treatments-xml/data/03/81/87/03818787645EFFD21BD2FF44FC2470A8.xml

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<document id="620FE9F8EB88B204125F4E96FB6A5002" ID-DOI="10.11646/zootaxa.5506.1.1" ID-ISSN="1175-5326" ID-Zenodo-Dep="13745824" ID-ZooBank="AFDFAEE4-9B4A-4792-80E7-27DC9ECC23D8" IM.bibliography_approvedBy="carolina" IM.illustrations_approvedBy="carolina" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="carolina" IM.taxonomicNames_approvedBy="carolina" IM.treatments_approvedBy="carolina" checkinTime="1726052091569" checkinUser="plazi" docAuthor="Titus, Benjamin M., Bennett-Smith, Morgan F., Chiodo, Tommaso &amp; Rodríguez, Estefanía" docDate="2024" docId="03818787645EFFD21BD2FF44FC2470A8" docLanguage="en" docName="zootaxa.5506.1.1.pdf" docOrigin="Zootaxa 5506 (1)" docSource="http://dx.doi.org/10.11646/zootaxa.5506.1.1" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Radianthus crispa" docType="treatment" docVersion="4" lastPageNumber="19" masterDocId="FFB8FFFF644EFFC01B45FF9BFFDB7179" masterDocTitle="The clownfish-hosting sea anemones (Anthozoa: Actiniaria): updated nomenclature, biogeography, and practical field guide." masterLastPageNumber="34" masterPageNumber="1" pageNumber="17" updateTime="1726245086095" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
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<mods:title id="252BB7F7CFBE97BCE4081292A4B02EE8">The clownfish-hosting sea anemones (Anthozoa: Actiniaria): updated nomenclature, biogeography, and practical field guide.</mods:title>
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<mods:affiliation id="2635318AE8EE2F60788F834661448FAD">Department of Biological Sciences, University of Alabama, Tuscaloosa, AL, USA 35487 &amp; Dauphin Island Sea Lab, 101 Bienville Blvd, Dauphin Island, AL, USA 36528</mods:affiliation>
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<paragraph id="8B973691645EFFD01BD2FF44FBC87180" blockId="16.[151,1043,223,250]" box="[151,1043,223,250]" pageId="16" pageNumber="17">
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<taxonomicName id="4C284D12645EFFD01BD2FF44FCDE7180" ID-CoL="9GPNW" authority="(Hemprich &amp; Ehrenberg, 1834)" baseAuthorityName="Hemprich &amp; Ehrenberg" baseAuthorityYear="1834" box="[151,773,223,250]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="16" pageNumber="17" phylum="Cnidaria" rank="species" species="crispa">
Radianthus crispa (Hemprich &amp;
<bibRefCitation id="EFB94B60645EFFD0196EFF44FD267180" author="Ehrenberg, C. G." box="[555,765,223,249]" pageId="16" pageNumber="17" pagination="225 - 380" refId="ref16320" refString="Ehrenberg, C. G. (1834) Beitrage zur physiologischen Kenntniss der Corallenthiere im allgemeinen, und besonders des rothen Meeres, nebst einem Versuche zur physiologischen Systematik derselben. Abhandlungen der Koniglichen Akademie der Wissenschaften zu Berlin, 1, 225 - 380." type="journal article" year="1834">Ehrenberg, 1834</bibRefCitation>
)
</taxonomicName>
(
<figureCitation id="13132A14645EFFD01851FF44FC5A7180" box="[788,897,223,249]" captionStart="FIGURE 6" captionStartId="15.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@15.[226,1361,181,1728]" captionTargetPageId="15" captionText="FIGURE 6. Representative images of the leathery sea anemone Radianthus crispa encompassing a broad range of geographic and phenotypic variation. A) Yellow individual hosting juvenile Amphiprion chrysopterus on a typical fore reef environment in the Pacific Ocean. Note the densely packed tentacles with tapered tips coming to a point (Moorea, French Polynesia). B) White individual with pink tentacle tips (Moorea, French Polynesia). C) Tan/Brown individual with less densely packed tentacles that curl slightly (Saudi Arabia, Red Sea). D) Macro photograph underneath oral disc. i = well-defined raised verrucae that are similarly colored to the surrounding column (Okinawa, Japan). E) Small individual in stereotypical sandy microhabitat with slightly curled tentacled tips and horizontally striated tentacle pattern (Gulf of Oman, United Arab Emirates). Photographs by Benjamin M. Titus and James Reimer." figureDoi="http://doi.org/10.5281/zenodo.13745836" httpUri="https://zenodo.org/record/13745836/files/figure.png" pageId="16" pageNumber="17">Figure 6</figureCitation>
;
<figureCitation id="13132A14645EFFD018D5FF44FBD27180" box="[912,1033,223,249]" captionStart="FIGURE 3" captionStartId="12.[152,255,1779,1803]" captionTargetBox="[190,1386,1209,1756]" captionTargetId="figure-510@12.[189,1398,1209,1756]" captionTargetPageId="12" captionText="FIGURE 3. Confirmed geographic range of Entacmaea quadricolor in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry)." figureDoi="http://doi.org/10.5281/zenodo.13745830" httpUri="https://zenodo.org/record/13745830/files/figure.png" pageId="16" pageNumber="17">Figure S3</figureCitation>
)
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<subSubSection id="C332651A645EFFD21BD2FEBCFC2470A8" lastPageId="18" lastPageNumber="19" pageId="16" pageNumber="17" type="discussion">
<paragraph id="8B973691645EFFD01BD2FEBCFAC273B4" blockId="16.[151,1437,295,1114]" pageId="16" pageNumber="17">
The leathery sea anemone,
<taxonomicName id="4C284D12645EFFD01A94FEBCFD7B7038" baseAuthorityName="Hemprich &amp; Ehrenberg" baseAuthorityYear="1834" box="[465,672,295,321]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="16" pageNumber="17" phylum="Cnidaria" rank="species" species="crispa">Radianthus crispa</taxonomicName>
(
<figureCitation id="13132A14645EFFD019F5FEBCFCCE7038" box="[688,789,295,321]" captionStart="FIGURE 6" captionStartId="15.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@15.[226,1361,181,1728]" captionTargetPageId="15" captionText="FIGURE 6. Representative images of the leathery sea anemone Radianthus crispa encompassing a broad range of geographic and phenotypic variation. A) Yellow individual hosting juvenile Amphiprion chrysopterus on a typical fore reef environment in the Pacific Ocean. Note the densely packed tentacles with tapered tips coming to a point (Moorea, French Polynesia). B) White individual with pink tentacle tips (Moorea, French Polynesia). C) Tan/Brown individual with less densely packed tentacles that curl slightly (Saudi Arabia, Red Sea). D) Macro photograph underneath oral disc. i = well-defined raised verrucae that are similarly colored to the surrounding column (Okinawa, Japan). E) Small individual in stereotypical sandy microhabitat with slightly curled tentacled tips and horizontally striated tentacle pattern (Gulf of Oman, United Arab Emirates). Photographs by Benjamin M. Titus and James Reimer." figureDoi="http://doi.org/10.5281/zenodo.13745836" httpUri="https://zenodo.org/record/13745836/files/figure.png" pageId="16" pageNumber="17">Figure 6</figureCitation>
;
<figureCitation id="13132A14645EFFD01866FEBCFC4F7038" box="[803,916,295,321]" captionStart="FIGURE 3" captionStartId="12.[152,255,1779,1803]" captionTargetBox="[190,1386,1209,1756]" captionTargetId="figure-510@12.[189,1398,1209,1756]" captionTargetPageId="12" captionText="FIGURE 3. Confirmed geographic range of Entacmaea quadricolor in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry)." figureDoi="http://doi.org/10.5281/zenodo.13745830" httpUri="https://zenodo.org/record/13745830/files/figure.png" pageId="16" pageNumber="17">Figure S3</figureCitation>
), was originally described from the Red Sea. This species gets its common name from its thick body wall, which gives the animal a tough leathery texture. The tentacles in this species are all alike, elongate (up to
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), tapering gradually to a pointed tip (
<figureCitation id="13132A14645EFFD01F97FEF4FA9370F0" box="[1234,1352,367,393]" captionStart="FIGURE 6" captionStartId="15.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@15.[226,1361,181,1728]" captionTargetPageId="15" captionText="FIGURE 6. Representative images of the leathery sea anemone Radianthus crispa encompassing a broad range of geographic and phenotypic variation. A) Yellow individual hosting juvenile Amphiprion chrysopterus on a typical fore reef environment in the Pacific Ocean. Note the densely packed tentacles with tapered tips coming to a point (Moorea, French Polynesia). B) White individual with pink tentacle tips (Moorea, French Polynesia). C) Tan/Brown individual with less densely packed tentacles that curl slightly (Saudi Arabia, Red Sea). D) Macro photograph underneath oral disc. i = well-defined raised verrucae that are similarly colored to the surrounding column (Okinawa, Japan). E) Small individual in stereotypical sandy microhabitat with slightly curled tentacled tips and horizontally striated tentacle pattern (Gulf of Oman, United Arab Emirates). Photographs by Benjamin M. Titus and James Reimer." figureDoi="http://doi.org/10.5281/zenodo.13745836" httpUri="https://zenodo.org/record/13745836/files/figure.png" pageId="16" pageNumber="17">Figure 6A</figureCitation>
;
<figureCitation id="13132A14645EFFD01E16FEF4FF6C70D4" captionStart="FIGURE 3" captionStartId="12.[152,255,1779,1803]" captionTargetBox="[190,1386,1209,1756]" captionTargetId="figure-510@12.[189,1398,1209,1756]" captionTargetPageId="12" captionText="FIGURE 3. Confirmed geographic range of Entacmaea quadricolor in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry)." figureDoi="http://doi.org/10.5281/zenodo.13745830" httpUri="https://zenodo.org/record/13745830/files/figure.png" pageId="16" pageNumber="17">Figure S3</figureCitation>
). Tentacles are numerous (up to
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the largest individuals) and can be densely packed in some individuals. Tentacles frequently curl, yet only slightly, and not to the same degree as in
<taxonomicName id="4C284D12645EFFD01897FE23FB8170A8" baseAuthorityName="Quoy &amp; Gaimard" baseAuthorityYear="1883" box="[978,1114,439,465]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="16" pageNumber="17" phylum="Cnidaria" rank="species" species="doreensis">R. doreensis</taxonomicName>
(
<figureCitation id="13132A14645EFFD01F22FE2CFAD070AB" box="[1127,1291,439,466]" captionStart="FIGURE 6" captionStartId="15.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@15.[226,1361,181,1728]" captionTargetPageId="15" captionText="FIGURE 6. Representative images of the leathery sea anemone Radianthus crispa encompassing a broad range of geographic and phenotypic variation. A) Yellow individual hosting juvenile Amphiprion chrysopterus on a typical fore reef environment in the Pacific Ocean. Note the densely packed tentacles with tapered tips coming to a point (Moorea, French Polynesia). B) White individual with pink tentacle tips (Moorea, French Polynesia). C) Tan/Brown individual with less densely packed tentacles that curl slightly (Saudi Arabia, Red Sea). D) Macro photograph underneath oral disc. i = well-defined raised verrucae that are similarly colored to the surrounding column (Okinawa, Japan). E) Small individual in stereotypical sandy microhabitat with slightly curled tentacled tips and horizontally striated tentacle pattern (Gulf of Oman, United Arab Emirates). Photographs by Benjamin M. Titus and James Reimer." figureDoi="http://doi.org/10.5281/zenodo.13745836" httpUri="https://zenodo.org/record/13745836/files/figure.png" pageId="16" pageNumber="17">Figure 6C &amp; E</figureCitation>
,
<figureCitation id="13132A14645EFFD01E50FE2CFF76708C" captionStart="FIGURE 3" captionStartId="12.[152,255,1779,1803]" captionTargetBox="[190,1386,1209,1756]" captionTargetId="figure-510@12.[189,1398,1209,1756]" captionTargetPageId="12" captionText="FIGURE 3. Confirmed geographic range of Entacmaea quadricolor in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry)." figureDoi="http://doi.org/10.5281/zenodo.13745830" httpUri="https://zenodo.org/record/13745830/files/figure.png" pageId="16" pageNumber="17">Figure S3A, C</figureCitation>
). The body and tentacle color of this species range from white and light purple to tan/brown or yellow and may take on a matte appearance (
<figureCitation id="13132A14645EFFD01A8BFE64FDF47360" box="[462,559,511,537]" captionStart="FIGURE 6" captionStartId="15.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@15.[226,1361,181,1728]" captionTargetPageId="15" captionText="FIGURE 6. Representative images of the leathery sea anemone Radianthus crispa encompassing a broad range of geographic and phenotypic variation. A) Yellow individual hosting juvenile Amphiprion chrysopterus on a typical fore reef environment in the Pacific Ocean. Note the densely packed tentacles with tapered tips coming to a point (Moorea, French Polynesia). B) White individual with pink tentacle tips (Moorea, French Polynesia). C) Tan/Brown individual with less densely packed tentacles that curl slightly (Saudi Arabia, Red Sea). D) Macro photograph underneath oral disc. i = well-defined raised verrucae that are similarly colored to the surrounding column (Okinawa, Japan). E) Small individual in stereotypical sandy microhabitat with slightly curled tentacled tips and horizontally striated tentacle pattern (Gulf of Oman, United Arab Emirates). Photographs by Benjamin M. Titus and James Reimer." figureDoi="http://doi.org/10.5281/zenodo.13745836" httpUri="https://zenodo.org/record/13745836/files/figure.png" pageId="16" pageNumber="17">Figure 6</figureCitation>
;
<figureCitation id="13132A14645EFFD0197FFE64FD727360" box="[570,681,511,537]" captionStart="FIGURE 3" captionStartId="12.[152,255,1779,1803]" captionTargetBox="[190,1386,1209,1756]" captionTargetId="figure-510@12.[189,1398,1209,1756]" captionTargetPageId="12" captionText="FIGURE 3. Confirmed geographic range of Entacmaea quadricolor in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry)." figureDoi="http://doi.org/10.5281/zenodo.13745830" httpUri="https://zenodo.org/record/13745830/files/figure.png" pageId="16" pageNumber="17">Figure S3</figureCitation>
). Some individuals are bright pink/magenta, and it is not uncommon to encounter individuals with pink-tipped tentacles (
<figureCitation id="13132A14645EFFD0199DFDB8FC977344" box="[728,844,547,573]" captionStart="FIGURE 6" captionStartId="15.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@15.[226,1361,181,1728]" captionTargetPageId="15" captionText="FIGURE 6. Representative images of the leathery sea anemone Radianthus crispa encompassing a broad range of geographic and phenotypic variation. A) Yellow individual hosting juvenile Amphiprion chrysopterus on a typical fore reef environment in the Pacific Ocean. Note the densely packed tentacles with tapered tips coming to a point (Moorea, French Polynesia). B) White individual with pink tentacle tips (Moorea, French Polynesia). C) Tan/Brown individual with less densely packed tentacles that curl slightly (Saudi Arabia, Red Sea). D) Macro photograph underneath oral disc. i = well-defined raised verrucae that are similarly colored to the surrounding column (Okinawa, Japan). E) Small individual in stereotypical sandy microhabitat with slightly curled tentacled tips and horizontally striated tentacle pattern (Gulf of Oman, United Arab Emirates). Photographs by Benjamin M. Titus and James Reimer." figureDoi="http://doi.org/10.5281/zenodo.13745836" httpUri="https://zenodo.org/record/13745836/files/figure.png" pageId="16" pageNumber="17">Figure 6B</figureCitation>
;
<figureCitation id="13132A14645EFFD01812FDB8FC227344" box="[855,1017,547,573]" captionStart="FIGURE 3" captionStartId="12.[152,255,1779,1803]" captionTargetBox="[190,1386,1209,1756]" captionTargetId="figure-510@12.[189,1398,1209,1756]" captionTargetPageId="12" captionText="FIGURE 3. Confirmed geographic range of Entacmaea quadricolor in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry)." figureDoi="http://doi.org/10.5281/zenodo.13745830" httpUri="https://zenodo.org/record/13745830/files/figure.png" pageId="16" pageNumber="17">Figure S3C, E</figureCitation>
). Tentacle patterns vary, but generally have a mottled (
<figureCitation id="13132A14645EFFD01A0DFDDCFDD77318" box="[328,524,583,610]" captionStart="FIGURE 3" captionStartId="12.[152,255,1779,1803]" captionTargetBox="[190,1386,1209,1756]" captionTargetId="figure-510@12.[189,1398,1209,1756]" captionTargetPageId="12" captionText="FIGURE 3. Confirmed geographic range of Entacmaea quadricolor in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry)." figureDoi="http://doi.org/10.5281/zenodo.13745830" httpUri="https://zenodo.org/record/13745830/files/figure.png" pageId="16" pageNumber="17">Figure S3A, B, E</figureCitation>
), lightly striped (
<figureCitation id="13132A14645EFFD0198CFDDCFCE67318" box="[713,829,583,609]" captionStart="FIGURE 6" captionStartId="15.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@15.[226,1361,181,1728]" captionTargetPageId="15" captionText="FIGURE 6. Representative images of the leathery sea anemone Radianthus crispa encompassing a broad range of geographic and phenotypic variation. A) Yellow individual hosting juvenile Amphiprion chrysopterus on a typical fore reef environment in the Pacific Ocean. Note the densely packed tentacles with tapered tips coming to a point (Moorea, French Polynesia). B) White individual with pink tentacle tips (Moorea, French Polynesia). C) Tan/Brown individual with less densely packed tentacles that curl slightly (Saudi Arabia, Red Sea). D) Macro photograph underneath oral disc. i = well-defined raised verrucae that are similarly colored to the surrounding column (Okinawa, Japan). E) Small individual in stereotypical sandy microhabitat with slightly curled tentacled tips and horizontally striated tentacle pattern (Gulf of Oman, United Arab Emirates). Photographs by Benjamin M. Titus and James Reimer." figureDoi="http://doi.org/10.5281/zenodo.13745836" httpUri="https://zenodo.org/record/13745836/files/figure.png" pageId="16" pageNumber="17">Figure 6E</figureCitation>
), or speckled appearance (
<figureCitation id="13132A14645EFFD01F24FDDCFB3D7318" box="[1121,1254,583,609]" captionStart="FIGURE 3" captionStartId="12.[152,255,1779,1803]" captionTargetBox="[190,1386,1209,1756]" captionTargetId="figure-510@12.[189,1398,1209,1756]" captionTargetPageId="12" captionText="FIGURE 3. Confirmed geographic range of Entacmaea quadricolor in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry)." figureDoi="http://doi.org/10.5281/zenodo.13745830" httpUri="https://zenodo.org/record/13745830/files/figure.png" pageId="16" pageNumber="17">Figure S3B</figureCitation>
). The column of
<taxonomicName id="4C284D12645EFFD01BD2FDF7FF2673FC" baseAuthorityName="Hemprich &amp; Ehrenberg" baseAuthorityYear="1834" box="[151,253,620,645]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="16" pageNumber="17" phylum="Cnidaria" rank="species" species="crispa">R. crispa</taxonomicName>
is typically white/gray/tan in color and dotted with conspicuous raised verrucae which form longitudinal rows and are the same color as the column (
<figureCitation id="13132A14645EFFD0193AFD14FD2D73D0" box="[639,758,655,681]" captionStart="FIGURE 6" captionStartId="15.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@15.[226,1361,181,1728]" captionTargetPageId="15" captionText="FIGURE 6. Representative images of the leathery sea anemone Radianthus crispa encompassing a broad range of geographic and phenotypic variation. A) Yellow individual hosting juvenile Amphiprion chrysopterus on a typical fore reef environment in the Pacific Ocean. Note the densely packed tentacles with tapered tips coming to a point (Moorea, French Polynesia). B) White individual with pink tentacle tips (Moorea, French Polynesia). C) Tan/Brown individual with less densely packed tentacles that curl slightly (Saudi Arabia, Red Sea). D) Macro photograph underneath oral disc. i = well-defined raised verrucae that are similarly colored to the surrounding column (Okinawa, Japan). E) Small individual in stereotypical sandy microhabitat with slightly curled tentacled tips and horizontally striated tentacle pattern (Gulf of Oman, United Arab Emirates). Photographs by Benjamin M. Titus and James Reimer." figureDoi="http://doi.org/10.5281/zenodo.13745836" httpUri="https://zenodo.org/record/13745836/files/figure.png" pageId="16" pageNumber="17">Figure 6D</figureCitation>
). Verrucae are strongly adherent to the surrounding substrate and regularly hold debris, shells, and other sediment that can be seen when the oral disc is folded back.
</paragraph>
<paragraph id="8B973691645EFFD01B82FD4CFE237520" blockId="16.[151,1437,295,1114]" pageId="16" pageNumber="17">
A morphological and microhabitat shift seems to occur in this species between Indian and Pacific Ocean populations, and although this is not unilateral across all individuals, it is noticeable. Generally,
<taxonomicName id="4C284D12645EFFD01FAFFD67FA89726C" baseAuthorityName="Hemprich &amp; Ehrenberg" baseAuthorityYear="1834" box="[1258,1362,764,789]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="16" pageNumber="17" phylum="Cnidaria" rank="species" species="crispa">R. crispa</taxonomicName>
in the Indian Ocean occupy sand pocket microhabitats that are found adjacent to hard reef substrate and have the pedal disc and column burrowed deeply in the sand (
<figureCitation id="13132A14645EFFD01936FCD8FCD27227" box="[627,777,835,862]" captionStart="FIGURE 6" captionStartId="15.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@15.[226,1361,181,1728]" captionTargetPageId="15" captionText="FIGURE 6. Representative images of the leathery sea anemone Radianthus crispa encompassing a broad range of geographic and phenotypic variation. A) Yellow individual hosting juvenile Amphiprion chrysopterus on a typical fore reef environment in the Pacific Ocean. Note the densely packed tentacles with tapered tips coming to a point (Moorea, French Polynesia). B) White individual with pink tentacle tips (Moorea, French Polynesia). C) Tan/Brown individual with less densely packed tentacles that curl slightly (Saudi Arabia, Red Sea). D) Macro photograph underneath oral disc. i = well-defined raised verrucae that are similarly colored to the surrounding column (Okinawa, Japan). E) Small individual in stereotypical sandy microhabitat with slightly curled tentacled tips and horizontally striated tentacle pattern (Gulf of Oman, United Arab Emirates). Photographs by Benjamin M. Titus and James Reimer." figureDoi="http://doi.org/10.5281/zenodo.13745836" httpUri="https://zenodo.org/record/13745836/files/figure.png" pageId="16" pageNumber="17">Figure 6C, E</figureCitation>
;
<figureCitation id="13132A14645EFFD01853FCD8FC677224" box="[790,956,835,861]" captionStart="FIGURE 3" captionStartId="12.[152,255,1779,1803]" captionTargetBox="[190,1386,1209,1756]" captionTargetId="figure-510@12.[189,1398,1209,1756]" captionTargetPageId="12" captionText="FIGURE 3. Confirmed geographic range of Entacmaea quadricolor in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry)." figureDoi="http://doi.org/10.5281/zenodo.13745830" httpUri="https://zenodo.org/record/13745830/files/figure.png" pageId="16" pageNumber="17">Figure S3A, B</figureCitation>
). Indian Ocean specimens of
<taxonomicName id="4C284D12645EFFD01E49FCDFFAA97224" baseAuthorityName="Hemprich &amp; Ehrenberg" baseAuthorityYear="1834" box="[1292,1394,836,861]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="16" pageNumber="17" phylum="Cnidaria" rank="species" species="crispa">R. crispa</taxonomicName>
are typically found in calm patch reef or marginal reef habitats. In contrast, Pacific Ocean individuals more commonly occupy hard substrata directly in fore reef habitats and are more commonly found competing for space with stony corals (
<figureCitation id="13132A14645EFFD01BACFC34FE5A72B3" box="[233,385,943,970]" captionStart="FIGURE 6" captionStartId="15.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@15.[226,1361,181,1728]" captionTargetPageId="15" captionText="FIGURE 6. Representative images of the leathery sea anemone Radianthus crispa encompassing a broad range of geographic and phenotypic variation. A) Yellow individual hosting juvenile Amphiprion chrysopterus on a typical fore reef environment in the Pacific Ocean. Note the densely packed tentacles with tapered tips coming to a point (Moorea, French Polynesia). B) White individual with pink tentacle tips (Moorea, French Polynesia). C) Tan/Brown individual with less densely packed tentacles that curl slightly (Saudi Arabia, Red Sea). D) Macro photograph underneath oral disc. i = well-defined raised verrucae that are similarly colored to the surrounding column (Okinawa, Japan). E) Small individual in stereotypical sandy microhabitat with slightly curled tentacled tips and horizontally striated tentacle pattern (Gulf of Oman, United Arab Emirates). Photographs by Benjamin M. Titus and James Reimer." figureDoi="http://doi.org/10.5281/zenodo.13745836" httpUri="https://zenodo.org/record/13745836/files/figure.png" pageId="16" pageNumber="17">Figure 6A, B</figureCitation>
;
<figureCitation id="13132A14645EFFD01ACBFC34FDEE72B0" box="[398,565,943,969]" captionStart="FIGURE 3" captionStartId="12.[152,255,1779,1803]" captionTargetBox="[190,1386,1209,1756]" captionTargetId="figure-510@12.[189,1398,1209,1756]" captionTargetPageId="12" captionText="FIGURE 3. Confirmed geographic range of Entacmaea quadricolor in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry)." figureDoi="http://doi.org/10.5281/zenodo.13745830" httpUri="https://zenodo.org/record/13745830/files/figure.png" pageId="16" pageNumber="17">Figure S3C, D</figureCitation>
). Indian Ocean members of
<taxonomicName id="4C284D12645EFFD01831FC2BFC0172B0" baseAuthorityName="Hemprich &amp; Ehrenberg" baseAuthorityYear="1834" box="[884,986,944,969]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="16" pageNumber="17" phylum="Cnidaria" rank="species" species="crispa">R. crispa</taxonomicName>
have tentacles that are fewer in number and less densely packed than those from the Pacific, and are smaller in their body size and maximum oral disc diameter. Finally, Pacific Ocean members of
<taxonomicName id="4C284D12645EFFD019DDFC63FD247568" baseAuthorityName="Hemprich &amp; Ehrenberg" baseAuthorityYear="1834" box="[664,767,1016,1041]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="16" pageNumber="17" phylum="Cnidaria" rank="species" species="crispa">R. crispa</taxonomicName>
are more colorful in general, often have pink tentacle tips, and are more likely to take on body colors with pinks and yellows whereas Indian Ocean
<taxonomicName id="4C284D12645EFFD01F38FB87FB39754C" baseAuthorityName="Hemprich &amp; Ehrenberg" baseAuthorityYear="1834" box="[1149,1250,1052,1077]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="16" pageNumber="17" phylum="Cnidaria" rank="species" species="crispa">R. crispa</taxonomicName>
are typically tan or brown (
<figureCitation id="13132A14645EFFD01A49FBA4FEB67520" box="[268,365,1087,1113]" captionStart="FIGURE 6" captionStartId="15.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@15.[226,1361,181,1728]" captionTargetPageId="15" captionText="FIGURE 6. Representative images of the leathery sea anemone Radianthus crispa encompassing a broad range of geographic and phenotypic variation. A) Yellow individual hosting juvenile Amphiprion chrysopterus on a typical fore reef environment in the Pacific Ocean. Note the densely packed tentacles with tapered tips coming to a point (Moorea, French Polynesia). B) White individual with pink tentacle tips (Moorea, French Polynesia). C) Tan/Brown individual with less densely packed tentacles that curl slightly (Saudi Arabia, Red Sea). D) Macro photograph underneath oral disc. i = well-defined raised verrucae that are similarly colored to the surrounding column (Okinawa, Japan). E) Small individual in stereotypical sandy microhabitat with slightly curled tentacled tips and horizontally striated tentacle pattern (Gulf of Oman, United Arab Emirates). Photographs by Benjamin M. Titus and James Reimer." figureDoi="http://doi.org/10.5281/zenodo.13745836" httpUri="https://zenodo.org/record/13745836/files/figure.png" pageId="16" pageNumber="17">Figure 6</figureCitation>
;
<figureCitation id="13132A14645EFFD01A3FFBA4FE317520" box="[378,490,1087,1113]" captionStart="FIGURE 3" captionStartId="12.[152,255,1779,1803]" captionTargetBox="[190,1386,1209,1756]" captionTargetId="figure-510@12.[189,1398,1209,1756]" captionTargetPageId="12" captionText="FIGURE 3. Confirmed geographic range of Entacmaea quadricolor in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry)." figureDoi="http://doi.org/10.5281/zenodo.13745830" httpUri="https://zenodo.org/record/13745830/files/figure.png" pageId="16" pageNumber="17">Figure S3</figureCitation>
).
</paragraph>
<caption id="DF576619645EFFD01BDDF932FEEF7670" ID-DOI="http://doi.org/10.5281/zenodo.13745838" ID-Zenodo-Dep="13745838" httpUri="https://zenodo.org/record/13745838/files/figure.png" pageId="16" pageNumber="17" startId="16.[152,255,1705,1729]" targetBox="[189,1393,1137,1681]" targetPageId="16" targetType="figure">
<paragraph id="8B973691645EFFD01BDDF932FEEF7670" blockId="16.[151,1437,1705,1801]" pageId="16" pageNumber="17">
FIGURE 7. Confirmed geographic range of
<taxonomicName id="4C284D12645EFFD019CAF932FC9477B8" baseAuthorityName="Hemprich &amp; Ehrenberg" baseAuthorityYear="1834" box="[655,847,1705,1729]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="16" pageNumber="17" phylum="Cnidaria" rank="species" species="crispa">Radianthus crispa</taxonomicName>
in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry).
</paragraph>
</caption>
<caption id="DF576619645FFFD11BDDF885FC7E7693" ID-DOI="http://doi.org/10.5281/zenodo.13745840" ID-Zenodo-Dep="13745840" httpUri="https://zenodo.org/record/13745840/files/figure.png" pageId="17" pageNumber="18" startId="17.[152,255,1822,1846]" targetBox="[201,1387,181,1798]" targetPageId="17" targetType="figure">
<paragraph id="8B973691645FFFD11BDDF885FC7E7693" blockId="17.[151,1436,1822,2026]" pageId="17" pageNumber="18">
FIGURE 8. Representative images of the long-tentacled sea anemone
<taxonomicName id="4C284D12645FFFD1183DF885FB88764F" baseAuthorityName="Quoy &amp; Gaimard" baseAuthorityYear="1883" box="[888,1107,1822,1846]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="17" pageNumber="18" phylum="Cnidaria" rank="species" species="doreensis">Radianthus doreensis</taxonomicName>
encompassing a broad range of geographic and phenotypic variation. A) Whole individual in typical sandy microhabitat (Bali, Indonesia). B) Macro photograph of characteristic curling/spiraling tentacle pattern (Kwajalein Atoll, Marshall Islands). C) Partially retracted individual revealing highly contrasting white verrucae on brown column (arrow; Anilao, Philippines). D) Small purple individual with longitudinally striped tentacles and oral disc (Anilao, Philippines). E) Large individual with longitudinal “zebra-stripe” tentacle and oral disc pattern (Anilao, Philippines). All photographs by Scott and Jeanette Johnson.
</paragraph>
</caption>
<paragraph id="8B973691645CFFD21B82FF0CFC2470A8" blockId="18.[151,1437,151,465]" pageId="18" pageNumber="19">
Geographically,
<taxonomicName id="4C284D12645CFFD21AC7FF0CFE3271C9" baseAuthorityName="Hemprich &amp; Ehrenberg" baseAuthorityYear="1834" box="[386,489,151,176]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="18" pageNumber="19" phylum="Cnidaria" rank="species" species="crispa">R. crispa</taxonomicName>
is the most widespread host anemone species, ranging from the Northern Red Sea throughout the entire Indian Ocean, Coral Triangle, and extending East all the way to
<collectingCountry id="F33F7601645CFFD21F39FF20FA9B71AC" box="[1148,1344,187,213]" name="French Polynesia" pageId="18" pageNumber="19">French Polynesia</collectingCountry>
(
<figureCitation id="13132A14645CFFD21E14FF20FF7C7180" captionStart="FIGURE 7" captionStartId="16.[152,255,1705,1729]" captionTargetBox="[189,1393,1137,1681]" captionTargetId="figure-455@16.[189,1397,1137,1681]" captionTargetPageId="16" captionText="FIGURE 7. Confirmed geographic range of Radianthus crispa in the Indo-West Pacific. Red dots represent species observations from the Global Biodiversity Information Facility (GBIF). Blue shaded area represents shallow water habitat (60 m bathymetry)." figureDoi="http://doi.org/10.5281/zenodo.13745838" httpUri="https://zenodo.org/record/13745838/files/figure.png" pageId="18" pageNumber="19">Figure 7</figureCitation>
). Like
<taxonomicName id="4C284D12645CFFD21BBEFF44FE797180" baseAuthorityName="Leuckart in Ruppell &amp; Leuckart" baseAuthorityYear="1828" box="[251,418,223,249]" class="Anthozoa" family="Actiniidae" genus="Entacmaea" kingdom="Animalia" order="Actiniaria" pageId="18" pageNumber="19" phylum="Cnidaria" rank="species" species="quadricolor">E. quadricolor</taxonomicName>
,
<taxonomicName id="4C284D12645CFFD21AF7FF44FDC27181" baseAuthorityName="Hemprich &amp; Ehrenberg" baseAuthorityYear="1834" box="[434,537,223,248]" class="Anthozoa" family="Stichodactylidae" genus="Radianthus" kingdom="Animalia" order="Actiniaria" pageId="18" pageNumber="19" phylum="Cnidaria" rank="species" species="crispa">R. crispa</taxonomicName>
extends into high latitudes, nearly reaching temperate habitats in the Japanese Archipelago to the North, as well as marginal reef habitats in
<collectingCountry id="F33F7601645CFFD2187FFE98FC7A7064" box="[826,929,259,285]" name="Australia" pageId="18" pageNumber="19">Australia</collectingCountry>
(Solitary Islands) and
<collectingCountry id="F33F7601645CFFD21FDFFE98FAF07064" box="[1178,1323,259,285]" name="South Africa" pageId="18" pageNumber="19">South Africa</collectingCountry>
.Although this anemone forms associations with nearly as many clownfish species as
<taxonomicName id="4C284D12645CFFD218AFFEBCFB4B7038" baseAuthorityName="Leuckart in Ruppell &amp; Leuckart" baseAuthorityYear="1828" box="[1002,1168,295,321]" class="Anthozoa" family="Actiniidae" genus="Entacmaea" kingdom="Animalia" order="Actiniaria" pageId="18" pageNumber="19" phylum="Cnidaria" rank="species" species="quadricolor">E. quadricolor</taxonomicName>
, in our experience, it is found without clownfish symbionts more frequently than
<taxonomicName id="4C284D12645CFFD21852FED0FC67701C" baseAuthorityName="Leuckart in Ruppell &amp; Leuckart" baseAuthorityYear="1828" box="[791,956,331,357]" class="Anthozoa" family="Actiniidae" genus="Entacmaea" kingdom="Animalia" order="Actiniaria" pageId="18" pageNumber="19" phylum="Cnidaria" rank="species" species="quadricolor">E. quadricolor</taxonomicName>
. Solitary individuals that do not host fishes are common in the Red Sea and around the Arabian Peninsula. This species regularly hosts juvenile rather than adult clownfishes. It is not known to reproduce asexually and does not form extensive aggregations, but it is not uncommon to find this species adjacent to other clownfish-hosting anemones.
</paragraph>
</subSubSection>
</treatment>
</document>