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<mods:title id="5D1913F0B1C5AB8BC7B2CEAD173676A2">The clownfish-hosting sea anemones (Anthozoa: Actiniaria): updated nomenclature, biogeography, and practical field guide.</mods:title>
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<paragraph id="8B9736916444FFCA1BD2F8B1FB1B763A" blockId="10.[151,1216,1833,1860]" box="[151,1216,1833,1860]" pageId="10" pageNumber="11">
<heading id="D0DF81FD6444FFCA1BD2F8B1FB1B763A" bold="true" box="[151,1216,1833,1860]" fontSize="11" level="1" pageId="10" pageNumber="11" reason="1">
<taxonomicName id="4C284D126444FFCA1BD2F8B1FE6E763D" ID-CoL="39X4C" authority="(Leuckart in Ruppell &amp; Leuckart, 1828)" baseAuthorityName="Leuckart in Ruppell &amp; Leuckart" baseAuthorityYear="1828" box="[151,437,1833,1860]" class="Anthozoa" family="Actiniidae" genus="Entacmaea" kingdom="Animalia" order="Actiniaria" pageId="10" pageNumber="11" phylum="Cnidaria" rank="species" species="quadricolor">Entacmaea quadricolor</taxonomicName>
(Leuckart in
<bibRefCitation id="EFB94B606444FFCA1924F8B1FC71763A" author="Ruppell, E. &amp; Leuckart, F. S." box="[609,938,1833,1860]" pageId="10" pageNumber="11" refId="ref17510" refString="Ruppell, E. &amp; Leuckart, F. S. (1828) Atlas zu der Reise im Nordlichen Afrika von Eduard Ru ¨ ppell, Neue Wirbellose Thiere des Rothen Meers. Heinr. Ludw. Brvnner, Frankfurt am Main, 47 pp." type="book" year="1828">Ruppell &amp; Leuckart, 1828</bibRefCitation>
) (
<figureCitation id="13132A146444FFCA1887F8B2FBF5763D" box="[962,1070,1833,1860]" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="10" pageNumber="11">Figure 2</figureCitation>
;
<figureCitation id="13132A146444FFCA1F78F8B2FB6C763A" box="[1085,1207,1833,1859]" captionStart="FIGURE 1" captionStartId="9.[152,255,733,757]" captionTargetBox="[304,1282,184,706]" captionTargetId="figure-17@9.[300,1287,181,709]" captionTargetPageId="9" captionText="FIGURE 1. Labeled illustration of Mertens carpet anemone Stichodactyla mertensii." figureDoi="http://doi.org/10.5281/zenodo.13745826" httpUri="https://zenodo.org/record/13745826/files/figure.png" pageId="10" pageNumber="11">Figure S1</figureCitation>
)
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<paragraph id="8B9736916444FFCC1BD2F8E9FED47360" blockId="10.[151,1436,1906,2040]" lastBlockId="12.[151,1437,151,1185]" lastPageId="12" lastPageNumber="13" pageId="10" pageNumber="11">
The bubble-tip sea anemone
<taxonomicName id="4C284D126444FFCA1A95F8E8FD0376F5" baseAuthorityName="Leuckart in Ruppell &amp; Leuckart" baseAuthorityYear="1828" box="[464,728,1906,1932]" class="Anthozoa" family="Actiniidae" genus="Entacmaea" kingdom="Animalia" order="Actiniaria" pageId="10" pageNumber="11" phylum="Cnidaria" rank="species" species="quadricolor">Entacmaea quadricolor</taxonomicName>
is among the most iconic and phenotypically variable clownfish-hosting sea anemone species in terms of color, pattern, tentacle morphology, and growth patterns (
<figureCitation id="13132A146444FFCA1FA1F80DFA9D76C9" box="[1252,1350,1942,1968]" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="10" pageNumber="11">Figure 2</figureCitation>
;
<figureCitation id="13132A146444FFCA1E11F80DFF6D76AD" captionStart="FIGURE 1" captionStartId="9.[152,255,733,757]" captionTargetBox="[304,1282,184,706]" captionTargetId="figure-17@9.[300,1287,181,709]" captionTargetPageId="9" captionText="FIGURE 1. Labeled illustration of Mertens carpet anemone Stichodactyla mertensii." figureDoi="http://doi.org/10.5281/zenodo.13745826" httpUri="https://zenodo.org/record/13745826/files/figure.png" pageId="10" pageNumber="11">Figure S1</figureCitation>
). This species gets its common name from the characteristic bulbous swellings that commonly form at, or near, the tentacle tips. When bulbs are present, this species is easy to identify underwater. However, tentacle shape is highly variable both intra-specifically and even intra-individually. It is not uncommon to encounter individuals whose tentacles are simultaneously bulbed and digitiform (smooth, not-bulbed, uniformly shaped;
<figureCitation id="13132A146442FFCC1FD2FF20FAD671AC" box="[1175,1293,187,213]" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="12" pageNumber="13">Figure 2A</figureCitation>
), completely bulbous (
<figureCitation id="13132A146442FFCC1BB8FF44FE697180" box="[253,434,223,249]" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="12" pageNumber="13">Figure 2B, C, D</figureCitation>
), or completely digitiform and long (up to
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;
<figureCitation id="13132A146442FFCC18B1FF44FBBD7180" box="[1012,1126,223,249]" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="12" pageNumber="13">Figure 2E</figureCitation>
). Tentacle tips, as a rule, are blunt ended. Tentacle color is typically brown/tan or green (
<figureCitation id="13132A146442FFCC1861FE98FC5A7064" box="[804,897,259,285]" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="12" pageNumber="13">Figure 2</figureCitation>
A-D) but can also be bright red or orange (
<figureCitation id="13132A146442FFCC1E17FE98FF617038" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="12" pageNumber="13">Figure 2E</figureCitation>
). Tentacle tips are frequently purple/magenta. Tentacle patterns are variable. Some tentacles take on a dull matte appearance (
<figureCitation id="13132A146442FFCC1A60FED0FE44701C" box="[293,415,331,357]" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="12" pageNumber="13">Figure 2A</figureCitation>
;
<figureCitation id="13132A146442FFCC1AE9FED0FDE9701C" box="[428,562,331,357]" captionStart="FIGURE 1" captionStartId="9.[152,255,733,757]" captionTargetBox="[304,1282,184,706]" captionTargetId="figure-17@9.[300,1287,181,709]" captionTargetPageId="9" captionText="FIGURE 1. Labeled illustration of Mertens carpet anemone Stichodactyla mertensii." figureDoi="http://doi.org/10.5281/zenodo.13745826" httpUri="https://zenodo.org/record/13745826/files/figure.png" pageId="12" pageNumber="13">Figure S1D</figureCitation>
), but can also be striated (
<figureCitation id="13132A146442FFCC1824FED0FC03701C" box="[865,984,331,357]" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="12" pageNumber="13">Figure 2C</figureCitation>
), translucent (
<figureCitation id="13132A146442FFCC1F3CFED0FAF8701C" box="[1145,1315,331,357]" captionStart="FIGURE 1" captionStartId="9.[152,255,733,757]" captionTargetBox="[304,1282,184,706]" captionTargetId="figure-17@9.[300,1287,181,709]" captionTargetPageId="9" captionText="FIGURE 1. Labeled illustration of Mertens carpet anemone Stichodactyla mertensii." figureDoi="http://doi.org/10.5281/zenodo.13745826" httpUri="https://zenodo.org/record/13745826/files/figure.png" pageId="12" pageNumber="13">Figure S1A, C</figureCitation>
), speckled (
<figureCitation id="13132A146442FFCC1BDAFEF4FE9870F0" box="[159,323,367,393]" captionStart="FIGURE 1" captionStartId="9.[152,255,733,757]" captionTargetBox="[304,1282,184,706]" captionTargetId="figure-17@9.[300,1287,181,709]" captionTargetPageId="9" captionText="FIGURE 1. Labeled illustration of Mertens carpet anemone Stichodactyla mertensii." figureDoi="http://doi.org/10.5281/zenodo.13745826" httpUri="https://zenodo.org/record/13745826/files/figure.png" pageId="12" pageNumber="13">Figure S1A, C</figureCitation>
), or some combination thereof. In many individuals the equator of the bulb is a mottled white pattern (
<figureCitation id="13132A146442FFCC1BDAFE08FEDB70D4" box="[159,256,403,429]" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="12" pageNumber="13">Figure 2</figureCitation>
;
<figureCitation id="13132A146442FFCC1A4EFE08FEA370D4" box="[267,376,403,429]" captionStart="FIGURE 1" captionStartId="9.[152,255,733,757]" captionTargetBox="[304,1282,184,706]" captionTargetId="figure-17@9.[300,1287,181,709]" captionTargetPageId="9" captionText="FIGURE 1. Labeled illustration of Mertens carpet anemone Stichodactyla mertensii." figureDoi="http://doi.org/10.5281/zenodo.13745826" httpUri="https://zenodo.org/record/13745826/files/figure.png" pageId="12" pageNumber="13">Figure S1</figureCitation>
). Tentacles are densely packed, typically obscuring the oral disc and mouth. Verrucae are absent on the column
<taxonomicName id="4C284D126442FFCC1A7AFE2CFE3E70A8" baseAuthorityName="Leuckart in Ruppell &amp; Leuckart" baseAuthorityYear="1828" box="[319,485,439,465]" class="Anthozoa" family="Actiniidae" genus="Entacmaea" kingdom="Animalia" order="Actiniaria" pageId="12" pageNumber="13" phylum="Cnidaria" rank="species" species="quadricolor">E. quadricolor</taxonomicName>
(
<figureCitation id="13132A146442FFCC1AB6FE2CFDA370A8" box="[499,632,439,465]" captionStart="FIGURE 1" captionStartId="9.[152,255,733,757]" captionTargetBox="[304,1282,184,706]" captionTargetId="figure-17@9.[300,1287,181,709]" captionTargetPageId="9" captionText="FIGURE 1. Labeled illustration of Mertens carpet anemone Stichodactyla mertensii." figureDoi="http://doi.org/10.5281/zenodo.13745826" httpUri="https://zenodo.org/record/13745826/files/figure.png" pageId="12" pageNumber="13">Figure S1C</figureCitation>
). Column color is usually bright red or magenta. The column and pedal disc are not typically visible and burrowed deep in a crevice or hole in the reef. The body wall is thin and tentacles tear easily.
</paragraph>
</subSubSection>
<caption id="DF5766196445FFCB1BDDF943FAB47695" ID-DOI="http://doi.org/10.5281/zenodo.13745828" ID-Zenodo-Dep="13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="11" pageNumber="12" startId="11.[152,255,1752,1776]" targetBox="[226,1361,181,1728]" targetPageId="11" targetType="figure">
<paragraph id="8B9736916445FFCB1BDDF943FAB47695" blockId="11.[151,1437,1752,2029]" pageId="11" pageNumber="12">
FIGURE 2. Representative images of the bubble-tip sea anemone
<taxonomicName id="4C284D126445FFCB181CF942FB967789" baseAuthorityName="Leuckart in Ruppell &amp; Leuckart" baseAuthorityYear="1828" box="[857,1101,1752,1776]" class="Anthozoa" family="Actiniidae" genus="Entacmaea" kingdom="Animalia" order="Actiniaria" pageId="11" pageNumber="12" phylum="Cnidaria" rank="species" species="quadricolor">Entacmaea quadricolor</taxonomicName>
encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of
<taxonomicName id="4C284D126445FFCB1A21F8F3FD8976F9" authorityName="Bleeker" authorityYear="1852" box="[356,594,1896,1920]" family="Pomacentridae" genus="Amphiprion" kingdom="Animalia" order="Perciformes" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="melanopus">Amphiprion melanopus</taxonomicName>
(Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson.
</paragraph>
</caption>
<subSubSection id="C332651A6442FFCC1B82FDB8FB9475D8" pageId="12" pageNumber="13" type="description">
<paragraph id="8B9736916442FFCC1B82FDB8FBF172DC" blockId="12.[151,1437,151,1185]" pageId="12" pageNumber="13">
<taxonomicName id="4C284D126442FFCC1B82FDB8FE087344" baseAuthorityName="Leuckart in Ruppell &amp; Leuckart" baseAuthorityYear="1828" box="[199,467,547,573]" class="Anthozoa" family="Actiniidae" genus="Entacmaea" kingdom="Animalia" order="Actiniaria" pageId="12" pageNumber="13" phylum="Cnidaria" rank="species" species="quadricolor">Entacmaea quadricolor</taxonomicName>
exhibits two primary growth patterns and can be found as either large (up to
<quantity id="4CD09B746442FFCC1E7AFDB8FA467344" box="[1343,1437,547,573]" metricMagnitude="-1" metricUnit="m" metricValue="4.0" pageId="12" pageNumber="13" unit="mm" value="400.0">400 mm</quantity>
in oral disc diameter) solitary individuals (
<figureCitation id="13132A146442FFCC19C1FDDCFD267318" box="[644,765,583,609]" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="12" pageNumber="13">Figure 2A</figureCitation>
) or clonal aggregations of smaller individuals. Depending on habitat and geography, clonal aggregations can be comprised of small clusters of medium-sized individuals (typically 2-6 anemones per cluster), or dense aggregations of small individuals that can form extensive fields of anemones (dozens to hundreds of individuals;
<figureCitation id="13132A146442FFCC19ECFD28FCC573B4" box="[681,798,691,717]" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="12" pageNumber="13">Figure 2B</figureCitation>
). This latter growth form is typically found in the Coral Triangle to Central Pacific Ocean in shallow habitats where anemones grow on tops of reefs or among the branches of shallow stony corals (
<figureCitation id="13132A146442FFCC1AE9FD60FDFF726C" box="[428,548,763,789]" captionStart="FIGURE 2" captionStartId="11.[152,255,1752,1776]" captionTargetBox="[226,1361,181,1728]" captionTargetId="figure-17@11.[226,1361,181,1728]" captionTargetPageId="11" captionText="FIGURE 2. Representative images of the bubble-tip sea anemone Entacmaea quadricolor encompassing a broad range of geographic and phenotypic variation. A) Wide angle photograph of large solitary individual highlighting typical microhabitat requirements for the species. Note the tentacles are both bulbous and digitiform within this individual (Saudi Arabia, Red Sea). B) Dense aggregation of hundreds of small clonal sea anemones growing among a shallow branching stony coral serving as hosts to a colony of Amphiprion melanopus (Kwajalein Atoll, Marshall Islands). C) Macro photograph of characteristic bubble tips. Note the striated pattern of the tentacle (Fares-Maathodaa, Maldives). D) Individual anemone with all tentacles exhibiting characteristic bulbous tentacle morphology (Fares-Maathodaa, Maldives). E) Red solitary individual with long digitiform tentacles hosting (Saudi Arabia, Red Sea). Photographs by Morgan Bennett-Smith, Benjamin M. Titus, and Scott Johnson." figureDoi="http://doi.org/10.5281/zenodo.13745828" httpUri="https://zenodo.org/record/13745828/files/figure.png" pageId="12" pageNumber="13">Figure 2A</figureCitation>
). Both shallow aggregations and large solitary individuals can be found on the same reefs and typically segregate by depth, with the large solitary individuals occurring in deeper water. Regardless of growth form,
<taxonomicName id="4C284D126442FFCC1A12FCD8FE257224" baseAuthorityName="Leuckart in Ruppell &amp; Leuckart" baseAuthorityYear="1828" box="[343,510,835,861]" class="Anthozoa" family="Actiniidae" genus="Entacmaea" kingdom="Animalia" order="Actiniaria" pageId="12" pageNumber="13" phylum="Cnidaria" rank="species" species="quadricolor">E. quadricolor</taxonomicName>
requires hard stable substrate and is never found with its pedal disc and column burrowed in the sand. This species generally requires calm habitats with low wave exposure. When disturbed,
<taxonomicName id="4C284D126442FFCC1EC1FCFCFEC672DC" baseAuthorityName="Leuckart in Ruppell &amp; Leuckart" baseAuthorityYear="1828" class="Anthozoa" family="Actiniidae" genus="Entacmaea" kingdom="Animalia" order="Actiniaria" pageId="12" pageNumber="13" phylum="Cnidaria" rank="species" species="quadricolor">E. quadricolor</taxonomicName>
can disappear fully from view by withdrawing into the reef structure.
</paragraph>
<paragraph id="8B9736916442FFCC1B82FC34FB9475D8" blockId="12.[151,1437,151,1185]" pageId="12" pageNumber="13">
The geographic range of this species is broad, ranging from the very Northern Red Sea, throughout the Indian Ocean, Coral Triangle, and into the Central Pacific reaching the
<collectingCountry id="F33F76016442FFCC1829FC48FBFC7294" box="[876,1063,979,1005]" name="Marshall Islands" pageId="12" pageNumber="13">Marshall Islands</collectingCountry>
but not East to
<collectingCountry id="F33F76016442FFCC1F9FFC48FA477294" box="[1242,1436,979,1005]" name="French Polynesia" pageId="12" pageNumber="13">French Polynesia</collectingCountry>
(
<figureCitation id="13132A146442FFCC1BDAFC6CFED87568" box="[159,259,1015,1041]" 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="12" pageNumber="13">Figure 3</figureCitation>
). High-latitude populations of this species are also common, nearly reaching temperate habitats in the Japanese Archipelago to the North, as well as marginal reef habitats in
<collectingCountry id="F33F76016442FFCC18E3FB80FBD6754C" box="[934,1037,1051,1077]" name="Australia" pageId="12" pageNumber="13">Australia</collectingCountry>
(Solitary Islands) and
<collectingCountry id="F33F76016442FFCC1E4DFB80FA42754C" box="[1288,1433,1051,1077]" name="South Africa" pageId="12" pageNumber="13">South Africa</collectingCountry>
. This is among the most common host anemones encountered throughout its range and almost always is found in association with clownfishes. However, in marginal reef habitats in high latitudes it is not uncommon for this species to be found without fish symbionts, particularly the small clonal populations.
</paragraph>
</subSubSection>
</treatment>
</document>