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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Cleaning up the 'Bigmessidae': Molecular phylogeny of scleractinian corals from Faviidae, Merulinidae, Pectiniidae and Trachyphylliidae</title><author><name sortKey="Huang, Danwei" sort="Huang, Danwei" uniqKey="Huang D" first="Danwei" last="Huang">Danwei Huang</name><affiliation><nlm:aff id="I1">Scripps Institution of Oceanography, University of California, San Diego, La Jolla, CA 92093, USA</nlm:aff></affiliation><affiliation><nlm:aff id="I2">Department of Biological Sciences, National University of Singapore, Singapore 117543, Singapore</nlm:aff></affiliation></author><author><name sortKey="Licuanan, Wilfredo Y" sort="Licuanan, Wilfredo Y" uniqKey="Licuanan W" first="Wilfredo Y" last="Licuanan">Wilfredo Y. Licuanan</name><affiliation><nlm:aff id="I3">Br. Alfred Shields Marine Station and Biology Department, De La Salle University, Manila 1004, The Philippines</nlm:aff></affiliation></author><author><name sortKey="Baird, Andrew H" sort="Baird, Andrew H" uniqKey="Baird A" first="Andrew H" last="Baird">Andrew H. Baird</name><affiliation><nlm:aff id="I4">ARC Center of Excellence for Coral Reef Studies, James Cook University, Townsville, Queensland 4811, Australia</nlm:aff></affiliation></author><author><name sortKey="Fukami, Hironobu" sort="Fukami, Hironobu" uniqKey="Fukami H" first="Hironobu" last="Fukami">Hironobu Fukami</name><affiliation><nlm:aff id="I5">Department of Marine Biology and Environmental Science, University of Miyazaki, Miyazaki 889-2192, Japan</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">21299898</idno><idno type="pmc">3042006</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3042006</idno><idno type="RBID">PMC:3042006</idno><idno type="doi">10.1186/1471-2148-11-37</idno><date when="2011">2011</date><idno type="wicri:Area/Pmc/Corpus">000490</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Cleaning up the 'Bigmessidae': Molecular phylogeny of scleractinian corals from Faviidae, Merulinidae, Pectiniidae and Trachyphylliidae</title><author><name sortKey="Huang, Danwei" sort="Huang, Danwei" uniqKey="Huang D" first="Danwei" last="Huang">Danwei Huang</name><affiliation><nlm:aff id="I1">Scripps Institution of Oceanography, University of California, San Diego, La Jolla, CA 92093, USA</nlm:aff></affiliation><affiliation><nlm:aff id="I2">Department of Biological Sciences, National University of Singapore, Singapore 117543, Singapore</nlm:aff></affiliation></author><author><name sortKey="Licuanan, Wilfredo Y" sort="Licuanan, Wilfredo Y" uniqKey="Licuanan W" first="Wilfredo Y" last="Licuanan">Wilfredo Y. Licuanan</name><affiliation><nlm:aff id="I3">Br. Alfred Shields Marine Station and Biology Department, De La Salle University, Manila 1004, The Philippines</nlm:aff></affiliation></author><author><name sortKey="Baird, Andrew H" sort="Baird, Andrew H" uniqKey="Baird A" first="Andrew H" last="Baird">Andrew H. Baird</name><affiliation><nlm:aff id="I4">ARC Center of Excellence for Coral Reef Studies, James Cook University, Townsville, Queensland 4811, Australia</nlm:aff></affiliation></author><author><name sortKey="Fukami, Hironobu" sort="Fukami, Hironobu" uniqKey="Fukami H" first="Hironobu" last="Fukami">Hironobu Fukami</name><affiliation><nlm:aff id="I5">Department of Marine Biology and Environmental Science, University of Miyazaki, Miyazaki 889-2192, Japan</nlm:aff></affiliation></author></analytic><series><title level="j">BMC Evolutionary Biology</title><idno type="eISSN">1471-2148</idno><imprint><date when="2011">2011</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><sec><title>Background</title><p>Molecular phylogenetic studies on scleractinian corals have shown that most taxa are not reflective of their evolutionary histories. Based principally on gross morphology, traditional taxonomy suffers from the lack of well-defined and homologous characters that can sufficiently describe scleractinian diversity. One of the most challenging clades recovered by recent analyses is 'Bigmessidae', an informal grouping that comprises four conventional coral families, Faviidae, Merulinidae, Pectiniidae and Trachyphylliidae, interspersed among one another with no apparent systematic pattern. There is an urgent need for taxonomic revisions in this clade, but it is vital to first establish phylogenetic relationships within the group. In this study, we reconstruct the evolutionary history of 'Bigmessidae' based on five DNA sequence markers gathered from 76 of the 132 currently recognized species collected from five reef regions in the central Indo-Pacific and the Atlantic.</p></sec><sec><title>Results</title><p>We present a robust molecular phylogeny of 'Bigmessidae' based on the combined five-gene data, achieving a higher degree of resolution compared to previous analyses. Two Pacific species presumed to be in 'Bigmessidae' are more closely related to outgroup clades, suggesting that other unsampled taxa have unforeseen affinities. As expected, nested within 'Bigmessidae' are four conventional families as listed above, and relationships among them generally corroborate previous molecular analyses. Our more resolved phylogeny supports a close association of <italic>Hydnophora </italic>(Merulinidae) with <italic>Favites </italic>+ <italic>Montastraea </italic>(Faviidae), rather than with the rest of Merulinidae, i.e., <italic>Merulina </italic>and <italic>Scapophyllia</italic>. <italic>Montastraea annularis</italic>, the only Atlantic 'Bigmessidae' is sister to <italic>Cyphastrea</italic>, a grouping that can be reconciled by their septothecal walls, a microstructural feature of the skeleton determined by recent morphological work. Characters at the subcorallite scale appear to be appropriate synapomorphies for other subclades, which cannot be explained using macromorphology. Indeed, wide geographic sampling here has revealed more instances of possible cryptic taxa confused by evolutionary convergence of gross coral morphology.</p></sec><sec><title>Conclusions</title><p>Numerous examples of cryptic taxa determined in this study support the assertion that diversity estimates of scleractinian corals are erroneous. Fortunately, the recovery of most 'Bigmessidae' genera with only minor degrees of paraphyly offers some hope for impending taxonomic amendments. Subclades are well defined and supported by subcorallite morphological features, providing a robust framework for further systematic work.</p></sec></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Romano, Sl" uniqKey="Romano S">SL Romano</name></author><author><name sortKey="Palumbi, Sr" uniqKey="Palumbi S">SR Palumbi</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Romano, Sl" uniqKey="Romano S">SL Romano</name></author><author><name sortKey="Palumbi, Sr" uniqKey="Palumbi S">SR Palumbi</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Chen, Ca" uniqKey="Chen C">CA Chen</name></author><author><name sortKey="Wallace, Cc" uniqKey="Wallace C">CC Wallace</name></author><author><name sortKey="Wolstenholme, Jk" uniqKey="Wolstenholme J">JK Wolstenholme</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Wells, Jw" uniqKey="Wells J">JW 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<front><journal-meta><journal-id journal-id-type="nlm-ta">BMC Evol Biol</journal-id><journal-title-group><journal-title>BMC Evolutionary Biology</journal-title></journal-title-group><issn pub-type="epub">1471-2148</issn><publisher><publisher-name>BioMed Central</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">21299898</article-id><article-id pub-id-type="pmc">3042006</article-id><article-id pub-id-type="publisher-id">1471-2148-11-37</article-id><article-id pub-id-type="doi">10.1186/1471-2148-11-37</article-id><article-categories><subj-group subj-group-type="heading"><subject>Research Article</subject></subj-group></article-categories><title-group><article-title>Cleaning up the 'Bigmessidae': Molecular phylogeny of scleractinian corals from Faviidae, Merulinidae, Pectiniidae and Trachyphylliidae</article-title></title-group><contrib-group><contrib contrib-type="author" corresp="yes" id="A1"><name><surname>Huang</surname><given-names>Danwei</given-names></name><xref ref-type="aff" rid="I1">1</xref><xref ref-type="aff" rid="I2">2</xref><email>huangdanwei@ucsd.edu</email></contrib><contrib contrib-type="author" id="A2"><name><surname>Licuanan</surname><given-names>Wilfredo Y</given-names></name><xref ref-type="aff" rid="I3">3</xref><email>licuananw@dlsu.edu.ph</email></contrib><contrib contrib-type="author" id="A3"><name><surname>Baird</surname><given-names>Andrew H</given-names></name><xref ref-type="aff" rid="I4">4</xref><email>andrew.baird@jcu.edu.au</email></contrib><contrib contrib-type="author" id="A4"><name><surname>Fukami</surname><given-names>Hironobu</given-names></name><xref ref-type="aff" rid="I5">5</xref><email>hirofukami@cc.miyazaki-u.ac.jp</email></contrib></contrib-group><aff id="I1"><label>1</label>Scripps Institution of Oceanography, University of California, San Diego, La Jolla, CA 92093, USA</aff><aff id="I2"><label>2</label>Department of Biological Sciences, National University of Singapore, Singapore 117543, Singapore</aff><aff id="I3"><label>3</label>Br. Alfred Shields Marine Station and Biology Department, De La Salle University, Manila 1004, The Philippines</aff><aff id="I4"><label>4</label>ARC Center of Excellence for Coral Reef Studies, James Cook University, Townsville, Queensland 4811, Australia</aff><aff id="I5"><label>5</label>Department of Marine Biology and Environmental Science, University of Miyazaki, Miyazaki 889-2192, Japan</aff><pub-date pub-type="collection"><year>2011</year></pub-date><pub-date pub-type="epub"><day>7</day><month>2</month><year>2011</year></pub-date><volume>11</volume><fpage>37</fpage><lpage>37</lpage><history><date date-type="received"><day>15</day><month>6</month><year>2010</year></date><date date-type="accepted"><day>7</day><month>2</month><year>2011</year></date></history><permissions><copyright-statement>Copyright ©2011 Huang et al; licensee BioMed Central Ltd.</copyright-statement><copyright-year>2011</copyright-year><copyright-holder>Huang et al; licensee BioMed Central Ltd.</copyright-holder><license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by/2.0"><license-p>This is an Open Access article distributed under the terms of the Creative Commons Attribution License (<ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/2.0">http://creativecommons.org/licenses/by/2.0</ext-link>), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.</license-p></license></permissions><self-uri xlink:href="http://www.biomedcentral.com/1471-2148/11/37"></self-uri><abstract><sec><title>Background</title><p>Molecular phylogenetic studies on scleractinian corals have shown that most taxa are not reflective of their evolutionary histories. Based principally on gross morphology, traditional taxonomy suffers from the lack of well-defined and homologous characters that can sufficiently describe scleractinian diversity. One of the most challenging clades recovered by recent analyses is 'Bigmessidae', an informal grouping that comprises four conventional coral families, Faviidae, Merulinidae, Pectiniidae and Trachyphylliidae, interspersed among one another with no apparent systematic pattern. There is an urgent need for taxonomic revisions in this clade, but it is vital to first establish phylogenetic relationships within the group. In this study, we reconstruct the evolutionary history of 'Bigmessidae' based on five DNA sequence markers gathered from 76 of the 132 currently recognized species collected from five reef regions in the central Indo-Pacific and the Atlantic.</p></sec><sec><title>Results</title><p>We present a robust molecular phylogeny of 'Bigmessidae' based on the combined five-gene data, achieving a higher degree of resolution compared to previous analyses. Two Pacific species presumed to be in 'Bigmessidae' are more closely related to outgroup clades, suggesting that other unsampled taxa have unforeseen affinities. As expected, nested within 'Bigmessidae' are four conventional families as listed above, and relationships among them generally corroborate previous molecular analyses. Our more resolved phylogeny supports a close association of <italic>Hydnophora </italic>(Merulinidae) with <italic>Favites </italic>+ <italic>Montastraea </italic>(Faviidae), rather than with the rest of Merulinidae, i.e., <italic>Merulina </italic>and <italic>Scapophyllia</italic>. <italic>Montastraea annularis</italic>, the only Atlantic 'Bigmessidae' is sister to <italic>Cyphastrea</italic>, a grouping that can be reconciled by their septothecal walls, a microstructural feature of the skeleton determined by recent morphological work. Characters at the subcorallite scale appear to be appropriate synapomorphies for other subclades, which cannot be explained using macromorphology. Indeed, wide geographic sampling here has revealed more instances of possible cryptic taxa confused by evolutionary convergence of gross coral morphology.</p></sec><sec><title>Conclusions</title><p>Numerous examples of cryptic taxa determined in this study support the assertion that diversity estimates of scleractinian corals are erroneous. Fortunately, the recovery of most 'Bigmessidae' genera with only minor degrees of paraphyly offers some hope for impending taxonomic amendments. Subclades are well defined and supported by subcorallite morphological features, providing a robust framework for further systematic work.</p></sec></abstract></article-meta></front><body><sec><title>Background</title><p>For the last two decades, coral systematists have been untangling the complex evolutionary relationships among scleractinian species using DNA sequence data. Seminal molecular phylogenetic work by Romano and Palumbi [<xref ref-type="bibr" rid="B1">1</xref>,<xref ref-type="bibr" rid="B2">2</xref>] divided the Scleractinia into two major clades, the robust and complex groups, and indicated many problems with traditional taxonomy based on morphology (see also [<xref ref-type="bibr" rid="B3">3</xref>]). For instance, <italic>Leptastrea </italic>was recovered within a Fungiina clade rather than the suborder Faviina, where morphological studies had placed it (e.g., [<xref ref-type="bibr" rid="B4">4</xref>,<xref ref-type="bibr" rid="B5">5</xref>]). Gradually, using more genetic loci, further evidence was uncovered to show that non-monophyly of coral taxa is widespread in Scleractinia (e.g., [<xref ref-type="bibr" rid="B6">6</xref>-<xref ref-type="bibr" rid="B11">11</xref>]). This culminated in a comprehensive survey of the entire taxon by Fukami et al. [<xref ref-type="bibr" rid="B12">12</xref>], which showed that while Scleractinia is monophyletic, most taxonomic groups within it are not. In fact, a staggering 11 of 16 conventional families are polyphyletic.</p><p>Undoubtedly, one of the most challenging clades that have been recovered by recent analyses is a group of robust corals in clade XVII [<xref ref-type="bibr" rid="B12">12</xref>]. The disarray within the clade is epitomized by its informal name 'Bigmessidae' [<xref ref-type="bibr" rid="B13">13</xref>,<xref ref-type="bibr" rid="B14">14</xref>]. This clade contains species from four traditional coral families, Faviidae, Merulinidae, Pectiniidae and Trachyphylliidae, interspersed among one another in the tree based on mitochondrial cytochrome oxidase I (COI) and cytochrome b gene sequences [<xref ref-type="bibr" rid="B12">12</xref>]. With the exception of the <italic>Montastraea annularis </italic>complex, all members of this clade are from the Indo-Pacific. Families with all species included within clade XVII are Trachyphylliidae (monospecific) and Merulinidae, the latter being polyphyletic, while Faviidae and Pectiniidae have representatives present within and outside clade XVII. Although the clade has not been examined in detail, Huang et al. [<xref ref-type="bibr" rid="B15">15</xref>] showed that representatives from other families (Merulinidae and Mussidae) are also nested within it, and several genera are not monophyletic (i.e., <italic>Echinopora</italic>, <italic>Favia</italic>, <italic>Favites, Goniastrea and Montastraea</italic>). In addition, Fukami et al. [<xref ref-type="bibr" rid="B12">12</xref>] found para- or polyphyly in <italic>Leptoria</italic>, <italic>Oulophyllia </italic>and <italic>Platygyra </italic>for at least one marker.</p><p>Clearly, there exists an urgent need for taxonomic revisions in this clade, amidst the ongoing disarray in the Scleractinia. But in order to begin any form of revision for clade XVII, it is first necessary to determine which subclades are problematic, using as complete a morphological and genetic coverage as possible. Up to this point, the largest number of markers used for analysis of this group has been derived from Fukami et al. [<xref ref-type="bibr" rid="B12">12</xref>], who used the aforementioned mitochondrial genes, as well as the nuclear β-tubulin and 28S rDNA separately. However, only 33 species represented by 38 terminals were analyzed for clade XVII, and several subclades were not resolved due to their short branches. Resolution was improved in Huang et al. [<xref ref-type="bibr" rid="B15">15</xref>], which included 85 terminals from 43 species, but that study used only COI and a noncoding intergenic mitochondrial region (IGR).</p><p>In this study, we present data for five molecular markers—two mitochondrial and three nuclear loci—from 76 of the 132 currently recognized species in clade XVII [<xref ref-type="bibr" rid="B12">12</xref>]. We also included seven species from other robust corals as outgroups. Corals were sequenced from five reef regions—the central and northern Great Barrier Reef in Australia, Wakayama in Japan, Batangas in the Philippines, Singapore and the Caribbean. We reconstruct the evolutionary history of clade XVII and identify subclade placement of species that have not been studied in a molecular phylogenetic context. As some species were sampled from multiple locations, we also test if these corals were as widespread as previously recorded.</p></sec><sec sec-type="methods"><title>Methods</title><sec><title>Specimen collection and DNA extraction</title><p>Specimens were collected from coral reefs in five regions—Singapore, Wakayama (Japan), Queensland (Australia), Batangas (The Philippines), and the Caribbean. To ensure consistency in identifications among localities, each coral was sampled by at least two authors, based on morphological features that can be recognized in the field. The identity was later confirmed in the laboratory after examining skeletal traits [<xref ref-type="bibr" rid="B5">5</xref>,<xref ref-type="bibr" rid="B16">16</xref>-<xref ref-type="bibr" rid="B21">21</xref>]. In total, 124 specimens from 83 species in clades XIV-XXI have been included in the present analysis (Table <xref ref-type="table" rid="T1">1</xref>; see Additional file <xref ref-type="supplementary-material" rid="S1">1</xref>). We photographed each colony in the field and collected between 10 and 100 cm<sup>2 </sup>of coral from each colony using a hammer and chisel, with ~2cm<sup>2 </sup>of tissue preserved in 100% ethanol.</p><table-wrap id="T1" position="float"><label>Table 1</label><caption><p>Species and DNA sequences examined in this study.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="center">No.</th><th align="left">Species</th><th align="left">Voucher</th><th align="center">28S rDNA</th><th align="center">histone H3</th><th align="center">ITS rDNA</th><th align="center">mt COI</th><th align="center">mt IGR</th></tr></thead><tbody><tr><td align="center">1</td><td align="left"><italic>Acanthastrea echinata </italic>(XX; Mussidae)</td><td align="left">S031</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203399">HQ203399</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203520">HQ203520</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203308">HQ203308</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371658">EU371658</ext-link></td><td></td></tr><tr><td align="center">2</td><td align="left"><italic>Barabattoia amicorum</italic></td><td align="left">S047</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203400">HQ203400</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203521">HQ203521</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203309">HQ203309</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345412">FJ345412</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345480">FJ345480</ext-link></td></tr><tr><td align="center">3</td><td align="left"><italic>Caulastraea echinulata</italic></td><td align="left">S041</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203401">HQ203401</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203522">HQ203522</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345414">FJ345414</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345496">FJ345496</ext-link></td></tr><tr><td align="center">4</td><td align="left"><italic>Caulastraea furcata</italic></td><td align="left">P108</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203402">HQ203402</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203523">HQ203523</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203248">HQ203248</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203639">HQ203639</ext-link></td></tr><tr><td align="center">5</td><td align="left"><bold><italic>Caulastraea tumida</italic></bold></td><td align="left">G61875</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203403">HQ203403</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203524">HQ203524</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203310">HQ203310</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203249">HQ203249</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203640">HQ203640</ext-link></td></tr><tr><td align="center">6</td><td align="left"><italic>Cyphastrea chalcidicum</italic></td><td align="left">G61902</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203404">HQ203404</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203525">HQ203525</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203311">HQ203311</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203250">HQ203250</ext-link></td><td></td></tr><tr><td align="center">7</td><td align="left"><italic>Cyphastrea chalcidicum</italic></td><td align="left">S103</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203405">HQ203405</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203526">HQ203526</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203312">HQ203312</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345415">FJ345415</ext-link></td><td></td></tr><tr><td align="center">8</td><td align="left"><italic>Cyphastrea microphthalma</italic></td><td align="left">S069</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203406">HQ203406</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203527">HQ203527</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345416">FJ345416</ext-link></td><td></td></tr><tr><td align="center">9</td><td align="left"><italic>Cyphastrea serailia</italic></td><td align="left">G61889</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203407">HQ203407</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203528">HQ203528</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203313">HQ203313</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203251">HQ203251</ext-link></td><td></td></tr><tr><td align="center">10</td><td align="left"><italic>Cyphastrea serailia</italic></td><td align="left">S024</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203408">HQ203408</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203529">HQ203529</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203314">HQ203314</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371659">EU371659</ext-link></td><td></td></tr><tr><td align="center">11</td><td align="left"><italic>Cyphastrea serailia</italic></td><td align="left">P120</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203409">HQ203409</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203530">HQ203530</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203252">HQ203252</ext-link></td><td></td></tr><tr><td align="center">12</td><td align="left"><italic>Diploastrea heliopora </italic>(XV)</td><td align="left">S048</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203410">HQ203410</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203531">HQ203531</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203315">HQ203315</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371660">EU371660</ext-link></td><td></td></tr><tr><td align="center">13</td><td align="left"><italic>Echinopora gemmacea</italic></td><td align="left">S120</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203411">HQ203411</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203532">HQ203532</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203316">HQ203316</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345418">FJ345418</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345457">FJ345457</ext-link></td></tr><tr><td align="center">14</td><td align="left"><bold><italic>Echinopora horrida</italic></bold></td><td align="left">G61907</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203412">HQ203412</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203533">HQ203533</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203317">HQ203317</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203253">HQ203253</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203641">HQ203641</ext-link></td></tr><tr><td align="center">15</td><td align="left"><italic>Echinopora lamellosa</italic></td><td align="left">S109</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203413">HQ203413</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203534">HQ203534</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203318">HQ203318</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345419">FJ345419</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345458">FJ345458</ext-link></td></tr><tr><td align="center">16</td><td align="left"><bold><italic>Echinopora mammiformis</italic></bold></td><td align="left">G61884</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203414">HQ203414</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203535">HQ203535</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203319">HQ203319</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203254">HQ203254</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203642">HQ203642</ext-link></td></tr><tr><td align="center">17</td><td align="left"><italic>Echinopora pacificus</italic></td><td align="left">S110</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203415">HQ203415</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203536">HQ203536</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203320">HQ203320</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345420">FJ345420</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345459">FJ345459</ext-link></td></tr><tr><td align="center">18</td><td align="left"><italic>Favia danae</italic></td><td align="left">G61885</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203416">HQ203416</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203537">HQ203537</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203321">HQ203321</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203643">HQ203643</ext-link></td></tr><tr><td align="center">19</td><td align="left"><italic>Favia danae</italic></td><td align="left">S092</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203417">HQ203417</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203538">HQ203538</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371663">EU371663</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345476">FJ345476</ext-link></td></tr><tr><td align="center">20</td><td align="left"><italic>Favia favus</italic></td><td align="left">G61880</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203418">HQ203418</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203539">HQ203539</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203322">HQ203322</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203255">HQ203255</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203644">HQ203644</ext-link></td></tr><tr><td align="center">21</td><td align="left"><italic>Favia favus</italic></td><td align="left">G61915</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203419">HQ203419</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203540">HQ203540</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203323">HQ203323</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203256">HQ203256</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203645">HQ203645</ext-link></td></tr><tr><td align="center">22</td><td align="left"><italic>Favia favus</italic></td><td align="left">S003</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203420">HQ203420</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203541">HQ203541</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203324">HQ203324</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371710">EU371710</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345511">FJ345511</ext-link></td></tr><tr><td align="center">23</td><td align="left"><italic>Favia favus</italic></td><td align="left">S025</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203421">HQ203421</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203542">HQ203542</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371664">EU371664</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345465">FJ345465</ext-link></td></tr><tr><td align="center">24</td><td align="left"><italic>Favia favus</italic></td><td align="left">S040</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203422">HQ203422</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203543">HQ203543</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203325">HQ203325</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371665">EU371665</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345466">FJ345466</ext-link></td></tr><tr><td align="center">25</td><td align="left"><italic>Favia favus</italic></td><td align="left">P105</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203423">HQ203423</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203544">HQ203544</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203257">HQ203257</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203646">HQ203646</ext-link></td></tr><tr><td align="center">26</td><td align="left"><italic>Favia fragum </italic>(XXI)</td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="AF549222">AF549222</ext-link></td><td></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="AB117222">AB117222</ext-link></td><td></td></tr><tr><td align="center">27</td><td align="left"><italic>Favia </italic>cf. <italic>laxa</italic></td><td align="left">S013</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203424">HQ203424</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203545">HQ203545</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371707">EU371707</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345508">FJ345508</ext-link></td></tr><tr><td align="center">28</td><td align="left"><italic>Favia </italic>cf. <italic>laxa</italic></td><td align="left">S014</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203425">HQ203425</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203546">HQ203546</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203326">HQ203326</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371708">EU371708</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345509">FJ345509</ext-link></td></tr><tr><td align="center">29</td><td align="left"><italic>Favia lizardensis</italic></td><td align="left">G61872</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203426">HQ203426</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203547">HQ203547</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203327">HQ203327</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203647">HQ203647</ext-link></td></tr><tr><td align="center">30</td><td align="left"><italic>Favia lizardensis</italic></td><td align="left">S072</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203427">HQ203427</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203548">HQ203548</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203328">HQ203328</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371668">EU371668</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345484">FJ345484</ext-link></td></tr><tr><td align="center">31</td><td align="left"><italic>Favia lizardensis</italic></td><td align="left">P136</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203428">HQ203428</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203549">HQ203549</ext-link></td><td></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203648">HQ203648</ext-link></td></tr><tr><td align="center">32</td><td align="left"><bold><italic>Favia </italic>cf. <italic>maritima</italic></bold></td><td align="left">G61912</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203429">HQ203429</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203550">HQ203550</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203329">HQ203329</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203258">HQ203258</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203649">HQ203649</ext-link></td></tr><tr><td align="center">33</td><td align="left"><italic>Favia matthaii</italic></td><td align="left">G61881</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203430">HQ203430</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203551">HQ203551</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203330">HQ203330</ext-link></td><td></td><td></td></tr><tr><td align="center">34</td><td align="left"><italic>Favia matthaii</italic></td><td align="left">G61883</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203431">HQ203431</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203552">HQ203552</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203331">HQ203331</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203259">HQ203259</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203650">HQ203650</ext-link></td></tr><tr><td align="center">35</td><td align="left"><italic>Favia matthaii</italic></td><td align="left">S005</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203432">HQ203432</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203553">HQ203553</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203332">HQ203332</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371669">EU371669</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345471">FJ345471</ext-link></td></tr><tr><td align="center">36</td><td align="left"><italic>Favia matthaii</italic></td><td align="left">S029</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203433">HQ203433</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203554">HQ203554</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203333">HQ203333</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371671">EU371671</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345473">FJ345473</ext-link></td></tr><tr><td align="center">37</td><td align="left"><italic>Favia maxima</italic></td><td align="left">S052</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203434">HQ203434</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203555">HQ203555</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203334">HQ203334</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371674">EU371674</ext-link></td><td></td></tr><tr><td align="center">38</td><td align="left"><italic>Favia maxima</italic></td><td align="left">P142</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203435">HQ203435</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203556">HQ203556</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203260">HQ203260</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203651">HQ203651</ext-link></td></tr><tr><td align="center">39</td><td align="left"><italic>Favia </italic>cf. <italic>maxima</italic></td><td align="left">P134</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203436">HQ203436</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203557">HQ203557</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203335">HQ203335</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203261">HQ203261</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203652">HQ203652</ext-link></td></tr><tr><td align="center">40</td><td align="left"><italic>Favia pallida</italic></td><td align="left">G61898</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203437">HQ203437</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203558">HQ203558</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203336">HQ203336</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203653">HQ203653</ext-link></td></tr><tr><td align="center">41</td><td align="left"><italic>Favia pallida</italic></td><td align="left">S036</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203438">HQ203438</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203559">HQ203559</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203337">HQ203337</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371675">EU371675</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345482">FJ345482</ext-link></td></tr><tr><td align="center">42</td><td align="left"><bold><italic>Favia rosaria</italic></bold></td><td align="left">G61911</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203439">HQ203439</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203560">HQ203560</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203338">HQ203338</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203262">HQ203262</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203654">HQ203654</ext-link></td></tr><tr><td align="center">43</td><td align="left"><italic>Favia rotumana</italic></td><td align="left">S068</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203440">HQ203440</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203561">HQ203561</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203339">HQ203339</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345427">FJ345427</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345485">FJ345485</ext-link></td></tr><tr><td align="center">44</td><td align="left"><bold><italic>Favia rotundata</italic></bold></td><td align="left">G61874</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203441">HQ203441</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203562">HQ203562</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203340">HQ203340</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203263">HQ203263</ext-link></td><td></td></tr><tr><td align="center">45</td><td align="left"><bold><italic>Favia rotundata</italic></bold></td><td align="left">P132</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203442">HQ203442</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203563">HQ203563</ext-link></td><td></td><td></td><td></td></tr><tr><td align="center">46</td><td align="left"><italic>Favia speciosa</italic></td><td align="left">S001</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203443">HQ203443</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203564">HQ203564</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203341">HQ203341</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371677">EU371677</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345505">FJ345505</ext-link></td></tr><tr><td align="center">47</td><td align="left"><italic>Favia speciosa</italic></td><td align="left">S026</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203444">HQ203444</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203565">HQ203565</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371680">EU371680</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345506">FJ345506</ext-link></td></tr><tr><td align="center">48</td><td align="left"><italic>Favia speciosa</italic></td><td align="left">P103</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203445">HQ203445</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203566">HQ203566</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203342">HQ203342</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203264">HQ203264</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203655">HQ203655</ext-link></td></tr><tr><td align="center">49</td><td align="left"><italic>Favia stelligera</italic></td><td align="left">P141</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203446">HQ203446</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203567">HQ203567</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203343">HQ203343</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203265">HQ203265</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203656">HQ203656</ext-link></td></tr><tr><td align="center">50</td><td align="left"><bold><italic>Favia truncatus</italic></bold></td><td align="left">G61897</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203447">HQ203447</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203568">HQ203568</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203344">HQ203344</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203266">HQ203266</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203657">HQ203657</ext-link></td></tr><tr><td align="center">51</td><td align="left"><bold><italic>Favites abdita</italic></bold></td><td align="left">S002</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203448">HQ203448</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203569">HQ203569</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203345">HQ203345</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203267">HQ203267</ext-link></td><td></td></tr><tr><td align="center">52</td><td align="left"><italic>Favites chinensis</italic></td><td align="left">S084</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203449">HQ203449</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203570">HQ203570</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203346">HQ203346</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203268">HQ203268</ext-link></td><td></td></tr><tr><td align="center">53</td><td align="left"><italic>Favites complanata</italic></td><td align="left">S007</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203450">HQ203450</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203571">HQ203571</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203347">HQ203347</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371689">EU371689</ext-link></td><td></td></tr><tr><td align="center">54</td><td align="left"><bold><italic>Favites flexuosa</italic></bold></td><td align="left">P116</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203451">HQ203451</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203572">HQ203572</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203348">HQ203348</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203269">HQ203269</ext-link></td><td></td></tr><tr><td align="center">55</td><td align="left"><italic>Favites halicora</italic></td><td align="left">S115</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203452">HQ203452</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203573">HQ203573</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203349">HQ203349</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203270">HQ203270</ext-link></td><td></td></tr><tr><td align="center">56</td><td align="left"><italic>Favites paraflexuosa</italic></td><td align="left">S100</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203453">HQ203453</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203574">HQ203574</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203350">HQ203350</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371694">EU371694</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345521">FJ345521</ext-link></td></tr><tr><td align="center">57</td><td align="left"><italic>Favites pentagona</italic></td><td align="left">S086</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203454">HQ203454</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203575">HQ203575</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203351">HQ203351</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371695">EU371695</ext-link></td><td></td></tr><tr><td align="center">58</td><td align="left"><italic>Favites pentagona</italic></td><td align="left">P111</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203455">HQ203455</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203576">HQ203576</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203271">HQ203271</ext-link></td><td></td></tr><tr><td align="center">59</td><td align="left"><bold><italic>Favites russelli</italic></bold></td><td align="left">G61895</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203456">HQ203456</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203577">HQ203577</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203352">HQ203352</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203272">HQ203272</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203658">HQ203658</ext-link></td></tr><tr><td align="center">60</td><td align="left"><bold><italic>Favites stylifera</italic></bold></td><td align="left">P128</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203457">HQ203457</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203578">HQ203578</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203353">HQ203353</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203273">HQ203273</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203659">HQ203659</ext-link></td></tr><tr><td align="center">61</td><td align="left"><italic>Goniastrea aspera</italic></td><td align="left">S107</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203458">HQ203458</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203579">HQ203579</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203354">HQ203354</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345430">FJ345430</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345487">FJ345487</ext-link></td></tr><tr><td align="center">62</td><td align="left"><italic>Goniastrea australensis</italic></td><td align="left">G61876</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203459">HQ203459</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203580">HQ203580</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203355">HQ203355</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203274">HQ203274</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203660">HQ203660</ext-link></td></tr><tr><td align="center">63</td><td align="left"><italic>Goniastrea australensis</italic></td><td align="left">S088</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203460">HQ203460</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203581">HQ203581</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203356">HQ203356</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345431">FJ345431</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345490">FJ345490</ext-link></td></tr><tr><td align="center">64</td><td align="left"><italic>Goniastrea australensis</italic></td><td align="left">S098</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203461">HQ203461</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203582">HQ203582</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371696">EU371696</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345491">FJ345491</ext-link></td></tr><tr><td align="center">65</td><td align="left"><italic>Goniastrea edwardsi</italic></td><td align="left">S045</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203462">HQ203462</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203583">HQ203583</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203357">HQ203357</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371697">EU371697</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345492">FJ345492</ext-link></td></tr><tr><td align="center">66</td><td align="left"><italic>Goniastrea edwardsi</italic></td><td align="left">S117</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203463">HQ203463</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203584">HQ203584</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345432">FJ345432</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345493">FJ345493</ext-link></td></tr><tr><td align="center">67</td><td align="left"><italic>Goniastrea favulus</italic></td><td align="left">G61877</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203464">HQ203464</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203585">HQ203585</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203358">HQ203358</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203661">HQ203661</ext-link></td></tr><tr><td align="center">68</td><td align="left"><italic>Goniastrea favulus</italic></td><td align="left">S022</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203465">HQ203465</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203586">HQ203586</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371698">EU371698</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345494">FJ345494</ext-link></td></tr><tr><td align="center">69</td><td align="left"><italic>Goniastrea palauensis</italic></td><td align="left">S021</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203466">HQ203466</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203587">HQ203587</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203359">HQ203359</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371699">EU371699</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345488">FJ345488</ext-link></td></tr><tr><td align="center">70</td><td align="left"><italic>Goniastrea pectinata</italic></td><td align="left">G61879</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203467">HQ203467</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203588">HQ203588</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203360">HQ203360</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203662">HQ203662</ext-link></td></tr><tr><td align="center">71</td><td align="left"><italic>Goniastrea pectinata</italic></td><td align="left">S043</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203468">HQ203468</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203589">HQ203589</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345434">FJ345434</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345489">FJ345489</ext-link></td></tr><tr><td align="center">72</td><td align="left"><italic>Goniastrea pectinata</italic></td><td align="left">P110</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203469">HQ203469</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203590">HQ203590</ext-link></td><td></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203663">HQ203663</ext-link></td></tr><tr><td align="center">73</td><td align="left"><italic>Goniastrea retiformis</italic></td><td align="left">S083</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203470">HQ203470</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203591">HQ203591</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203361">HQ203361</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371700">EU371700</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345527">FJ345527</ext-link></td></tr><tr><td align="center">74</td><td align="left"><italic>Goniastrea retiformis</italic></td><td align="left">P119</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203471">HQ203471</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203592">HQ203592</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203275">HQ203275</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203664">HQ203664</ext-link></td></tr><tr><td align="center">75</td><td align="left"><italic>Hydnophora exesa </italic>(Merulinidae)</td><td align="left">P127</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203472">HQ203472</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203593">HQ203593</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203362">HQ203362</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203276">HQ203276</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203665">HQ203665</ext-link></td></tr><tr><td align="center">76</td><td align="left"><bold><italic>Hydnophora microconos </italic></bold>(Merulinidae)</td><td align="left">P121</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203473">HQ203473</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203594">HQ203594</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203363">HQ203363</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203277">HQ203277</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203666">HQ203666</ext-link></td></tr><tr><td align="center">77</td><td align="left"><bold><italic>Hydnophora pilosa</italic></bold> (Merulinidae)</td><td align="left">P138</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203474">HQ203474</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203595">HQ203595</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203364">HQ203364</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203278">HQ203278</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203667">HQ203667</ext-link></td></tr><tr><td align="center">78</td><td align="left"><italic>Leptoria irregularis</italic></td><td align="left">P133</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203475">HQ203475</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203596">HQ203596</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203279">HQ203279</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203668">HQ203668</ext-link></td></tr><tr><td align="center">79</td><td align="left"><italic>Leptoria phrygia</italic></td><td align="left">S081</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203476">HQ203476</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203597">HQ203597</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203365">HQ203365</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371705">EU371705</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345529">FJ345529</ext-link></td></tr><tr><td align="center">80</td><td align="left"><italic>Lobophyllia corymbosa </italic>(XIX; Mussidae)</td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="AF549237">AF549237</ext-link></td><td></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="AB117241">AB117241</ext-link></td><td></td></tr><tr><td align="center">81</td><td align="left"><italic>Merulina ampliata </italic>(Merulinidae)</td><td align="left">P106</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203477">HQ203477</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203598">HQ203598</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203280">HQ203280</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203669">HQ203669</ext-link></td></tr><tr><td align="center">82</td><td align="left"><italic>Merulina scabricula </italic>(Merulinidae)</td><td align="left">P114</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203478">HQ203478</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203599">HQ203599</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203366">HQ203366</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203281">HQ203281</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203670">HQ203670</ext-link></td></tr><tr><td align="center">83</td><td align="left"><italic>Montastraea annularis</italic></td><td align="left">A622</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203479">HQ203479</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203600">HQ203600</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203367">HQ203367</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203282">HQ203282</ext-link></td><td></td></tr><tr><td align="center">84</td><td align="left"><italic>Montastraea </italic>cf. <italic>annuligera</italic></td><td align="left">P117</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203481">HQ203481</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203602">HQ203602</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203369">HQ203369</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203671">HQ203671</ext-link></td></tr><tr><td align="center">85</td><td align="left"><italic>Montastraea cavernosa </italic>(XVI)</td><td align="left">A005</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203480">HQ203480</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203601">HQ203601</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203368">HQ203368</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203283">HQ203283</ext-link></td><td></td></tr><tr><td align="center">86</td><td align="left"><bold><italic>Montastraea colemani</italic></bold></td><td align="left">P118</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203482">HQ203482</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203603">HQ203603</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203284">HQ203284</ext-link></td><td></td></tr><tr><td align="center">87</td><td align="left"><italic>Montastraea curta</italic></td><td align="left">G61882</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203483">HQ203483</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203604">HQ203604</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203370">HQ203370</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203285">HQ203285</ext-link></td><td></td></tr><tr><td align="center">88</td><td align="left"><italic>Montastraea curta</italic></td><td align="left">P122</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203484">HQ203484</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203605">HQ203605</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203286">HQ203286</ext-link></td><td></td></tr><tr><td align="center">89</td><td align="left"><italic>Montastraea magnistellata</italic></td><td align="left">G61896</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203485">HQ203485</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203606">HQ203606</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203371">HQ203371</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203287">HQ203287</ext-link></td><td></td></tr><tr><td align="center">90</td><td align="left"><italic>Montastraea magnistellata</italic></td><td align="left">P109</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203486">HQ203486</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203607">HQ203607</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203288">HQ203288</ext-link></td><td></td></tr><tr><td align="center">91</td><td align="left"><bold><italic>Montastraea multipunctata</italic></bold></td><td align="left">P131</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203487">HQ203487</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203608">HQ203608</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203372">HQ203372</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203289">HQ203289</ext-link></td><td></td></tr><tr><td align="center">92</td><td align="left"><bold><italic>Montastraea salebrosa</italic></bold></td><td align="left">P139</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203488">HQ203488</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203609">HQ203609</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203373">HQ203373</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203290">HQ203290</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203672">HQ203672</ext-link></td></tr><tr><td align="center">93</td><td align="left"><italic>Montastraea valenciennesi</italic></td><td align="left">G61904</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203489">HQ203489</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203610">HQ203610</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203291">HQ203291</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203673">HQ203673</ext-link></td></tr><tr><td align="center">94</td><td align="left"><italic>Montastraea valenciennesi</italic></td><td align="left">S006</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203490">HQ203490</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203611">HQ203611</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203374">HQ203374</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371713">EU371713</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345514">FJ345514</ext-link></td></tr><tr><td align="center">95</td><td align="left"><italic>Montastraea valenciennesi</italic></td><td align="left">S008</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203491">HQ203491</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203612">HQ203612</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371714">EU371714</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345515">FJ345515</ext-link></td></tr><tr><td align="center">96</td><td align="left"><italic>Montastraea valenciennesi</italic></td><td align="left">P102</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203492">HQ203492</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203613">HQ203613</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203375">HQ203375</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203292">HQ203292</ext-link></td><td></td></tr><tr><td align="center">97</td><td align="left"><bold><italic>Moseleya latistellata</italic></bold></td><td align="left">G61909</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203493">HQ203493</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203614">HQ203614</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203376">HQ203376</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203293">HQ203293</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203674">HQ203674</ext-link></td></tr><tr><td align="center">98</td><td align="left"><italic>Mussa angulosa </italic>(XXI; Mussidae)</td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="AF549236">AF549236</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="AB441402">AB441402</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="NC_008163">NC_008163</ext-link></td><td></td></tr><tr><td align="center">99</td><td align="left"><italic>Mycedium elephantotus </italic>(Pectiniidae)</td><td align="left">S121</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203494">HQ203494</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203615">HQ203615</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203377">HQ203377</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203294">HQ203294</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203675">HQ203675</ext-link></td></tr><tr><td align="center">100</td><td align="left"><bold><italic>Mycedium robokaki </italic></bold>(Pectiniidae)</td><td align="left">S126</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203495">HQ203495</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203616">HQ203616</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203378">HQ203378</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203295">HQ203295</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203676">HQ203676</ext-link></td></tr><tr><td align="center">101</td><td align="left"><italic>Oulophyllia bennettae</italic></td><td align="left">G61873</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203496">HQ203496</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203617">HQ203617</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203296">HQ203296</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203677">HQ203677</ext-link></td></tr><tr><td align="center">102</td><td align="left"><italic>Oulophyllia bennettae</italic></td><td align="left">S033</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203497">HQ203497</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203618">HQ203618</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203379">HQ203379</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345436">FJ345436</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345497">FJ345497</ext-link></td></tr><tr><td align="center">103</td><td align="left"><italic>Oulophyllia </italic>aff. <italic>bennettae</italic></td><td align="left">P140</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203498">HQ203498</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203619">HQ203619</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203380">HQ203380</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203297">HQ203297</ext-link></td><td></td></tr><tr><td align="center">104</td><td align="left"><italic>Oulophyllia crispa</italic></td><td align="left">S055</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203499">HQ203499</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203620">HQ203620</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203381">HQ203381</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371721">EU371721</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345500">FJ345500</ext-link></td></tr><tr><td align="center">105</td><td align="left"><italic>Pectinia alcicornis </italic>(Pectiniidae)</td><td align="left">P124</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203500">HQ203500</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203621">HQ203621</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203382">HQ203382</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203298">HQ203298</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203678">HQ203678</ext-link></td></tr><tr><td align="center">106</td><td align="left"><bold><italic>Pectinia ayleni</italic></bold> (Pectiniidae)</td><td align="left">S122</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203501">HQ203501</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203622">HQ203622</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203383">HQ203383</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203299">HQ203299</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203679">HQ203679</ext-link></td></tr><tr><td align="center">107</td><td align="left"><bold><italic>Pectinia lactuca </italic></bold>(Pectiniidae)</td><td align="left">P115</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203502">HQ203502</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203623">HQ203623</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203384">HQ203384</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203300">HQ203300</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203680">HQ203680</ext-link></td></tr><tr><td align="center">108</td><td align="left"><italic>Pectinia paeonia </italic>(Pectiniidae)</td><td align="left">P126</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203503">HQ203503</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203624">HQ203624</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203385">HQ203385</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203301">HQ203301</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203681">HQ203681</ext-link></td></tr><tr><td align="center">109</td><td align="left"><bold><italic>Platygyra acuta</italic></bold></td><td align="left">P123</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203504">HQ203504</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203625">HQ203625</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203386">HQ203386</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203682">HQ203682</ext-link></td></tr><tr><td align="center">110</td><td align="left"><bold><italic>Platygyra contorta</italic></bold></td><td align="left">P112</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203505">HQ203505</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203626">HQ203626</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203387">HQ203387</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203683">HQ203683</ext-link></td></tr><tr><td align="center">111</td><td align="left"><italic>Platygyra daedalea</italic></td><td align="left">G61878</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203506">HQ203506</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203627">HQ203627</ext-link></td><td></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203684">HQ203684</ext-link></td></tr><tr><td align="center">112</td><td align="left"><italic>Platygyra daedalea</italic></td><td align="left">S116</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203507">HQ203507</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203628">HQ203628</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203388">HQ203388</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345440">FJ345440</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345530">FJ345530</ext-link></td></tr><tr><td align="center">113</td><td align="left"><italic>Platygyra lamellina</italic></td><td align="left">G61887</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203508">HQ203508</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203629">HQ203629</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203389">HQ203389</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203302">HQ203302</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203685">HQ203685</ext-link></td></tr><tr><td align="center">114</td><td align="left"><italic>Platygyra lamellina</italic></td><td align="left">S114</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203509">HQ203509</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203630">HQ203630</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345441">FJ345441</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345531">FJ345531</ext-link></td></tr><tr><td align="center">115</td><td align="left"><italic>Platygyra pini</italic></td><td align="left">G61899</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203510">HQ203510</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203631">HQ203631</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203390">HQ203390</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203303">HQ203303</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203686">HQ203686</ext-link></td></tr><tr><td align="center">116</td><td align="left"><italic>Platygyra pini</italic></td><td align="left">S035</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203511">HQ203511</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203632">HQ203632</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203391">HQ203391</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345443">FJ345443</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345535">FJ345535</ext-link></td></tr><tr><td align="center">117</td><td align="left"><bold><italic>Platygyra ryukyuensis</italic></bold></td><td align="left">P101</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203512">HQ203512</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203633">HQ203633</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203392">HQ203392</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203304">HQ203304</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203687">HQ203687</ext-link></td></tr><tr><td align="center">118</td><td align="left"><italic>Platygyra sinensis</italic></td><td align="left">S118</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203513">HQ203513</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203634">HQ203634</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203393">HQ203393</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345442">FJ345442</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345534">FJ345534</ext-link></td></tr><tr><td align="center">119</td><td align="left"><italic>Platygyra sinensis</italic></td><td align="left">P130</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203514">HQ203514</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203635">HQ203635</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203305">HQ203305</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203688">HQ203688</ext-link></td></tr><tr><td align="center">120</td><td align="left"><italic>Platygyra </italic>cf. <italic>verweyi</italic></td><td align="left">S037</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203515">HQ203515</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203636">HQ203636</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203394">HQ203394</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="EU371722">EU371722</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345532">FJ345532</ext-link></td></tr><tr><td align="center">121</td><td align="left"><italic>Plesiastrea versipora </italic>(XIV)</td><td align="left">S127</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203397">HQ203397</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203518">HQ203518</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203307">HQ203307</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203246">HQ203246</ext-link></td><td></td></tr><tr><td align="center">122</td><td align="left"><italic>Plesiastrea versipora </italic>(XIV)</td><td align="left">P137</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203398">HQ203398</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203519">HQ203519</ext-link></td><td></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203247">HQ203247</ext-link></td><td></td></tr><tr><td align="center">123</td><td align="left"><italic>Scapophyllia cylindrica </italic>(Merulinidae)</td><td align="left">S060</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203516">HQ203516</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203637">HQ203637</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203395">HQ203395</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345444">FJ345444</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="FJ345502">FJ345502</ext-link></td></tr><tr><td align="center">124</td><td align="left"><italic>Trachyphyllia geoffroyi </italic>(Trachyphylliidae)</td><td align="left">J001</td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203517">HQ203517</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203638">HQ203638</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203396">HQ203396</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203306">HQ203306</ext-link></td><td align="center"><ext-link ext-link-type="gen" xlink:href="HQ203689">HQ203689</ext-link></td></tr></tbody></table><table-wrap-foot><p>Unless indicated by roman numerals and/or family names in parentheses, all species belong to clade XVII and Faviidae, respectively. Species placed in a molecular phylogenetic context for the first time are in bold. Specimens with voucher numbers starting with 'G' are from Great Barrier Reef (Australia), 'S' from Singapore, 'J' from Japan, 'P' from the Philippines, and 'A' from the Atlantic. GenBank accession numbers are displayed for each molecular marker.</p></table-wrap-foot></table-wrap><p>For each colony from Singapore, Japan and the Caribbean, DNA was extracted from ~2 cm<sup>2 </sup>of tissue digested in twice their volume of CHAOS solution (not an acronym; 4 M guanidine thiocyanate, 0.1% N-lauroyl sarcosine sodium, 10 mM Tris pH 8, 0.1 M 2-mercaptoethanol) for at least three days at room temperature before DNA extraction using a phenol-chloroform based method with a phenol extraction buffer (100 mM TrisCl pH 8, 10 mM EDTA, 0.1% SDS) [<xref ref-type="bibr" rid="B15">15</xref>,<xref ref-type="bibr" rid="B22">22</xref>-<xref ref-type="bibr" rid="B24">24</xref>]. For specimens from Australia and the Philippines, genomic DNA was extracted from the tissues preserved in ethanol using the Qiagen DNeasy kit, following the manufacturer's instructions.</p><p>The rest of the colony was sprayed with a powerful water jet to remove as much tissue as possible before being bleached in 5-10% sodium hypochlorite solution. The skeletons were rinsed in fresh water, dried, and deposited in the Raffles Museum of Biodiversity Research (Singapore), Seto Marine Biological Laboratory (Wakayama, Japan), Museum of Tropical Queensland (Australia), and De La Salle University (Manila, The Philippines) (Table <xref ref-type="table" rid="T1">1</xref>).</p></sec><sec><title>PCR amplification and sequencing</title><p>A total of five molecular markers were amplified for a majority of the samples (Tables <xref ref-type="table" rid="T1">1</xref> and <xref ref-type="table" rid="T2">2</xref>). They consist of three nuclear and two mitochondrial loci: (1) 28S rDNA D1 and D2 fragments; (2) histone H3; (3) internal transcribed spacers 1 and 2, including 5.8S rDNA (ITS in short); (4) cytochrome oxidase subunit I (COI); and (5) noncoding intergenic region situated between COI and the formylmethionine transfer RNA gene (IGR in short) [<xref ref-type="bibr" rid="B8">8</xref>,<xref ref-type="bibr" rid="B23">23</xref>,<xref ref-type="bibr" rid="B25">25</xref>-<xref ref-type="bibr" rid="B27">27</xref>].</p><table-wrap id="T2" position="float"><label>Table 2</label><caption><p>Molecular markers utilized for phylogenetic reconstruction.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left">Marker</th><th align="left">Primer pairs</th><th align="left">Total characters (informative)</th><th align="left">Model</th><th align="left">Source</th></tr></thead><tbody><tr><td align="left">28S rDNA</td><td align="left">C1': 5'-ACC CGC TGA ATT TAA GCA T-3'<break></break>D2MAD: 5'-GAC GAT CGA TTT GCA CGT CA-3'</td><td align="left">861 (135)</td><td align="left">HKY+Γ</td><td align="left">[<xref ref-type="bibr" rid="B25">25</xref>]</td></tr><tr><td align="left">histone H3</td><td align="left">H3F: 5'-ATG GCT CGT ACC AAG CAG ACV GC-3'<break></break>H3R: 5'-ATA TCC TTR GGC ATR ATR GTG AC-3'</td><td align="left">374 (73)</td><td align="left">HKY+Γ</td><td align="left">[<xref ref-type="bibr" rid="B26">26</xref>]</td></tr><tr><td align="left">ITS rDNA</td><td align="left">A18S: 5'-GATCGAACGGTTTAGTGAGG-3'<break></break>ITS-4: 5'-TCCTCCGCTTATTGATATGC-3'</td><td align="left">1137 (425)</td><td align="left">SYM+Γ</td><td align="left">[<xref ref-type="bibr" rid="B27">27</xref>]</td></tr><tr><td align="left">mt COI</td><td align="left">MCOIF: 5'-TCTACAAATCATAAAGACATAGG-3'<break></break>MCOIR: 5'-GAGAAATTATACCAAAACCAGG-3'</td><td align="left">719 (71)</td><td align="left">HKY+I</td><td align="left">[<xref ref-type="bibr" rid="B8">8</xref>]</td></tr><tr><td align="left">mt IGR</td><td align="left">MNC1f: 5'-GAGCTGGGCTTCTTTAGAGTG-3'<break></break>MNC1r: 5'-GTGAGACTCGAACTCACTTTTC-3'</td><td align="left">1509 (763)</td><td align="left">SYM+I</td><td align="left">[<xref ref-type="bibr" rid="B23">23</xref>]</td></tr></tbody></table></table-wrap><p>The mitochondrial intergenic region (IGR) was too variable to be aligned across the entire clade, so only alignable sequences were included in the analysis. ITS comprises multiple copies in the nuclear genome, but the primers we used have shown high fidelity for a single copy, precluding the need to clone the amplicons [<xref ref-type="bibr" rid="B27">27</xref>-<xref ref-type="bibr" rid="B33">33</xref>]. Nevertheless, in the unlikely case that paralogs were sequenced, our analyses could be confused by incomplete lineage sorting [<xref ref-type="bibr" rid="B7">7</xref>]. We therefore sequenced the ITS locus from at most one representative of each species, unless analyses of the other four markers did not recover its sequences as a clade. In the latter case, sequences may actually belong to separate cryptic species that have been obscured by gross morphological similarities. For COI, not all specimens of each species were necessarily sequenced since intraspecific variation of this gene is limited [<xref ref-type="bibr" rid="B15">15</xref>,<xref ref-type="bibr" rid="B24">24</xref>].</p><p>PCR products were purified with ExoSAP-IT (GE Healthcare, Uppsala, Sweden) and sequencing was performed by Advanced Studies in Genomics, Proteomics and Bioinformatics (ASGPB) at the University of Hawaii at Manoa using the Applied Biosystems BigDye Terminator kit and an ABI 3730XL sequencer. New sequences were deposited in GenBank under accession numbers <ext-link ext-link-type="gen" xlink:href="HQ203246">HQ203246</ext-link>-<ext-link ext-link-type="gen" xlink:href="HQ203689">HQ203689</ext-link> (Table <xref ref-type="table" rid="T1">1</xref>).</p></sec><sec><title>Phylogenetic analyses</title><p>Sequences were organized into five separate data matrices using Mesquite 2.72 [<xref ref-type="bibr" rid="B34">34</xref>], and each aligned with the accurate alignment option (E-INS-i) in MAFFT 6.7 [<xref ref-type="bibr" rid="B35">35</xref>-<xref ref-type="bibr" rid="B37">37</xref>] under default parameters. Substitution saturation of protein-coding genes was assessed via DAMBE [<xref ref-type="bibr" rid="B38">38</xref>,<xref ref-type="bibr" rid="B39">39</xref>], where we found histone H3 and COI to be unsaturated at the third codon positions for tree inference. Consequently, we concatenated the five gene matrices into a single partitioned matrix consisting of 4600 characters, 1467 of which were parsimony informative. This was analyzed using maximum parsimony, Bayesian likelihood, and maximum likelihood methods. We also carried out these analyses on a four-gene dataset omitting the ITS partition to determine if the phylogenetic reconstruction was sensitive to the ITS sampling strategy.</p><p>Under a maximum parsimony framework, we utilized new search technologies [<xref ref-type="bibr" rid="B40">40</xref>,<xref ref-type="bibr" rid="B41">41</xref>] in the software TNT 1.1 [<xref ref-type="bibr" rid="B42">42</xref>,<xref ref-type="bibr" rid="B43">43</xref>]. Tree searches consisted of 50000 random addition sequence replicates under the default sectorial, ratchet, drift and tree fusing parameters. Gaps were treated as missing data and clade stability was inferred using 1000 bootstrap replicates each employing 100 random addition sequences.</p><p>For maximum likelihood, neighbor-joining and Bayesian analyses, we determined the most suitable model of molecular evolution for each gene partition and the concatenated matrix using jModelTest 0.1.1 [<xref ref-type="bibr" rid="B44">44</xref>,<xref ref-type="bibr" rid="B45">45</xref>] to test for a total of 24 models, following the Akaike Information Criterion (AIC). The maximum likelihood tree for each partition and the combined dataset was inferred using RAxML 7.2.3 [<xref ref-type="bibr" rid="B46">46</xref>,<xref ref-type="bibr" rid="B47">47</xref>] at the Cyberinfrastructure for Phylogenetic Research (CIPRES; <ext-link ext-link-type="uri" xlink:href="http://www.phylo.org">http://www.phylo.org</ext-link>), employing the GTRGAMMA model. The proportion of invariable sites and gamma distribution shape parameter for variable sites were estimated during the maximum likelihood analysis. Multiparametric bootstrap analysis was carried out using 1000 bootstrap replicates. Maximum likelihood analysis was also carried out with PhyML 3.0 [<xref ref-type="bibr" rid="B45">45</xref>] on the combined data, utilizing the AIC-chosen model (GTR+I+Γ), and generating 1000 bootstrap replicates. The neighbor-joining tree of the combined data was calculated in PAUP*4.0b10 [<xref ref-type="bibr" rid="B48">48</xref>] with 1000 bootstrap replicates, employing the evolutionary model selected above.</p><p>Bayesian inference was carried out in MrBayes 3.1.2 [<xref ref-type="bibr" rid="B49">49</xref>,<xref ref-type="bibr" rid="B50">50</xref>], using the resources of the Computational Biology Service Unit from Cornell University, with each partition modeled (Table <xref ref-type="table" rid="T2">2</xref>) but unlinked for separate parameter estimations. Four Markov chains of 10 million generations were implemented in twelve runs, saving a tree every 100th generation. MCMC convergence among the runs was monitored using Tracer 1.5 [<xref ref-type="bibr" rid="B51">51</xref>], where we ascertained that only four of the twelve runs converged on the shortest trees (only two runs converged for the four-gene analysis; see [<xref ref-type="bibr" rid="B52">52</xref>-<xref ref-type="bibr" rid="B54">54</xref>]), and the first 40001 trees were to be discarded as burn-in.</p><p>Additionally, compensatory base changes because of the secondary structure of the ITS rDNA loci may lead to non-independence and increased homoplasy of characters [<xref ref-type="bibr" rid="B55">55</xref>-<xref ref-type="bibr" rid="B57">57</xref>]. Hence, analysis of the secondary structure of this region may result in a more rigorous phylogeny [<xref ref-type="bibr" rid="B58">58</xref>-<xref ref-type="bibr" rid="B61">61</xref>]. Using the ITS2 segment of each ITS sequence, secondary structure was predicted by searching the ITS2 database [<xref ref-type="bibr" rid="B62">62</xref>] for the best match template and then modeling its structure based on free energy minimization. The ITS2 sequences and their associated structural information were aligned using 4SALE 1.5 [<xref ref-type="bibr" rid="B63">63</xref>,<xref ref-type="bibr" rid="B64">64</xref>], and then exported for analysis in ProfDistS 0.9.8 [<xref ref-type="bibr" rid="B65">65</xref>-<xref ref-type="bibr" rid="B68">68</xref>]. The profile neighbor-joining algorithm was executed with 10000 bootstrap replicates on the RNA structural alignment, using the GTR model and rate matrix 'Q_ITS2.txt' for distance correction. ITS2 could not be amplified from <italic>Hydnophora microconos</italic>, <italic>H. pilosa </italic>and <italic>Merulina scabricula</italic>. Consequently these species were excluded from the analysis.</p></sec></sec><sec><title>Results and Discussion</title><p>In this study, the evolutionary history of the 'Bigmessidae' corals was robustly reconstructed using five genes. Relations among other clade representatives chosen as outgroups were also inferred. The maximum likelihood reconstructions carried out by RAxML 7.2.3 and PhyML 3.0 had log likelihood values of -36224.67 and -36995.48, respectively. As they were identical when considering nodes with bootstrap values ≥50, we present the RAxML tree that garnered a higher likelihood score (Figures <xref ref-type="fig" rid="F1">1</xref> and <xref ref-type="fig" rid="F2">2</xref>). A total of 182 most parsimonious trees (tree length = 6178) were obtained. No conflicts between tree optimization procedures (including Bayesian inference and the neighbor-joining algorithm) were apparent when considering only the supported nodes (bootstrap ≥50 and posterior probability ≥0.9) (see Additional file <xref ref-type="supplementary-material" rid="S2">2</xref>). Analyses excluding the ITS partition also gave congruent results. Several clades were consistent and well supported among maximum likelihood, parsimony and Bayesian inferences. We named some of these groups within clade XVII from A to I, consistent with the classification in Budd and Stolarski [<xref ref-type="bibr" rid="B69">69</xref>]. On the other hand, the neighbor-joining method generated a relatively unresolved tree—subclades A, C, F and I did not achieve bootstrap values of ≥50 (see Additional file <xref ref-type="supplementary-material" rid="S2">2</xref>).</p><fig id="F1" position="float"><label>Figure 1</label><caption><p><bold>Maximum likelihood tree of the combined molecular data</bold>. Species have been summarized into genera where possible. One asterisk denotes paraphyletic genus, two asterisks polyphyly, and three represents a genus that is both para- and polyphyletic. All taxa from conventional family Faviidae unless otherwise indicated. Clade designations XIV to XXI shown; clade XVII divided into well-supported subclades. Numbers adjacent to branches/taxa are support values (maximum likelihood bootstrap ≥50, maximum parsimony bootstrap ≥50, followed by Bayesian posterior probability ≥0.9). Filled circles indicate well-supported clades (bootstrap values ≥98 and posterior probability of 1).</p></caption><graphic xlink:href="1471-2148-11-37-1"></graphic></fig><fig id="F2" position="float"><label>Figure 2</label><caption><p><bold>Maximum likelihood topologies of each subclade</bold>. Numbers above branches are maximum likelihood bootstrap ≥50 and Bayesian posterior probability ≥0.9, while number below denotes maximum parsimony bootstrap ≥50. Family classification follows definitions given for Figure 1. Type species of genera are in bold.</p></caption><graphic xlink:href="1471-2148-11-37-2"></graphic></fig><p>The combined five-gene data yielded the most resolved phylogeny hitherto of clade XVII, with most branches garnering high support values. However, most partitions gave fairly unresolved trees when analyzed individually (see Additional file <xref ref-type="supplementary-material" rid="S3">3</xref>). By examining the support of subclades among trees obtained via different partitions, we found that nuclear markers contributed a greater extent to the final tree topology (Table <xref ref-type="table" rid="T3">3</xref>). Histone H3, for instance, supported all higher-level groupings and all subclades except D/E (Figure <xref ref-type="fig" rid="F1">1</xref>). The 28S and ITS rDNA gene trees had moderate resolution within clade XVII, with only two unresolved subclades each. Surprisingly, the tree based on ITS2 rDNA secondary structure had less resolution than the primary sequence alignment. Indeed, the former has demonstrated potential for resolving intrageneric phylogenies in other anthozoans [<xref ref-type="bibr" rid="B70">70</xref>,<xref ref-type="bibr" rid="B71">71</xref>], but it is less informative for relationships at higher taxonomic levels [<xref ref-type="bibr" rid="B72">72</xref>,<xref ref-type="bibr" rid="B73">73</xref>]. Evidently, the COI tree was poorly resolved, with ≥50 bootstrap support for few relationships among major clades and only one subclade. The slow evolution of the mitochondrial COI gene among anthozoans is certainly the reason behind this [<xref ref-type="bibr" rid="B24">24</xref>,<xref ref-type="bibr" rid="B74">74</xref>,<xref ref-type="bibr" rid="B75">75</xref>]. While the intergenic marker (IGR) adjacent to COI on the mitochondrial genome has shown promise for phylogenetic reconstruction among Faviidae and Mussidae [<xref ref-type="bibr" rid="B15">15</xref>,<xref ref-type="bibr" rid="B23">23</xref>,<xref ref-type="bibr" rid="B76">76</xref>], it cannot be unambiguously aligned between the major clades. We urge the development of more nuclear phylogenetic markers that can be reliably applied across diverse scleractinian clades.</p><table-wrap id="T3" position="float"><label>Table 3</label><caption><p>Clades supported by maximum likelihood analysis for each partition.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left">Clade</th><th align="center">Nuclear DNA</th><th align="center">mt DNA</th><th align="center">28S rDNA</th><th align="center">histone H3</th><th align="center">ITS<break></break>sequence</th><th align="center">ITS<break></break>structure</th><th align="center">mt COI</th><th align="center">mt IGR</th></tr></thead><tbody><tr><td align="left">XV to XXI</td><td align="center">√√</td><td align="center">√√</td><td align="center">√√</td><td align="center">√√</td><td align="center">√√</td><td align="center">√√</td><td align="center">√√</td><td></td></tr><tr><td align="left">XV+XVI</td><td align="center">√√</td><td align="center">X</td><td align="center">√√</td><td align="center">√√</td><td align="center">√</td><td align="center">√√</td><td align="center">XX</td><td></td></tr><tr><td align="left">XVII to XXI</td><td align="center">√√</td><td align="center">√√</td><td align="center">√</td><td align="center">√√</td><td align="center">√√</td><td align="center">√</td><td align="center">√√</td><td></td></tr><tr><td align="left">XXI</td><td align="center">√√</td><td align="center">√√</td><td align="center">√</td><td></td><td></td><td></td><td align="center">√√</td><td></td></tr><tr><td align="left">XIX+XX<sup>1</sup></td><td align="center">√√</td><td align="center">√</td><td align="center">√</td><td align="center">√√</td><td align="center">X</td><td align="center">√√</td><td align="center">√</td><td></td></tr><tr><td align="left">XVII</td><td align="center">√√</td><td align="center">X</td><td align="center">√</td><td align="center">√√</td><td align="center">√</td><td align="center">X</td><td align="center">X</td><td align="center">√√</td></tr><tr><td align="left">XVII-A</td><td align="center">√√</td><td align="center">X</td><td align="center">√√</td><td align="center">√√</td><td align="center">√√</td><td align="center">X</td><td align="center">X</td><td align="center">X</td></tr><tr><td align="left">XVII-B</td><td align="center">√√</td><td align="center">X</td><td align="center">X</td><td align="center">√√</td><td align="center">√√</td><td align="center">√√</td><td align="center">X</td><td align="center">√</td></tr><tr><td align="left">XVII-C</td><td align="center">√√</td><td align="center">XX</td><td align="center">√√</td><td align="center">√√</td><td align="center">√√</td><td align="center">X</td><td align="center">X</td><td></td></tr><tr><td align="left">XVII-D/E</td><td align="center">√√</td><td align="center">XX</td><td align="center">X</td><td align="center">X</td><td align="center">√√</td><td align="center">√</td><td align="center">XX</td><td align="center">√√</td></tr><tr><td align="left">XVII-F</td><td align="center">√√</td><td align="center">X</td><td align="center">√√</td><td align="center">√√</td><td align="center">X</td><td align="center">√√</td><td align="center">XX</td><td></td></tr><tr><td align="left">XVII-G</td><td align="center">√√</td><td align="center">√√</td><td align="center">√√</td><td align="center">√√</td><td align="center">X</td><td align="center">X</td><td align="center">√</td><td align="center">√√</td></tr><tr><td align="left">XVII-H</td><td align="center">√√</td><td align="center">X</td><td align="center">√√</td><td align="center">√√</td><td align="center">√√</td><td></td><td align="center">√√</td><td align="center">√√</td></tr><tr><td align="left">XVII-I<sup>2</sup></td><td align="center">√√</td><td align="center">X</td><td align="center">√√</td><td align="center">√√</td><td align="center">√√</td><td align="center">√</td><td align="center">X</td><td align="center">X</td></tr></tbody></table><table-wrap-foot><p><sup>1</sup><italic>Montastraea multipunctata </italic>and <italic>Moseleya latistellata </italic>are herein considered as part of clade XIX+XX.</p><p><sup>2</sup>Subclade I is expanded to include <italic>Montastraea salebrosa</italic>.</p><p>'√√': clade present with ≥50 bootstrap support; '√': clade present but not supported (<50 bootstrap); 'XX': contradicted clade with ≥50 bootstrap support; and 'X': contradicted clade not supported. Empty cells indicate insufficient data.</p></table-wrap-foot></table-wrap><p>Most relationships among clades XV to XXI obtained in this study corroborate results of Fukami et al. [<xref ref-type="bibr" rid="B12">12</xref>] (Figure <xref ref-type="fig" rid="F1">1</xref>). The only difference occurs in the sister grouping of <italic>Diploastrea heliopora </italic>(XV) and <italic>Montastraea cavernosa </italic>(XVI) (supported by all analyses except Bayesian likelihood) that form a grade in Fukami et al. [<xref ref-type="bibr" rid="B12">12</xref>]. The monophyly of the clade XVII+XIX+XX (Pacific faviids and mussids) is recovered but not well supported. <italic>Montastraea multipunctata </italic>and <italic>Moseleya latistellata </italic>are Pacific faviids, and therefore presumably in clade XVII. But as a result of superficial similarities, they have historically been associated with the Pacific mussids <italic>Blastomussa merleti </italic>(clade XIV) [<xref ref-type="bibr" rid="B77">77</xref>] and <italic>Acanthastrea hillae </italic>(clade XVIII) [<xref ref-type="bibr" rid="B5">5</xref>,<xref ref-type="bibr" rid="B18">18</xref>], respectively. Here, we find them to be more closely related to clades XIX and XX instead, revealing a taxonomic situation more challenging than anticipated. Pacific faviids other than <italic>Diploastrea heliopora </italic>can no longer be restricted to clade XVII, and the possibility exists that yet-to-be sampled taxa provisionally placed in clade XVII—particularly the monotypic genera, <italic>Australogyra</italic>, <italic>Erythrastrea</italic>, <italic>Boninastrea </italic>and <italic>Paraclavarina</italic>—have unexpected affinities.</p><p>Nested within clade XVII are four conventional families—Faviidae, Merulinidae, Pectiniidae and Trachyphylliidae (Figure <xref ref-type="fig" rid="F1">1</xref>). Two Pectiniidae genera, <italic>Pectinia </italic>and <italic>Mycedium </italic>(XVII-E) form the sister clade to <italic>Oulophyllia</italic>. This is a similar relationship to the results of Fukami et al. [<xref ref-type="bibr" rid="B12">12</xref>], although here we also show with reasonable support that <italic>Oulophyllia </italic>is monophyletic, and <italic>Caulastraea </italic>is an outgroup rather than nested within <italic>Oulophyllia </italic>(XVII-D). Merulinidae is represented by <italic>Hydnophora</italic>, <italic>Merulina </italic>and <italic>Scapophyllia</italic>. <italic>Hydnophora </italic>is more closely related to <italic>Favites </italic>and Pacific <italic>Montastraea </italic>spp. than <italic>Merulina </italic>and <italic>Scapophyllia</italic>, which form a grade within the clade dominated by <italic>Goniastrea</italic>. The monospecific Trachyphylliidae is nested within the clade consisting primarily of <italic>Favia </italic>spp., and is sister to <italic>Favia lizardensis </italic>and <italic>F. truncatus </italic>(Figure <xref ref-type="fig" rid="F2">2</xref>). Work is ongoing to redescribe clade XVII by incorporating the above families and applying a new taxon name since the type species of Faviidae, <italic>Favia fragum </italic>(Esper, 1797), belongs to clade XXI [<xref ref-type="bibr" rid="B12">12</xref>].</p><p>The genetic affiliation of <italic>Hydnophora </italic>and <italic>Trachyphyllia </italic>with Faviidae has previously been proposed by Fukami et al. [<xref ref-type="bibr" rid="B8">8</xref>,<xref ref-type="bibr" rid="B12">12</xref>]. However, this is not exclusively a molecular hypothesis. Based on a combination of colony, corallite and subcorallite characters (e.g., polyp budding; wall, septal and columellar structures), Vaughan and Wells, 1943 [<xref ref-type="bibr" rid="B78">78</xref>], placed the two taxa within Faviidae. But later, Chevalier, 1975 [<xref ref-type="bibr" rid="B79">79</xref>], attempted to distinguish <italic>Trachyphyllia </italic>from Faviidae based on minor differences in wall and septal structures by elevating it to the rank of family. Correspondingly, Veron, 1985 [<xref ref-type="bibr" rid="B17">17</xref>], moved <italic>Hydnophora </italic>into Merulinidae because of <italic>Hydnophora </italic>species' macromorphological similarities (i.e., colony growth form and polyp structure) with <italic>Merulina ampliata </italic>and <italic>Scapophyllia cylindrica</italic>, which are genetically in the same lineage (subclade A) as several <italic>Goniastrea </italic>spp. and <italic>Favia stelligera </italic>(Figures <xref ref-type="fig" rid="F1">1</xref> and <xref ref-type="fig" rid="F2">2</xref>; see also [<xref ref-type="bibr" rid="B8">8</xref>,<xref ref-type="bibr" rid="B12">12</xref>]).</p><p><italic>Montastraea annularis </italic>and likely other members of the species complex (<italic>M. faveolata </italic>and <italic>M. franksi</italic>) are the only Atlantic species in clade XVII (see also [<xref ref-type="bibr" rid="B8">8</xref>,<xref ref-type="bibr" rid="B12">12</xref>]). Most significantly here, <italic>M. annularis </italic>is sister to <italic>Cyphastrea</italic>, forming clade XVII-C (Figure <xref ref-type="fig" rid="F1">1</xref>). This placement may seem bizarre in the context of traditional macromorphological characters used to classify scleractinians (e.g., [<xref ref-type="bibr" rid="B4">4</xref>,<xref ref-type="bibr" rid="B78">78</xref>]). However, recent work at the microstructural scale (centers of rapid accretion and thickening deposits) has suggested that their septothecal walls (formed by fusion of outer margins of septa) may unite the two taxa [<xref ref-type="bibr" rid="B69">69</xref>] (see also [<xref ref-type="bibr" rid="B80">80</xref>]). These subcorallite features appear to be appropriate synapomorphies for other subclades. For instance, clade XVII-A consists of <italic>Merulina</italic>, <italic>Scapophyllia</italic>, <italic>Goniastrea </italic>A and <italic>Favia stelligera </italic>(Figure <xref ref-type="fig" rid="F2">2</xref>). At the corallite level, these corals cannot be reconciled within the same taxon, since <italic>Favia stelligera </italic>corallites have single centers with separate walls (plocoid), <italic>Goniastrea </italic>spp. have fused walls (cerioid) and may form valleys (meandroid), while <italic>Merulina </italic>and <italic>Scapophyllia </italic>are composed predominantly of elongated valleys (see Additional file <xref ref-type="supplementary-material" rid="S1">1</xref>). On the other hand, they share the apomorphy of having septothecal walls with abortive septa (thin bands between normal septa with their own centers of rapid accretion).</p><p>The use of macromorphology for identifying 'Bigmessidae' species is known for being problematic as most of these characters are homoplasious [<xref ref-type="bibr" rid="B15">15</xref>,<xref ref-type="bibr" rid="B80">80</xref>,<xref ref-type="bibr" rid="B81">81</xref>]. The ability to distinguish clades based on microstructural features is encouraging for scleractinian systematics. Micromorphology, at the scale of septal teeth and granules, has also exhibited promise as phylogenetic characters [<xref ref-type="bibr" rid="B25">25</xref>,<xref ref-type="bibr" rid="B80">80</xref>,<xref ref-type="bibr" rid="B82">82</xref>-<xref ref-type="bibr" rid="B85">85</xref>]. Interestingly, in light of recent molecular hypotheses, other biological traits, in particular, sexuality and to a lesser extent, breeding mode appear highly conserved and could be further developed as phylogenetic markers [<xref ref-type="bibr" rid="B86">86</xref>,<xref ref-type="bibr" rid="B87">87</xref>].</p><p>Prior to the use of molecular data to build evolutionary trees, it was a great challenge to determine which morphological characters could be useful for classification, given their intraspecific variability [<xref ref-type="bibr" rid="B32">32</xref>,<xref ref-type="bibr" rid="B88">88</xref>] and phenotypic plasticity [<xref ref-type="bibr" rid="B89">89</xref>-<xref ref-type="bibr" rid="B94">94</xref>]. Indeed, the general anthozoan body plan is relatively simple, and scleractinians in particular have few discrete morphological characters that are known to be phylogenetically informative at the polyp level [<xref ref-type="bibr" rid="B4">4</xref>,<xref ref-type="bibr" rid="B95">95</xref>-<xref ref-type="bibr" rid="B97">97</xref>]. As a result of the recent disarray in coral systematics, morphological taxonomies of scleractinians have been heavily criticized (e.g., [<xref ref-type="bibr" rid="B8">8</xref>,<xref ref-type="bibr" rid="B12">12</xref>,<xref ref-type="bibr" rid="B98">98</xref>,<xref ref-type="bibr" rid="B99">99</xref>]). Molecular characters, which are much more numerous and arguably neutrally evolving, can certainly aid our understanding of evolutionary relationships. However, morphological evidence supporting various molecular clades in the present analysis suggests that morphology at novel scales will play an essential role in the taxonomy of 'Bigmessidae' [<xref ref-type="bibr" rid="B80">80</xref>].</p><p>Widespread sampling in this study has shown that corals thought to belong to the same species across the central Indo-Pacific are actually from distinct lineages. Consider <italic>Goniastrea australensis </italic>(Milne Edwards and Haime, 1857), which occurs in two clades (Figures <xref ref-type="fig" rid="F1">1</xref> and <xref ref-type="fig" rid="F2">2</xref>; see also Additional file <xref ref-type="supplementary-material" rid="S1">1</xref>). Since this species was first described from Australia, the Australian specimen that clustered with <italic>Favites russelli </italic>and <italic>Montastraea curta </italic>should be considered <italic>G. australensis</italic>, while the two specimens from Singapore (S088 and S098, subclade A) probably represent new species yet to be described. This is certainly not an isolated case. A similar situation is revealed for <italic>Montastraea valenciennesi</italic>. Specimens from Australia (G61904) and Singapore (S006 and S008) are in subclade B of mostly <italic>Favia </italic>spp., while the representative from the Philippines (P102) is in subclade F, a distant clade comprising mainly <italic>Favites </italic>species. Interestingly, two reproductively isolated morphotypes of <italic>M. valenciennesi </italic>were recently found to co-occur in Wakayama (Japan), distinguished by the degree of wall fusion among corallites [<xref ref-type="bibr" rid="B100">100</xref>]. Chevalier, 1971 [<xref ref-type="bibr" rid="B101">101</xref>], upon examination of the holotype, placed the species in <italic>Favia </italic>on the basis of corallites possessing separate walls and budding intratentacularly (see also [<xref ref-type="bibr" rid="B102">102</xref>-<xref ref-type="bibr" rid="B108">108</xref>]). This suggests that the name <italic>Favia valenciennesi </italic>(Milne Edwards and Haime, 1848) could be applied to the Australian and Singaporean specimens in subclade B, while P102 (subclade F) is a new species.</p><p>Less extensive issues occur among <italic>Goniastrea </italic>and <italic>Favia </italic>species. For instance, <italic>G. pectinata </italic>(subclade A), collected from three locations, is clearly paraphyletic, with <italic>G. australensis </italic>and <italic>G. favulus </italic>nested within them (Figure <xref ref-type="fig" rid="F2">2</xref>). For <italic>Favia </italic>(subclade B), of six <italic>F. favus </italic>specimens collected from three localities, only three of these form a supported clade while the rest are dispersed within clade XVII-B with no apparent biogeographical pattern. The nesting of <italic>Barabattoia amicorum </italic>among <italic>Favia </italic>spp. has been consistently recovered in recent molecular phylogenies [<xref ref-type="bibr" rid="B12">12</xref>,<xref ref-type="bibr" rid="B15">15</xref>], but this affinity was in fact the dominant hypothesis [<xref ref-type="bibr" rid="B5">5</xref>,<xref ref-type="bibr" rid="B107">107</xref>-<xref ref-type="bibr" rid="B109">109</xref>] until Veron, 1986 [<xref ref-type="bibr" rid="B18">18</xref>], included the species in its current genus. Conversely, <italic>Favia rotundata </italic>clusters with <italic>Favites </italic>spp. rather than its congeners, but it was indeed originally described as <italic>Favites rotundata </italic>Veron, Pichon and Wijsman-Best, 1977 [<xref ref-type="bibr" rid="B5">5</xref>] (see also [<xref ref-type="bibr" rid="B109">109</xref>,<xref ref-type="bibr" rid="B110">110</xref>]).</p><p>The polyphyly of most 'Bigmessidae' genera seems to confer a bleak outlook for revisionary work. However, as we have shown in Figure <xref ref-type="fig" rid="F1">1</xref>, several genera can be clearly grouped as clades with limited name changes. For instance, subclade F is composed of species from <italic>Favites </italic>Link, 1807, <italic>Montastraea </italic>de Blainville, 1830, and <italic>Favia </italic>Ehrenberg, 1834 (Figure <xref ref-type="fig" rid="F2">2</xref>). While the remaining <italic>Favites </italic>spp. (i.e., <italic>F. pentagona</italic>, <italic>F. russelli</italic>, and <italic>F. stylifera</italic>) are not included within this subclade, the type species of this genus is <italic>Favites abdita </italic>(Ellis and Solander, 1786, type locality 'Probablement les mers des Grandes-Indes', Lamarck, 1816 [<xref ref-type="bibr" rid="B111">111</xref>]). The representative of the latter we used falls well within subclade F. Since no other type species were recovered and with <italic>Favites </italic>Link, 1807, being the oldest valid genus in the subclade, <italic>Favites </italic>should be expanded to include the other species, while <italic>F. pentagona</italic>, <italic>F. russelli </italic>and <italic>F. stylifera </italic>will have to be subsumed within other genera. Several other multi-species genera in fact appear stable: <italic>Caulastraea</italic>, <italic>Cyphastrea</italic>, <italic>Echinopora</italic>, <italic>Hydnophora</italic>, <italic>Leptoria</italic>, <italic>Merulina </italic>and <italic>Oulophyllia</italic>. Name changes are certainly not necessary for <italic>Favites </italic>and <italic>Platygyra</italic>, since they host their respective type species in the subclades shown in Figure <xref ref-type="fig" rid="F2">2</xref>.</p></sec><sec><title>Conclusions</title><p>Numerous instances of cryptic taxa determined in this study support the assertion that coral diversity estimates have been fraught with errors [<xref ref-type="bibr" rid="B8">8</xref>]. Traits relating to the gross skeletal morphology of corals are unreliable for species description and identification because of their potential for intraspecific variability [<xref ref-type="bibr" rid="B32">32</xref>,<xref ref-type="bibr" rid="B88">88</xref>] and environment-induced plasticity [<xref ref-type="bibr" rid="B89">89</xref>-<xref ref-type="bibr" rid="B94">94</xref>]. Yet, these characters have served as the foundation for scleractinian taxonomy (e.g., [<xref ref-type="bibr" rid="B4">4</xref>,<xref ref-type="bibr" rid="B5">5</xref>]). Fortunately, using molecular data, the recovery of most genera within the 'Bigmessidae' with only minor degrees of paraphyly spells hope for impending taxonomic amendments. Our results show that most genera only require slight revisions, and most major changes are necessary only at the level of the major clades described in Fukami et al. [<xref ref-type="bibr" rid="B12">12</xref>]. Certainly, broad taxonomic sampling within Faviidae has revealed more species with unexpected affinities, such as <italic>Moseleya latistellata </italic>and <italic>Montastraea multipunctata</italic>. Clade XVII may consequently have to be redefined to exclude them.</p><p>Nevertheless, 'Bigmessidae' subclades are well defined and will no doubt provide a robust framework for taxonomic revisions. The fact that microstructural features support 'Bigmessidae' subclades also offers hope for the morphological approach. Evolutionary relationships among subclades are still provisional due to insufficient statistical support, but they can be clarified with further sampling of nuclear sequences. Eventually, a well-resolved tree of a redescribed clade XVII will be available to reconstruct the morphological evolution of 'Bigmessidae' at various scales.</p></sec><sec><title>Authors' contributions</title><p>DH obtained the DNA sequences in the laboratory, performed the phylogenetic analyses, and had a major role in writing the manuscript. All authors collected the specimens examined, contributed to and approved the final manuscript.</p></sec><sec sec-type="supplementary-material"><title>Supplementary Material</title><supplementary-material content-type="local-data" id="S1"><caption><title>Additional file 1</title><p><bold>'Bigmessidae' corals</bold>. Photographs of most coral specimens sequenced in this study. More photographs are available from the authors.</p></caption><media xlink:href="1471-2148-11-37-S1.PDF" mimetype="application" mime-subtype="pdf"><caption><p>Click here for file</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="S2"><caption><title>Additional file 2</title><p><bold>Maximum likelihood tree topology of the combined molecular data</bold>. Numbers above branches are maximum likelihood bootstrap ≥50 and Bayesian posterior probability ≥0.9, while numbers below denote maximum parsimony bootstrap ≥50 and neighbor-joining bootstrap ≥50. Family classification follows definitions given for Figure <xref ref-type="fig" rid="F1">1</xref>.</p></caption><media xlink:href="1471-2148-11-37-S2.PDF" mimetype="application" mime-subtype="pdf"><caption><p>Click here for file</p></caption></media></supplementary-material><supplementary-material content-type="local-data" id="S3"><caption><title>Additional file 3</title><p><bold>Maximum likelihood tree topology of each partition</bold>. Numbers adjacent to branches are bootstrap support values ≥50. Definitions for family classification follow Figure <xref ref-type="fig" rid="F1">1</xref>.</p></caption><media xlink:href="1471-2148-11-37-S3.PDF" mimetype="application" mime-subtype="pdf"><caption><p>Click here for file</p></caption></media></supplementary-material></sec></body><back><sec><title>Acknowledgements</title><p>We thank all who helped with the field collections, including Zeehan Jaafar, Ywee Chieh Tay, Katrina Luzon, Norievill Espana, Eznairah-Jeung Narida and Monica Orquieza. Flavia Nunes kindly provided the Atlantic specimens. We acknowledge Ann Budd for critical discussions on coral morphology; Carmen Ablan-Lagman and Glenn Oyong for lab support at De La Salle University; Rudolf Meier, Loke Ming Chou and Peter Todd for lab support at National University of Singapore; Carden Wallace, Paul Muir and Barbara Done for museum support at Museum of Tropical Queensland; and staff of Orpheus Island Research Station for field support at Orpheus Island. Special thanks go to Gregory Rouse and Nancy Knowlton for valuable advice and support. For comments on this manuscript, we thank Liz Borda, Tito Lotufo, Yun Lei Tan, three anonymous reviewers and the Associate Editor. Collections were made in Australia under Great Barrier Reef Marine Park Authority permit G09/29715.1, and in the Philippines under Department of Agriculture gratuitous permit FBP-0027-09. 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