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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.854.34549" ID-GBIF-Dataset="c8c8de10-f8f0-45b7-98a5-e42961c3afca" ID-PMC="PMC6580842" ID-Pensoft-Pub="1313-2970-854-145" ID-PubMed="31239821" ID-ZBK="F870F6860B4B4B24858373757950FDB1" ModsDocAuthor="" ModsDocDate="2019" ModsDocID="1313-2970-854-145" ModsDocOrigin="ZooKeys 854" ModsDocTitle="Molecular data reveal a new species of Rhopalias Stiles &amp; Hassall, 1898 (Digenea, Echinostomatidae) in the Common opossum, Didelphismarsupialis L. (Mammalia, Didelphidae) in the Yucatán Peninsula, Mexico" checkinTime="1560202670226" checkinUser="pensoft" docAuthor="Lopez-Caballero, Jorge, Mata-Lopez, Rosario &amp; Leon, Gerardo Perez-Ponce de" docDate="2019" docId="17D66EACFB821A1F9C764B3B971454AB" docLanguage="en" docName="ZooKeys 854: 145-163" docOrigin="ZooKeys 854" docSource="http://dx.doi.org/10.3897/zookeys.854.34549" docTitle="Rhopalias oochi Lopez-Caballero, Mata-Lopez &amp; Leon, 2019, sp. nov." docType="treatment" docUuid="2AFA9155-52CE-4436-A95F-2B0E85F93C72" docUuidSource="ZooBank" docVersion="5" lastPageNumber="152" masterDocId="FFD7D3667F1D117DFFFED172FFF21245" masterDocTitle="Molecular data reveal a new species of Rhopalias Stiles &amp; Hassall, 1898 (Digenea, Echinostomatidae) in the Common opossum, Didelphismarsupialis L. (Mammalia, Didelphidae) in the Yucatan Peninsula, Mexico" masterLastPageNumber="163" masterPageNumber="145" pageNumber="151" updateTime="1668167377334" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>Molecular data reveal a new species of Rhopalias Stiles &amp; Hassall, 1898 (Digenea, Echinostomatidae) in the Common opossum, Didelphismarsupialis L. (Mammalia, Didelphidae) in the Yucatan Peninsula, Mexico</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Lopez-Caballero, Jorge</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Mata-Lopez, Rosario</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Leon, Gerardo Perez-Ponce de</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>2019</mods:date>
<mods:detail type="volume">
<mods:number>854</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>145</mods:start>
<mods:end>163</mods:end>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.854.34549</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.854.34549</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-854-145</mods:identifier>
<mods:identifier type="ZBK">F870F6860B4B4B24858373757950FDB1</mods:identifier>
<mods:identifier type="ZooBank">F870F6860B4B4B24858373757950FDB1</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="157500986" LSID="urn:lsid:zoobank.org:act:2AFA9155-52CE-4436-A95F-2B0E85F93C72" httpUri="http://treatment.plazi.org/id/17D66EACFB821A1F9C764B3B971454AB" lastPageId="8" lastPageNumber="152" pageId="6" pageNumber="151">
<subSubSection pageId="6" pageNumber="151" type="nomenclature">
<paragraph pageId="6" pageNumber="151">
<taxonomicName LSID="http://zoobank.org/2AFA9155-52CE-4436-A95F-2B0E85F93C72" class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Rhopalias oochi" order="Plagiorchiida" pageId="6" pageNumber="151" phylum="Platyhelminthes" rank="species" species="oochi">Rhopalias oochi</taxonomicName>
<taxonomicNameLabel pageId="6" pageNumber="151">sp. nov.</taxonomicNameLabel>
Figures 3
<normalizedToken originalValue="AD">A-D</normalizedToken>
, 4 A, D, E, 5A, D
</paragraph>
</subSubSection>
<subSubSection pageId="6" pageNumber="151" type="synonym">
<paragraph pageId="6" pageNumber="151">Synonym.</paragraph>
<paragraph pageId="6" pageNumber="151">
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias coronatus" order="Plagiorchiida" pageId="6" pageNumber="151" phylum="Platyhelminthes" rank="species" species="coronatus">Rhopalias coronatus</taxonomicName>
of
<bibRefCitation author="Acosta-Virgen, K" journalOrPublisher="ZooKeys" pageId="12" pageNumber="157" pagination="131 - 152" title="Helminths of three species of opossums (Mammalia, Didelphidae) from Mexico." url="https://doi.org/10.3897/zookeys.511.9571" volume="511" year="2015">Acosta-Virgen et al. (2015)</bibRefCitation>
. Specimens deposited in the CNHE (9504).
</paragraph>
</subSubSection>
<subSubSection pageId="6" pageNumber="151" type="type host">
<paragraph pageId="6" pageNumber="151">Type host.</paragraph>
<paragraph pageId="6" pageNumber="151">
<taxonomicName class="Mammalia" family="Didelphidae" genus="Didelphis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Didelphis marsupialis" order="Didelphimorphia" pageId="6" pageNumber="151" phylum="Chordata" rank="species" species="marsupialis">Didelphis marsupialis</taxonomicName>
Linnaeus, Common
<taxonomicName genus="Echinostomatoidea" lsidName="Echinostomatoidea opossum" pageId="6" pageNumber="151" rank="species" species="opossum">opossum</taxonomicName>
(
<taxonomicName class="Mammalia" lsidName="Echinostomatoidea" pageId="6" pageNumber="151" rank="class">Mammalia</taxonomicName>
:
<taxonomicName family="Didelphidae" lsidName="" pageId="6" pageNumber="151" rank="family">Didelphidae</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="152" type="type locality">
<paragraph pageId="7" pageNumber="152">
<pageBreakToken pageId="7" pageNumber="152" start="start">Type</pageBreakToken>
locality.
</paragraph>
<paragraph pageId="7" pageNumber="152">
Rancho Hobonil, Tzucacab,
<normalizedToken originalValue="Yucatán">Yucatan</normalizedToken>
state, Mexico (
<geoCoordinate direction="north" orientation="latitude" precision="15" value="20.016111">20°00'58&quot;N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="15" value="-89.020004">89°01'12&quot;W</geoCoordinate>
).
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="152" type="site in host">
<paragraph pageId="7" pageNumber="152">Site in host.</paragraph>
<paragraph pageId="7" pageNumber="152">Small intestine.</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="152" type="prevalence and intesity of infection">
<paragraph pageId="7" pageNumber="152">Prevalence and intesity of infection.</paragraph>
<paragraph pageId="7" pageNumber="152">
100% (1 of 1
<taxonomicName genus="Echinostomatoidea" lsidName="Echinostomatoidea opossum" pageId="7" pageNumber="152" rank="species" species="opossum">opossum</taxonomicName>
), infected with 15 trematodes.
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="152" type="type specimens">
<paragraph pageId="7" pageNumber="152">Type specimens.</paragraph>
<paragraph pageId="7" pageNumber="152">Holotype: CNHE 9504; paratypes: CNHE 10926 (3 specimens) and hologenophores CNHE 11069 (4 specimens).</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="152" type="etymology">
<paragraph pageId="7" pageNumber="152">Etymology.</paragraph>
<paragraph pageId="7" pageNumber="152">
The specific epithet refers to the common name of the host where the new species was found. In the Mayan language,
<normalizedToken originalValue="“ooch”">&quot;ooch&quot;</normalizedToken>
means
<taxonomicName genus="Echinostomatoidea" lsidName="Echinostomatoidea opossum" pageId="7" pageNumber="152" rank="species" species="opossum">opossum</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="152" type="description">
<paragraph pageId="7" pageNumber="152">Description.</paragraph>
<paragraph pageId="7" pageNumber="152">Based on 11 adult specimens (including 4 hologenophores). Measurements are given in Table 3. Trematodes with a long body, forebody concave, wider than hindbody, with a pair of armed retractile tentacles with 4-7 spines (Fig. 3A, B, C). Tegument covered with spines reaching posterior end of body (Fig. 4A). Tegument spines U-shaped, with distal tip pectinated (Fig. 4D, E). Oral and flanking spines present (Figs 3C, 5A, 5D). Muscular sacs long, reaching far beyond posterior margin of pharynx, and may or may not reach the anterior margin of ventral sucker (Fig. 3A, B). Oral sucker subterminal, rounded, well-developed, short prepharynx, pharynx muscular, and relatively long oesophagus; caecal bifurcation at short distance anterior to genital pore (Fig. 3B); long caeca extending to the posterior end of body (Fig. 3A). Ventral sucker muscular and subspherical, in the first third of body, larger than oral sucker (Figs 3A, B, 4A). Testes two, elongated, in tandem, contiguous, no overlapped, located in mid-body; anterior testis shorter than posterior testis (Fig. 3B). Cirrus sac long, claviform, containing a well-developed prostate complex and seminal vesicle, extending beyond ventral sucker and terminating near anterior border of ovary. Genital pore between ventral sucker and caecal bifurcation. Ovary slightly oval, postacetabular, pretesticular. Uterus intercaecal, between ovary and genital pore. Metraterm long (Fig. 4D). Vitelline follicles in lateral fields, beginning at mid-level between ventral sucker and ovary, ending at posterior end. Gravid specimens with few eggs, oval-shaped, operculated; embryonated eggs with thin shell.</paragraph>
<caption pageId="7" pageNumber="152">
<paragraph pageId="7" pageNumber="152">
Figure 3.
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias oochi" order="Plagiorchiida" pageId="7" pageNumber="152" phylum="Platyhelminthes" rank="species" species="oochi">Rhopalias oochi</taxonomicName>
sp. nov., parasite of
<taxonomicName class="Mammalia" family="Didelphidae" genus="Didelphis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Didelphis marsupialis" order="Didelphimorphia" pageId="7" pageNumber="152" phylum="Chordata" rank="species" species="marsupialis">Didelphis marsupialis</taxonomicName>
from
<normalizedToken originalValue="Yucatán">Yucatan</normalizedToken>
, Mexico. A Microphotograph, ventral view of entire body B Line drawing, ventral view C Detail of the oral, flanking and tentacle spines, ventral view D Cirrus sac, vagina, and eggs, ventral view. Scale bars: 10
<normalizedToken originalValue="μm">μm</normalizedToken>
(
<normalizedToken originalValue="AC">A-C</normalizedToken>
); 400
<normalizedToken originalValue="μm">μm</normalizedToken>
(D).
</paragraph>
</caption>
<caption pageId="7" pageNumber="152">
<paragraph pageId="7" pageNumber="152">
Figure 4. SEM micrographs of
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias" order="Plagiorchiida" pageId="7" pageNumber="152" phylum="Platyhelminthes" rank="genus">Rhopalias</taxonomicName>
species.
<normalizedToken originalValue="AC">A-C</normalizedToken>
Ventral view of the body of adult specimens showing the distribution of spines
<normalizedToken originalValue="DI">D-I</normalizedToken>
Detail of the spines shape on the ventral surface of hindbody A, D, E
<taxonomicName lsidName="R. oochi" pageId="7" pageNumber="152" rank="species" species="oochi">R. oochi</taxonomicName>
sp. nov. B, F, G
<taxonomicName lsidName="R. coronatus" pageId="7" pageNumber="152" rank="species" species="coronatus">R. coronatus</taxonomicName>
C, H, I
<taxonomicName lsidName="R. macracanthus" pageId="7" pageNumber="152" rank="species" species="macracanthus">R. macracanthus</taxonomicName>
. Scale bars: 100
<normalizedToken originalValue="μm">μm</normalizedToken>
(
<normalizedToken originalValue="AC">A-C</normalizedToken>
); 50
<normalizedToken originalValue="μm">μm</normalizedToken>
(D, E); 10
<normalizedToken originalValue="μm">μm</normalizedToken>
(F, G); 20
<normalizedToken originalValue="μm">μm</normalizedToken>
(H, I).
</paragraph>
</caption>
</subSubSection>
<subSubSection lastPageId="8" lastPageNumber="153" pageId="7" pageNumber="152" type="remarks">
<paragraph pageId="7" pageNumber="152">Remarks.</paragraph>
<paragraph lastPageId="8" lastPageNumber="153" pageId="7" pageNumber="152">
The genus
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias" order="Plagiorchiida" pageId="7" pageNumber="152" phylum="Platyhelminthes" rank="genus">Rhopalias</taxonomicName>
currently contains six species as parasites of New World marsupials (
<bibRefCitation author="Haverkost, TR" journalOrPublisher="Journal of Parasitology" pageId="12" pageNumber="157" pagination="716 - 726" title="A review of species in the genus Rhopalias (Rudolphi, 1819)." url="https://doi.org/10.1645/GE-1423.1" volume="94" year="2008">Haverkost and Gardner 2008</bibRefCitation>
), i.e.,
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias coronatus" order="Plagiorchiida" pageId="7" pageNumber="152" phylum="Platyhelminthes" rank="species" species="coronatus">Rhopalias coronatus</taxonomicName>
, the type species;
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias horridus" order="Plagiorchiida" pageId="7" pageNumber="152" phylum="Platyhelminthes" rank="species" species="horridus">Rhopalias horridus</taxonomicName>
(Diesing, 1850)
<bibRefCitation author="Stiles, CW" journalOrPublisher="Archives de Parasitologie" pageId="14" pageNumber="159" pagination="81 - 99" title="Notes on parasites, an inventory of the genera and subgenera of the trematode family Fasciolidae." volume="1" year="1898">Stiles and Hassall 1898</bibRefCitation>
;
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias baculifer" order="Plagiorchiida" pageId="7" pageNumber="152" phylum="Platyhelminthes" rank="species" species="baculifer">Rhopalias baculifer</taxonomicName>
Braun, 1901;
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias macracanthus" order="Plagiorchiida" pageId="7" pageNumber="152" phylum="Platyhelminthes" rank="species" species="macracanthus">Rhopalias macracanthus</taxonomicName>
Chandler, 1932;
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias caballeroi" order="Plagiorchiida" pageId="7" pageNumber="152" phylum="Platyhelminthes" rank="species" species="caballeroi">Rhopalias caballeroi</taxonomicName>
Kifune &amp; Uyema, 1982; and
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias caucensis" order="Plagiorchiida" pageId="7" pageNumber="152" phylum="Platyhelminthes" rank="species" species="caucensis">Rhopalias caucensis</taxonomicName>
Rivillas, Caro, Carvajal &amp;
<normalizedToken originalValue="Vélez">Velez</normalizedToken>
, 2004.
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias oochi" order="Plagiorchiida" pageId="7" pageNumber="152" phylum="Platyhelminthes" rank="species" species="oochi">Rhopalias oochi</taxonomicName>
sp. nov. represents the seventh described species and is readily distinguished from five of the congeneric species, excepting
<taxonomicName lsidName="R. coronatus" pageId="7" pageNumber="152" rank="species" species="coronatus">R. coronatus</taxonomicName>
by having tentacle sacs extending far beyond the posterior margin of pharynx. In their detailed morphological revision of the genus
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias" order="Plagiorchiida" pageId="7" pageNumber="152" phylum="Platyhelminthes" rank="genus">Rhopalias</taxonomicName>
,
<bibRefCitation author="Haverkost, TR" journalOrPublisher="Journal of Parasitology" pageId="12" pageNumber="157" pagination="716 - 726" title="A review of species in the genus Rhopalias (Rudolphi, 1819)." url="https://doi.org/10.1645/GE-1423.1" volume="94" year="2008">Haverkost and Gardner (2008)</bibRefCitation>
provided a key to species of the genus and found that this character is reliable to distinguish between two groups of species. In
<taxonomicName lsidName="R. horridus" pageId="7" pageNumber="152" rank="species" species="horridus">R. horridus</taxonomicName>
, the tentacle sacs surpass the posterior margin of pharynx to reach the mid-level of oesophagus; however, they never extend beyond the caecal bifurcation. In
<taxonomicName lsidName="R. oochi" pageId="7" pageNumber="152" rank="species" species="oochi">R. oochi</taxonomicName>
sp. nov. the tentacle sacs extend to reach the anterior margin of ventral sucker.
<taxonomicName class="Trematoda" family="Echinostomatidae" genus="Rhopalias" higherTaxonomySource="GBIF" kingdom="Metazoa" lsidName="Rhopalias horridus" order="Plagiorchiida" pageId="7" pageNumber="152" phylum="Platyhelminthes" rank="species" species="horridus">Rhopalias horridus</taxonomicName>
further differ from the new species, and the remaining congeners by the large number of tentacle spines (&gt; 30) (see key in
<bibRefCitation author="Haverkost, TR" journalOrPublisher="Journal of Parasitology" pageId="12" pageNumber="157" pagination="716 - 726" title="A review of species in the genus Rhopalias (Rudolphi, 1819)." url="https://doi.org/10.1645/GE-1423.1" volume="94" year="2008">Haverkost and Gardner 2008</bibRefCitation>
).
<pageBreakToken pageId="8" pageNumber="153" start="start">The</pageBreakToken>
new species most closely resembles the type species,
<taxonomicName lsidName="R. coronatus" pageId="8" pageNumber="153" rank="species" species="coronatus">R. coronatus</taxonomicName>
; in fact the specimens upon which the description of the new species is based were originally recorded as
<taxonomicName lsidName="R. coronatus" pageId="8" pageNumber="153" rank="species" species="coronatus">R. coronatus</taxonomicName>
(see
<bibRefCitation author="Acosta-Virgen, K" journalOrPublisher="ZooKeys" pageId="12" pageNumber="157" pagination="131 - 152" title="Helminths of three species of opossums (Mammalia, Didelphidae) from Mexico." url="https://doi.org/10.3897/zookeys.511.9571" volume="511" year="2015">Acosta-Virgen et al. 2015</bibRefCitation>
). Genetic data revealed that they might actually represent a different species. In
<taxonomicName lsidName="R. coronatus" pageId="8" pageNumber="153" rank="species" species="coronatus">R. coronatus</taxonomicName>
as in the new species tentacle sacs extend far beyond the posterior border of pharynx; in
<taxonomicName lsidName="R. coronatus" pageId="8" pageNumber="153" rank="species" species="coronatus">R. coronatus</taxonomicName>
, both sacs usually surpass the anterior border of ventral sucker (although our observation of numerous voucher specimens indicate that the extension is variable among individuals); in the new species, the tentacle sacs consistently extend to almost reach the anterior border of ventral sucker, but never surpass it. Additionally,
<taxonomicName lsidName="R. coronatus" pageId="8" pageNumber="153" rank="species" species="coronatus">R. coronatus</taxonomicName>
differ from the new species by having larger oral spines (at least four times larger), and by the distribution of vitelline follicles. In
<taxonomicName lsidName="R. coronatus" pageId="8" pageNumber="153" rank="species" species="coronatus">R. coronatus</taxonomicName>
follicles extend between the posterior end of body and the posterior border of ventral sucker; instead, in
<taxonomicName lsidName="R. oochi" pageId="8" pageNumber="153" rank="species" species="oochi">R. oochi</taxonomicName>
sp. nov. vitelline follicles do not reach the posterior end of body, and in the forebody, they reach the mid-level of cirrus sac. Morphometrically, most measurements are clearly overlapped between the new species and
<taxonomicName lsidName="R. coronatus" pageId="8" pageNumber="153" rank="species" species="coronatus">R. coronatus</taxonomicName>
(except in oral spines size). We observed and measured 21 individuals of
<taxonomicName lsidName="R. coronatus" pageId="8" pageNumber="153" rank="species" species="coronatus">R. coronatus</taxonomicName>
deposited at the CNHE. Our measurements, and those reported in other studies (see Table 3) indicate that
<taxonomicName lsidName="R. coronatus" pageId="8" pageNumber="153" rank="species" species="coronatus">R. coronatus</taxonomicName>
is morphometrically quite variable. Furthermore, SEM study of the ultrastructure of the body surface provided additional evidence to distinguish the new species from
<taxonomicName lsidName="R. coronatus" pageId="8" pageNumber="153" rank="species" species="coronatus">R. coronatus</taxonomicName>
and
<taxonomicName lsidName="R. macracanthus" pageId="8" pageNumber="153" rank="species" species="macracanthus">R. macracanthus</taxonomicName>
; for these two species we collected specimens and some were fixed for a comparative SEM study (Fig. 4). In the three species body spines are denser in the forebody; however, the extension, size, and shape of spines clearly distinguish the three species. In
<taxonomicName lsidName="R. oochi" pageId="8" pageNumber="153" rank="species" species="oochi">R. oochi</taxonomicName>
sp. nov., spines are robust and pectinate and they extend along the body almost to the posterior end (Fig. 4A, D, E); meanwhile, in
<taxonomicName lsidName="R. coronatus" pageId="8" pageNumber="153" rank="species" species="coronatus">R. coronatus</taxonomicName>
spines are distributed to the mid-level of hindbody and are entire and tongue-shaped (Fig. 4B, F, G) and in
<taxonomicName lsidName="R. macracanthus" pageId="8" pageNumber="153" rank="species" species="macracanthus">R. macracanthus</taxonomicName>
, even though spines extend more posteriorly, they are not as robust as in the new species; spines in
<taxonomicName lsidName="R. macracanthus" pageId="8" pageNumber="153" rank="species" species="macracanthus">R. macracanthus</taxonomicName>
are pectinate and arrow-shaped (Fig. 4C, H, I).
</paragraph>
</subSubSection>
</treatment>
</document>