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<mods:title id="68B162B4E1311924166D88E5940B1FF4">A review of gorgonian coral species (Cnidaria, Octocorallia, Alcyonacea) held in the Santa Barbara Museum of Natural History research collection: focus on species from Scleraxonia, Holaxonia, and Calcaxonia - Part I: Introduction, species of Scleraxonia and Holaxonia (Family Acanthogorgiidae)</mods:title>
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<subSubSection id="184DFCED0053993568FA4816E0ABFD90" pageId="0" pageNumber="1" type="nomenclature">
<paragraph id="509A82D21223E613E40DE4AD5C93EBC8" pageId="0" pageNumber="1">
<taxonomicName id="1F3288EE3F47E12BA17FCBBB89F85877" ID-CoL="6LPM6" authority="Bayer, 1956" authorityName="Bayer" authorityYear="1956" class="Anthozoa" family="Coralliidae" genus="Hemicorallium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hemicorallium regale" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="species" species="regale">Hemicorallium regale (Bayer, 1956)</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="D051009BCE7E6CB039B6227ABAF0931A" pageId="0" pageNumber="1" type="reference_group">
<paragraph id="A9DBC13A2D29910E1B60FF811AACF220" pageId="0" pageNumber="1">
<taxonomicName id="4778297135ADA74E930DF15732E5FE91" class="Anthozoa" family="Coralliidae" genus="Corallium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Corallium regale" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="species" species="regale">Corallium regale</taxonomicName>
Bayer, 1956b: 70, 73-76; 77-78; figs 5c; 7 
<normalizedToken id="6EACA29AC8B5B5622C9D65F3D5CF56C7" originalValue="e–g">e-g</normalizedToken>
. 
<bibRefCitation id="509F421C91BB65F84A4E7953C6CB978A" author="Bayer, FM" journalOrPublisher="Proceedings of the Biological Society of Washington" pageId="0" pageNumber="1" pagination="222 - 228" title="A new genus of the scleraxonian family Coralliidae (Octocorallia: Gorgonacea)." url="https://repository.si.edu/handle/10088/1240" volume="116" year="2003">Bayer and Cairns 2003</bibRefCitation>
: 224.
</paragraph>
</subSubSection>
<subSubSection id="1E01DABE88C385D75C2295E3601E0E3A" pageId="0" pageNumber="1" type="synonyms">
<paragraph id="2FED70FED3ACF8F2C9CE00553941DCE1" pageId="0" pageNumber="1">Synonyms.</paragraph>
<paragraph id="3FE79144046F62DFE2A565593E5FC71A" pageId="0" pageNumber="1">(see Remarks section below.)</paragraph>
</subSubSection>
<subSubSection id="0F9A5FFB622A2114E10F5757312DAD29" pageId="0" pageNumber="1" type="materials_examined">
<paragraph id="B4F8CFA7333F0CB86A4905FE7ECF345E" pageId="0" pageNumber="1">Material examined.</paragraph>
<paragraph id="5FD2D62D153C89D2B41E357A98038557" pageId="0" pageNumber="1">No material in SBMNH collection (see Appendix 1: List of material examined).</paragraph>
</subSubSection>
<subSubSection id="01FCB2305B8404DDD9B1A645E4896129" pageId="0" pageNumber="1" type="remarks">
<paragraph id="82F4919CADC03FC428405EA4AADA0E68" pageId="0" pageNumber="1">Remarks.</paragraph>
<paragraph id="680A2A535F0E0F3C6CF386C48516DF17" pageId="0" pageNumber="1">
Around the Pacific, from Hawaii (holotype, USNM 49520) to offshore seamounts some miles west of California coast (outer edge of Bight western boundary, USNM 94460), certainly at substantial depth (based on specimens housed in NMNH collection). Not enough specimens examined (or collected with attention to specific collection locations) to determine extent of north-south range. Bayer commented (1956b), &quot;of all the Hawaiian precious corals, 
<taxonomicName id="1A0B15FA36FC69898CA9C87E3C41507D" lsidName="C. regale" pageId="0" pageNumber="1" rank="species" species="regale">C. regale</taxonomicName>
has the best color and might be of commercial value if it could be fished in quantity.&quot; Thin, calcareous extensions of axis extending outward to thick coenenchyme can support expansion of coenenchyme near sides of branches as recurved flaps, a distinctive feature. These can form tunnels inhabited by polychaete commensals (
<bibRefCitation id="5FD6A7977616500172CFC78517D1ABDB" pageId="0" pageNumber="1">Baco and Shirley 2005</bibRefCitation>
, 
<bibRefCitation id="93317C2FDD28B975AFE62AC852233DC7" author="Baco, AR" journalOrPublisher="Oceanography" pageId="0" pageNumber="1" pagination="108 - 117" title="Exploration for Deep-Sea Corals on North Pacific Seamounts and Islands." url="https://doi.org/10.5670/oceanog.2007.11" volume="20" year="2007">Baco 2007</bibRefCitation>
). Could be given the common name &quot;Regal red coral.&quot;
</paragraph>
<paragraph id="3AD043C94653F801109BC637530DD7E3" pageId="0" pageNumber="1">
Several species in the now updated genus 
<taxonomicName id="6E9C0499A0D18CE99B0392732565BF33" class="Anthozoa" family="Coralliidae" genus="Hemicorallium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hemicorallium" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="genus">Hemicorallium</taxonomicName>
, including this one, found within the Hawaiian archipelago (and elsewhere in the western Pacific, including international waters) have been the focus of commercial exploitation (then recognized as species in the genus 
<taxonomicName id="B09C7F983CC7CF93675B82474F621B54" class="Anthozoa" family="Coralliidae" genus="Corallium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Corallium" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="genus">Corallium</taxonomicName>
). Based on reports made public by CITES, regarding &quot;consideration of proposals for amendment of Appendices I and II&quot; (2007, 2010) and 
<bibRefCitation id="B130A5981AE9EA6CBB2EAEE2FC41800C" author="Bruckner, A" journalOrPublisher="Kungliga Svenska Vetenskapsakademiens Handlingar" pageId="0" pageNumber="1" url="https://www.livingoceansfoundation.org/wpcontent/uploads/2013/08/Bruckner_DiseasePrevalence.pdf" year="2009">Bruckner (2009)</bibRefCitation>
, there was much discussion, and confusion, as to whether this species (using older genus designation) was valid and/or whether it could be synonymous with 
<taxonomicName id="165E142B7AC3ECDC24871AC73D586791" class="Anthozoa" family="Coralliidae" genus="Corallium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Corallium laauense" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="species" species="laauense">Corallium laauense</taxonomicName>
(misspelled as 
<taxonomicName id="392EAA22F9E022198178F4F2E2E817A1" lsidName="C. lauuense" pageId="0" pageNumber="1" rank="species" species="lauuense">C. lauuense</taxonomicName>
). 
<bibRefCitation id="7ECB31F874B5DE971B59A968C6F360BC" author="Grigg, RW" journalOrPublisher="Pacific Science" pageId="0" pageNumber="1" pagination="167 - 175" title="Present knowledge of the systematics and zoogeography of the order Gorgonacea in Hawaii." volume="30" year="1976">Grigg and Bayer (1976</bibRefCitation>
: 169) and 
<bibRefCitation id="2A34AC95CE8A2F1EBF5CF0FC5BD11553" author="Grigg, RW" journalOrPublisher="Marine Fisheries Review" pageId="0" pageNumber="1" pagination="13 - 20" title="Precious corals in Hawaii: discovery of a new bed and revised management measures for existing beds." volume="64" year="2002">Grigg (2002</bibRefCitation>
: 17-19) specifically mentioned 
<taxonomicName id="3E242B31055ADC20B318900662D6E7AE" lsidName="C. regale" pageId="0" pageNumber="1" rank="species" species="regale">C. regale</taxonomicName>
and/or 
<taxonomicName id="6A84872D0A63C428A141B4B99EE6D60C" lsidName="C. laauense" pageId="0" pageNumber="1" rank="species" species="laauense">C. laauense</taxonomicName>
as separate species. CoP14 Prop. 21 (CITES, 2007) listed this species as a potential synonym for 
<taxonomicName id="31CFEAE417C9BD90D174EB6CE02F888B" lsidName="C. laauense" pageId="0" pageNumber="1" rank="species" species="laauense">C. laauense</taxonomicName>
; this based on 
<bibRefCitation id="B86A554AAAC008B09DB621AAD115AB45" author="Baco, AR" editor="Freiwald, A" journalOrPublisher="Oceanography" pageId="0" pageNumber="1" title="Springer-Verlag, Berlin-Heidelberg" url="https://doi.org/10.1007/3-540-27673-4_33" year="2005">Baco and Shank (2005</bibRefCitation>
: 664). However, 
<bibRefCitation id="637E8AA4011F667ACAADE158258AAE1A" author="Baco, AR" editor="Freiwald, A" journalOrPublisher="Oceanography" pageId="0" pageNumber="1" title="Springer-Verlag, Berlin-Heidelberg" url="https://doi.org/10.1007/3-540-27673-4_33" year="2005">Baco and Shank (2005)</bibRefCitation>
did not treat this species as a synonym of 
<taxonomicName id="B4BD9E902E53B00D6EC556C16BE69174" lsidName="C. lauuense" pageId="0" pageNumber="1" rank="species" species="lauuense">C. lauuense</taxonomicName>
(note the misspelling), but a comment they made regarding the work of 
<bibRefCitation id="A55986F19E901499A631EE579D21E272" author="Grigg, RW" journalOrPublisher="Marine Fisheries Review" pageId="0" pageNumber="1" pagination="13 - 20" title="Precious corals in Hawaii: discovery of a new bed and revised management measures for existing beds." volume="64" year="2002">Grigg (2002)</bibRefCitation>
may have inadvertently lead some to assume that was the case. Cairns (CITES, 2007) did not consider these two taxa to be synonymous and 
<bibRefCitation id="68A0F96B30EA9010520E25C30D135FEA" author="Bruckner, A" journalOrPublisher="Kungliga Svenska Vetenskapsakademiens Handlingar" pageId="0" pageNumber="1" url="https://www.livingoceansfoundation.org/wpcontent/uploads/2013/08/Bruckner_DiseasePrevalence.pdf" year="2009">Bruckner (2009</bibRefCitation>
: 321) also discussed these taxa as two separate species. Bruckner indicated that although he discussed them as two separate species, &quot;these 2 species may be synonymous.&quot; In the document, Proposal, CoP15 Prop. 21 (CITES 2010), 
<taxonomicName id="1BFC5E44D22B36F09043AA722F88DA04" lsidName="C. regale" pageId="0" pageNumber="1" rank="species" species="regale">C. regale</taxonomicName>
was still shown as a synonym of 
<taxonomicName id="BC3DF908142E22E5D52FBE68FB71417A" lsidName="C. lauuense" pageId="0" pageNumber="1" rank="species" species="lauuense">C. lauuense</taxonomicName>
(sic); note the following statement: &quot; 
<taxonomicName id="6E36C920D50A49EB88D642FE6B838561" lsidName="C. lauuense" pageId="0" pageNumber="1" rank="species" species="lauuense">C. lauuense</taxonomicName>
and 
<taxonomicName id="6C86ABDD029A35A3907F6402E6EB23DC" lsidName="C. regale" pageId="0" pageNumber="1" rank="species" species="regale">C. regale</taxonomicName>
are listed as separate species in the US Precious Coral Fishery Management plan, but these species are usually considered synonymous (
<bibRefCitation id="CF91165E9D7D5C13F09AA7360584E8F2" author="Parrish, FA" editor="George, RY" journalOrPublisher="[pages 1271 - 1312] In: Rossi S, Bramanti L, Gori A, Orejas C (Eds) The Ecology of Benthic Biodiversity Hotspots. Springer International Publishing" pageId="0" pageNumber="1" title="Bulletin of Marine Science 81 (Supplement 1)" url="http://www.pifsc.noaa.gov/library/pubs/Parrish_5477_2007.pdf" year="2007">Parrish 2007</bibRefCitation>
)&quot;. Again, 
<taxonomicName id="009D55ED59E7B57948B33BE6B0F655B5" lsidName="C. laauense" pageId="0" pageNumber="1" rank="species" species="laauense">C. laauense</taxonomicName>
and 
<taxonomicName id="A73B1915437C1DF702ED2CA0D9BE8513" lsidName="C. regale" pageId="0" pageNumber="1" rank="species" species="regale">C. regale</taxonomicName>
were treated as synonymous; however, regarding 
<taxonomicName id="FFEDE79D2C34C2931B367403F3677121" lsidName="C. regale" pageId="0" pageNumber="1" rank="species" species="regale">C. regale</taxonomicName>
, 
<bibRefCitation id="F56C286174A898044DB99BBE459A744A" author="Baco, AR" editor="Freiwald, A" journalOrPublisher="Oceanography" pageId="0" pageNumber="1" title="Springer-Verlag, Berlin-Heidelberg" url="https://doi.org/10.1007/3-540-27673-4_33" year="2005">Baco and Shank (2005)</bibRefCitation>
stated: &quot; 
<taxonomicName id="B5003604A17F9F0FDF3761D36900E43A" lsidName="C. lauuense" pageId="0" pageNumber="1" rank="species" species="lauuense">C. lauuense</taxonomicName>
was previously misidentified and referred to as 
<taxonomicName id="4A909539BD32DB5F07BB9281ABFBBD92" lsidName="C. regale" pageId="0" pageNumber="1" rank="species" species="regale">C. regale</taxonomicName>
which is not an indication of synonymy. There may still be some unresolved taxonomic problems concerning these two species.&quot; Additionally, &quot; 
<bibRefCitation id="3295D4FFD915BDE00687056486B7BCB4" author="Bayer, FM" journalOrPublisher="Proceedings of the Biological Society of Washington" pageId="0" pageNumber="1" pagination="222 - 228" title="A new genus of the scleraxonian family Coralliidae (Octocorallia: Gorgonacea)." url="https://repository.si.edu/handle/10088/1240" volume="116" year="2003">Bayer and Cairns (2003)</bibRefCitation>
differs from the SS species list in a number of ways: 
<taxonomicName id="D7B3BFE9415FC4F27F56507456DCF859" lsidName="C. boshuense" pageId="0" pageNumber="1" rank="species" species="boshuense">C. boshuense</taxonomicName>
, 
<taxonomicName id="11008D75EA4AFBFB408B444DACAB4DDA" lsidName="C. niveum" pageId="0" pageNumber="1" rank="species" species="niveum">C. niveum</taxonomicName>
, 
<taxonomicName id="6B3ED86B00ACF994F6C82CBFEF421AEC" lsidName="C. porcellanum" pageId="0" pageNumber="1" rank="species" species="porcellanum">C. porcellanum</taxonomicName>
, 
<taxonomicName id="1756515C02AE7F804B5BD5D3EFE65625" lsidName="C. pusillum" pageId="0" pageNumber="1" rank="species" species="pusillum">C. pusillum</taxonomicName>
, 
<taxonomicName id="E21DD829CD16BBC84D1D35A44C5F6A38" lsidName="C. vanderbilti" pageId="0" pageNumber="1" rank="species" species="vanderbilti">C. vanderbilti</taxonomicName>
, and 
<taxonomicName id="AD54436742D4951EC43FAAF817A976F4" lsidName="C. variabile" pageId="0" pageNumber="1" rank="species" species="variabile">C. variabile</taxonomicName>
are not mentioned; 
<taxonomicName id="C2926E4408B1A206386F4EDD774B5182" lsidName="C. regale" pageId="0" pageNumber="1" rank="species" species="regale">C. regale</taxonomicName>
is treated as valid.&quot; (CITES 2010); of these species, 
<taxonomicName id="831047F5082A35CB8C74744B48580EE9" lsidName="C. boshuense" pageId="0" pageNumber="1" rank="species" species="boshuense">C. boshuense</taxonomicName>
and 
<taxonomicName id="3754DFA2AC692E29521AB0C81F0C23D9" lsidName="C. variabile" pageId="0" pageNumber="1" rank="species" species="variabile">C. variabile</taxonomicName>
have been moved to the genus 
<taxonomicName id="01CEA7DC0C626CC739930913341364B3" class="Anthozoa" family="Coralliidae" genus="Hemicorallium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hemicorallium" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="genus">Hemicorallium</taxonomicName>
(
<bibRefCitation id="4099EB65748DA430E230AA73D222CADD" author="Cordeiro, R" journalOrPublisher="American Fisheries Society, Bethesda, Maryland" pageId="0" pageNumber="1" url="http://www.marinespecies.org/aphia.php?p=taxdetails&amp;id=718773" year="2019">Cordeiro et al. 2019</bibRefCitation>
) in the WoRMS Database. The final implication was that 
<taxonomicName id="37476C92751B27723F321CCE126E70DD" class="Anthozoa" family="Coralliidae" genus="Corallium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Corallium" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="genus">Corallium</taxonomicName>
sp. was found throughout &quot;the Hawaiian archipelago and into the Emperor Seamount Chain&quot; (
<bibRefCitation id="C61D7BC08006FD9AE5DF1D878158CA0F" author="Baco, AR" journalOrPublisher="Oceanography" pageId="0" pageNumber="1" pagination="108 - 117" title="Exploration for Deep-Sea Corals on North Pacific Seamounts and Islands." url="https://doi.org/10.5670/oceanog.2007.11" volume="20" year="2007">Baco 2007</bibRefCitation>
), but that the certainty of species identification was still in question.
</paragraph>
<paragraph id="A03FB68E070B7A1A7A8B6C3419065455" pageId="0" pageNumber="1">
In studying 
<normalizedToken id="CF0B952A889D2C3A02F85245EF2446FA" originalValue="Bayer’s">Bayer's</normalizedToken>
original 1956 description of 
<taxonomicName id="F3A1A06D83BBD55A99C1A9DB638EBA90" lsidName="C. laauense" pageId="0" pageNumber="1" rank="species" species="laauense">C. laauense</taxonomicName>
(correctly spelled) and 
<taxonomicName id="B00EABDB715A47799DBBC73DC7588F93" lsidName="C. regale" pageId="0" pageNumber="1" rank="species" species="regale">C. regale</taxonomicName>
, no determination could be made as to why these two were linked as synonymous. The now recognized species 
<taxonomicName id="AE2BD61D09E3CE32813F54B04B8273DE" class="Anthozoa" family="Coralliidae" genus="Hemicorallium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hemicorallium regale" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="species" species="regale">Hemicorallium regale</taxonomicName>
has many double club sclerites, while 
<taxonomicName id="BF5D96E539B267387084D3F52242789E" lsidName="H. laauense" pageId="0" pageNumber="1" rank="species" species="laauense">H. laauense</taxonomicName>
has none. As well, the entire colony of 
<taxonomicName id="E0F172728C4E79A62586EA06E35C9663" lsidName="H. regale" pageId="0" pageNumber="1" rank="species" species="regale">H. regale</taxonomicName>
, as well as the axis, is pink, while the colony color of 
<taxonomicName id="8A3417D0E28A0EB99BF1F7857E760D76" lsidName="H. laauense" pageId="0" pageNumber="1" rank="species" species="laauense">H. laauense</taxonomicName>
is white or very faintly pink with a white axis. SEM images are on file (
<normalizedToken id="2FAEC679366BB73706937BD47B85FF23" originalValue="Bayer’s">Bayer's</normalizedToken>
personal collection), SEM #2283, USNM 94460.
</paragraph>
<paragraph id="B67477933ED187559960B62D63FF9771" pageId="0" pageNumber="1">
The work by 
<bibRefCitation id="AA1F70BD334216608D3DD6A825C8F985" author="Tu, T-H" journalOrPublisher="Zootaxa" pageId="0" pageNumber="1" pagination="301 - 328" title="Coralliidae (Anthozoa: Octocorallia) from the INDEMARES 2010 expedition to north and northwest Spain (northeast Atlantic), with delimitation of a new species using both morphological and molecular approaches." url="https://doi.org/10.11646/zootaxa.3926.3.1" volume="3926" year="2015 a">Tu et al. (2015a</bibRefCitation>
, 
<bibRefCitation id="3EA22A1065B523E7C39CD4ED6EEE0F58" author="Tu, T-H" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="1" pagination="173 - 184" title="Phylogeny and systematics of deep-sea precious corals (Anthozoa: Octocorallia: Coralliidae)." url="https://doi.org/10.1016/j.ympev.2014.09.031" volume="84" year="2015 b">b</bibRefCitation>
; 
<bibRefCitation id="ADE10E20DA98B94C6D09765177CDB51B" author="Tu, T-H" journalOrPublisher="Marine Biology Research" pageId="0" pageNumber="1" pagination="1003 - 1038" title="Taxonomic revision of Corallidae with descriptions of new species from New Caledonia and the Hawaiian Archipelago." url="https://doi.org/10.1080/17451000.2016.1241411" volume="12" year="2016">2016</bibRefCitation>
) resulted in most species of 
<taxonomicName id="E81B8A91D467DFF19D39C90220F2D4B7" class="Anthozoa" family="Coralliidae" genus="Corallium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Corallium" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="genus">Corallium</taxonomicName>
being placed in the genus 
<taxonomicName id="D05ADC6FF7EC9F6489CBD758A04EA387" class="Anthozoa" family="Coralliidae" genus="Hemicorallium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Hemicorallium" order="Alcyonacea" pageId="0" pageNumber="1" phylum="Cnidaria" rank="genus">Hemicorallium</taxonomicName>
. WoRMS (
<bibRefCitation id="41B7EE6B0435A4C3F6A456FA50DF0016" author="Cordeiro, R" journalOrPublisher="American Fisheries Society, Bethesda, Maryland" pageId="0" pageNumber="1" url="http://www.marinespecies.org/aphia.php?p=taxdetails&amp;id=718773" year="2019">Cordeiro et al. 2019</bibRefCitation>
) confirms the placement of the species discussed here: 
<taxonomicName id="E514CA56222D072ADD9F1639056BC37B" lsidName="H. ducale" pageId="0" pageNumber="1" rank="species" species="ducale">H. ducale</taxonomicName>
, 
<taxonomicName id="B22B6D5FD0E3E78009704D0852047E16" lsidName="H. imperiale" pageId="0" pageNumber="1" rank="species" species="imperiale">H. imperiale</taxonomicName>
, 
<taxonomicName id="DF176F119FC44AF8DC503E7A8B17F1BB" lsidName="H. regale" pageId="0" pageNumber="1" rank="species" species="regale">H. regale</taxonomicName>
(as well as 
<taxonomicName id="65010DD08ED1F8E8B51A88724F89F247" lsidName="H. laauense" pageId="0" pageNumber="1" rank="species" species="laauense">H. laauense</taxonomicName>
); all are listed as accepted, separate species.
</paragraph>
</subSubSection>
</treatment>
</document>