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<document id="AEA1C86C599EE5884850D7176B6E1A31" ID-CLB-Dataset="2385" ID-DOI="10.11646/zootaxa.5093.3.4" ID-GBIF-Dataset="e7f4500d-24b9-4dfb-9967-46d032dad854" ID-ISSN="1175-5326" ID-Zenodo-Dep="5909954" ID-ZooBank="58B73298-6A33-4040-B898-ED3D8A69DA81" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1643276591335" checkinUser="plazi" docAuthor="Watling, Les, Saucier, Esprit Heestand &amp; France, Scott C." docDate="2022" docId="8E2A87918464FFE35CE0F9D5FD6EF8D9" docLanguage="en" docName="zootaxa.5093.3.4.pdf" docOrigin="Zootaxa 5093 (3)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Keratoisididae Gray 1870" docType="treatment" docVersion="7" lastPageNumber="349" masterDocId="7213FFE9846EFFEF5C77FF8FFF8EFF94" masterDocTitle="Towards a revision of the bamboo corals (Octocorallia): Part 4, delineating the family Keratoisididae" masterLastPageNumber="375" masterPageNumber="337" pageNumber="347" updateTime="1699304541045" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
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<mods:title id="0271C4A9ABB94AE67339348F891AE430">Towards a revision of the bamboo corals (Octocorallia): Part 4, delineating the family Keratoisididae</mods:title>
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<mods:namePart id="01006670D24477758856AA3DD60B12B1">Watling, Les</mods:namePart>
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<mods:namePart id="E5D486C91DD1335A02AD820B474EFB52">Saucier, Esprit Heestand</mods:namePart>
<mods:affiliation id="BA3B59EBB6D2AE934E87E25F1AC76D35">Faculty of Science, Brigham Young University - Hawaii, 55 - 220 Kulanui St, Laie, Hawaii, 96762, USA</mods:affiliation>
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<mods:namePart id="9AEBC7BBBB9A8A1A0AEECAFBB7050CA3">France, Scott C.</mods:namePart>
<mods:affiliation id="967378AB4DA8135FBE051A6DA2A0BE05">Department of Biology, University of Louisiana at Lafayette, Lafayette, Louisiana, USA</mods:affiliation>
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<paragraph id="063C36878464FFE55CE0F9D5FD0AF9E0" blockId="10.[151,644,1626,1652]" box="[151,644,1626,1652]" pageId="10" pageNumber="347">
<heading id="5D7481EB8464FFE55CE0F9D5FD0AF9E0" bold="true" box="[151,644,1626,1652]" fontSize="11" level="1" pageId="10" pageNumber="347" reason="1">
<emphasis id="34F7EA958464FFE55CE0F9D5FD0AF9E0" bold="true" box="[151,644,1626,1652]" pageId="10" pageNumber="347">
Details regarding
<taxonomicName id="C1834D048464FFE55D0FF9D5FDBEF9E0" ID-CoL="8QC4N" authorityName="Gray" authorityYear="1870" box="[376,560,1626,1652]" class="Anthozoa" family="Keratoisididae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Alcyonacea" pageId="10" pageNumber="347" phylum="Cnidaria" rank="family">Keratoisididae</taxonomicName>
clades
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="4E99650C8464FFE35CE0F92DFD6EF8D9" lastPageId="12" lastPageNumber="349" pageId="10" pageNumber="347" type="description">
<paragraph id="063C36878464FFE45CE0F92DFED7FE89" blockId="10.[151,1437,1698,2013]" lastBlockId="11.[151,1437,151,2013]" lastPageId="11" lastPageNumber="348" pageId="10" pageNumber="347">
Clade
<emphasis id="34F7EA958464FFE55C97F92DFF60F928" bold="true" box="[224,238,1698,1724]" pageId="10" pageNumber="347">3</emphasis>
(
<emphasis id="34F7EA958464FFE55C88F92DFE93F928" bold="true" box="[255,285,1698,1724]" pageId="10" pageNumber="347">S1</emphasis>
) encompasses sequences derived from the
<typeStatus id="D93888258464FFE55F73F92DFCE8F928" box="[772,870,1698,1724]" pageId="10" pageNumber="347" type="holotype">holotype</typeStatus>
specimen of
<taxonomicName id="C1834D048464FFE55F89F92DFA1BF929" authority="Watling, 2015" authorityName="Watling" authorityYear="2015" box="[1022,1429,1698,1725]" class="Anthozoa" family="Isididae" genus="Cladarisis" kingdom="Animalia" order="Alcyonacea" pageId="10" pageNumber="347" phylum="Cnidaria" rank="species" species="nouvianae">
<emphasis id="34F7EA958464FFE55F89F92DFB7EF928" box="[1022,1264,1698,1724]" italics="true" pageId="10" pageNumber="347">Cladarisis nouvianae</emphasis>
Watling, 2015
</taxonomicName>
, two undescribed taxa, and a specimen (USNM 75083) identified as
<taxonomicName id="C1834D048464FFE55FE2F949FB7CF974" authority="(Esper, 1788)" baseAuthorityName="Esper" baseAuthorityYear="1788" box="[917,1266,1734,1760]" class="Anthozoa" family="Isididae" genus="Isidella" kingdom="Animalia" order="Alcyonacea" pageId="10" pageNumber="347" phylum="Cnidaria" rank="species" species="elongata">
<emphasis id="34F7EA958464FFE55FE2F949FBDDF974" box="[917,1107,1734,1760]" italics="true" pageId="10" pageNumber="347">Isidella elongata</emphasis>
(Esper, 1788)
</taxonomicName>
by F.M. Bayer and collected from near the type locality in the Mediterranean Sea. Our sequence for that specimen is restricted to the mtMutS-5 and a short 18S region. We were able to obtain a partial sequence (654bp for
<emphasis id="34F7EA958464FFE558B1F880FAA8F8BD" box="[1222,1318,1807,1833]" italics="true" pageId="10" pageNumber="347">mtMutS-</emphasis>
5: matching positions
<quantity id="C17B9B628464FFE55D47F8BDFE2DF8D8" box="[304,419,1842,1868]" metricMagnitude="1" metricUnit="m" metricValue="1.01727" metricValueMax="1.8465799999999999" metricValueMin="0.18796" pageId="10" pageNumber="347" unit="in" value="400.5" valueMax="727.0" valueMin="74.0">74727 in</quantity>
the GenBank sequence
<accessionNumber id="19D0AB648464FFE55EC1F8BDFCBDF8D9" box="[694,819,1842,1869]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/KC984577" pageId="10" pageNumber="347">KC984577</accessionNumber>
) from the
<typeStatus id="D93888258464FFE55FDCF8BDFB83F8D8" box="[939,1037,1842,1868]" pageId="10" pageNumber="347" type="holotype">holotype</typeStatus>
specimen of
<taxonomicName id="C1834D048464FFE558D1F8BCFAB4F8D8" authorityName="Bayer" authorityYear="1990" box="[1190,1338,1842,1868]" class="Anthozoa" family="Isididae" genus="Isidella" kingdom="Animalia" order="Alcyonacea" pageId="10" pageNumber="347" phylum="Cnidaria" rank="species" species="trichotoma">
<emphasis id="34F7EA958464FFE558D1F8BCFAB4F8D8" box="[1190,1338,1842,1868]" italics="true" pageId="10" pageNumber="347">I. trichotoma</emphasis>
</taxonomicName>
(USNM 56715) and found it to uniquely match a portion of the haplotype of USNM 75083; however, we have been unable to replicate these results as subsequent PCR amplification attempts of the
<typeStatus id="D93888258464FFE55F8FF8F5FBD4F800" box="[1016,1114,1914,1940]" pageId="10" pageNumber="347" type="holotype">holotype</typeStatus>
specimen failed to produce an amplicon. Divergence among the 4 haplotypes of clade
<emphasis id="34F7EA958464FFE55F48F811FCC3F82C" bold="true" box="[831,845,1950,1976]" pageId="10" pageNumber="347">3</emphasis>
is greater than within clade
<emphasis id="34F7EA958464FFE558EDF811FB26F82C" bold="true" box="[1178,1192,1950,1976]" pageId="10" pageNumber="347">2</emphasis>
, with intra-clade p =
<geoCoordinate id="63B750408464FFE55CE0F84DFF6AF849" box="[151,228,1986,2013]" degrees="0.0005" direction="north" orientation="latitude" pageId="10" pageNumber="347" precision="5" value="5.0E-4">0.0005</geoCoordinate>
<geoCoordinate id="63B750408464FFE55C93F84DFEB1F849" box="[228,319,1986,2013]" degrees="0.0033" direction="west" orientation="longitude" pageId="10" pageNumber="347" precision="5" value="-0.0033">0.0033</geoCoordinate>
for the
<emphasis id="34F7EA958464FFE55DEDF84CFE74F849" box="[410,506,1987,2013]" italics="true" pageId="10" pageNumber="347">mtMutS-</emphasis>
5 region. Morphological characters of this clade include (
<tableCitation id="4B01033C8464FFE558FCF84DFB6AF848" box="[1163,1252,1986,2012]" captionStart="TABLE 4" captionStartId="17.[151,239,484,510]" captionTargetPageId="17" captionText="TABLE 4. Summary of morphological characters associated with each molecular subclade or indel type." httpUri="http://table.plazi.org/id/52FC660F847FFFFE5CE0FE6BFB78FE6A" pageId="10" pageNumber="347" tableUuid="52FC660F847FFFFE5CE0FE6BFB78FE6A">Table 4</tableCitation>
;
<figureCitation id="9EB82A028464FFE55886F84DFABBF848" box="[1265,1333,1986,2012]" captionStart="FIGURE 2" captionStartId="21.[151,250,1645,1669]" captionTargetBox="[280,1308,181,1621]" captionTargetId="figure-19@21.[280,1308,181,1621]" captionTargetPageId="21" captionText="FIGURE 2. A selection of colony types from the Family Keratoisididae. Clade names as noted in the text, followed by species name (if known), specimen number, approximate colony size, and location. S1, Cladarisis nouvianae, CAT 207-1, 30 cm, Cat Island, Bahamas; M1, Orstomisis sp., NMHN IK-2008-14, 45 cm across branched area, New Caledonia; I1, Lepidisis caryophyllia, GIL 101-1, 17 cm, Gilbert Canyon, northeastern USA; A1, Acanella arbuscula, MAN 803-1, 30 cm height, Manning Seamount, New England Seamounts; J3-1, KEL 604-2, 30 cm, Kelvin Seamounts, New England Seamounts; J3-2, KUK 218-1, approx. 75 cm width of fan, Kükenthal Peak, Caloosahatchee Seamount, Corner Rise Seamounts; J3-3, NAS 101-3, 30 cm field of view, individual colony approx. 50-75 cm, Nashville Seamount, New England Seamounts; I4, BAL 205-1, 50 cm height, Balanus Seamount, New England Seamounts; F1, ANN 209-1, longest branch approx. 40 cm, Annei Seamount, Emperor Seamounts; C1, BAL 211-3, approx. 2 m length, Balanus Seamount; D1, GOO 101-1, approx. 1 m. length, Goode Peak, Caloosahatchee Seamount; D2, Eknomisis dalioi, KEL 406-1, colony approx. 1 m tall by 1.5 m wide, Kelvin Seamount; B1, KEL 408-1, approx. 2 to 3 m length, Kelvin Seamount." figureDoi="http://doi.org/10.5281/zenodo.5909958" httpUri="https://zenodo.org/record/5909958/files/figure.png" pageId="10" pageNumber="347">Fig. 2</figureCitation>
, S
<figureCitation id="9EB82A028464FFE55921F84DFAECF848" box="[1366,1378,1986,2012]" captionStart="FIGURE 1" captionStartId="20.[151,250,1882,1906]" captionTargetBox="[367,1212,181,1844]" captionTargetId="figure-21@20.[363,1225,179,1844]" captionTargetPageId="20" captionText="FIGURE 1. RAxML phylogenetic tree based on a concatenated alignment of mtMutS-5, mtMutS-3 and 18S molecular sequences of 92 total taxa (86 keratoisids + 6 out group). Bootstrap values&gt;50 are shown above the nodes. Bayesian posterior probability values&gt;0.5 are shown below the node. Numbers 1 to 5 refer to nodes discussed in the text. Clades delimited by vertical bars on the right side are named following the convention of France (2007) and are discussed in detail in the text." figureDoi="http://doi.org/10.5281/zenodo.5909956" httpUri="https://zenodo.org/record/5909956/files/figure.png" pageId="10" pageNumber="347">1</figureCitation>
;
<figureCitation id="9EB82A028464FFE45907F84DFF28FF25" captionStart="FIGURE 3" captionStartId="22.[151,250,1637,1661]" captionTargetBox="[346,1241,181,1614]" captionTargetId="figure-21@22.[346,1241,181,1614]" captionTargetPageId="22" captionText="FIGURE 3. A selection of polyp types from the Family Keratoisididae. Clade names as noted in the text, followed by species name (if known), specimen number, special treatment (if any), and location. All polyps are less than 1 cm in length or height and are in contracted state due to preservation. S1, Cladarisis nouvianae, CAT 207-1, Cat Island, Bahamas; M1, Orstomisis sp., NMHN IK-2008-14, polyp cleared with clove oil, New Caledonia; I1, Lepidisis caryophyllia, GIL 101-1, SEM view with external tissue removed, Gilbert Canyon, northeast USA; A1, Acanella arbuscula, KUK 103-2, Kükenthal Peak, Caloosahatchee Seamount, Corner Rise Seamounts; J3-1, KEL 604-2, Kelvin Seamount, New England Seamounts; J3-2, KUK 218-1, Kükenthal Peak; I4, BAL 205-1, polyp cleared with clove oil, Balanus Seamount, New England seamounts; F1, KEL 104-1, Kelvin Seamount; H1, Keratoisis magnifica, TMAG K3830, Tasmania Seamounts; C1, BAL 111-1, Balanus Seamount; G1, TMAG K3856, Tasmanian Seamounts; D1, MAN 707-2, Manning Seamount, New England Seamounts; D2, Eknomisis dalioi, KEL 406-1, Kelvin Seamount; B1-1, PIC 102-1, Picket Seamount, New England Seamounts; B1-2, J2098-6-4, Aleutian Islands slope, Alaska, USA." figureDoi="http://doi.org/10.5281/zenodo.5909960" httpUri="https://zenodo.org/record/5909960/files/figure.png" lastPageId="11" lastPageNumber="348" pageId="10" pageNumber="347">Fig. 3</figureCitation>
, S
<figureCitation id="9EB82A028465FFE45CBFFF18FF5AFF25" box="[200,212,151,177]" captionStart="FIGURE 1" captionStartId="20.[151,250,1882,1906]" captionTargetBox="[367,1212,181,1844]" captionTargetId="figure-21@20.[363,1225,179,1844]" captionTargetPageId="20" captionText="FIGURE 1. RAxML phylogenetic tree based on a concatenated alignment of mtMutS-5, mtMutS-3 and 18S molecular sequences of 92 total taxa (86 keratoisids + 6 out group). Bootstrap values&gt;50 are shown above the nodes. Bayesian posterior probability values&gt;0.5 are shown below the node. Numbers 1 to 5 refer to nodes discussed in the text. Clades delimited by vertical bars on the right side are named following the convention of France (2007) and are discussed in detail in the text." figureDoi="http://doi.org/10.5281/zenodo.5909956" httpUri="https://zenodo.org/record/5909956/files/figure.png" pageId="11" pageNumber="348">1</figureCitation>
): a solid, relatively narrow-diameter axis subdivided by organic nodes; branches are sparse and originate at nodes that are moderately flexible; polyp sclerites are all rods, often thick and curved, and arranged in an irregular fashion over the surface of the polyp; on contraction, the polyp shortens and the tentacles fold over or into the apical part of the polyp.
</paragraph>
<paragraph id="063C36878465FFE45CB0FEA8FBFAFDA9" blockId="11.[151,1437,151,2013]" pageId="11" pageNumber="348">
Clade
<emphasis id="34F7EA958465FFE45D78FEA9FE93FED4" bold="true" box="[271,285,294,320]" pageId="11" pageNumber="348">4</emphasis>
contains most of the diversity in the tree and most closely matches the former
<taxonomicName id="C1834D048465FFE458F0FEA8FB6EFED5" box="[1159,1248,295,321]" class="Anthozoa" family="Isididae" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="family">Isididae</taxonomicName>
subfamily
<taxonomicName id="C1834D048465FFE4592BFEA8FE89FEF1" authorityName="Gray" authorityYear="1870" class="Anthozoa" family="Isididae" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="subFamily" subFamily="Keratoisidinae">Keratoisidinae</taxonomicName>
, which has now been elevated to the rank of family (
<bibRefCitation id="62124B768465FFE45F1AFEC4FB33FEF1" author="Heestand Saucier, E. &amp; France, S. C. &amp; Watling, L." box="[877,1213,331,357]" pageId="11" pageNumber="348" pagination="247 - 272" refId="ref14195" refString="Heestand Saucier, E., France, S. C. &amp; Watling, L. (2021) Toward a revision of the bamboo corals: Part 3, deconstructing the Family Isididae. Zootaxa, 5047 (3), 247 - 272. https: // doi. org / 10.11646 / zootaxa. 5047.3.2" type="journal article" year="2021">
Heestand Saucier
<emphasis id="34F7EA958465FFE45849FEC4FBF6FEF1" box="[1086,1144,331,357]" italics="true" pageId="11" pageNumber="348">et al.</emphasis>
2021
</bibRefCitation>
). Clade
<emphasis id="34F7EA958465FFE4596AFEC5FAA5FEF0" bold="true" box="[1309,1323,330,356]" pageId="11" pageNumber="348">4</emphasis>
is further characterized by several well-supported subclades (
<emphasis id="34F7EA958465FFE45EA1FEE1FD70FE1C" bold="true" box="[726,766,366,392]" pageId="11" pageNumber="348">M1</emphasis>
,
<emphasis id="34F7EA958465FFE45F7CFEE1FCAAFE1C" bold="true" box="[779,804,366,392]" pageId="11" pageNumber="348">I1</emphasis>
,
<emphasis id="34F7EA958465FFE45F45FEE1FCDAFE1C" bold="true" box="[818,852,366,392]" pageId="11" pageNumber="348">A1</emphasis>
,
<emphasis id="34F7EA958465FFE45F16FEE1FCF3FE1C" bold="true" box="[865,893,366,392]" pageId="11" pageNumber="348">J3</emphasis>
) and a large group (
<emphasis id="34F7EA958465FFE4582AFEE0FBE5FE1D" bold="true" box="[1117,1131,367,393]" pageId="11" pageNumber="348">5</emphasis>
) containing subclades with varying levels of support (
<emphasis id="34F7EA958465FFE45DC9FE1DFE59FE38" bold="true" box="[446,471,402,428]" pageId="11" pageNumber="348">I4</emphasis>
,
<emphasis id="34F7EA958465FFE45D92FE1DFD8DFE38" bold="true" box="[485,515,402,428]" pageId="11" pageNumber="348">F1</emphasis>
,
<emphasis id="34F7EA958465FFE45E65FE1DFDBBFE38" bold="true" box="[530,565,402,428]" pageId="11" pageNumber="348">H1</emphasis>
,
<emphasis id="34F7EA958465FFE45E34FE1DFDE8FE38" bold="true" box="[579,614,402,428]" pageId="11" pageNumber="348">C1</emphasis>
,
<emphasis id="34F7EA958465FFE45E04FE1DFD19FE38" bold="true" box="[627,663,402,428]" pageId="11" pageNumber="348">G1</emphasis>
,
<emphasis id="34F7EA958465FFE45ED2FE1DFD49FE38" bold="true" box="[677,711,402,428]" pageId="11" pageNumber="348">D1</emphasis>
&amp;
<emphasis id="34F7EA958465FFE45EAAFE1DFD71FE38" bold="true" box="[733,767,402,428]" pageId="11" pageNumber="348">D2</emphasis>
, and
<emphasis id="34F7EA958465FFE45F4BFE1CFCE9FE38" bold="true" box="[828,871,402,429]" pageId="11" pageNumber="348">B1)</emphasis>
(see
<figureCitation id="9EB82A028465FFE45FD6FE1CFC6FFE39" box="[929,993,403,429]" captionStart="FIGURE 1" captionStartId="20.[151,250,1882,1906]" captionTargetBox="[367,1212,181,1844]" captionTargetId="figure-21@20.[363,1225,179,1844]" captionTargetPageId="20" captionText="FIGURE 1. RAxML phylogenetic tree based on a concatenated alignment of mtMutS-5, mtMutS-3 and 18S molecular sequences of 92 total taxa (86 keratoisids + 6 out group). Bootstrap values&gt;50 are shown above the nodes. Bayesian posterior probability values&gt;0.5 are shown below the node. Numbers 1 to 5 refer to nodes discussed in the text. Clades delimited by vertical bars on the right side are named following the convention of France (2007) and are discussed in detail in the text." figureDoi="http://doi.org/10.5281/zenodo.5909956" httpUri="https://zenodo.org/record/5909956/files/figure.png" pageId="11" pageNumber="348">Fig. 1</figureCitation>
); as noted earlier these subclade names follow a convention established by
<bibRefCitation id="62124B768465FFE45E55FE38FD31FE45" author="France, S. C." box="[546,703,439,465]" pageId="11" pageNumber="348" pagination="323 - 333" refId="ref13930" refString="France, S. C. (2007) Genetic analysis of bamboo corals (Cnidaria: Octocorallia: Isididae): Does lack of colony branching distinguish Lepidisis from Keratoisis? Bulletin of Marine Science, 81, 323 - 333." type="journal article" year="2007">
<collectingCountry id="7E9476178465FFE45E55FE38FDE0FE45" box="[546,622,439,465]" name="France" pageId="11" pageNumber="348">France</collectingCountry>
(2007)
</bibRefCitation>
based on pattern of unique indels and have been used by other investigators, so we retain them here. More genetic data will be needed to boost support for internal nodes and resolve the sister-taxon relationships among these subclades. Morphological features of these subclades are summarized in
<tableCitation id="4B01033C8465FFE45CE0FDACFF62FDA9" box="[151,236,547,573]" captionStart="TABLE 4" captionStartId="17.[151,239,484,510]" captionTargetPageId="17" captionText="TABLE 4. Summary of morphological characters associated with each molecular subclade or indel type." httpUri="http://table.plazi.org/id/52FC660F847FFFFE5CE0FE6BFB78FE6A" pageId="11" pageNumber="348" tableUuid="52FC660F847FFFFE5CE0FE6BFB78FE6A">Table 4</tableCitation>
, and selected examples of colony and polyp form are illustrated in
<figureCitation id="9EB82A028465FFE45FABFDACFBA6FDA9" box="[988,1064,547,573]" captionStart="FIGURE 2" captionStartId="21.[151,250,1645,1669]" captionTargetBox="[280,1308,181,1621]" captionTargetId="figure-19@21.[280,1308,181,1621]" captionTargetPageId="21" captionText="FIGURE 2. A selection of colony types from the Family Keratoisididae. Clade names as noted in the text, followed by species name (if known), specimen number, approximate colony size, and location. S1, Cladarisis nouvianae, CAT 207-1, 30 cm, Cat Island, Bahamas; M1, Orstomisis sp., NMHN IK-2008-14, 45 cm across branched area, New Caledonia; I1, Lepidisis caryophyllia, GIL 101-1, 17 cm, Gilbert Canyon, northeastern USA; A1, Acanella arbuscula, MAN 803-1, 30 cm height, Manning Seamount, New England Seamounts; J3-1, KEL 604-2, 30 cm, Kelvin Seamounts, New England Seamounts; J3-2, KUK 218-1, approx. 75 cm width of fan, Kükenthal Peak, Caloosahatchee Seamount, Corner Rise Seamounts; J3-3, NAS 101-3, 30 cm field of view, individual colony approx. 50-75 cm, Nashville Seamount, New England Seamounts; I4, BAL 205-1, 50 cm height, Balanus Seamount, New England Seamounts; F1, ANN 209-1, longest branch approx. 40 cm, Annei Seamount, Emperor Seamounts; C1, BAL 211-3, approx. 2 m length, Balanus Seamount; D1, GOO 101-1, approx. 1 m. length, Goode Peak, Caloosahatchee Seamount; D2, Eknomisis dalioi, KEL 406-1, colony approx. 1 m tall by 1.5 m wide, Kelvin Seamount; B1, KEL 408-1, approx. 2 to 3 m length, Kelvin Seamount." figureDoi="http://doi.org/10.5281/zenodo.5909958" httpUri="https://zenodo.org/record/5909958/files/figure.png" pageId="11" pageNumber="348">Figs. 2</figureCitation>
and
<figureCitation id="9EB82A028465FFE45829FDACFBE0FDA9" box="[1118,1134,547,573]" captionStart="FIGURE 3" captionStartId="22.[151,250,1637,1661]" captionTargetBox="[346,1241,181,1614]" captionTargetId="figure-21@22.[346,1241,181,1614]" captionTargetPageId="22" captionText="FIGURE 3. A selection of polyp types from the Family Keratoisididae. Clade names as noted in the text, followed by species name (if known), specimen number, special treatment (if any), and location. All polyps are less than 1 cm in length or height and are in contracted state due to preservation. S1, Cladarisis nouvianae, CAT 207-1, Cat Island, Bahamas; M1, Orstomisis sp., NMHN IK-2008-14, polyp cleared with clove oil, New Caledonia; I1, Lepidisis caryophyllia, GIL 101-1, SEM view with external tissue removed, Gilbert Canyon, northeast USA; A1, Acanella arbuscula, KUK 103-2, Kükenthal Peak, Caloosahatchee Seamount, Corner Rise Seamounts; J3-1, KEL 604-2, Kelvin Seamount, New England Seamounts; J3-2, KUK 218-1, Kükenthal Peak; I4, BAL 205-1, polyp cleared with clove oil, Balanus Seamount, New England seamounts; F1, KEL 104-1, Kelvin Seamount; H1, Keratoisis magnifica, TMAG K3830, Tasmania Seamounts; C1, BAL 111-1, Balanus Seamount; G1, TMAG K3856, Tasmanian Seamounts; D1, MAN 707-2, Manning Seamount, New England Seamounts; D2, Eknomisis dalioi, KEL 406-1, Kelvin Seamount; B1-1, PIC 102-1, Picket Seamount, New England Seamounts; B1-2, J2098-6-4, Aleutian Islands slope, Alaska, USA." figureDoi="http://doi.org/10.5281/zenodo.5909960" httpUri="https://zenodo.org/record/5909960/files/figure.png" pageId="11" pageNumber="348">3</figureCitation>
.
</paragraph>
<paragraph id="063C36878465FFE45CB0FDC8FB5BFC5D" blockId="11.[151,1437,151,2013]" pageId="11" pageNumber="348">
Subclade
<emphasis id="34F7EA958465FFE45D42FDC9FED0FDF4" bold="true" box="[309,350,582,608]" pageId="11" pageNumber="348">M1</emphasis>
comprises 2 haplotypes from
<specimenCount id="1085FD0E8465FFE45EC6FDC8FCD9FDF5" box="[689,855,583,609]" pageId="11" pageNumber="348" type="generic">two specimens</specimenCount>
both identified as
<taxonomicName id="C1834D048465FFE4585FFDC8FA12FDF5" authority="Bayer, 1990" authorityName="Bayer" authorityYear="1990" box="[1064,1436,583,609]" class="Anthozoa" family="Isididae" genus="Orstomisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="crosnieri">
<emphasis id="34F7EA958465FFE4585FFDC8FA83FDF4" box="[1064,1293,583,609]" italics="true" pageId="11" pageNumber="348">Orstomisis crosnieri</emphasis>
<bibRefCitation id="62124B768465FFE45963FDC8FA12FDF5" author="Bayer, F. M." box="[1300,1436,583,609]" pageId="11" pageNumber="348" pagination="205 - 228" refId="ref13270" refString="Bayer, F. M. (1990) A new isidid octocoral (Anthozoa: Gorgonacea) from New Caledonia, with descriptions of other new species from elsewhere in the Pacific Ocean. Proceedings of the Biological Society of Washington, 103, 205 - 228." type="journal article" year="1990">Bayer, 1990</bibRefCitation>
</taxonomicName>
and collected from near the type locality (
<collectingCountry id="7E9476178465FFE45E0FFDE4FCA4FD12" box="[632,810,619,646]" name="New Caledonia" pageId="11" pageNumber="348">New Caledonia</collectingCountry>
) for that species. The 18S sequence for
<taxonomicName id="C1834D048465FFE4588DFDE4FA0CFD10" authorityName="Bayer" authorityYear="1990" box="[1274,1410,619,645]" class="Anthozoa" family="Isididae" genus="Orstomisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="crosnieri">
<emphasis id="34F7EA958465FFE4588DFDE4FA0CFD10" box="[1274,1410,619,645]" italics="true" pageId="11" pageNumber="348">O. crosnieri</emphasis>
</taxonomicName>
A does not differ from the 18S sequences of the
<typeStatus id="D93888258465FFE45EC7FD00FC89FD3D" box="[688,775,655,681]" pageId="11" pageNumber="348" type="syntype">syntype</typeStatus>
(
<accessionNumber id="19D0AB648465FFE45F6EFD01FC18FD3D" box="[793,918,654,681]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/AF052916" pageId="11" pageNumber="348">AF052916</accessionNumber>
; Berntson
<emphasis id="34F7EA958465FFE45865FD00FBC3FD3D" box="[1042,1101,655,681]" italics="true" pageId="11" pageNumber="348">et al.</emphasis>
2001). Both specimens were collected on the same cruise but from different stations and the sequences were derived in separate labs (A by the current authors and B from
<bibRefCitation id="62124B768465FFE45DB0FD58FD14FD65" author="Duenas, L. F. &amp; Alderslade, P. &amp; Sanchez, J. A." box="[455,666,727,753]" pageId="11" pageNumber="348" pagination="15 - 28" refId="ref13800" refString="Duenas, L. F., Alderslade, P. &amp; Sanchez, J. A. (2014) Molecular systematics of the deep-sea bamboo corals (Octocorallia: Isididae: Keratoisidinae) from New Zealand with descriptions of two new species of Keratoisis. Molecular Phylogenetics and Evolution, 74, 15 - 28. https: // doi. org / 10.1016 / j. ympev. 2014.01.031" type="journal article" year="2014">
Dueñas
<emphasis id="34F7EA958465FFE45E56FD58FDD6FD65" box="[545,600,727,753]" italics="true" pageId="11" pageNumber="348">et al.</emphasis>
2014
</bibRefCitation>
); the
<emphasis id="34F7EA958465FFE45EA3FD58FCBAFD65" box="[724,820,727,753]" italics="true" pageId="11" pageNumber="348">mtMutS-</emphasis>
5 haplotypes are identical.A third haplotype associated with a specimen (VOL1) identified as
<taxonomicName id="C1834D048465FFE45E27FD74FD56FC80" authorityName="Bayer" authorityYear="1990" box="[592,728,763,789]" class="Anthozoa" family="Isididae" genus="Orstomisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="crosnieri">
<emphasis id="34F7EA958465FFE45E27FD74FD56FC80" box="[592,728,763,789]" italics="true" pageId="11" pageNumber="348">O. crosnieri</emphasis>
</taxonomicName>
and also collected from near the type locality is on an unsupported branch, which, if the unsupported nodes were collapsed, would result in a trichotomy with the
<emphasis id="34F7EA958465FFE45956FC91FAC4FCAC" bold="true" box="[1313,1354,798,824]" pageId="11" pageNumber="348">M1</emphasis>
and
<emphasis id="34F7EA958465FFE459F4FC91FA12FCAC" bold="true" box="[1411,1436,798,824]" pageId="11" pageNumber="348">I1</emphasis>
clades.
<taxonomicName id="C1834D048465FFE45C9BFCCCFE62FCC9" authority="C" box="[236,492,835,861]" class="Anthozoa" family="Isididae" genus="Orstomisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="343" phylum="Cnidaria" rank="species" species="crosnieri">
<emphasis id="34F7EA958465FFE45C9BFCCCFE5FFCC8" box="[236,465,835,861]" italics="true" pageId="11" pageNumber="348">Orstomisis crosnieri</emphasis>
C
</taxonomicName>
differs from
<taxonomicName id="C1834D048465FFE45EF6FCCCFC86FCC8" box="[641,776,835,861]" class="Anthozoa" family="Isididae" genus="Orstomisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="crosnieri">
<emphasis id="34F7EA958465FFE45EF6FCCCFC86FCC8" box="[641,776,835,861]" italics="true" pageId="11" pageNumber="348">O. crosnieri</emphasis>
</taxonomicName>
A and B by [only] p = 0.0074 at
<emphasis id="34F7EA958465FFE458F5FCCCFB6CFCC9" box="[1154,1250,835,861]" italics="true" pageId="11" pageNumber="348">mtMutS-</emphasis>
5, and might be an undescribed
<taxonomicName id="C1834D048465FFE45D3EFCE8FDADFC15" box="[329,547,871,897]" class="Anthozoa" family="Isididae" genus="Orstomisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="undetermined">
<emphasis id="34F7EA958465FFE45D3EFCE8FE4CFC15" box="[329,450,871,897]" italics="true" pageId="11" pageNumber="348">Orstomisis</emphasis>
species.
</taxonomicName>
Taxa in subclade
<emphasis id="34F7EA958465FFE45E98FCE9FC99FC14" bold="true" box="[751,791,870,896]" pageId="11" pageNumber="348">M1</emphasis>
, typified by
<taxonomicName id="C1834D048465FFE45FDEFCE8FBBEFC14" box="[937,1072,871,897]" class="Anthozoa" family="Isididae" genus="Orstomisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="crosnieri">
<emphasis id="34F7EA958465FFE45FDEFCE8FBBEFC14" box="[937,1072,871,897]" italics="true" pageId="11" pageNumber="348">O. crosnieri</emphasis>
</taxonomicName>
, have a solid axis; dichotomous or pseudodichotomous branching that originates at the organic nodes; polyp sclerites that are all rod-shaped; and a polyp that does not shorten on contraction but has tentacles that simply fold over the mouth area.
</paragraph>
<paragraph id="063C36878465FFE45CB0FC5CFD84FAED" blockId="11.[151,1437,151,2013]" pageId="11" pageNumber="348">
Subclade
<emphasis id="34F7EA958465FFE45D41FC5DFEC0FC78" bold="true" box="[310,334,978,1004]" pageId="11" pageNumber="348">I1</emphasis>
is one of the more genetically divergent of the clade
<emphasis id="34F7EA958465FFE45FDDFC5DFC36FC78" bold="true" box="[938,952,978,1004]" pageId="11" pageNumber="348">4</emphasis>
subclades (along with the group at clade
<emphasis id="34F7EA958465FFE459F9FC5CFA12FC79" bold="true" box="[1422,1436,979,1005]" pageId="11" pageNumber="348">5</emphasis>
from
<emphasis id="34F7EA958465FFE45CA1FC79FF7BFB84" bold="true" box="[214,245,1014,1040]" pageId="11" pageNumber="348">F1</emphasis>
to
<emphasis id="34F7EA958465FFE45D6BFC78FEB2FB84" bold="true" box="[284,316,1014,1041]" pageId="11" pageNumber="348">B1</emphasis>
), in terms of genetic distance between haplotypes, with an intra-subclade p =
<geoCoordinate id="63B750408465FFE458CFFC79FA8BFB85" box="[1208,1285,1014,1041]" degrees="0.0005" direction="north" orientation="latitude" pageId="11" pageNumber="348" precision="5" value="5.0E-4">0.0005</geoCoordinate>
<geoCoordinate id="63B750408465FFE45972FC79FADFFB85" box="[1285,1361,1014,1041]" degrees="0.013" direction="west" orientation="longitude" pageId="11" pageNumber="348" precision="55" value="-0.013">0.013</geoCoordinate>
at
<emphasis id="34F7EA958465FFE45900FC78FF55FBA1" italics="true" pageId="11" pageNumber="348">mt- MutS-</emphasis>
5. It includes specimens initially identified in 3 different genera (
<taxonomicName id="C1834D048465FFE45FA4FB94FBA9FBA1" box="[979,1063,1051,1077]" class="Anthozoa" family="Isididae" genus="Isidella" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE45FA4FB94FBA9FBA1" box="[979,1063,1051,1077]" italics="true" pageId="11" pageNumber="348">Isidella</emphasis>
</taxonomicName>
,
<taxonomicName id="C1834D048465FFE4584FFB94FB25FBA0" authorityName="Wright" authorityYear="1869" box="[1080,1195,1051,1076]" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE4584FFB94FB25FBA0" box="[1080,1195,1051,1076]" italics="true" pageId="11" pageNumber="348">Keratoisis</emphasis>
</taxonomicName>
,
<taxonomicName id="C1834D048465FFE458CCFB94FAAEFBA1" authorityName="Verrill" authorityYear="1883" box="[1211,1312,1051,1077]" class="Anthozoa" family="Isididae" genus="Lepidisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE458CCFB94FAAEFBA1" box="[1211,1312,1051,1077]" italics="true" pageId="11" pageNumber="348">Lepidisis</emphasis>
</taxonomicName>
) and three different haplotypes from specimens identified as
<taxonomicName id="C1834D048465FFE45EB2FBB0FBCEFBCD" authority="Etnoyer, 2008" authorityName="Etnoyer" authorityYear="2008" box="[709,1088,1087,1113]" class="Anthozoa" family="Isididae" genus="Isidella" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="tentaculum">
<emphasis id="34F7EA958465FFE45EB2FBB0FC12FBCD" box="[709,924,1087,1113]" italics="true" pageId="11" pageNumber="348">Isidella tentaculum</emphasis>
Etnoyer, 2008
</taxonomicName>
(one of which is the
<typeStatus id="D93888258465FFE4595DFBB0FA00FBCE" box="[1322,1422,1087,1114]" pageId="11" pageNumber="348" type="holotype">holotype</typeStatus>
). Sequences from three of the
<taxonomicName id="C1834D048465FFE45DAEFBECFD2AFBE9" box="[473,676,1123,1149]" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="undetermined">
<emphasis id="34F7EA958465FFE45DAEFBECFDC2FBE8" box="[473,588,1123,1148]" italics="true" pageId="11" pageNumber="348">Keratoisis</emphasis>
species
</taxonomicName>
are derived from
<typeStatus id="D93888258465FFE45F1AFBECFC41FBE9" box="[877,975,1123,1149]" pageId="11" pageNumber="348" type="holotype">holotype</typeStatus>
specimens, but, as discussed below, they almost certainly should not be in that genus. Members of this subclade possess a hollow, thin axis, at least in the distal parts of the colony; when present, branches originate at the nodes; polyp sclerites, when present, are rods, often in the form of needles, with long, sometimes thick needles positioned between the bases of the tentacles (referred to as intertentacular needles due to their mesenterial, or aligned with the septa, arrangement); and the polyp on contraction does not shorten but the tentacles fold over the mouth. This subclade contains
<taxonomicName id="C1834D048465FFE458D4FA98FEA6FAC1" authority="Verrill, 1883" authorityName="Verrill" authorityYear="1883" class="Anthozoa" family="Isididae" genus="Lepidisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="caryophyllia">
<emphasis id="34F7EA958465FFE458D4FA98FA13FAA5" box="[1187,1437,1303,1329]" italics="true" pageId="11" pageNumber="348">Lepidisis caryophyllia</emphasis>
<bibRefCitation id="62124B768465FFE45CE0FAB4FEA6FAC1" author="Verrill, A. E." box="[151,296,1339,1365]" pageId="11" pageNumber="348" pagination="1 - 72" refId="ref15349" refString="Verrill, A. E. (1883) Report on the Anthozoa, and on some additional species dredged by the &quot; Blake &quot; in 1877 - 1879, and by the U. S. Fish Commission Steamer &quot; Fish Hawk &quot; in 1880 - 82. Bulletin of the Museum of Comparative Zoology, 11, 1 - 72." type="journal article" year="1883">Verrill, 1883</bibRefCitation>
</taxonomicName>
, one of the oldest known keratoisids and the type species in one of the most commonly mis-identified genera (
<bibRefCitation id="62124B768465FFE45C87FAD0FE73FAED" author="Watling, L. &amp; France, S. C." box="[240,509,1375,1401]" pageId="11" pageNumber="348" pagination="97 - 110" refId="ref15518" refString="Watling, L. &amp; France, S. C. (2021) Towards a revision of the bamboo corals: Part 2, untangling the genus Lepidisis (Octocorallia: Isididae). Bulletin of the Peabody Museum of Natural History, 62, 97 - 110. https: // doi. org / 10.3374 / 014.062.0202" type="journal article" year="2021">
Watling &amp;
<collectingCountry id="7E9476178465FFE45D19FAD0FE34FAED" box="[366,442,1375,1401]" name="France" pageId="11" pageNumber="348">France</collectingCountry>
2021
</bibRefCitation>
).
</paragraph>
<paragraph id="063C36878465FFE45CB0FA0CFD8BF90D" blockId="11.[151,1437,151,2013]" pageId="11" pageNumber="348">
Subclade
<emphasis id="34F7EA958465FFE45D43FA0DFED8FA08" bold="true" box="[308,342,1410,1436]" pageId="11" pageNumber="348">A1</emphasis>
contains all taxa identified as
<emphasis id="34F7EA958465FFE45EDFFA0CFC9CFA09" box="[680,786,1411,1437]" italics="true" pageId="11" pageNumber="348">
<taxonomicName id="C1834D048465FFE45EDFFA0CFC80FA09" box="[680,782,1411,1437]" class="Anthozoa" family="Isididae" genus="Acanella" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">Acanella</taxonomicName>
,
</emphasis>
along with a single haplotype downloaded from GenBank and identified there as
<taxonomicName id="C1834D048465FFE45DE0FA28FDDAFA55" baseAuthorityName="Esper" baseAuthorityYear="1788" box="[407,596,1447,1473]" class="Anthozoa" family="Isididae" genus="Isidella" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="elongata">
<emphasis id="34F7EA958465FFE45DE0FA28FDDAFA55" box="[407,596,1447,1473]" italics="true" pageId="11" pageNumber="348">Isidella elongata</emphasis>
</taxonomicName>
.
<bibRefCitation id="62124B768465FFE45E15FA28FC39FA55" author="Heestand Saucier, E. &amp; Sajjadi, A. &amp; France, S. C." box="[610,951,1447,1473]" pageId="11" pageNumber="348" pagination="359 - 390" refId="ref14254" refString="Heestand Saucier, E., Sajjadi, A. &amp; France, S. C. (2017) A taxonomic review of the genus Acanella (Cnidaria: Octocorallia: Isididae) in the North Atlantic Ocean, with descriptions of two new species. Zootaxa, 4323 (3), 359 - 390. https: // doi. org / 10.11646 / zootaxa. 4323.3.2" type="journal article" year="2017">
Heestand Saucier
<emphasis id="34F7EA958465FFE45F5AFA28FCEBFA55" box="[813,869,1447,1473]" italics="true" pageId="11" pageNumber="348">et al.</emphasis>
(2017)
</bibRefCitation>
noted that at least some colonies identified as
<taxonomicName id="C1834D048465FFE45CC1FA44FEBEFA71" baseAuthorityName="Esper" baseAuthorityYear="1788" box="[182,304,1483,1509]" class="Anthozoa" family="Isididae" genus="Isidella" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="elongata">
<emphasis id="34F7EA958465FFE45CC1FA44FEBEFA71" box="[182,304,1483,1509]" italics="true" pageId="11" pageNumber="348">I. elongata</emphasis>
</taxonomicName>
could be considered to be conspecific with
<taxonomicName id="C1834D048465FFE45F68FA44FC74FA71" box="[799,1018,1483,1509]" class="Anthozoa" family="Isididae" genus="Acanella" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="arbuscula">
<emphasis id="34F7EA958465FFE45F68FA44FC74FA71" box="[799,1018,1483,1509]" italics="true" pageId="11" pageNumber="348">Acanella arbuscula</emphasis>
</taxonomicName>
. Divergence among the 8 haplotypes of subclade
<emphasis id="34F7EA958465FFE45D57FA61FECCF99C" bold="true" box="[288,322,1518,1544]" pageId="11" pageNumber="348">A1</emphasis>
is on par with that among the 4 haplotypes of subclade
<emphasis id="34F7EA958465FFE45FC8FA61FC52F99C" bold="true" box="[959,988,1518,1544]" pageId="11" pageNumber="348">S1</emphasis>
, with intra-subclade p =
<geoCoordinate id="63B750408465FFE4588EFA61FAC9F99D" box="[1273,1351,1518,1545]" degrees="0.0005" direction="north" orientation="latitude" pageId="11" pageNumber="348" precision="5" value="5.0E-4">0.0005</geoCoordinate>
<geoCoordinate id="63B750408465FFE45930FA61FA12F99D" box="[1351,1436,1518,1545]" degrees="0.0083" direction="west" orientation="longitude" pageId="11" pageNumber="348" precision="5" value="-0.0083">-0.0083</geoCoordinate>
at
<emphasis id="34F7EA958465FFE45CC5F99CFE9CF9B9" box="[178,274,1555,1581]" italics="true" pageId="11" pageNumber="348">mtMutS-</emphasis>
5. Morphologically, members of this subclade are characterized by a solid axis, with branches originating at the organic nodes, usually in whorls, often of up to 5 branches; sclerites of the polyps are needles and other rods, occasionally intertentacular in position; and a polyp that does not shorten on contraction but has tentacles that simply fold over the mouth area.
</paragraph>
<paragraph id="063C36878465FFE35CB0F92CFA85FEF1" blockId="11.[151,1437,151,2013]" lastBlockId="12.[151,1437,151,1869]" lastPageId="12" lastPageNumber="349" pageId="11" pageNumber="348">
Subclade
<emphasis id="34F7EA958465FFE45D43F92DFEDFF928" bold="true" box="[308,337,1698,1724]" pageId="11" pageNumber="348">J3</emphasis>
includes specimens identified to 5 different genera (
<taxonomicName id="C1834D048465FFE45FEFF92CFC62F929" box="[920,1004,1699,1725]" class="Anthozoa" family="Isididae" genus="Isidella" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE45FEFF92CFC62F929" box="[920,1004,1699,1725]" italics="true" pageId="11" pageNumber="348">Isidella</emphasis>
</taxonomicName>
,
<taxonomicName id="C1834D048465FFE45F8DF92CFBEFF928" authorityName="Alderslade &amp; McFadden" authorityYear="2012" box="[1018,1121,1699,1724]" class="Anthozoa" family="Isididae" genus="Jasonisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE45F8DF92CFBEFF928" box="[1018,1121,1699,1724]" italics="true" pageId="11" pageNumber="348">Jasonisis</emphasis>
</taxonomicName>
,
<taxonomicName id="C1834D048465FFE45819F92CFB6FF928" authorityName="Wright" authorityYear="1869" box="[1134,1249,1699,1724]" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE45819F92CFB6FF928" box="[1134,1249,1699,1724]" italics="true" pageId="11" pageNumber="348">Keratoisis</emphasis>
</taxonomicName>
,
<emphasis id="34F7EA958465FFE45898F92CFF51F974" italics="true" pageId="11" pageNumber="348">
<taxonomicName id="C1834D048465FFE45898F92CFAD9F929" authorityName="Verrill" authorityYear="1883" box="[1263,1367,1699,1725]" class="Anthozoa" family="Isididae" genus="Lepidisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">Lepidisis</taxonomicName>
,
<taxonomicName id="C1834D048465FFE45916F92CFF51F974" authorityName="Bayer" authorityYear="1990" class="Anthozoa" family="Isididae" genus="Orstomisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">Orstomisis</taxonomicName>
</emphasis>
) as well as unidentified taxa that are putatively new species. Specimens assigned to the genera
<taxonomicName id="C1834D048465FFE4588DF948FAC0F975" box="[1274,1358,1735,1761]" class="Anthozoa" family="Isididae" genus="Isidella" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE4588DF948FAC0F975" box="[1274,1358,1735,1761]" italics="true" pageId="11" pageNumber="348">Isidella</emphasis>
</taxonomicName>
,
<taxonomicName id="C1834D048465FFE4592CF948FF5CF890" authorityName="Wright" authorityYear="1869" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE4592CF948FF5CF890" italics="true" pageId="11" pageNumber="348">Keratoisis</emphasis>
</taxonomicName>
,
<taxonomicName id="C1834D048465FFE45C95F964FEC9F891" authorityName="Verrill" authorityYear="1883" box="[226,327,1771,1797]" class="Anthozoa" family="Isididae" genus="Lepidisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE45C95F964FEC9F891" box="[226,327,1771,1797]" italics="true" pageId="11" pageNumber="348">Lepidisis</emphasis>
</taxonomicName>
and
<taxonomicName id="C1834D048465FFE45DF5F964FE75F891" authorityName="Bayer" authorityYear="1990" box="[386,507,1771,1797]" class="Anthozoa" family="Isididae" genus="Orstomisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE45DF5F964FE75F891" box="[386,507,1771,1797]" italics="true" pageId="11" pageNumber="348">Orstomisis</emphasis>
</taxonomicName>
were provisionally identified before the genus
<taxonomicName id="C1834D048465FFE4586DF964FB0FF890" authorityName="Alderslade &amp; McFadden" authorityYear="2012" box="[1050,1153,1771,1796]" class="Anthozoa" family="Isididae" genus="Jasonisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE4586DF964FB0FF890" box="[1050,1153,1771,1796]" italics="true" pageId="11" pageNumber="348">Jasonisis</emphasis>
</taxonomicName>
was described (in 2012) and are almost certainly incorrectly identified. One of the included haplotypes in Subclade
<emphasis id="34F7EA958465FFE458E3F881FB3EF8BC" bold="true" box="[1172,1200,1806,1832]" pageId="11" pageNumber="348">J3</emphasis>
comes from the
<typeStatus id="D93888258465FFE4591BF880FA12F8BD" box="[1388,1436,1807,1833]" pageId="11" pageNumber="348">type</typeStatus>
specimen for the genus
<taxonomicName id="C1834D048465FFE45DEBF8BCFD8DF8D8" authorityName="Alderslade &amp; McFadden" authorityYear="2012" box="[412,515,1843,1868]" class="Anthozoa" family="Isididae" genus="Jasonisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE45DEBF8BCFD8DF8D8" box="[412,515,1843,1868]" italics="true" pageId="11" pageNumber="348">Jasonisis</emphasis>
</taxonomicName>
(
<taxonomicName id="C1834D048465FFE45E66F8BCFC7EF8D9" authority="Alderslade &amp; McFadden, 2012" authorityName="Alderslade &amp; McFadden" authorityYear="2012" box="[529,1008,1843,1869]" class="Anthozoa" family="Isididae" genus="Jasonisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="thresheri">
<emphasis id="34F7EA958465FFE45E66F8BCFD00F8D9" box="[529,654,1843,1869]" italics="true" pageId="11" pageNumber="348">J. thresheri</emphasis>
<bibRefCitation id="62124B768465FFE45EE3F8BCFC7EF8D9" author="Alderslade, P. &amp; McFadden, C. S." box="[660,1008,1843,1869]" pageId="11" pageNumber="348" pagination="21 - 39" refId="ref13137" refString="Alderslade, P. &amp; McFadden, C. S. (2012) A new genus and species of the family Isididae (Coelenterata: Octocorallia) from a CMAR Biodiversity study, and a discussion on the subfamilial placement of some nominal isidid genera. Zootaxa, 3154 (1), 21 - 39. https: // doi. org / 10.11646 / zootaxa. 3154.1.2" type="journal article" year="2012">Alderslade &amp; McFadden, 2012</bibRefCitation>
</taxonomicName>
). The
<taxonomicName id="C1834D048465FFE45846F8BCFB16F8D8" authorityName="Alderslade &amp; McFadden" authorityYear="2012" box="[1073,1176,1843,1868]" class="Anthozoa" family="Isididae" genus="Jasonisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE45846F8BCFB16F8D8" box="[1073,1176,1843,1868]" italics="true" pageId="11" pageNumber="348">Jasonisis</emphasis>
</taxonomicName>
<typeStatus id="D93888258465FFE458EAF8BCFB43F8D9" box="[1181,1229,1843,1869]" pageId="11" pageNumber="348">type</typeStatus>
specimen and tentatively identified
<taxonomicName id="C1834D048465FFE45D14F8D8FE44F8E4" authorityName="Alderslade &amp; McFadden" authorityYear="2012" box="[355,458,1879,1904]" class="Anthozoa" family="Isididae" genus="Jasonisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE45D14F8D8FE44F8E4" box="[355,458,1879,1904]" italics="true" pageId="11" pageNumber="348">Jasonisis</emphasis>
</taxonomicName>
specimens do not share the same indel structure (
<taxonomicName id="C1834D048465FFE45F86F8D8FBE1F8E5" authorityName="Alderslade &amp; McFadden" authorityYear="2012" box="[1009,1135,1879,1905]" class="Anthozoa" family="Isididae" genus="Jasonisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="species" species="thresheri">
<emphasis id="34F7EA958465FFE45F86F8D8FBE1F8E5" box="[1009,1135,1879,1905]" italics="true" pageId="11" pageNumber="348">J. thresheri</emphasis>
</taxonomicName>
has 18 base pairs (6 amino acids) fewer than the other tentative
<taxonomicName id="C1834D048465FFE45E43F8F4FD15F800" authorityName="Alderslade &amp; McFadden" authorityYear="2012" box="[564,667,1915,1940]" class="Anthozoa" family="Isididae" genus="Jasonisis" kingdom="Animalia" order="Alcyonacea" pageId="11" pageNumber="348" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958465FFE45E43F8F4FD15F800" box="[564,667,1915,1940]" italics="true" pageId="11" pageNumber="348">Jasonisis</emphasis>
</taxonomicName>
specimens). Members of this clade show a wide diversity of colony and polyp form. Three colony forms and two polyp
<typeStatus id="D93888258465FFE45EA9F810FC97F82D" box="[734,793,1951,1977]" pageId="11" pageNumber="348">types</typeStatus>
are illustrated in
<figureCitation id="9EB82A028465FFE45FA9F810FBA5F82D" box="[990,1067,1951,1977]" captionStart="FIGURE 2" captionStartId="21.[151,250,1645,1669]" captionTargetBox="[280,1308,181,1621]" captionTargetId="figure-19@21.[280,1308,181,1621]" captionTargetPageId="21" captionText="FIGURE 2. A selection of colony types from the Family Keratoisididae. Clade names as noted in the text, followed by species name (if known), specimen number, approximate colony size, and location. S1, Cladarisis nouvianae, CAT 207-1, 30 cm, Cat Island, Bahamas; M1, Orstomisis sp., NMHN IK-2008-14, 45 cm across branched area, New Caledonia; I1, Lepidisis caryophyllia, GIL 101-1, 17 cm, Gilbert Canyon, northeastern USA; A1, Acanella arbuscula, MAN 803-1, 30 cm height, Manning Seamount, New England Seamounts; J3-1, KEL 604-2, 30 cm, Kelvin Seamounts, New England Seamounts; J3-2, KUK 218-1, approx. 75 cm width of fan, Kükenthal Peak, Caloosahatchee Seamount, Corner Rise Seamounts; J3-3, NAS 101-3, 30 cm field of view, individual colony approx. 50-75 cm, Nashville Seamount, New England Seamounts; I4, BAL 205-1, 50 cm height, Balanus Seamount, New England Seamounts; F1, ANN 209-1, longest branch approx. 40 cm, Annei Seamount, Emperor Seamounts; C1, BAL 211-3, approx. 2 m length, Balanus Seamount; D1, GOO 101-1, approx. 1 m. length, Goode Peak, Caloosahatchee Seamount; D2, Eknomisis dalioi, KEL 406-1, colony approx. 1 m tall by 1.5 m wide, Kelvin Seamount; B1, KEL 408-1, approx. 2 to 3 m length, Kelvin Seamount." figureDoi="http://doi.org/10.5281/zenodo.5909958" httpUri="https://zenodo.org/record/5909958/files/figure.png" pageId="11" pageNumber="348">Figs. 2</figureCitation>
and
<figureCitation id="9EB82A028465FFE45815F810FBFCF82D" box="[1122,1138,1951,1977]" captionStart="FIGURE 3" captionStartId="22.[151,250,1637,1661]" captionTargetBox="[346,1241,181,1614]" captionTargetId="figure-21@22.[346,1241,181,1614]" captionTargetPageId="22" captionText="FIGURE 3. A selection of polyp types from the Family Keratoisididae. Clade names as noted in the text, followed by species name (if known), specimen number, special treatment (if any), and location. All polyps are less than 1 cm in length or height and are in contracted state due to preservation. S1, Cladarisis nouvianae, CAT 207-1, Cat Island, Bahamas; M1, Orstomisis sp., NMHN IK-2008-14, polyp cleared with clove oil, New Caledonia; I1, Lepidisis caryophyllia, GIL 101-1, SEM view with external tissue removed, Gilbert Canyon, northeast USA; A1, Acanella arbuscula, KUK 103-2, Kükenthal Peak, Caloosahatchee Seamount, Corner Rise Seamounts; J3-1, KEL 604-2, Kelvin Seamount, New England Seamounts; J3-2, KUK 218-1, Kükenthal Peak; I4, BAL 205-1, polyp cleared with clove oil, Balanus Seamount, New England seamounts; F1, KEL 104-1, Kelvin Seamount; H1, Keratoisis magnifica, TMAG K3830, Tasmania Seamounts; C1, BAL 111-1, Balanus Seamount; G1, TMAG K3856, Tasmanian Seamounts; D1, MAN 707-2, Manning Seamount, New England Seamounts; D2, Eknomisis dalioi, KEL 406-1, Kelvin Seamount; B1-1, PIC 102-1, Picket Seamount, New England Seamounts; B1-2, J2098-6-4, Aleutian Islands slope, Alaska, USA." figureDoi="http://doi.org/10.5281/zenodo.5909960" httpUri="https://zenodo.org/record/5909960/files/figure.png" pageId="11" pageNumber="348">3</figureCitation>
. Colony form ranges from simple fans with sparse branching to complex fans with multiple branches, but in the NW Atlantic at least, colonies can also take on the form perhaps best described as brambles, forming low-growing tangled arrays of mostly thin branches spreading over wide areas of the seafloor. Subclade
<emphasis id="34F7EA958462FFE35F26FF35FCE3FF40" bold="true" box="[849,877,186,212]" pageId="12" pageNumber="349">J3</emphasis>
colonies may have the following characters: axis hollow, sometimes thick, solid near the colony base; branches almost always present, may originate at the nodes, immediately distal to the nodes, or, in the bramble forms, at the internodes; polyp sclerites are spindles, flat rods and scales (occasionally with a fluted margin), needles and long rods; on contraction the polyp shortens slightly, and the tentacles fold over the mouth; polyps are often covered with a layer of tissue containing many cnidae.
</paragraph>
<paragraph id="063C36878462FFE35CB0FEE0FCFFFD3D" blockId="12.[151,1437,151,1869]" pageId="12" pageNumber="349">
Subclade
<emphasis id="34F7EA958462FFE35D43FEE1FEC3FE1C" bold="true" box="[308,333,366,392]" pageId="12" pageNumber="349">I4</emphasis>
is perhaps better referred to as “lineage
<emphasis id="34F7EA958462FFE35F78FEE1FCA9FE1C" bold="true" box="[783,807,366,392]" pageId="12" pageNumber="349">I4</emphasis>
”, being comprised of only a single haplotype, which we detected in
<specimenCount id="1085FD0E8462FFE35D60FE1CFE20FE39" box="[279,430,403,429]" pageId="12" pageNumber="349" type="generic">10 specimens</specimenCount>
(Supplemental
<tableCitation id="4B01033C8462FFE35E2BFE1CFD31FE39" box="[604,703,403,429]" captionStart="TABLE 1" captionStartId="3.[151,239,463,489]" captionTargetPageId="3" captionText="TABLE 1. Primers used in successful PCR and cycle sequencing. Most primer combinations were selected to cross a gene boundary, shown in the first two columns. Primers" httpUri="http://table.plazi.org/id/52FC660F846DFFEC5CE0FE40FE8CFDA5" pageId="12" pageNumber="349" tableUuid="52FC660F846DFFEC5CE0FE40FE8CFDA5">Table S1</tableCitation>
). It was assigned a subclade name (first published in
<bibRefCitation id="62124B768462FFE3597AFE1CFE9EFE45" author="van der Ham, J. L. &amp; Brugler, M. R. &amp; France, S. C." pageId="12" pageNumber="349" pagination="183 - 192" refId="ref14126" refString="van der Ham, J. L., Brugler, M. R. &amp; France, S. C. (2009) Exploring the utility of an indel-rich mitochondrial region as a molecular barcode for bamboo corals (Octocorallia: Isididae). Marine Genomics, 2, 183 - 192. https: // doi. org / 10.1016 / j. margen. 2009.10.002" type="journal article" year="2009">
van der Ham
<emphasis id="34F7EA958462FFE35CE0FE38FF46FE45" box="[151,200,439,465]" italics="true" pageId="12" pageNumber="349">et al</emphasis>
. 2009
</bibRefCitation>
) because the haplotype has a unique pattern of indels in
<emphasis id="34F7EA958462FFE35FFDFE38FC6FFE45" box="[906,993,439,465]" italics="true" pageId="12" pageNumber="349">mtMutS</emphasis>
and
<emphasis id="34F7EA958462FFE3586FFE38FBC9FE45" box="[1048,1095,439,465]" italics="true" pageId="12" pageNumber="349">igr4</emphasis>
relative to other keratoisidids. In our analysis, lineage
<emphasis id="34F7EA958462FFE35DD2FE55FE30FE60" bold="true" box="[421,446,474,500]" pageId="12" pageNumber="349">I4</emphasis>
arises from a polytomy with subclade
<emphasis id="34F7EA958462FFE35F02FE55FC1CFE60" bold="true" box="[885,914,474,500]" pageId="12" pageNumber="349">J3</emphasis>
and clade
<emphasis id="34F7EA958462FFE3587BFE54FB94FE61" bold="true" box="[1036,1050,475,501]" pageId="12" pageNumber="349">5</emphasis>
.
<emphasis id="34F7EA958462FFE3585EFE55FBCCFE60" bold="true" box="[1065,1090,474,500]" pageId="12" pageNumber="349">I4</emphasis>
colony form is in the shape of a candelabra arising from a moderately elongate stalk, e.g. no branching in the lower part of the colony; the axis is solid in the lower part of the colony but hollow in the upper reaches of the branches; branching above the stalk begins as a trident, additional branches being added laterally in pairs in one plane; polyps are tall, sclerites are all rods, mostly needle-like, and sclerites are generally absent from the basal part; due to lack of proximal sclerites polyps can shorten on contraction with tentacles folding over the mouth.
</paragraph>
<paragraph id="063C36878462FFE35CB0FD3CFD6EF8D9" blockId="12.[151,1437,151,1869]" pageId="12" pageNumber="349">
Clade
<emphasis id="34F7EA958462FFE35D66FD3CFE91FD59" bold="true" box="[273,287,691,717]" pageId="12" pageNumber="349">5</emphasis>
(containing subclades
<emphasis id="34F7EA958462FFE35E52FD3DFDCAFD58" bold="true" box="[549,580,690,716]" pageId="12" pageNumber="349">F1</emphasis>
to
<emphasis id="34F7EA958462FFE35E1CFD3CFD02FD58" bold="true" box="[619,652,690,717]" pageId="12" pageNumber="349">B1</emphasis>
) encompasses the greatest number of haplotypes and includes specimens identified to the genera
<emphasis id="34F7EA958462FFE35D99FD58FC6DFD64" box="[494,995,727,753]" italics="true" pageId="12" pageNumber="349">
<taxonomicName id="C1834D048462FFE35D99FD58FDF2FD65" authorityName="Wright" authorityYear="1885" box="[494,636,727,753]" class="Anthozoa" family="Isididae" genus="Bathygorgia" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">Bathygorgia</taxonomicName>
,
<taxonomicName id="C1834D048462FFE35EFEFD58FD71FD65" authorityName="Watling &amp; France" authorityYear="2011" box="[649,767,727,753]" class="Anthozoa" family="Isididae" genus="Eknomisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">Eknomisis</taxonomicName>
,
<taxonomicName id="C1834D048462FFE35F7BFD58FCEDFD65" box="[780,867,727,753]" class="Anthozoa" family="Isididae" genus="Isidella" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">Isidella</taxonomicName>
,
<taxonomicName id="C1834D048462FFE35F07FD58FC6DFD64" authorityName="Wright" authorityYear="1869" box="[880,995,727,752]" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">Keratoisis</taxonomicName>
</emphasis>
, and
<taxonomicName id="C1834D048462FFE35852FD58FB04FD65" authorityName="Verrill" authorityYear="1883" box="[1061,1162,727,753]" class="Anthozoa" family="Isididae" genus="Lepidisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958462FFE35852FD58FB04FD65" box="[1061,1162,727,753]" italics="true" pageId="12" pageNumber="349">Lepidisis</emphasis>
</taxonomicName>
. Specimens assigned to the genera
<taxonomicName id="C1834D048462FFE35D61FD74FEE4FC81" box="[278,362,763,789]" class="Anthozoa" family="Isididae" genus="Isidella" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958462FFE35D61FD74FEE4FC81" box="[278,362,763,789]" italics="true" pageId="12" pageNumber="349">Isidella</emphasis>
</taxonomicName>
and
<taxonomicName id="C1834D048462FFE35DD2FD74FD84FC81" authorityName="Verrill" authorityYear="1883" box="[421,522,763,789]" class="Anthozoa" family="Isididae" genus="Lepidisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958462FFE35DD2FD74FD84FC81" box="[421,522,763,789]" italics="true" pageId="12" pageNumber="349">Lepidisis</emphasis>
</taxonomicName>
are almost certainly incorrectly identified and are likely undescribed species of
<taxonomicName id="C1834D048462FFE35CE0FC90FE84FCAC" authorityName="Wright" authorityYear="1869" box="[151,266,799,824]" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958462FFE35CE0FC90FE84FCAC" box="[151,266,799,824]" italics="true" pageId="12" pageNumber="349">Keratoisis</emphasis>
</taxonomicName>
,
<taxonomicName id="C1834D048462FFE35D6EFC90FE02FCAD" authorityName="Watling &amp; France" authorityYear="2011" box="[281,396,799,825]" class="Anthozoa" family="Isididae" genus="Eknomisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958462FFE35D6EFC90FE02FCAD" box="[281,396,799,825]" italics="true" pageId="12" pageNumber="349">Eknomisis</emphasis>
</taxonomicName>
, or
<taxonomicName id="C1834D048462FFE35DCEFC90FDCAFCAD" authorityName="Wright" authorityYear="1885" box="[441,580,799,825]" class="Anthozoa" family="Isididae" genus="Bathygorgia" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958462FFE35DCEFC90FDCAFCAD" box="[441,580,799,825]" italics="true" pageId="12" pageNumber="349">Bathygorgia</emphasis>
</taxonomicName>
, or an undescribed genus. Clade
<emphasis id="34F7EA958462FFE35FCDFC90FC46FCAD" bold="true" box="[954,968,799,825]" pageId="12" pageNumber="349">5</emphasis>
includes 7 smaller clades, although some are only weakly supported, each with
<emphasis id="34F7EA958462FFE35E3CFCCCFD2CFCC9" box="[587,674,835,861]" italics="true" pageId="12" pageNumber="349">mtMutS</emphasis>
-5 haplotypes of a unique indel structure, e.g. there are 7 different patterns of indel structure within clade
<emphasis id="34F7EA958462FFE35E22FCE8FDEDFC15" bold="true" box="[597,611,871,897]" pageId="12" pageNumber="349">5</emphasis>
that are not shared with other named clades of keratoisidids (clades
<emphasis id="34F7EA958462FFE35907FCE9FAF0FC14" bold="true" box="[1392,1406,870,896]" pageId="12" pageNumber="349">3</emphasis>
&amp;
<emphasis id="34F7EA958462FFE35CE0FC05FF2BFC30" bold="true" box="[151,165,906,932]" pageId="12" pageNumber="349">4</emphasis>
). Genetic distances among
<emphasis id="34F7EA958462FFE35DAAFC04FDBAFC31" box="[477,564,907,933]" italics="true" pageId="12" pageNumber="349">mtMutS</emphasis>
-5 sequences within each of the 7 subclades range from p =
<geoCoordinate id="63B750408462FFE358A8FC04FAA2FC31" box="[1247,1324,907,933]" degrees="0.0120" direction="north" orientation="latitude" pageId="12" pageNumber="349" precision="5" value="0.012">0.0120</geoCoordinate>
<geoCoordinate id="63B750408462FFE35944FC04FA17FC31" box="[1331,1433,907,933]" degrees="0.0948" direction="west" orientation="longitude" pageId="12" pageNumber="349" precision="5" value="-0.0948"> 0.0948</geoCoordinate>
. Included in clade
<emphasis id="34F7EA958462FFE35D16FC20FEE1FC5D" bold="true" box="[353,367,943,969]" pageId="12" pageNumber="349">5</emphasis>
are the two species of the newly resurrected genus
<taxonomicName id="C1834D048462FFE35FC1FC20FBCFFC5D" authorityName="Wright" authorityYear="1885" box="[950,1089,943,969]" class="Anthozoa" family="Isididae" genus="Bathygorgia" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958462FFE35FC1FC20FBCFFC5D" box="[950,1089,943,969]" italics="true" pageId="12" pageNumber="349">Bathygorgia</emphasis>
</taxonomicName>
, although these did not cluster together and showed a high level of divergence from each other (p = 0.10189 over the first half of
<emphasis id="34F7EA958462FFE35883FC5CFAC5FC79" box="[1268,1355,979,1005]" italics="true" pageId="12" pageNumber="349">mtMutS</emphasis>
-5, the only region we were able to sequence for
<taxonomicName id="C1834D048462FFE35E00FC78FC83FB85" box="[631,781,1015,1041]" class="Anthozoa" family="Isididae" genus="Bathygorgia" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="species" species="abyssicola">
<emphasis id="34F7EA958462FFE35E00FC78FC83FB85" box="[631,781,1015,1041]" italics="true" pageId="12" pageNumber="349">B. abyssicola</emphasis>
</taxonomicName>
); each species of
<taxonomicName id="C1834D048462FFE35FABFC78FBE9FB85" authorityName="Wright" authorityYear="1885" box="[988,1127,1015,1041]" class="Anthozoa" family="Isididae" genus="Bathygorgia" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958462FFE35FABFC78FBE9FB85" box="[988,1127,1015,1041]" italics="true" pageId="12" pageNumber="349">Bathygorgia</emphasis>
</taxonomicName>
also has a unique
<emphasis id="34F7EA958462FFE3594BFC78FA1DFB85" box="[1340,1427,1015,1041]" italics="true" pageId="12" pageNumber="349">mtMutS</emphasis>
- 5 haplotype with an indel structure not seen in any of the other specimens (n=398) analyzed for this study. The other nominal genera identified within clade
<emphasis id="34F7EA958462FFE35EF8FBB0FD13FBCD" bold="true" box="[655,669,1087,1113]" pageId="12" pageNumber="349">5</emphasis>
for the most part do not form monophyletic groups. The
<typeStatus id="D93888258462FFE3596BFBB0FAF0FBCD" box="[1308,1406,1087,1113]" pageId="12" pageNumber="349" type="holotype">holotype</typeStatus>
of the monotypic genus
<taxonomicName id="C1834D048462FFE35DFAFBECFD8EFBE9" authorityName="Watling &amp; France" authorityYear="2011" box="[397,512,1123,1149]" class="Anthozoa" family="Isididae" genus="Eknomisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958462FFE35DFAFBECFD8EFBE9" box="[397,512,1123,1149]" italics="true" pageId="12" pageNumber="349">Eknomisis</emphasis>
</taxonomicName>
is found nested within a strongly supported subclade (
<emphasis id="34F7EA958462FFE35804FBEDFB18FBE8" bold="true" box="[1139,1174,1122,1148]" pageId="12" pageNumber="349">D2</emphasis>
) dominated by species tentatively assigned to
<taxonomicName id="C1834D048462FFE35DEBFB08FD81FB34" authorityName="Wright" authorityYear="1869" box="[412,527,1159,1184]" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958462FFE35DEBFB08FD81FB34" box="[412,527,1159,1184]" italics="true" pageId="12" pageNumber="349">Keratoisis</emphasis>
</taxonomicName>
. This subclade also contains a haplotype from a specimen identified as
<taxonomicName id="C1834D048462FFE35948FB08FA12FB34" authorityName="Wright" authorityYear="1869" box="[1343,1436,1159,1184]" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="species" species="grayi">
<emphasis id="34F7EA958462FFE35948FB08FA12FB34" box="[1343,1436,1159,1184]" italics="true" pageId="12" pageNumber="349">K. grayi</emphasis>
</taxonomicName>
(an identification influenced by the collection location), which is the type species of
<taxonomicName id="C1834D048462FFE35822FB24FB46FB50" authorityName="Wright" authorityYear="1869" box="[1109,1224,1195,1220]" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958462FFE35822FB24FB46FB50" box="[1109,1224,1195,1220]" italics="true" pageId="12" pageNumber="349">Keratoisis</emphasis>
</taxonomicName>
and for the family
<taxonomicName id="C1834D048462FFE35CE0FB40FEB0FB7D" authorityName="Gray" authorityYear="1870" box="[151,318,1231,1257]" class="Anthozoa" family="Keratoisididae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="family">Keratoisididae</taxonomicName>
. Subclade
<emphasis id="34F7EA958462FFE35DCCFB41FE50FB7C" bold="true" box="[443,478,1230,1256]" pageId="12" pageNumber="349">G1</emphasis>
contains only two haplotypes, representing the
<typeStatus id="D93888258462FFE35870FB40FBE7FB7D" box="[1031,1129,1231,1257]" pageId="12" pageNumber="349" type="holotype">holotype</typeStatus>
specimens of
<taxonomicName id="C1834D048462FFE35966FB40FEA6FA98" class="Anthozoa" family="Isididae" genus="Bathygorgia" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="species" species="tasmaniensis">
<emphasis id="34F7EA958462FFE35966FB40FEA6FA98" italics="true" pageId="12" pageNumber="349">Bathygorgia tasmaniensis</emphasis>
</taxonomicName>
and
<taxonomicName id="C1834D048462FFE35D16FB7CFC9FFA99" authority="(Duenas et al. 2014)" baseAuthorityName="Duenas" baseAuthorityYear="2014" box="[353,785,1267,1293]" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="species" species="peara">
<emphasis id="34F7EA958462FFE35D16FB7CFD93FA98" box="[353,541,1267,1292]" italics="true" pageId="12" pageNumber="349">Keratoisis peara</emphasis>
(
<bibRefCitation id="62124B768462FFE35E59FB7CFC87FA99" author="Duenas, L. F. &amp; Alderslade, P. &amp; Sanchez, J. A." box="[558,777,1267,1293]" pageId="12" pageNumber="349" pagination="15 - 28" refId="ref13800" refString="Duenas, L. F., Alderslade, P. &amp; Sanchez, J. A. (2014) Molecular systematics of the deep-sea bamboo corals (Octocorallia: Isididae: Keratoisidinae) from New Zealand with descriptions of two new species of Keratoisis. Molecular Phylogenetics and Evolution, 74, 15 - 28. https: // doi. org / 10.1016 / j. ympev. 2014.01.031" type="journal article" year="2014">Dueñas et al. 2014</bibRefCitation>
)
</taxonomicName>
. Subclade
<emphasis id="34F7EA958462FFE35FFBFB7DFC21FA98" bold="true" box="[908,943,1266,1292]" pageId="12" pageNumber="349">G1</emphasis>
is only weakly supported and the two haplotypes are divergent (as shown by the relatively long branch to
<taxonomicName id="C1834D048462FFE35F10FA98FB99FAA4" box="[871,1047,1303,1328]" class="Anthozoa" family="Isididae" genus="Bathygorgia" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="species" species="tasmaniensis">
<emphasis id="34F7EA958462FFE35F10FA98FB99FAA4" box="[871,1047,1303,1328]" italics="true" pageId="12" pageNumber="349">B. tasmaniensis</emphasis>
</taxonomicName>
) and also differ by a 9bp (3 amino acid) insertion in
<taxonomicName id="C1834D048462FFE35D2BFAB4FE33FAC0" box="[348,445,1339,1364]" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="species" species="peara">
<emphasis id="34F7EA958462FFE35D2BFAB4FE33FAC0" box="[348,445,1339,1364]" italics="true" pageId="12" pageNumber="349">K. peara</emphasis>
</taxonomicName>
. Subclade
<emphasis id="34F7EA958462FFE35E4EFAB4FDD4FAC0" bold="true" box="[569,602,1338,1365]" pageId="12" pageNumber="349">B1</emphasis>
encompasses 15 different haplotypes that share the same
<emphasis id="34F7EA958462FFE35896FAB4FAC6FAC0" box="[1249,1352,1339,1365]" italics="true" pageId="12" pageNumber="349">mtMutS -</emphasis>
5 indel structure and are only weakly divergent from each other. Within Clade
<emphasis id="34F7EA958462FFE35FCCFAD0FC47FAED" bold="true" box="[955,969,1375,1401]" pageId="12" pageNumber="349">5</emphasis>
there is a wide diversity of colony types, from unbranched whips, to sparsely branched colonies, to large bushes with multiple branches. Branches (when present) are consistently internodal. Most of the haplotypes in subclade
<emphasis id="34F7EA958462FFE35FB0FA28FC69FA54" bold="true" box="[967,999,1446,1473]" pageId="12" pageNumber="349">B1</emphasis>
come from colonies that are generally whip-like (unbranched), although one haplotype from the eastern North Pacific is associated with bushes or planar fans and two others from the central and south Pacific are branched. Conversely, most of the haplotype diversity within subclade
<emphasis id="34F7EA958462FFE35D25F99DFE20F9B8" bold="true" box="[338,430,1554,1580]" pageId="12" pageNumber="349">D1&amp;D2</emphasis>
, is associated with colonies that are branching bushes or fans, with only a couple of unbranched forms in subclade
<emphasis id="34F7EA958462FFE35DB8F9B9FE7FF9C4" bold="true" box="[463,497,1590,1616]" pageId="12" pageNumber="349">D1</emphasis>
. Subclade
<emphasis id="34F7EA958462FFE35E1CF9B9FD00F9C4" bold="true" box="[619,654,1590,1616]" pageId="12" pageNumber="349">C1</emphasis>
is the only clade with multiple haplotypes to comprise exclusively unbranched colonies,
<taxonomicName id="C1834D048462FFE35D06F9D4FD7CF9E1" authority="(Nutting, 1910)" baseAuthorityName="Nutting" baseAuthorityYear="1910" box="[369,754,1627,1653]" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="species" species="wrighti">
<emphasis id="34F7EA958462FFE35D06F9D4FDB2F9E1" box="[369,572,1627,1653]" italics="true" pageId="12" pageNumber="349">Keratoisis wrighti</emphasis>
(Nutting, 1910)
</taxonomicName>
and
<taxonomicName id="C1834D048462FFE35F5EF9D4FBCCF9E1" authority="Grant, 1976" authorityName="Grant" authorityYear="1976" box="[809,1090,1627,1653]" class="Anthozoa" family="Isididae" genus="Keratoisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="species" species="zelandica">
<emphasis id="34F7EA958462FFE35F5EF9D4FC3BF9E1" box="[809,949,1627,1653]" italics="true" pageId="12" pageNumber="349">K. zelandica</emphasis>
<bibRefCitation id="62124B768462FFE35FCBF9D4FBCCF9E1" author="Grant, R." box="[956,1090,1627,1653]" pageId="12" pageNumber="349" refId="ref14089" refString="Grant, R. (1976) The marine fauna of New Zealand: Isididae (Octocorallia: Gorgonacea) from New Zealand and the Antarctic. New Zealand Oceangraphic Institute Memoir, 66,1 - 54." type="book" year="1976">Grant, 1976</bibRefCitation>
</taxonomicName>
having both been described as unbranched. Clade
<emphasis id="34F7EA958462FFE35D18F9F0FEF3F90D" bold="true" box="[367,381,1663,1689]" pageId="12" pageNumber="349">5</emphasis>
polyp sclerites range from rods and needles to flat rods and scales, usually with intertentacular (mesenterial) rods or needles. The presence of scales, or the absence of sclerites, in the proximal part of the polyp distinguishes most members of subclade
<emphasis id="34F7EA958462FFE35E29F948FDF1F974" bold="true" box="[606,639,1734,1761]" pageId="12" pageNumber="349">B1</emphasis>
from the other subclades. Polyps are generally tall and thin with little to no contraction, the tentacles folding over the mouth, but in
<emphasis id="34F7EA958462FFE35F5BF965FC07F890" bold="true" box="[812,905,1770,1796]" pageId="12" pageNumber="349">D1&amp;D2</emphasis>
the genus
<taxonomicName id="C1834D048462FFE35877F964FBFDF891" authorityName="Watling &amp; France" authorityYear="2011" box="[1024,1139,1771,1797]" class="Anthozoa" family="Isididae" genus="Eknomisis" kingdom="Animalia" order="Alcyonacea" pageId="12" pageNumber="349" phylum="Cnidaria" rank="genus">
<emphasis id="34F7EA958462FFE35877F964FBFDF891" box="[1024,1139,1771,1797]" italics="true" pageId="12" pageNumber="349">Eknomisis</emphasis>
</taxonomicName>
is reminiscent of subclade
<emphasis id="34F7EA958462FFE35CE0F881FF3BF8BC" bold="true" box="[151,181,1806,1832]" pageId="12" pageNumber="349">S1</emphasis>
, with the tentacles folding deep into the oral area, and intertentacular needles are absent, a form that is also seen in some specimens of subclades
<emphasis id="34F7EA958462FFE35E73F8BDFDA6F8D8" bold="true" box="[516,552,1842,1868]" pageId="12" pageNumber="349">H1</emphasis>
and
<emphasis id="34F7EA958462FFE35E29F8BCFDF1F8D8" bold="true" box="[606,639,1842,1869]" pageId="12" pageNumber="349">B1</emphasis>
(
<figureCitation id="9EB82A028462FFE35EF9F8BCFD5CF8D9" box="[654,722,1843,1869]" captionStart="FIGURE 3" captionStartId="22.[151,250,1637,1661]" captionTargetBox="[346,1241,181,1614]" captionTargetId="figure-21@22.[346,1241,181,1614]" captionTargetPageId="22" captionText="FIGURE 3. A selection of polyp types from the Family Keratoisididae. Clade names as noted in the text, followed by species name (if known), specimen number, special treatment (if any), and location. All polyps are less than 1 cm in length or height and are in contracted state due to preservation. S1, Cladarisis nouvianae, CAT 207-1, Cat Island, Bahamas; M1, Orstomisis sp., NMHN IK-2008-14, polyp cleared with clove oil, New Caledonia; I1, Lepidisis caryophyllia, GIL 101-1, SEM view with external tissue removed, Gilbert Canyon, northeast USA; A1, Acanella arbuscula, KUK 103-2, Kükenthal Peak, Caloosahatchee Seamount, Corner Rise Seamounts; J3-1, KEL 604-2, Kelvin Seamount, New England Seamounts; J3-2, KUK 218-1, Kükenthal Peak; I4, BAL 205-1, polyp cleared with clove oil, Balanus Seamount, New England seamounts; F1, KEL 104-1, Kelvin Seamount; H1, Keratoisis magnifica, TMAG K3830, Tasmania Seamounts; C1, BAL 111-1, Balanus Seamount; G1, TMAG K3856, Tasmanian Seamounts; D1, MAN 707-2, Manning Seamount, New England Seamounts; D2, Eknomisis dalioi, KEL 406-1, Kelvin Seamount; B1-1, PIC 102-1, Picket Seamount, New England Seamounts; B1-2, J2098-6-4, Aleutian Islands slope, Alaska, USA." figureDoi="http://doi.org/10.5281/zenodo.5909960" httpUri="https://zenodo.org/record/5909960/files/figure.png" pageId="12" pageNumber="349">Fig. 3</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>