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<document id="3404191C4247B60B2AD66C878272C54D" ID-CLB-Dataset="51143" ID-DOI="10.11646/zootaxa.4067.4.7" ID-GBIF-Dataset="9e5fc3f8-f756-47f8-a9aa-6406169b1c62" ID-ISSN="1175-5326" ID-Zenodo-Dep="270632" ID-ZooBank="4FE9F56D-6208-4B05-8EC1-A44612DD4946" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1459697758614" checkinUser="plazi" docAuthor="Rothman, Shevy B. S., Stern, Nir &amp; Goren, Menachem" docDate="2016" docId="03C187FFFFC4FFE2EBD0F96FFBF4FD2A" docLanguage="en" docName="zootaxa.4067.4.7.pdf" docOrigin="Zootaxa 4067 (4)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Epinephelus areolatus Forsskal 1775" docType="treatment" docVersion="8" lastPageNumber="481" masterDocId="FFF8FF87FFC4FFE0EB47FFC0FF90FFF3" masterDocTitle="First record of the Indo-Pacific areolate grouper Epinephelus areolatus (Forsskål, 1775) (Perciformes: Epinephelidae) in the Mediterranean Sea" masterLastPageNumber="483" masterPageNumber="479" pageNumber="479" updateTime="1698209852827" updateUser="ExternalLinkService">
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<mods:title id="C17FDAF54A700FCC2798BEF55DC02F3B">First record of the Indo-Pacific areolate grouper Epinephelus areolatus (Forsskål, 1775) (Perciformes: Epinephelidae) in the Mediterranean Sea</mods:title>
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<mods:namePart id="2CF15A88D2B58CD7623A1B4CE0984AFC">Rothman, Shevy B. S.</mods:namePart>
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<mods:namePart id="0B6076DA48F41BCB75F177FF7DF24489">Stern, Nir</mods:namePart>
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<subSubSection id="C3726562FFC4FFE0EBD0F96FFC8AF93A" box="[151,794,1711,1737]" pageId="0" pageNumber="479" type="nomenclature">
<paragraph id="8BD736E9FFC4FFE0EBD0F96FFC8AF93A" blockId="0.[151,1064,1677,1768]" box="[151,794,1711,1737]" pageId="0" pageNumber="479">
<heading id="D09F8185FFC4FFE0EBD0F96FFC8AF93A" bold="true" box="[151,794,1711,1737]" fontSize="11" level="1" pageId="0" pageNumber="479" reason="1">
<emphasis id="B91CEAFBFFC4FFE0EBD0F96FFC8AF93A" bold="true" box="[151,794,1711,1737]" pageId="0" pageNumber="479">
<taxonomicName id="4C684D6AFFC4FFE0EBD0F96FFDE5F93A" ID-CoL="6G5M7" authority="Forsskal, 1775" authorityName="Forsskal" authorityYear="1775" box="[151,629,1711,1737]" class="Actinopterygii" family="Serranidae" genus="Epinephelus" kingdom="Animalia" order="Perciformes" pageId="0" pageNumber="479" phylum="Chordata" rank="species" species="areolatus">
<emphasis id="B91CEAFBFFC4FFE0EBD0F96FFE36F93A" bold="true" box="[151,422,1711,1737]" italics="true" pageId="0" pageNumber="479">Epinephelus areolatus</emphasis>
(
<bibRefCitation id="EFF94B18FFC4FFE0EAF2F96FFDFDF93A" author="Forsskal" box="[437,621,1711,1737]" pageId="0" pageNumber="482" refString="Forsskal, P. (1775) Descriptiones animalium, avium, amphibiorum, piscium, insectorum, vermium, quae in itinere orientali observavit Petrus Forskal ... edidit Carsten Niebuhr. Adjuncta est materia medica kahirina atque tabula Maris Rubri geographica. exofficina Molleri, Hauniae, 20 + xxxiv + 164 pp." type="book" year="1775">Forsskål, 1775</bibRefCitation>
)
</taxonomicName>
(Figures 1,2)
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C3726562FFC4FFE0EBD0F911FBB8F91B" box="[151,1064,1744,1768]" pageId="0" pageNumber="479" type="reference_group">
<paragraph id="8BD736E9FFC4FFE0EBD0F911FBB8F91B" blockId="0.[151,1064,1677,1768]" box="[151,1064,1744,1768]" pageId="0" pageNumber="479">
<taxonomicName id="4C684D6AFFC4FFE0EBD0F911FE45F91B" authority="Forsskal 1775" authorityName="Forsskal" authorityYear="1775" box="[151,469,1744,1768]" class="Actinopterygii" family="Percidae" genus="Perca" kingdom="Animalia" order="Perciformes" pageId="0" pageNumber="479" phylum="Chordata" rank="species" species="areolata">
<emphasis id="B91CEAFBFFC4FFE0EBD0F911FEA5F91B" box="[151,309,1744,1768]" italics="true" pageId="0" pageNumber="479">Perca areolata</emphasis>
<bibRefCitation id="EFF94B18FFC4FFE0EA7BF911FE45F91B" author="Forsskal" box="[316,469,1745,1768]" pageId="0" pageNumber="482" refString="Forsskal, P. (1775) Descriptiones animalium, avium, amphibiorum, piscium, insectorum, vermium, quae in itinere orientali observavit Petrus Forskal ... edidit Carsten Niebuhr. Adjuncta est materia medica kahirina atque tabula Maris Rubri geographica. exofficina Molleri, Hauniae, 20 + xxxiv + 164 pp." type="book" year="1775">Forsskål 1775</bibRefCitation>
</taxonomicName>
:XI, 42 δ.
<typeStatus id="54D3884BFFC4FFE0E979F911FDE3F91B" box="[574,627,1745,1768]" pageId="0" pageNumber="479">Type</typeStatus>
locality: Jeddah,
<collectingCountry id="F37F7679FFC4FFE0E868F911FC2EF91B" box="[815,958,1745,1768]" name="Saudi Arabia" pageId="0" pageNumber="479">Saudi Arabia</collectingCountry>
, Red Sea.
</paragraph>
</subSubSection>
<subSubSection id="C3726562FFC4FFE0EBD0F8D3FAC4F801" pageId="0" pageNumber="479" type="materials_examined">
<paragraph id="8BD736E9FFC4FFE0EBD0F8D3FB02F8BF" blockId="0.[151,1436,1811,2034]" pageId="0" pageNumber="479">
<emphasis id="B91CEAFBFFC4FFE0EBD0F8D3FEE9F8D9" bold="true" box="[151,377,1811,1834]" pageId="0" pageNumber="479">Examined material.</emphasis>
SMNHTAU—P 15834, Standard length:
<quantity id="4C909B0CFFC4FFE0E87DF8D3FC05F8D9" box="[826,917,1811,1834]" metricMagnitude="-1" metricUnit="m" metricValue="3.02" pageId="0" pageNumber="479" unit="mm" value="302.0">302 mm</quantity>
, Off Tirat-Ha'Carmel, Mediterranean Sea,
<collectingCountry id="F37F7679FFC4FFE0EE27F8D3FA0BF8D9" box="[1376,1435,1811,1834]" name="Israel" pageId="0" pageNumber="479">Israel</collectingCountry>
(
<geoCoordinate id="EE5C502EFFC4FFE0EBD8F8F5FE8FF8BF" box="[159,287,1845,1868]" direction="north" orientation="latitude" pageId="0" pageNumber="479" precision="15" value="32.78389">32°47'02&quot;N</geoCoordinate>
,
<geoCoordinate id="EE5C502EFFC4FFE0EA6DF8F5FE38F8BF" box="[298,424,1845,1868]" direction="east" orientation="longitude" pageId="0" pageNumber="479" precision="15" value="34.90056">34°54'02&quot;E</geoCoordinate>
),
<date id="FFD61029FFC4FFE0EAFCF8F5FDBAF8BF" box="[443,554,1845,1868]" pageId="0" pageNumber="479" value="2015-08-30">30/8/2015</date>
, Coll. Ali Drawshi, at a depth of
<quantity id="4C909B0CFFC4FFE0E8CCF8F5FC2DF8BF" box="[907,957,1845,1868]" metricMagnitude="1" metricUnit="m" metricValue="3.7" pageId="0" pageNumber="479" unit="m" value="37.0">37m</quantity>
, with a fishing lure.
</paragraph>
<paragraph id="8BD736E9FFC4FFE0EB80F896FAC4F801" blockId="0.[151,1436,1811,2034]" pageId="0" pageNumber="479">
<emphasis id="B91CEAFBFFC4FFE0EB80F896FDCDF89E" bold="true" box="[199,605,1878,1901]" pageId="0" pageNumber="479">A brief description of the specimen.</emphasis>
An
<taxonomicName id="4C684D6AFFC4FFE0E9CAF895FC9FF89E" box="[653,783,1877,1901]" class="Actinopterygii" family="Serranidae" genus="Epinephelus" kingdom="Animalia" order="Perciformes" pageId="0" pageNumber="479" phylum="Chordata" rank="genus">
<emphasis id="B91CEAFBFFC4FFE0E9CAF895FC9FF89E" box="[653,783,1877,1901]" italics="true" pageId="0" pageNumber="479">Epinephelus</emphasis>
</taxonomicName>
species with emarginate caudal fin, 84 series of scales along the body; dorsal fin with 11 spines and 14 rays; anal fin with 3 spines and 8 rays; pectoral fins with 17 rays; pelvic fins with a spine and 5 rays. Gill rakers on lower limb of first gill arch 12, 1 at angle of the arch and
<docIssue id="CF0B4906FFC4FFE0EFC2F859FB02F843" box="[1157,1170,1945,1968]" pageId="0" pageNumber="479">4</docIssue>
rakers on upper limb + 4 rudimentary gill rakers. Rear margin of preopercle serrate, with small spines at its lower angle (
<figureCitation id="13532A6CFFC4FFE0EF3BF87AFB2DF822" box="[1148,1213,1978,2001]" captionStart="FIGURE 2" captionStartId="1.[151,244,1911,1933]" captionTargetBox="[186,1436,921,1889]" captionTargetId="figure@1.[151,1436,921,1890]" captionTargetPageId="1" captionText="FIGURE 2. Epinephelus areolatus (Forsskål, 1775) from the Mediterranean. The arrow indicates the small spines on the lower angle preopercle. SMNHTAU—P 15834. Photo O. Rittner." httpUri="https://zenodo.org/record/270634/files/figure.png" pageId="0" pageNumber="479">Fig. 2</figureCitation>
). First dorsal spine is above the origin of upper pectoral ray. Pelvic fin begins below a vertical line from the origin of lowest pectoral ray.
</paragraph>
</subSubSection>
<caption id="DF176661FFC5FFE1EBD0FC93FA9CFC9A" box="[151,1292,851,873]" httpUri="https://zenodo.org/record/270633/files/figure.png" pageId="1" pageNumber="480" targetBox="[151,1434,194,827]" targetPageId="1">
<paragraph id="8BD736E9FFC5FFE1EBD0FC93FA9CFC9A" blockId="1.[151,1292,851,873]" box="[151,1292,851,873]" pageId="1" pageNumber="480">
<emphasis id="B91CEAFBFFC5FFE1EBD0FC93FE9CFC9A" bold="true" box="[151,268,851,873]" pageId="1" pageNumber="480">FIGURE 1.</emphasis>
<taxonomicName id="4C684D6AFFC5FFE1EA55FC94FD1BFC9A" authority="Forsskal, 1775" authorityName="Forsskal" authorityYear="1775" box="[274,651,852,873]" class="Actinopterygii" family="Serranidae" genus="Epinephelus" kingdom="Animalia" order="Perciformes" pageId="1" pageNumber="480" phylum="Chordata" rank="species" species="areolatus">
<emphasis id="B91CEAFBFFC5FFE1EA55FC94FE78FC9A" box="[274,488,852,873]" italics="true" pageId="1" pageNumber="480">Epinephelus areolatus</emphasis>
(Forsskål, 1775)
</taxonomicName>
from the Mediterranean. SMNHTAU—P 15834. Photo O. Rittner.
</paragraph>
</caption>
<caption id="DF176661FFC5FFE1EBD0F8B7FD03F85A" httpUri="https://zenodo.org/record/270634/files/figure.png" pageId="1" pageNumber="480" targetBox="[186,1436,921,1889]" targetPageId="1">
<paragraph id="8BD736E9FFC5FFE1EBD0F8B7FD03F85A" blockId="1.[151,1436,1911,1961]" pageId="1" pageNumber="480">
<emphasis id="B91CEAFBFFC5FFE1EBD0F8B7FE9DF87E" bold="true" box="[151,269,1911,1933]" pageId="1" pageNumber="480">FIGURE 2.</emphasis>
<taxonomicName id="4C684D6AFFC5FFE1EA54F8B7FD1DF87F" authority="Forsskal, 1775" authorityName="Forsskal" authorityYear="1775" box="[275,653,1911,1932]" class="Actinopterygii" family="Serranidae" genus="Epinephelus" kingdom="Animalia" order="Perciformes" pageId="1" pageNumber="480" phylum="Chordata" rank="species" species="areolatus">
<emphasis id="B91CEAFBFFC5FFE1EA54F8B7FE79F87F" box="[275,489,1911,1932]" italics="true" pageId="1" pageNumber="480">Epinephelus areolatus</emphasis>
(Forsskål, 1775)
</taxonomicName>
from the Mediterranean. The arrow indicates the small spines on the lower angle preopercle. SMNHTAU—P 15834. Photo O. Rittner.
</paragraph>
</caption>
<subSubSection id="C3726562FFC6FFE2EB80FF58FDDAFE52" pageId="2" pageNumber="481" type="description">
<paragraph id="8BD736E9FFC6FFE2EB80FF58FBACFF07" blockId="2.[151,1436,152,729]" pageId="2" pageNumber="481">
<emphasis id="B91CEAFBFFC6FFE2EB80FF58FE9DFF5C" bold="true" box="[199,269,152,175]" pageId="2" pageNumber="481">Color.</emphasis>
Body reticulated with whitish background and brown spots that gradually become smaller and orange on head and belly (
<figureCitation id="13532A6CFFC6FFE2EA0AFF7AFE00FF22" box="[333,400,186,209]" captionStart="FIGURE 1" captionStartId="1.[151,244,851,873]" captionTargetBox="[151,1434,194,827]" captionTargetId="figure@1.[151,1436,194,828]" captionTargetPageId="1" captionText="FIGURE 1. Epinephelus areolatus (Forsskål, 1775) from the Mediterranean. SMNHTAU—P 15834. Photo O. Rittner." httpUri="https://zenodo.org/record/270633/files/figure.png" pageId="2" pageNumber="481">Fig. 1</figureCitation>
). The spots become darker towards rear part of body, caudal fin and soft part of dorsal fin. Posterior edge of caudal fin and upper edge of soft part of dorsal fin with white margin.
</paragraph>
<paragraph id="8BD736E9FFC6FFE2EB80FEC0FDDAFE52" blockId="2.[151,1436,152,729]" pageId="2" pageNumber="481">
<emphasis id="B91CEAFBFFC6FFE2EB80FEC0FE69FEE4" bold="true" box="[199,505,256,279]" pageId="2" pageNumber="481">Selected body proportions.</emphasis>
<collectingCountry id="F37F7679FFC6FFE2E946FEC0FD8FFEE4" box="[513,543,256,279]" name="American Samoa" pageId="2" pageNumber="481">As</collectingCountry>
percentage of standard length: head length, 37%; body depth, 35%; longest dorsal spine, 12%; longest dorsal ray, 12%; longest anal spine, 6%; longest anal ray, 16%; distance snout, pelvic fin, 42%; distance snout, base of pectoral fin, 35%; distance snout, dorsal fin, 36%; distance snout, anal fin, 72%; pectoral fin length, 22%; pelvic fin length of standard length, 17%.
<collectingCountry id="F37F7679FFC6FFE2E84AFEA8FCBBFE8C" box="[781,811,360,383]" name="American Samoa" pageId="2" pageNumber="481">As</collectingCountry>
percentage of head length: upper jaw length, 37%; eye diameter, 15%; distance snout-eye, 26%.
</paragraph>
</subSubSection>
<subSubSection id="C3726562FFC6FFE2EB80FE6DFBF4FD2A" pageId="2" pageNumber="481" type="distribution">
<paragraph id="8BD736E9FFC6FFE2EB80FE6DFB2BFE14" blockId="2.[151,1436,152,729]" pageId="2" pageNumber="481">
<emphasis id="B91CEAFBFFC6FFE2EB80FE6DFEC9FE37" bold="true" box="[199,345,429,452]" pageId="2" pageNumber="481">Distribution.</emphasis>
<taxonomicName id="4C684D6AFFC6FFE2EA18FE6DFE70FE37" box="[351,480,428,452]" class="Actinopterygii" family="Serranidae" genus="Epinephelus" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="481" phylum="Chordata" rank="species" species="areolatus">
<emphasis id="B91CEAFBFFC6FFE2EA18FE6DFE70FE37" box="[351,480,428,452]" italics="true" pageId="2" pageNumber="481">E. areolatus</emphasis>
</taxonomicName>
is widely distributed in the coastal waters of the Indian Ocean,
<collectingCountry id="F37F7679FFC6FFE2EFC4FE6DFB23FE37" box="[1155,1203,429,452]" name="India" pageId="2" pageNumber="481">Indo</collectingCountry>
Pacific Archipelago, north-west
<collectingCountry id="F37F7679FFC6FFE2EA48FE10FEE3FE14" box="[271,371,464,487]" name="Australia" pageId="2" pageNumber="481">Australia</collectingCountry>
, South
<collectingCountry id="F37F7679FFC6FFE2EA84FE10FD93FE14" box="[451,515,464,487]" name="China" pageId="2" pageNumber="481">China</collectingCountry>
sea,
<collectingCountry id="F37F7679FFC6FFE2E97EFE10FD1AFE14" box="[569,650,464,487]" name="Taiwan" pageId="2" pageNumber="481">Taiwan</collectingCountry>
,
<collectingCountry id="F37F7679FFC6FFE2E9D3FE10FD41FE14" box="[660,721,464,487]" name="Japan" pageId="2" pageNumber="481">Japan</collectingCountry>
and the Red Sea (
<bibRefCitation id="EFF94B18FFC6FFE2E8D5FE10FB3EFE14" author="Heemstra" box="[914,1198,464,487]" pageId="2" pageNumber="482" refString="Heemstra, P. C. &amp; Randall, J. E. (1993) FAO species catalogue vol. 16 groupers of the world (family Serranidae, subfamily Epinephelinae): An annotated and illustrated catalogue of the grouper, rockcod, hind, coral grouper, and lyretail species known to date. Food and Agriculture Organization of the United Nations, Rome, 382 pp." type="book" year="1993">Heemstra &amp; Randall 1993</bibRefCitation>
).
</paragraph>
<paragraph id="8BD736E9FFC6FFE2EB80FE32FBF4FD2A" blockId="2.[151,1436,152,729]" pageId="2" pageNumber="481">
<emphasis id="B91CEAFBFFC6FFE2EB80FE32FE76FDFA" bold="true" box="[199,486,498,521]" pageId="2" pageNumber="481">Genetic characterization.</emphasis>
Intra-specific ML (Maximum likelihood) phylogenetic tree of previously-published
<emphasis id="B91CEAFBFFC6FFE2EE29FE32FA0CFDFA" box="[1390,1436,498,521]" italics="true" pageId="2" pageNumber="481">COI</emphasis>
sequences has revealed a distinct biogeographic separation between populations located east to the Indo-Malay Peninsula, i.e., Western Indian Ocean (WIO), and west, i.e., West Pacific (WP,
<figureCitation id="13532A6CFFC6FFE2E8AAFDF8FBBEFDBC" box="[1005,1070,568,591]" captionStart="FIGURE 3" captionStartId="2.[151,244,1504,1526]" captionTargetBox="[163,1420,787,1479]" captionTargetId="figure@2.[163,1424,783,1483]" captionTargetPageId="2" captionText="FIGURE 3. Intra-specific phylogenetic relationships of Epinephleus areolatus from the Indo-west Pacific, based on Maximum likelihood analysis (ML) with 500 replications of COI sequences mined from GenBank database. * - indicates sequenced mined from BOLD database; ** - unpublished data." httpUri="https://zenodo.org/record/270635/files/figure.png" pageId="2" pageNumber="481">Fig. 3</figureCitation>
). Pairwise comparisons, based on K2P model, have shown an average genetic divergence of 3.49±0.61% between these clades and an average of 0.85±0.21% within the WIO clade and 0.46±0.16% within the WP. In addition, our alien specimen from the Mediterranean aligned within the WIO clade, and presented an identical haplotype with two specimens from the Gulf of Aqaba, Red Sea (R. Holtzman, unpublished data, 2015) thus confirming its Red Sea origin.
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