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<mods:title id="A24BCDF39DA6A0BCE2ACB0763082260B">The genera and species of Comatulidae (Comatulida: Crinoidea): taxonomic revisions and a molecular and morphological guide</mods:title>
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<subSubSection id="C32D3DEDFFB0A16EFF66DBEF115EFEC7" pageId="14" pageNumber="165" type="nomenclature">
<paragraph id="8B886E66FFB0A16EFF66DBEF11CBFF25" blockId="14.[151,500,222,282]" box="[151,500,222,248]" pageId="14" pageNumber="165">
<heading id="D0C0D90AFFB0A16EFF66DBEF11CBFF25" bold="true" box="[151,500,222,248]" fontSize="11" level="1" pageId="14" pageNumber="165" reason="1">
<emphasis id="B943B274FFB0A16EFF66DBEF11CBFF25" bold="true" box="[151,500,222,248]" pageId="14" pageNumber="165">
<taxonomicName id="4C3715E5FFB0A16EFF66DBEF1124FF25" box="[151,283,222,248]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="genus">
<emphasis id="B943B274FFB0A16EFF66DBEF1124FF25" bold="true" box="[151,283,222,248]" italics="true" pageId="14" pageNumber="165">Capillaster</emphasis>
</taxonomicName>
AH
<bibRefCitation id="EFA61397FFB0A16EFEA4DBEF11CBFF25" author="Clark" box="[341,500,222,248]" pageId="14" pageNumber="165" refString="Clark, A. H. (1909 e) The type of the genus Comaster. Proceedings of the Biological Society of Washington, 22, 87 - 90." type="journal article" year="1909" yearSuffix="e">Clark, 1909e</bibRefCitation>
</emphasis>
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</paragraph>
<paragraph id="8B886E66FFB0A16EFF66DA30115EFEC7" blockId="14.[151,500,222,282]" box="[151,353,257,282]" pageId="14" pageNumber="165">
<tableCitation id="C6B55BDDFFB0A16EFF66DA3010D4FEC4" box="[151,235,257,282]" captionStart="TABLE 1" captionStartId="2.[125,197,537,558]" captionText="TABLE 1. Diagnostic morphological characteristics of genera of Comatulidae." pageId="14" pageNumber="165">Table 1</tableCitation>
;
<figureCitation id="130C72E3FFB0A16EFF09DA30115EFEC7" box="[248,353,257,282]" captionStart-0="FIGURE 5" captionStart-1="FIGURE 6" captionStartId-0="14.[151,250,1813,1835]" captionStartId-1="15.[151,250,1888,1910]" captionTargetBox-0="[151,1436,667,1796]" captionTargetBox-1="[159,1411,207,1867]" captionTargetId-0="figure@14.[151,1436,667,1796]" captionTargetId-1="figure@15.[150,1437,193,1867]" captionTargetPageId-0="14" captionTargetPageId-1="15" captionText-0="FIGURE 5. Capillaster. (A C, F I) Capillaster multiradiatus— (A, B, G I) live, on soft coral (A) and hard coral (B) during the day, and (G H) with centrodorsal hidden and arms exposed at night; (C, I) aboral surface, showing rounded pentagonal centrodorsal and row of cirri; (F) comb with single, triangular teeth confluent with outer edge of pinnular. (D E) Capillaster sentosus— (D) live, exposed for photograph. (E) ray base, preserved, showing rounded pentagonal centrodorsal with one row of cirri. Madang, Papua New Guinea: A. MNHN-IE- 2013 - 8105; B. MNHN-IE- 2013 - 8079; D. MNHN-IE- 2013 - 8066; G, I. MNHN-IE- 2013 - 8066; H. not vouchered. Raja Ampat, Indonesia: C. not vouchered (RA 219); F. SIO-BIC E 6179. Australia or Timor: E. MNHN EcCs 1006." captionText-1="FIGURE 6. Maximum likelihood tree of Capillaster inferred from COI. Symbols and labels as in Figure 2. Location abbreviations—Great Barrier Reef, Australia (GBR); Madang, Papua New Guinea (PNG); Raja Ampat, Indonesia (Raja); Singapore Strait, Singapore (Sing). New COI sequences are Genbank KR 010307 - KR 010329." httpUri-0="https://zenodo.org/record/580178/files/figure.png" httpUri-1="https://zenodo.org/record/580179/files/figure.png" pageId="14" pageNumber="165">Figs. 56</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="C32D3DEDFFB0A16EFF66DA731402FE1A" pageId="14" pageNumber="165" type="materials_examined">
<paragraph id="8B886E66FFB0A16EFF66DA7312DFFE86" blockId="14.[151,1436,322,635]" box="[151,736,322,347]" pageId="14" pageNumber="165">
<emphasis id="B943B274FFB0A16EFF66DA73110DFE86" bold="true" box="[151,306,322,347]" pageId="14" pageNumber="165">Type species.</emphasis>
<taxonomicName id="4C3715E5FFB0A16EFEC9DA7212E4FE86" authority="Carpenter, 1888" authorityName="Carpenter" authorityYear="1888" box="[312,731,323,347]" class="Crinoidea" family="Comatulidae" genus="Actinometra" higherTaxonomySource="CoL" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="species" species="sentosa">
<emphasis id="B943B274FFB0A16EFEC9DA721221FE87" box="[312,542,323,346]" italics="true" pageId="14" pageNumber="165">Actinometra sentosa</emphasis>
<bibRefCitation id="EFA61397FFB0A16EFDD4DA7212E4FE86" author="Carpenter" box="[549,731,323,347]" pageId="14" pageNumber="165" refString="Carpenter, P. H. (1888) Report on the Crinoidea collected during the Voyage of H. M. S. Challenger, during the years 1873 - 1876. Part II-The Comatulae. In: Thomson, C. W. &amp; Murray, J. (Eds.), Report of the Scientific Results of the Voyage of H. M. S. Challenger during the years 1873 - 1876, Zoology-Vol. XXVI. Neill &amp; Company, Edinburgh, pp. 1 - 400." type="book chapter" year="1888">Carpenter, 1888</bibRefCitation>
</taxonomicName>
.
</paragraph>
<paragraph id="8B886E66FFB0A16EFF36DA571402FE1A" blockId="14.[151,1436,322,635]" pageId="14" pageNumber="165">
<emphasis id="B943B274FFB0A16EFF36DA5711D4FEA2" bold="true" box="[199,491,358,383]" pageId="14" pageNumber="165">Other included taxa (7).</emphasis>
<taxonomicName id="4C3715E5FFB0A16EFE05DA5613D9FEA2" authority="Hartlaub, 1890" authorityName="Hartlaub" authorityYear="1890" box="[500,998,358,383]" class="Crinoidea" family="Comatulidae" genus="Actinometra" higherTaxonomySource="CoL" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="species" species="macrobrachius">
<emphasis id="B943B274FFB0A16EFE05DA56130FFEA3" box="[500,816,358,382]" italics="true" pageId="14" pageNumber="165">Actinometra macrobrachius</emphasis>
<bibRefCitation id="EFA61397FFB0A16EFCC8DA5713D9FEA2" author="Hartlaub" box="[825,998,358,383]" pageId="14" pageNumber="165" refString="Hartlaub, C. (1890) Beitrag zur Kenntniss der Comatuliden-Fauna des indischen Archipels. Nachrichten von der Koniglichen Gesellschaft der Wissenschaften und der Georgs-August-Universitat zu Gottingen, 5, 168 - 187." type="journal article" year="1890">Hartlaub, 1890</bibRefCitation>
</taxonomicName>
;
<taxonomicName id="4C3715E5FFB0A16EFC05DA5714EBFEA3" box="[1012,1236,358,382]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="species" species="asterias">
<emphasis id="B943B274FFB0A16EFC05DA5714EBFEA3" box="[1012,1236,358,382]" italics="true" pageId="14" pageNumber="165">Capillaster asterias</emphasis>
</taxonomicName>
AH
<bibRefCitation id="EFA61397FFB0A16EFAFFDA5715A8FEA2" author="Clark" box="[1294,1431,358,383]" pageId="14" pageNumber="165" refString="Clark, A. H. (1931) A monograph of the existing crinoids 1 (3). Bulletin of the United States National Museum, 82, 1 - 916." type="journal article" year="1931">Clark, 1931</bibRefCitation>
;
<taxonomicName id="4C3715E5FFB0A16EFF66DABA11A0FE7E" box="[151,415,395,419]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="species" species="gracilicirra">
<emphasis id="B943B274FFB0A16EFF66DABA11A0FE7E" box="[151,415,395,419]" italics="true" pageId="14" pageNumber="165">Capillaster gracilicirra</emphasis>
</taxonomicName>
AH
<bibRefCitation id="EFA61397FFB0A16EFE26DABB1253FE7E" author="Clark" box="[471,620,394,419]" pageId="14" pageNumber="165" refString="Clark, A. H. (1912 a) Seventeen new East Indian crinoids belonging to the families Comasteridae and Zygometridae. Proceedings of the Biological Society of Washington, 25, 17 - 28." type="journal article" year="1912" yearSuffix="a">Clark, 1912a</bibRefCitation>
;
<taxonomicName id="4C3715E5FFB0A16EFD89DABA141FFE7E" authority="Linnaeus, 1758" authorityName="Linnaeus" authorityYear="1758" box="[632,1056,394,419]" class="Asteroidea" family="Asteriidae" genus="Asterias" kingdom="Animalia" order="Forcipulatida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="species" species="multiradiata">
<emphasis id="B943B274FFB0A16EFD89DABA1357FE7E" box="[632,872,395,419]" italics="true" pageId="14" pageNumber="165">Asterias multiradiata</emphasis>
<bibRefCitation id="EFA61397FFB0A16EFC81DABB141FFE7E" author="Linnaeus" box="[880,1056,394,419]" pageId="14" pageNumber="165" refString="Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata. Laurentius Salvius, Holmiae, 824 pp." type="book" year="1758">Linnaeus, 1758</bibRefCitation>
</taxonomicName>
;
<taxonomicName id="4C3715E5FFB0A16EFBDCDABA10ECFE1A" authority="Messing, 2003" authorityName="Messing" authorityYear="2003" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="species" species="squarrosus">
<emphasis id="B943B274FFB0A16EFBDCDABA1511FE7F" box="[1069,1326,395,419]" italics="true" pageId="14" pageNumber="165">Capillaster squarrosus</emphasis>
Messing, 2003
</taxonomicName>
;
<taxonomicName id="4C3715E5FFB0A16EFF2CDA9F11ECFE1B" box="[221,467,430,454]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="species" species="tenuicirra">
<emphasis id="B943B274FFB0A16EFF2CDA9F11ECFE1B" box="[221,467,430,454]" italics="true" pageId="14" pageNumber="165">Capillaster tenuicirra</emphasis>
</taxonomicName>
AH
<bibRefCitation id="EFA61397FFB0A16EFDF8DA9F12A2FE1A" author="Clark" box="[521,669,430,455]" pageId="14" pageNumber="165" refString="Clark, A. H. (1912 a) Seventeen new East Indian crinoids belonging to the families Comasteridae and Zygometridae. Proceedings of the Biological Society of Washington, 25, 17 - 28." type="journal article" year="1912" yearSuffix="a">Clark, 1912a</bibRefCitation>
;
<taxonomicName id="4C3715E5FFB0A16EFD59DA9F1352FE1B" box="[680,877,430,454]" class="Crinoidea" family="Comasteridae" genus="Comatula" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="species" species="mariae">
<emphasis id="B943B274FFB0A16EFD59DA9F1352FE1B" box="[680,877,430,454]" italics="true" pageId="14" pageNumber="165">Comatula mariae</emphasis>
</taxonomicName>
AH
<bibRefCitation id="EFA61397FFB0A16EFC55DA9F1406FE1A" author="Clark" box="[932,1081,430,455]" pageId="14" pageNumber="165" refString="Clark, A. H. (1907 b) Description of new species of recent unstalked crinoids from the coast of northeastern Asia. Proceedings of the United States National Museum, 33, 127 - 156." type="journal article" year="1907" yearSuffix="b">Clark, 1907b</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C32D3DEDFFB0A171FF36DAE313DFFE0D" lastPageId="17" lastPageNumber="168" pageId="14" pageNumber="165" type="diagnosis">
<paragraph id="8B886E66FFB0A16EFF36DAE3156BFDA6" blockId="14.[151,1436,322,635]" pageId="14" pageNumber="165">
<emphasis id="B943B274FFB0A16EFF36DAE3117EFE36" bold="true" box="[199,321,466,491]" pageId="14" pageNumber="165">Diagnosis.</emphasis>
Mouth excentric in fully developed individuals; up to 110 arms; IBr2 series united by synarthry; first syzygy at 3+4 on arms arising from IBr; IIBr 2 or 4(3+4); IIIBr and beyond 3(2+3); arms arising from IIBr and beyond with first syzygy at 2+3 and first pinnule on br1; distal intersyzygial interval 427; distalmost pinnule comb on P3P21 (in
<taxonomicName id="4C3715E5FFB0A16EFE7CD90E1214FD8B" box="[397,555,575,598]" class="Crinoidea" family="Comasteridae" genus="Comatula" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="species" species="tenuicirrus">
<emphasis id="B943B274FFB0A16EFE7CD90E1214FD8B" box="[397,555,575,598]" italics="true" pageId="14" pageNumber="165">C. tenuicirrus</emphasis>
</taxonomicName>
at intervals as far as mid-arm); comb teeth single, triangular, confluent with outer edge of pinnule; comb not tapering significantly distally, or tapering to a blunt tip (in
<taxonomicName id="4C3715E5FFB0A16EFB64D952157CFDA6" box="[1173,1347,611,635]" class="Crinoidea" family="Comasteridae" genus="Comatula" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="species" species="gracilicirrus">
<emphasis id="B943B274FFB0A16EFB64D952157CFDA6" box="[1173,1347,611,635]" italics="true" pageId="14" pageNumber="165">C. gracilicirrus</emphasis>
</taxonomicName>
).
</paragraph>
<caption id="DF483EEEFFB0A16EFF66DC2411F7F83E" httpUri="https://zenodo.org/record/580178/files/figure.png" pageId="14" pageNumber="165" targetBox="[151,1436,667,1796]" targetPageId="14">
<paragraph id="8B886E66FFB0A16EFF66DC2411F7F83E" blockId="14.[151,1436,1813,2019]" pageId="14" pageNumber="165">
<emphasis id="B943B274FFB0A16EFF66DC24112DF8F6" bold="true" box="[151,274,1813,1835]" pageId="14" pageNumber="165">FIGURE 5.</emphasis>
<taxonomicName id="4C3715E5FFB0A16EFEE9DC2411B6F8F6" box="[280,393,1813,1835]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="genus">
<emphasis id="B943B274FFB0A16EFEE9DC2411B6F8F6" box="[280,393,1813,1835]" italics="true" pageId="14" pageNumber="165">Capillaster</emphasis>
</taxonomicName>
. (AC, FI)
<emphasis id="B943B274FFB0A16EFDFBDC24131EF8F6" box="[522,801,1813,1835]" italics="true" pageId="14" pageNumber="165">
<taxonomicName id="4C3715E5FFB0A16EFDFBDC241243F8F6" box="[522,636,1813,1835]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="genus">Capillaster</taxonomicName>
multiradiatus—
</emphasis>
(A, B, GI) live, on soft coral (A) and hard coral (B) during the day, and (GH) with centrodorsal hidden and arms exposed at night; (C, I) aboral surface, showing rounded pentagonal centrodorsal and row of cirri; (F) comb with single, triangular teeth confluent with outer edge of pinnular. (DE)
<emphasis id="B943B274FFB0A16EFADBDC62113CF85A" italics="true" pageId="14" pageNumber="165">
<taxonomicName id="4C3715E5FFB0A16EFADBDC6215A3F8B4" box="[1322,1436,1875,1897]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="14" pageNumber="165" phylum="Echinodermata" rank="genus">Capillaster</taxonomicName>
sentosus—
</emphasis>
(D) live, exposed for photograph. (E) ray base, preserved, showing rounded pentagonal centrodorsal with one row of cirri. Madang, Papua New Guinea: A. MNHN-IE-2013-8105; B. MNHN-IE-2013-8079; D. MNHN-IE-2013-8066; G, I. MNHN-IE-2013-8066; H. not vouchered. Raja Ampat, Indonesia: C. not vouchered (RA219); F. SIO-BIC E6179. Australia or Timor: E. MNHN EcCs 1006.
</paragraph>
</caption>
<caption id="DF483EEEFFB1A16FFF66DC511476F86E" httpUri="https://zenodo.org/record/580179/files/figure.png" pageId="15" pageNumber="166" targetBox="[159,1411,207,1867]" targetPageId="15">
<paragraph id="8B886E66FFB1A16FFF66DC511476F86E" blockId="15.[151,1436,1888,1971]" pageId="15" pageNumber="166">
<emphasis id="B943B274FFB1A16FFF66DC511127F8A8" bold="true" box="[151,280,1888,1910]" pageId="15" pageNumber="166">FIGURE 6.</emphasis>
Maximum likelihood tree of
<taxonomicName id="4C3715E5FFB1A16FFD90DC5112ECF8AB" box="[609,723,1888,1910]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="15" pageNumber="166" phylum="Echinodermata" rank="genus">
<emphasis id="B943B274FFB1A16FFD90DC5112ECF8AB" box="[609,723,1888,1910]" italics="true" pageId="15" pageNumber="166">Capillaster</emphasis>
</taxonomicName>
inferred from COI. Symbols and labels as in Figure 2. Location abbreviations—Great Barrier Reef, Australia (GBR); Madang, Papua New Guinea (PNG); Raja Ampat, Indonesia (Raja); Singapore Strait, Singapore (Sing). New COI sequences are Genbank
<accessionNumber id="9464F385FFB1A16FFCAADCAC13F2F86E" box="[859,973,1949,1971]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/KR010307" pageId="15" pageNumber="166">KR010307</accessionNumber>
-
<accessionNumber id="9464F385FFB1A16FFC25DCAC147AF86E" box="[980,1093,1949,1971]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/KR010329" pageId="15" pageNumber="166">KR010329</accessionNumber>
.
</paragraph>
</caption>
<caption id="DF483EEEFFAEA170FF66DC0A1246F871" httpUri="https://zenodo.org/record/580180/files/figure.png" pageId="16" pageNumber="167" targetBox="[159,1431,205,1819]" targetPageId="16">
<paragraph id="8B886E66FFAEA170FF66DC0A1246F871" blockId="16.[151,1436,1851,1964]" pageId="16" pageNumber="167">
<emphasis id="B943B274FFAEA170FF66DC0A1127F88D" bold="true" box="[151,280,1851,1873]" pageId="16" pageNumber="167">FIGURE 7.</emphasis>
Maximum likelihood tree of
<taxonomicName id="4C3715E5FFAEA170FDACDC0A12E7F88C" box="[605,728,1851,1873]" class="Crinoidea" family="Comatulidae" genus="Phanogenia" higherTaxonomySource="CoL" kingdom="Animalia" order="Comatulida" pageId="16" pageNumber="167" phylum="Echinodermata" rank="genus">
<emphasis id="B943B274FFAEA170FDACDC0A12E7F88C" box="[605,728,1851,1873]" italics="true" pageId="16" pageNumber="167">Phanogenia</emphasis>
</taxonomicName>
inferred from COI. Symbols and labels as in Figure 2. Location abbreviations—Borneo (Born); Great Barrier Reef, Australia (GBR); Malaysia (Malay); Papua New Guinea (PNG); Philippines (Philip); Raja Ampat, Indonesia (Raja); Singapore Strait, Singapore (Sing); Viligili, Maldives (Mald). New COI sequences are Genbank
<accessionNumber id="9464F385FFAEA170FE7BDCA711C3F871" box="[394,508,1942,1964]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/KR010294" pageId="16" pageNumber="167">KR010294</accessionNumber>
-
<accessionNumber id="9464F385FFAEA170FDF2DCA7124AF871" box="[515,629,1942,1964]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/KR010306" pageId="16" pageNumber="167">KR010306</accessionNumber>
.
</paragraph>
</caption>
<paragraph id="8B886E66FFAFA171FF36DBA613DFFE0D" blockId="17.[151,1437,151,933]" pageId="17" pageNumber="168">
<emphasis id="B943B274FFAFA171FF36DBA611A1FF6D" bold="true" box="[199,414,151,176]" pageId="17" pageNumber="168">Molecular results.</emphasis>
Specimens identified as
<taxonomicName id="4C3715E5FFAFA171FD45DBA913A7FF6D" box="[692,920,152,176]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="sentosus">
<emphasis id="B943B274FFAFA171FD45DBA913A7FF6D" box="[692,920,152,176]" italics="true" pageId="17" pageNumber="168">Capillaster sentosus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C3715E5FFAFA171FC3CDBA91444FF6D" box="[973,1147,152,176]" class="Crinoidea" family="Comasteridae" genus="Comatula" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="gracilicirrus">
<emphasis id="B943B274FFAFA171FC3CDBA91444FF6D" box="[973,1147,152,176]" italics="true" pageId="17" pageNumber="168">C. gracilicirrus</emphasis>
</taxonomicName>
and those identified as
<emphasis id="B943B274FFAFA171FA73DBA912B0FF09" italics="true" pageId="17" pageNumber="168">
<taxonomicName id="4C3715E5FFAFA171FA73DBA9110EFF09" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="multiradiatus">C. multiradiatus</taxonomicName>
,
<taxonomicName id="4C3715E5FFAFA171FECCDB8D11E1FF09" box="[317,478,188,212]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="squarrosus">C. squarrosus</taxonomicName>
,
<taxonomicName id="4C3715E5FFAFA171FE1ADB8D12B3FF09" box="[491,652,188,212]" class="Crinoidea" family="Comasteridae" genus="Comatula" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="tenuicirrus">C. tenuicirrus</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C3715E5FFAFA171FD38DB8D13ABFF09" box="[713,916,188,212]" class="Crinoidea" family="Comatulidae" genus="Actinometra" higherTaxonomySource="CoL" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="macrobrachius">
<emphasis id="B943B274FFAFA171FD38DB8D13ABFF09" box="[713,916,188,212]" italics="true" pageId="17" pageNumber="168">C. macrobrachius</emphasis>
</taxonomicName>
formed two monophyletic groups in both the parsimony and likelihood analyses (
<figureCitation id="130C72E3FFAFA171FDD9DBEE1253FF25" box="[552,620,223,248]" captionStart="FIGURE 6" captionStartId="15.[151,250,1888,1910]" captionTargetBox="[159,1411,207,1867]" captionTargetId="figure@15.[150,1437,193,1867]" captionTargetPageId="15" captionText="FIGURE 6. Maximum likelihood tree of Capillaster inferred from COI. Symbols and labels as in Figure 2. Location abbreviations—Great Barrier Reef, Australia (GBR); Madang, Papua New Guinea (PNG); Raja Ampat, Indonesia (Raja); Singapore Strait, Singapore (Sing). New COI sequences are Genbank KR 010307 - KR 010329." httpUri="https://zenodo.org/record/580179/files/figure.png" pageId="17" pageNumber="168">Fig. 6</figureCitation>
), with a minimum of 5.3% uncorrected COI distance between the clades.
<emphasis id="B943B274FFAFA171FF66DA3410F0FEC1" box="[151,207,261,284]" italics="true" pageId="17" pageNumber="168">Intra</emphasis>
specific divergence among specimens identified as
<taxonomicName id="4C3715E5FFAFA171FCD1DA35139CFEC1" box="[800,931,260,284]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="sentosus">
<emphasis id="B943B274FFAFA171FCD1DA35139CFEC1" box="[800,931,260,284]" italics="true" pageId="17" pageNumber="168">C. sentosus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C3715E5FFAFA171FC2EDA3514AFFEC1" box="[991,1168,260,284]" class="Crinoidea" family="Comasteridae" genus="Comatula" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="gracilicirrus">
<emphasis id="B943B274FFAFA171FC2EDA3514AFFEC1" box="[991,1168,260,284]" italics="true" pageId="17" pageNumber="168">C. gracilicirrus</emphasis>
</taxonomicName>
(here referred to as
<taxonomicName id="4C3715E5FFAFA171FA73DA3510C9FE9D" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="sentosus">
<emphasis id="B943B274FFAFA171FA73DA3510C9FE9D" italics="true" pageId="17" pageNumber="168">C. sentosus</emphasis>
</taxonomicName>
) was less than 1.5%. Specimens within the
<taxonomicName id="4C3715E5FFAFA171FD2EDA1913C4FE9D" box="[735,1019,296,320]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="multiradiatus">
<emphasis id="B943B274FFAFA171FD2EDA1913C4FE9D" box="[735,1019,296,320]" italics="true" pageId="17" pageNumber="168">Capillaster multiradiatus</emphasis>
</taxonomicName>
clade (identified as
<emphasis id="B943B274FFAFA171FB2CDA1911C1FEB9" italics="true" pageId="17" pageNumber="168">
<taxonomicName id="4C3715E5FFAFA171FB2CDA1915A7FE9D" box="[1245,1432,296,320]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="multiradiatus">C. multiradiatus</taxonomicName>
,
<taxonomicName id="4C3715E5FFAFA171FF66DA7D1101FEB9" box="[151,318,332,356]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="squarrosus">C. squarrosus</taxonomicName>
,
<taxonomicName id="4C3715E5FFAFA171FEA2DA7D11C5FEB9" box="[339,506,332,356]" class="Crinoidea" family="Comasteridae" genus="Comatula" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="tenuicirrus">C. tenuicirrus</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C3715E5FFAFA171FDB7DA7D1327FEB9" box="[582,792,332,356]" class="Crinoidea" family="Comatulidae" genus="Actinometra" higherTaxonomySource="CoL" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="macrobrachius">
<emphasis id="B943B274FFAFA171FDB7DA7D1327FEB9" box="[582,792,332,356]" italics="true" pageId="17" pageNumber="168">C. macrobrachius</emphasis>
</taxonomicName>
) showed an
<emphasis id="B943B274FFAFA171FC30DA7C13C8FEB9" box="[961,1015,333,356]" italics="true" pageId="17" pageNumber="168">intra</emphasis>
specific divergence of up to 7.3% uncorrected COI distance (n=20). A clear separation in
<emphasis id="B943B274FFAFA171FCE1DA411379FE5A" box="[784,838,368,391]" italics="true" pageId="17" pageNumber="168">intra</emphasis>
<typeStatus id="548CD0C4FFAFA171FCB6DA411348FE55" box="[839,887,368,392]" pageId="17" pageNumber="168">type</typeStatus>
versus
<emphasis id="B943B274FFAFA171FC3EDA41143BFE5A" box="[975,1028,368,391]" italics="true" pageId="17" pageNumber="168">inter</emphasis>
<typeStatus id="548CD0C4FFAFA171FBF5DA41140BFE55" box="[1028,1076,368,392]" pageId="17" pageNumber="168">type</typeStatus>
COI uncorrected distances was not recovered. Therefore, although this clade may represent multiple different species, we currently do not have molecular or morphological data allowing us to clearly distinguish species.
</paragraph>
</subSubSection>
<subSubSection id="C32D3DEDFFAFA171FF36DAED1403FD59" pageId="17" pageNumber="168" type="distribution">
<paragraph id="8B886E66FFAFA171FF36DAED1403FD59" blockId="17.[151,1437,151,933]" pageId="17" pageNumber="168">
<emphasis id="B943B274FFAFA171FF36DAED115FFE28" bold="true" box="[199,352,476,501]" pageId="17" pageNumber="168">Distribution.</emphasis>
Tropical Indo-western Pacific from northern
<collectingCountry id="F3202EF6FFAFA171FC9BDAED13EEFE28" box="[874,977,476,501]" name="Australia" pageId="17" pageNumber="168">Australia</collectingCountry>
(Fremantle, WA, to Lady Musgrave Is., QLD), west to East Africa and the Red Sea, east to
<collectingCountry id="F3202EF6FFAFA171FD2BD93113B7FDC5" box="[730,904,511,536]" name="New Caledonia" pageId="17" pageNumber="168">New Caledonia</collectingCountry>
and
<collectingCountry id="F3202EF6FFAFA171FC4ED9311421FDCA" box="[959,1054,512,535]" name="Vanuatu" pageId="17" pageNumber="168">Vanuatu</collectingCountry>
, and north to East
<collectingCountry id="F3202EF6FFAFA171FB1EDACE150DFDC5" box="[1263,1330,511,536]" name="China" pageId="17" pageNumber="168">China</collectingCountry>
Sea west of Amami-Oshima,
<collectingCountry id="F3202EF6FFAFA171FE8BD9141182FDE1" box="[378,445,549,572]" name="Japan" pageId="17" pageNumber="168">Japan</collectingCountry>
, and
<collectingRegion id="49F3A084FFAFA171FE0BD915127AFDE0" box="[506,581,548,573]" country="Micronesia" name="Chuuk" pageId="17" pageNumber="168">Chuuk</collectingRegion>
Atoll (AH
<bibRefCitation id="EFA61397FFAFA171FD39D9151375FDE1" author="Clark" box="[712,842,548,573]" pageId="17" pageNumber="168" refString="Clark, A. H. (1931) A monograph of the existing crinoids 1 (3). Bulletin of the United States National Museum, 82, 1 - 916." type="journal article" year="1931">Clark 1931</bibRefCitation>
;
<bibRefCitation id="EFA61397FFAFA171FCA6D914147EFDE0" author="Rowe" box="[855,1089,548,573]" pageId="17" pageNumber="168" refString="Rowe, F. W. E. &amp; Gates, J. (1995) Echinodermata. Zoological Catalogue of Australia. CSIRO, Melbourne, 510 pp." type="book" year="1995">Rowe &amp; Gates 1995</bibRefCitation>
;
<bibRefCitation id="EFA61397FFAFA171FBBFD91514C0FDE0" author="Messing" box="[1102,1279,548,573]" pageId="17" pageNumber="168" refString="Messing, C. G. (1998 b) An initial re-assessment of the distribution and diversity of the East Indian shallow-water crinoid fauna. Pp. 187 - 192. In: Mooi, R. &amp; Telford, M. (eds.) Echinoderms: San Francisco. Balkema, Rotterdam." type="book chapter" year="1998" yearSuffix="b">Messing 1998b</bibRefCitation>
;
<bibRefCitation id="EFA61397FFAFA171FAFAD91415A7FDE0" author="Meyer" box="[1291,1432,548,573]" pageId="17" pageNumber="168" refString="Meyer, D. L. (1986) Les crinoides. In: Guille, A., Laboute, P. &amp; Menou J. - L. (Eds.), Guide des etoiles de mer, oursins, et autres echinodermes du lagon de Nouvelle-Caledonie, Faune Tropicale. ORSTOM, Paris, pp. 199 - 225." type="book chapter" year="1986">Meyer 1986</bibRefCitation>
;
<bibRefCitation id="EFA61397FFAFA171FF66D9791127FDBD" author="Kogo" box="[151,280,584,608]" pageId="17" pageNumber="168" refString="Kogo, I. (1998) Crinoids from Japan and its adjacent waters. Special Publication of the Osaka Museum of National History, 30, 1 - 148." type="journal article" year="1998">Kogo 1998</bibRefCitation>
). Depth range: 0118 (?292) m. Except for
<taxonomicName id="4C3715E5FFAFA171FCF1D97913A3FD82" box="[768,924,584,607]" class="Crinoidea" family="Comasteridae" genus="Comatula" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="tenuicirrus">
<emphasis id="B943B274FFAFA171FCF1D97913A3FD82" box="[768,924,584,607]" italics="true" pageId="17" pageNumber="168">C. tenuicirrus</emphasis>
</taxonomicName>
(
<quantity id="4CCFC383FFAFA171FC5AD9791420FDBD" box="[939,1055,584,608]" metricMagnitude="2" metricUnit="m" metricValue="1.0" metricValueMax="1.18" metricValueMin="0.82" pageId="17" pageNumber="168" unit="m" value="100.0" valueMax="118.0" valueMin="82.0">82118 m</quantity>
), the great majority of records are from shallower than
<specimenCount id="9D31A5EFFFAFA171FE8ED95D1180FD59" box="[383,447,620,645]" pageId="17" pageNumber="168" type="generic">
<quantity id="4CCFC383FFAFA171FE8ED95D1184FD59" box="[383,443,620,645]" metricMagnitude="1" metricUnit="m" metricValue="5.0" pageId="17" pageNumber="168" unit="m" value="50.0">50 m</quantity>
.
</specimenCount>
A few records from greater than
<quantity id="4CCFC383FFAFA171FCC4D95D1343FD59" box="[821,892,620,645]" metricMagnitude="2" metricUnit="m" metricValue="1.5" pageId="17" pageNumber="168" unit="m" value="150.0">150 m</quantity>
are likely errors.
</paragraph>
</subSubSection>
<subSubSection id="C32D3DEDFFAFA171FF36D9BE131CFBCF" pageId="17" pageNumber="168" type="discussion">
<paragraph id="8B886E66FFAFA171FF36D9BE11AFFC78" blockId="17.[151,1437,151,933]" pageId="17" pageNumber="168">
<emphasis id="B943B274FFAFA171FF36D9BE1104FD75" bold="true" box="[199,315,655,680]" pageId="17" pageNumber="168">Remarks.</emphasis>
<taxonomicName id="4C3715E5FFAFA171FEB3D9A111FFFD75" box="[322,448,656,680]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="genus">
<emphasis id="B943B274FFAFA171FEB3D9A111FFFD75" box="[322,448,656,680]" italics="true" pageId="17" pageNumber="168">Capillaster</emphasis>
</taxonomicName>
is the only Indo-western Pacific genus with IIIBr and following brachiataxes of 3(2+3) and the first syzygy at 2+3 on free arms arising from IIBr and more distal brachitaxes, a trait shared with western Atlantic
<taxonomicName id="4C3715E5FFAFA171FEF1D9E91142FD2D" box="[256,381,728,752]" class="Crinoidea" family="Comasteridae" genus="Davidaster" higherTaxonomySource="GBIF" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="genus">
<emphasis id="B943B274FFAFA171FEF1D9E91142FD2D" box="[256,381,728,752]" italics="true" pageId="17" pageNumber="168">Davidaster</emphasis>
</taxonomicName>
and
<taxonomicName id="4C3715E5FFAFA171FE33D9E81211FD2D" box="[450,558,729,752]" class="Crinoidea" family="Comasteridae" genus="Nemaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="genus">
<emphasis id="B943B274FFAFA171FE33D9E81211FD2D" box="[450,558,729,752]" italics="true" pageId="17" pageNumber="168">Nemaster</emphasis>
</taxonomicName>
species. Trawled specimens in the collection at LIPI were attributed to
<taxonomicName id="4C3715E5FFAFA171FF66D9CD11EEFCC9" box="[151,465,764,789]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="gracilicirrus">
<emphasis id="B943B274FFAFA171FF66D9CD112AFCC9" box="[151,277,764,788]" italics="true" pageId="17" pageNumber="168">Capillaster</emphasis>
cf.
<emphasis id="B943B274FFAFA171FEB2D9CD11EEFCC9" box="[323,465,764,788]" italics="true" pageId="17" pageNumber="168">gracilicirrus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C3715E5FFAFA171FDFBD9CD12C6FCC9" box="[522,761,764,789]" class="Crinoidea" family="Comatulidae" genus="Actinometra" higherTaxonomySource="CoL" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="macrobrachius">
<emphasis id="B943B274FFAFA171FDFBD9CD121BFCCE" box="[522,548,764,787]" italics="true" pageId="17" pageNumber="168">C.</emphasis>
cf.
<emphasis id="B943B274FFAFA171FDA0D9CD12C6FCC9" box="[593,761,764,788]" italics="true" pageId="17" pageNumber="168">macrobrachius</emphasis>
</taxonomicName>
, and specimens dredged at
<collectingCountry id="F3202EF6FFAFA171FBC6D9CD1496FCC8" box="[1079,1193,764,789]" name="Singapore" pageId="17" pageNumber="168">Singapore</collectingCountry>
were attributed to
<taxonomicName id="4C3715E5FFAFA171FA73D9CD125FFCE5" authority="Messing &amp; Tay 2016" authorityName="Messing &amp; Tay" authorityYear="2016" class="Crinoidea" family="Comasteridae" genus="Comatula" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="tenuicirrus">
<emphasis id="B943B274FFAFA171FA73D9CD15A3FCCE" box="[1410,1436,764,787]" italics="true" pageId="17" pageNumber="168">C.</emphasis>
cf.
<emphasis id="B943B274FFAFA171FF30D8111103FCEA" box="[193,316,800,823]" italics="true" pageId="17" pageNumber="168">tenuicirrus</emphasis>
(
<bibRefCitation id="EFA61397FFAFA171FEA3D82E1267FCE5" author="Messing" box="[338,600,799,824]" pageId="17" pageNumber="168" refString="Messing, C. G. &amp; Tay, T. S. (2016) Extant Crinoidea (Echinodermata) of Singapore. Raffles Bulletin of Zoology Supplement, 34, 270 - 301." type="journal article" year="2016">Messing &amp; Tay 2016</bibRefCitation>
)
</taxonomicName>
, all on morphological grounds. As with
<taxonomicName id="4C3715E5FFAFA171FBA2D81114FEFCE5" box="[1107,1217,800,824]" class="Crinoidea" family="Comasteridae" genus="Comatula" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="genus">
<emphasis id="B943B274FFAFA171FBA2D81114FEFCE5" box="[1107,1217,800,824]" italics="true" pageId="17" pageNumber="168">Comatula</emphasis>
</taxonomicName>
and several other confamilial genera, species-level distinguishing features are often size- and proportion-related, reflecting possible oversplitting of the genus. The LIPI and
<collectingCountry id="F3202EF6FFAFA171FD77D85612C7FC5D" box="[646,760,871,896]" name="Singapore" pageId="17" pageNumber="168">Singapore</collectingCountry>
specimens are assigned here to either
<taxonomicName id="4C3715E5FFAFA171FB3DD859156DFC5D" box="[1228,1362,872,896]" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="sentosus">
<emphasis id="B943B274FFAFA171FB3DD859156DFC5D" box="[1228,1362,872,896]" italics="true" pageId="17" pageNumber="168">C. sentosus</emphasis>
</taxonomicName>
or
<taxonomicName id="4C3715E5FFAFA171FA73D8591111FC79" class="Crinoidea" family="Comasteridae" genus="Capillaster" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="species" species="multiradiatus">
<emphasis id="B943B274FFAFA171FA73D8591111FC79" italics="true" pageId="17" pageNumber="168">C. multiradiatus</emphasis>
</taxonomicName>
(
<figureCitation id="130C72E3FFAFA171FECCD8BD11BDFC78" box="[317,386,908,933]" captionStart="FIGURE 6" captionStartId="15.[151,250,1888,1910]" captionTargetBox="[159,1411,207,1867]" captionTargetId="figure@15.[150,1437,193,1867]" captionTargetPageId="15" captionText="FIGURE 6. Maximum likelihood tree of Capillaster inferred from COI. Symbols and labels as in Figure 2. Location abbreviations—Great Barrier Reef, Australia (GBR); Madang, Papua New Guinea (PNG); Raja Ampat, Indonesia (Raja); Singapore Strait, Singapore (Sing). New COI sequences are Genbank KR 010307 - KR 010329." httpUri="https://zenodo.org/record/580179/files/figure.png" pageId="17" pageNumber="168">Fig. 6</figureCitation>
).
</paragraph>
<paragraph id="8B886E66FFAFA171FF66D8C9131CFBCF" blockId="17.[151,803,1016,1042]" box="[151,803,1016,1042]" pageId="17" pageNumber="168">
<emphasis id="B943B274FFAFA171FF66D8C9131CFBCF" bold="true" box="[151,803,1016,1042]" pageId="17" pageNumber="168">
<taxonomicName id="4C3715E5FFAFA171FF66D8C91145FBCD" box="[151,378,1016,1042]" class="Crinoidea" family="Comatulidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Comatulida" pageId="17" pageNumber="168" phylum="Echinodermata" rank="tribe" tribe="Neocomatellini">NEOCOMATELLINI</taxonomicName>
<bibRefCitation id="EFA61397FFAFA171FE70D8C9131CFBCF" author="Summers" box="[385,803,1016,1042]" pageId="17" pageNumber="168" refString="Summers, M. M., Messing C. G. &amp; Rouse G. W. (2014 a). Phylogeny of Comatulidae (Echinodermata: Crinoidea: Comatulida). Molecular Phylogenetics &amp; Evolution, 80, 319 - 339." type="journal article" year="2014">Summers, Messing &amp; Rouse 2014</bibRefCitation>
</emphasis>
</paragraph>
</subSubSection>
</treatment>
</document>