (Tw, Taiwan; Ja, Japan). Numbers at the nodes represent the percentage of 1000 bootstrap values of> 70%. The outgroup (out) was a
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Paragordius" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paragordius" order="Gordioidea" pageId="10" pageNumber="11" phylum="Nematomorpha" rank="genus">Paragordius</taxonomicName>
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sp.
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection lastPageId="13" lastPageNumber="14" pageId="10" pageNumber="11" type="discussion">
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<paragraph pageId="10" pageNumber="11">Discussion.</paragraph>
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<paragraph pageId="10" pageNumber="11">
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In this article, a new species,
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="10" pageNumber="11" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
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sp. n., which parasitizes
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<taxonomicName class="Insecta" family="Mantidae" genus="Hierodula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hierodula formosana" order="Mantodea" pageId="10" pageNumber="11" phylum="Arthropoda" rank="species" species="formosana">Hierodula formosana</taxonomicName>
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and
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<taxonomicName class="Insecta" family="Mantidae" genus="Hierodula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hierodula patellifera" order="Mantodea" pageId="10" pageNumber="11" phylum="Arthropoda" rank="species" species="patellifera">Hierodula patellifera</taxonomicName>
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, was proposed and described including the morphology of the egg and larval stages. Because of a similar morphological description by
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<bibRefCitation author="Schmidt-Rhaesa, A" journalOrPublisher="Species Diversity" pageId="13" pageNumber="14" pagination="97 - 107" title="Japanese horsehair worms (Nematomorpha) from the Lake Biwa Museum." volume="9" year="2004">Schmidt-Rhaesa (2004)</bibRefCitation>
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, we categorized the species in that study into
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="10" pageNumber="11" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
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sp. n. and also limited the mantid host range of
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="10" pageNumber="11" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
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to the genus
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<taxonomicName class="Insecta" family="Mantidae" genus="Tenodera" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tenodera" order="Mantodea" pageId="10" pageNumber="11" phylum="Arthropoda" rank="genus">Tenodera</taxonomicName>
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.
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</paragraph>
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<subSection lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12" type="morphological comparison of c. formosanus sp. n. and c. japonensis">
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<paragraph pageId="11" pageNumber="12">
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<pageBreakToken pageId="11" pageNumber="12" start="start">Morphological</pageBreakToken>
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comparison of
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
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sp. n. and
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
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</paragraph>
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<paragraph lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">
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The two species,
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<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
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sp. n. and
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
, can be distinguished by the presence of dimorphism in male crowned areoles and the length of long filaments on female crowned areoles. A comparison of areolar types in
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
|
||
sp. n. and
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
is given in Table 2. Both
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
|
||
sp. n. and
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
have two forms of crowned areoles on their cuticles, one with moderate attachments on the top (short-crowned areoles) and the other with significantly longer attachments (long-crowned areoles). Both types of crowned areoles were found on these two species except for the male
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
|
||
sp. n. (the short-crowned areoles were not mentioned in the description of
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
by
|
||
<bibRefCitation author="Inoue, I" journalOrPublisher="Annotationes Zoologicae Japonenses" pageId="13" pageNumber="14" pagination="400 - 402" title="On a new species of Chordodes (Gordiacea) from Japan." volume="25" year="1952">Inoue (1952)</bibRefCitation>
|
||
), which indicates sexual dimorphism in
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
|
||
sp. n. We believe the female worm which was previously considered to be
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
by
|
||
<bibRefCitation author="Schmidt-Rhaesa, A" journalOrPublisher="Species Diversity" pageId="13" pageNumber="14" pagination="97 - 107" title="Japanese horsehair worms (Nematomorpha) from the Lake Biwa Museum." volume="9" year="2004">Schmidt-Rhaesa (2004)</bibRefCitation>
|
||
actually belongs to the species,
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="11" pageNumber="12" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
|
||
n. sp, described here. The reason why the sexual dimorphism was not mentioned by
|
||
<bibRefCitation author="Schmidt-Rhaesa, A" journalOrPublisher="Species Diversity" pageId="13" pageNumber="14" pagination="97 - 107" title="Japanese horsehair worms (Nematomorpha) from the Lake Biwa Museum." volume="9" year="2004">Schmidt-Rhaesa (2004)</bibRefCitation>
|
||
is probably that the free-living male is not the same species as the female. In addition, the length of the filaments on the long-crowned areoles is also a significant character differentiating these two species; they are
|
||
<pageBreakToken pageId="12" pageNumber="13" start="start">always</pageBreakToken>
|
||
> 200
|
||
<normalizedToken originalValue="μm">μm</normalizedToken>
|
||
in
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="12" pageNumber="13" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
|
||
sp. n. (or> 6-fold that of paired crowned areoles) but <200
|
||
<normalizedToken originalValue="μm">μm</normalizedToken>
|
||
in
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="12" pageNumber="13" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
(or <5-fold that of paired crowned areoles, and most of them are around 100
|
||
<normalizedToken originalValue="μm">μm</normalizedToken>
|
||
).
|
||
</paragraph>
|
||
<paragraph pageId="12" pageNumber="13">
|
||
In addition to the crowned areoles, other minor differences in Table 2 (thorn areoles and bulging areoles) are much more difficult to use in discriminating these two species. Although the presence of thorn areoles is always questionable due to their small number, it is still considered a key character for distinguishing different species (
|
||
<bibRefCitation author="Schmidt-Rhaesa, A" journalOrPublisher="Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg" pageId="13" pageNumber="14" pagination="37 - 114" title="Summary of Chordodes species (Nematomorpha, Gordiida), with a discussion of their diagnostic characters." volume="44" year="2008">Schmidt-Rhaesa et al. 2008</bibRefCitation>
|
||
). Thorn areoles were not reported by
|
||
<bibRefCitation author="Schmidt-Rhaesa, A" journalOrPublisher="Species Diversity" pageId="13" pageNumber="14" pagination="97 - 107" title="Japanese horsehair worms (Nematomorpha) from the Lake Biwa Museum." volume="9" year="2004">Schmidt-Rhaesa (2004)</bibRefCitation>
|
||
or
|
||
<bibRefCitation author="Baek, KM" journalOrPublisher="Korean Journal of Systematic Zoology" pageId="13" pageNumber="14" pagination="221 - 228" title="Two species of genus Chordodes (Gordioidea, Nematomorpha) from Korea." url="doi: 10.1038/387489a0" volume="9" year="1993">Baek (1993)</bibRefCitation>
|
||
but appeared in the description by
|
||
<bibRefCitation author="Inoue, I" journalOrPublisher="Annotationes Zoologicae Japonenses" pageId="13" pageNumber="14" pagination="400 - 402" title="On a new species of Chordodes (Gordiacea) from Japan." volume="25" year="1952">Inoue (1952)</bibRefCitation>
|
||
. In our study, thorn areoles were not found in nine
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="12" pageNumber="13" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
|
||
sp. n. but appeared in the other 31
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="12" pageNumber="13" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
|
||
sp. n. and one
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="12" pageNumber="13" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
. Contrary to the thorn areoles which can be easily distinguished from other types of areoles, bulging areoles are much easier to be confused with simple areoles (
|
||
<bibRefCitation author="Schmidt-Rhaesa, A" journalOrPublisher="Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg" pageId="13" pageNumber="14" pagination="37 - 114" title="Summary of Chordodes species (Nematomorpha, Gordiida), with a discussion of their diagnostic characters." volume="44" year="2008">Schmidt-Rhaesa et al. 2008</bibRefCitation>
|
||
). Although bulging areoles were consistently ignored in descriptions of
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="12" pageNumber="13" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
, we observed various heights of simple areoles, and some of them clustered in groups similar to bulging areoles as described by
|
||
<bibRefCitation author="Schmidt-Rhaesa, A" journalOrPublisher="Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg" pageId="13" pageNumber="14" pagination="37 - 114" title="Summary of Chordodes species (Nematomorpha, Gordiida), with a discussion of their diagnostic characters." volume="44" year="2008">Schmidt-Rhaesa et al. (2008)</bibRefCitation>
|
||
. This renders the presence of bulging areoles in
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="12" pageNumber="13" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
as well as
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="12" pageNumber="13" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
|
||
sp. n. questionable.
|
||
</paragraph>
|
||
</subSection>
|
||
<subSection pageId="12" pageNumber="13" type="molting and environmental effects on morphology">
|
||
<paragraph pageId="12" pageNumber="13">Molting and environmental effects on morphology</paragraph>
|
||
<paragraph pageId="12" pageNumber="13">
|
||
About 90 species belong to the genus
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes" order="Gordioidea" pageId="12" pageNumber="13" phylum="Nematomorpha" rank="genus">Chordodes</taxonomicName>
|
||
which were proposed using only five characteristic types of areoles (
|
||
<bibRefCitation author="Schmidt-Rhaesa, A" journalOrPublisher="Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg" pageId="13" pageNumber="14" pagination="37 - 114" title="Summary of Chordodes species (Nematomorpha, Gordiida), with a discussion of their diagnostic characters." volume="44" year="2008">Schmidt-Rhaesa et al. 2008</bibRefCitation>
|
||
). Unfortunately, most characters other than these areole types have seldom been mentioned. According to our observations, the complex ornamentations of the structure of the anterior end were previously considered to be smooth. A horsehair worm explores unknown environments with its head, and the head is also the first part which contacts outer environments when they emerge from a host.
|
||
<bibRefCitation author="Schmidt-Rhaesa, A" journalOrPublisher="Zoologischer Anzeiger" pageId="13" pageNumber="14" pagination="269 - 276" title="Cuticular ultrastructure of Chordodes nobilii Camerano, 1901, with a comparison of cuticular ultrastructure in horsehair worms (Nematomorpha)." url="doi: 10.1016/j.jcz.2006.08.001" volume="245" year="2006">Schmidt-Rhaesa and Gerke (2006)</bibRefCitation>
|
||
suggested that the apical filaments of the crowned areoles may have sensory functions. It is rational that the anterior and posterior ends of horsehair worms with the complex ornamentations play important roles in exploration and mating. Therefore, the smooth anterior ends in some descriptions (e.g.,
|
||
<bibRefCitation author="De Villalobos, C" journalOrPublisher="Systematic Parasitology" pageId="13" pageNumber="14" pagination="117 - 125" title="Scanning electron microscopy and intraspecific variation of Chordodes festae Camerano, 1897 and C. peraccae (Camerano, 1894) (Nematomorpha: Gordioidea)." url="doi: 10.1023/A:1011921326564" volume="50" year="2001">de Villalobos and Zanca 2001</bibRefCitation>
|
||
) and some of our samples may have been caused by damage from the environment. In addition to the damage, some samples were found to have been covered by residual juvenile skins, indicating that the worms had just molted. Shapes of the ornamentations on the ends under the larval cuticle were flat. The molting of hairworms was observed (
|
||
<bibRefCitation author="Schmidt-Rhaesa, A" journalOrPublisher="Proceedings of the Academy of Natural Sciences of Philadelphia" pageId="13" pageNumber="14" pagination="77 - 117" title="Redescription and compilation of Nearctic freshwater Nematomorpha (Gordiida), with the description of two new species." url="doi: 10.1635/0097-3157(2003)153[0077:RACONF]2.0.CO;2" volume="153" year="2003">Schmidt-Rhaesa et al. 2003</bibRefCitation>
|
||
), and its potential effects on the morphology should be carefully considered in the future studies. Residual skin was also found on newly hatched larvae, and on a larva still inside an egg, indicating that molting occurred before it emerged from the egg. As a group of Ecdysozoa (
|
||
<bibRefCitation author="Aguinaldo, AMA" journalOrPublisher="Nature" pageId="13" pageNumber="14" pagination="489 - 493" title="Evidence for a clade of nematodes, arthropods and other moulting animals." volume="387" year="1997">Aguinaldo et al. 1997</bibRefCitation>
|
||
), horsehair worms molt between each instar. According to our observations of metamorphosis from cysts to wormlike juveniles and from juveniles to adults, we suspect those horsehair worms may proceed through at least three molts before maturing.
|
||
</paragraph>
|
||
</subSection>
|
||
<subSection pageId="13" pageNumber="14" type="host specificity">
|
||
<paragraph pageId="13" pageNumber="14">
|
||
<pageBreakToken pageId="13" pageNumber="14" start="start">Host</pageBreakToken>
|
||
specificity
|
||
</paragraph>
|
||
<paragraph pageId="13" pageNumber="14">
|
||
Compared to the wide range of paratenic hosts, horsehair
|
||
<normalizedToken originalValue="worms’">worms'</normalizedToken>
|
||
definitive host range is limited to one or a few species. Because nematomorphs are sometimes found after they have emerged from their hosts, definitive information on hosts is unknown in some species.
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="13" pageNumber="14" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
was reported to be parasites of the mantids,
|
||
<taxonomicName class="Insecta" family="Mantidae" genus="Tenodera" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tenodera sinensis" order="Mantodea" pageId="13" pageNumber="14" phylum="Arthropoda" rank="species" species="sinensis">Tenodera sinensis</taxonomicName>
|
||
,
|
||
<taxonomicName class="Insecta" family="Mantidae" genus="Tenodera" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tenodera angustipennis" order="Mantodea" pageId="13" pageNumber="14" phylum="Arthropoda" rank="species" species="angustipennis">Tenodera angustipennis</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Inoue, I" journalOrPublisher="Annotationes Zoologicae Japonenses" pageId="13" pageNumber="14" pagination="400 - 402" title="On a new species of Chordodes (Gordiacea) from Japan." volume="25" year="1952">Inoue 1952</bibRefCitation>
|
||
),
|
||
<taxonomicName class="Insecta" family="Mantidae" genus="Hierodula" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hierodula patellifera" order="Mantodea" pageId="13" pageNumber="14" phylum="Arthropoda" rank="species" species="patellifera">Hierodula patellifera</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Schmidt-Rhaesa, A" journalOrPublisher="Species Diversity" pageId="13" pageNumber="14" pagination="97 - 107" title="Japanese horsehair worms (Nematomorpha) from the Lake Biwa Museum." volume="9" year="2004">Schmidt-Rhaesa 2004</bibRefCitation>
|
||
), and longhorn grasshoppers,
|
||
<taxonomicName class="Insecta" family="Tettigoniidae" genus="Hexacentrus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hexacentrus japonicus subsp. japonicus" order="Orthoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="subSpecies" species="japonicus" subSpecies="japonicus">Hexacentrus japonicus japonicus</taxonomicName>
|
||
(
|
||
<bibRefCitation pageId="13" pageNumber="14">Inoue 1955</bibRefCitation>
|
||
) in Japan. Since the hairworm described by
|
||
<bibRefCitation author="Schmidt-Rhaesa, A" journalOrPublisher="Species Diversity" pageId="13" pageNumber="14" pagination="97 - 107" title="Japanese horsehair worms (Nematomorpha) from the Lake Biwa Museum." volume="9" year="2004">Schmidt-Rhaesa (2004)</bibRefCitation>
|
||
was here considered to be
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes formosanus" order="Gordioidea" pageId="13" pageNumber="14" phylum="Nematomorpha" rank="species" species="formosanus">Chordodes formosanus</taxonomicName>
|
||
sp. n., the mantid host of
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="13" pageNumber="14" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
is now limited to two species of
|
||
<taxonomicName class="Insecta" family="Mantidae" genus="Tenodera" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tenodera" order="Mantodea" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">Tenodera</taxonomicName>
|
||
. Until now, we have no evidence to exclude
|
||
<taxonomicName class="Insecta" family="Tettigoniidae" genus="Hexacentrus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hexacentrus japonicus subsp. japonicus" order="Orthoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="subSpecies" species="japonicus" subSpecies="japonicus">Hexacentrus japonicus japonicus</taxonomicName>
|
||
from being a potential host of
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="13" pageNumber="14" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
. However, in our investigation, the host range of these two species seems to be limited to the generic level. Therefore, we still question the ability of
|
||
<taxonomicName class="Gordioida" family="Chordodidae" genus="Chordodes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Chordodes japonensis" order="Gordioidea" pageId="13" pageNumber="14" phylum="Nematomorpha" rank="species" species="japonensis">Chordodes japonensis</taxonomicName>
|
||
to develop inside hosts other than mantids.
|
||
</paragraph>
|
||
</subSection>
|
||
<subSection pageId="13" pageNumber="14" type="molecular approach and perspectives">
|
||
<paragraph pageId="13" pageNumber="14">Molecular approach and perspectives</paragraph>
|
||
<paragraph pageId="13" pageNumber="14">
|
||
In the 40 samples from Taiwan and Japan we examined, the taxonomic status was supported not only by their morphologies, but also by the partial COI sequences. COI sequences were used to study inter- and intraspecific relationships due to the high mutation rate (
|
||
<bibRefCitation author="Hebert, PDN" journalOrPublisher="Proceedings of the Royal Society of London Series B-Biological Sciences" pageId="13" pageNumber="14" pagination="313 - 321" title="Biological identifications through DNA barcodes." url="doi: 10.1098/rspb.2002.2218" volume="270" year="2003">Hebert et al. 2003</bibRefCitation>
|
||
). The low genetic divergence among our hairworm samples suggests their conspecific relationship, and our samples can also be separated from those hairworms that emerged from
|
||
<taxonomicName class="Insecta" family="Mantidae" genus="Tenodera" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tenodera" order="Mantodea" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">Tenodera</taxonomicName>
|
||
by theirsignificantly divergent sequences. Since species of immature stages can only be conjectured by adults in the same area (
|
||
<bibRefCitation author="Winterbourn, MJ" journalOrPublisher="Aquatic Insects" pageId="13" pageNumber="14" pagination="155 - 166" title="Dispersal, feeding and parasitism of adult stoneflies (Plecoptera) at a New Zealand forest stream." url="doi: 10.1080/01650420500062840" volume="27" year="2005">Winterbourn 2005</bibRefCitation>
|
||
), molecular data were herein proven to be a useful tool for identifying both adults and immatures. As molecular information for the phylum Nematomorpha is still limited, we believe more molecular data would be helpful and can be used to uncover uncertain relationships among horsehair worms in the near future.
|
||
</paragraph>
|
||
</subSection>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |